Annals of the Missouri Botanical Garden

ANNALS OF THE
MISSOURI BOTANICAL GARDEN

VOLUME 74
1987

The Annars, published quarterly, contains papers, primarily in systematic
botany, contributed from the Missouri Botanical Garden, St. Louis. Papers origi-
nating outside the Garden will also be accepted. Authors should write the Editor
for information concerning arrangements for publishing in the ANNALS.

EDITORIAL COMMITTEE
GroncE K. Rocers, Editor

Missouri Botanical Garden

MarsHALL R. CROSBY
Missouri Botanical Garden

GERRIT DAVIDSE
Missouri Botanical Garden

Jonn D. DWYER
Missouri Botanical Garden & St. Louis University

PETER GOLDBLATT
Missouri Botanical Garden

HENK VAN DER WERFF
Missouri Botanical Garden

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© Missouri Botanical Garden 1988

ISSN 0026-6493

Volume 74

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t

Volume 74, Number 4 | Annals of the
Spring 1987 Missouri Botanical Garden

The Annals, published quarterly, contains papers, primarily in systematic botany, con-
tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the
Garden will also be accepted. Authors should write the Editor for information concerning
arrangements for publishing in the ANNALS. Instructions to Authors are printed on the
inside back cover of the last issue of each volume.

Editorial Committee

George K. Rogers and Nancy R. Morin Marshall R. Crosby
Coeditors (this issue), Missouri Missouri Botanical Garden
Botanical Garden

Gerrit Davidse
Janice Wilson: sy 5.3 Missouri Botanical Garden
Editorial Assistant, Missouri Botanical
Gardens o es | John D. Dwyer -
S Missouri Botanical Garden &
Saint Louis University

m Peter Goldblatt ——
Missouri Botanical Garden

Ux Henk van der Werff

_ Missouri Botanical Garden

. Eleven, P.O. Box 299, St. Louis, MO 63166. Sub- SSN
ery charge, $35 per volume. Four issues per vol- -

Volume 74 Annals
Number 1 of the NZ
1987 Missouri

Botanical

Garden

SYSTEMATIC EMBRYOLOGY OF THE ANISOPHYLLEACEAE!
HIROSHI TOBE? AND PETER H. RAVEN?

ABSTRACT

An embryological study of Anisophylleaceae, which comprise Anisophyllea, Combretocarpus, Poga,
and Polygonanthus, and which have traditionally most often been referred as a tribe or subfamily to
: PARIE t

osome

of bretocarpus is reported as n = 8, that of the ~~ apii genera as n = 7. Embryologically
Anisophylleaceae are diversified and show differences from genus to genus, but they are clearly distinct
m raceae in having their combination of con RUE baies states, including persistent
nucellar tissue at least until i early stages of seed sp tng tou thin two cell-layered inner integument
Poga and P In trast to Rhizophoraceae, Anisophylleaceae
agree almost completely w ith Myrtales in their embryological features of the o rder. Embryological
of A e

Hd
=
`°

"a
=
°
°

evidence iode: supports the bonu ily and, decus

from other lines of evidence, suggests a Myrtale. n affinity for the family. Proposed assignments
Anisophylleaceae to Rosales or to Cornales a " not sup ported. An analysis of similarities in Character
states in the four genera the ra

one leading to Anisophyllea a nd Combretocarpus, and the other leading to Poga ápd Politak anthus.
Combretocarpus, with which Anisophyllea shares 3 few synapomorphies, is most specialized wit hema
the family in having many apomorphies. In contrast, om and Polygonanthus share many ples
morphies, most of which are also common to EE

Anisophylleaceae, as defined here, consists of referred Polygonanthus to Euphorbiaceae (Ducke,
four genera and 34 species, Anisophyllea (30 spp.), | 1932, 1933; Kuhlmann, 1940), Olacaceae (Croi-
Combretocarpus (1 sp.), Poga (1 sp.), and Polygo- zat, 1939), Saxifragaceae (Baehni & Dansereau,
nanthus (2 spp.) (Airy Shaw, 1973; Cronquist, 1939), or to its own family, Polygonanthaceae
1981, 1983). In contrast with the stable assign- (Croizat, 1943). Despite these, there is little doubt,
ment of three other genera, various authors have on the basis of morphological and wood anatom-

! Supported by grants to the junior author from the U.S. National Science Foundation and in part by the
Grant-in-Aid Scientific Research to the senior author from the Ministry of Education, Science and Culture,

Kalkman, Duncan W. Thomas, and Mohd Shah Bin Mohd Noor for their collection and arrangement of the
excellent material used in this study, and to Rolf Dahlgren, L. A. S. Johnson, Adrian Juncosa, and Richard
Keating for their useful comments regarding this manuscript.

? Biological Laboratory, Yoshida College, Kyoto University, Kyoto 606, Japan.

? Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166.

ANN. Missouni Bor. GARD. 74: 1-26. 1987.

TABLE |. Studied taxa and collections.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Taxa

Collections

Anisoph "d om
JB

(Jack in 1984, no voucher

Singapore. Bukit Timah Nature Reserve. Sidek Bin Kiah & Tan Yam Leong s.n.

Singapore. Botanic Garden, ides iei Sidek Bin Kiah s.n. in 1984, no voucher;
Mohd Shah s.n. in 1984, no voucher.

ass ag Maxwell Hill, Perak. B. C. Stone 15403, (KLU, MO).

Brunei. A. M. Juncosa s.n. in 1981, no voucher.

Anisophyllea sp.

Combretocarpus rotun-
Mi n Brunei

Poga oleosa Pierre

Cameroon. D. W. Thomas 3494, (MO).

nue ay Kuching, Sarawak. P. Chai s.n. in 1981, 1983, and 1985, no voucher.
. M. Juncosa s.n. in 1983, no voucher.

Cameroon. Korup Natl. Park. D. W. Thomas 2273, (MO).

Nigeria. Awi, Akamkpa. J. O. Ariwaodo s.n. in 1983, (FHI 99607).

Polygonanthus amazoni-

Brazil. Along the Rio Paca, Amazonas. J. Zarucchi 3138, 3184, (US).

ical evidence, that Polygonanthus fits well in An-
isophylleaceae, together with the three other gen-
era that have tra iar been placed there (see
Kuhlmann, 1944; Pir Rodrigues, 1971; Van
Vliet, 1976). Of the ur genera of this family,
Anisophyllea is relatively widely distributed in
tropical Africa and Asia, also occurring in trop-
ical South America; Combretocarpus is restricted
to West Malaysia, Poga to tropical West Africa;
and Polygonanthus to the Amazon Basin of Bra-
zil (Pires & Rodrigues, 1971)

The relationships of Anisophylleaceae have
been controversial. A traditional view, and the
one most widely accepted, is that Anisophylle-
aceae have close affinities with Rhizophoraceae,
and they often have been considered a tribe or
subfamily within a broadly conceived Rhizopho-
raceae (Bentham & Hooker, 1865; Baillon, 1877;

been treated as a distinct family, Anisophylle-
aceae have generally been considered closely re-

Thorne (1983) placed Rhizophoraceae (com-
posed of two subfamilies: Rhizophoroideae and
Anisophylleoideae) in the Cornales.

Recently, however, Cronquist (1981, 1983)
concluded that Anisophylleaceae were not closely
related to Rhizophoraceae and assigned them to
izophoraceae sensu

a

stri
Dahlgren (1983), who also denied any close re-

lationships between Anisophylleaceae and Rhi-
d idee pared „Anisophylleaceae in Cor-

s(C its own
; see also Dahl-

+

t
Dh; 1 1 (NÁ 4:4

or iis
gren & Thorne, 1984).

In the light of these diverse opinions, we have
attempted to determine whether Anisophylle-
aceae are actually closely related to Rhizophora-
ceae sensu stricto or not, or whether they might
even be grouped together as one family. If the
two groups are not closely related, what are their
respective affinities?

Anisophylleaceae have been studied to a very
limited extent, particularly regarding pes ana-
tomical characteristics. Their wood anatomy,
however, has been studied relatively intensive
(Marco, 1935; Geh & Keng, 1974; n Vliet,
1976). Based on a comparison of the Mog anat-
omy of the two groups, Van Vliet (1976) sup-
ported a broad definition of Rhizophoraceae in-
cluding Anisophylleaceae. In contrast, Behnke
(1984), basing his conclusions on the features of
their sieve-tube plastids, suggested Anisophyl-
leaceae were quite distinct from Rhizophoraceae
sensu stricto. He found that both Anisophyllea

and COMMIELOE HS pave S-type plastids (con-

J

plastids (containing protein) that are character-
istic of Rhizophoraceae sensu stricto. Another
interesting distinction between the two groups,
which has been known for some time, is that all
four genera of Anisophylleaceae are aluminum
accumulators; whereas the genera of Rhizopho-
raceae sensu stricto are not (Chenery, 1948;
Kukachka & Miller, 1980). Although embryo-
logical information has been extremely useful in
suggesting relationships at this level (see Tobe &

1987]

Raven, 1983), almost no information is available
on Anisophylleaceae. The only published data
on ovule morphology (of **Anisophylleia zeylan-
ica") is that of Karsten (1891) nearly 100 years
ago; however, most of these observations seem
to be incorrect, as we shall discuss subsequently.
Vaughan (1970) described the mature seed coat
structure of Poga, providing a drawing; Geh and
Keng (1974) reported on the endosperm in the
seeds of Anisophyllea and Combretocarpus. Ex-
cept for these fragments of information, appar-
ently nothing has been reported about the em-
bryological features of Anisophylleaceae.

In this paper, we present an overall study of
the embryology of Anisophylleaceae, which is
intended to provide information bearing on their
relationships and systematic position. We have
studied Anisophyllea and Combretocarpus in de-
tail, and Poga and Polygonanthus to a lesser de-
gree. Important features have been noted for all
genera and are presented here.

MATERIALS AND METHODS

All four genera, Anisophyllea, Combretocar-
pus, Poga, and Polygonanthus, were investigated
in this study. The species we studied are listed
in Table 1 together with their voucher infor-
mation. Flower buds and fruits in all stages of
development were collected and fixed in FAA (5
parts stock formalin; 5 parts glacial acetic acid;
90 parts 70% ethanol); however, female buds of
Poga oleosa and fruits of Polygonanthus ama-
zonicus were not available. Herbarium material
of Anisophyllea and Combretocarpus was studied
to supplement our observations of fruits and
seeds.

Preparations of microtome sections for obser-
vation were made following standard paraffin
techniques. After dehydration through a tertiary-
butyl alcohol series, the samples were embedded
in Paraplast with 56—58?C mp. Flower buds of
Anisophyllea disticha and fruits of Poga oleosa
were too hard to be sectioned without being soft-
ened initially. Therefore, after these structures
were trimmed to expose their tissues, the embed-
ded samples attached to blocks were soaked in
a mixture of a 10:3:90 glycerol: 10% Aerosol
OT: water (Schmid & Turner, 1977) for at least
several days at 20—25?C and then sectioned. Se-
rial sections 6-10 um thick were stained with
Heidenhain's hematoxylin, safranin, and fast-
green FCF and were mounted in Entellan. Ma-
ture seed coats of Anisophyllea sp. (D. W. Thom-
as 3494, MO) and Poga oleosa, which were too
thick and hard to be sectioned by standard par-

TOBE & RAVEN —ANISOPHYLLEACEAE 3

affin techniques, were embedded in a JB-4 plastic
and stained with 0.1% Toluidine Blue.

In order to count the number of cells in mature
pollen, we attempted to use safranin-staining of
the grains (Tobe & Raven, 1984). We failed to
obtain any staining ofthe pollen nuclei, however,
probably because a thick exine hinders the infil-
tration of dye. Consequently, we counted the
number of cells in the pollen using microtome-
sectioned pollen grains. The expressions we have
used for the frequency of different shapes of mi-
crospore tetrads follow those of Schmid (1982).

OBSERVATIONS
ANISOPHYLLEA R. BR.

The embryological characteristics were basi-
cally the same in the two species studied, one
from Africa and one from Asia, The features
reported in the foll
to be common to both species, , unless particular
vo dis are given.

Anther and microspores. The anther is tetra-
sporangiate. The wall prior to maturation com-
prises basically five cell layers: an epidermis, an
endothecium, two middle layers, and a tapetum
(Fig. 1); the wall formation therefore conforms
to be the Basic type (Davis, 1966: 10). The anther
wall, however, often has only one middle layer,
which shares a histogenetic origin with the ta-
petum. The tapetum is glandular (Fig. 2). At one
point in their development, the cells of the ta-
petum become 2-nucleate, but subsequently the
two nuclei fuse with each other. During matu-
ration, the middle layer(s) degenerate and the
epidermal cells are stretched tangentially while
the cells of the endothecium become more or less
enlarged (Fig. 2). Eventually, the endothecium
develops fibrous thickenings. Although the epi-
dermis persists, it is often collapsed on the en-
dothecium (Fig. 3). Anther dehiscence takes place
by longitudinal slits (Fig. 3). The connective tis-
sue between the two microsporangia of each the-
ca is completely disorganized before an anther
dehisces (Fig. 3).

Meiosis in a microspore mother cell is accom-
panied by simultaneous cytokinesis, and the re-
sultant microspore tetrads, on the basis of 50
selected tetrads (of Anisophyllea disticha), are
**usually" (92%) tetrahedral, **very occasionally"
(6%) decussate, and “rarely” (2%) isobilateral.
The pollen grains are two-celled at the time of
shedding (Fig. 4

Chromosomes. Since pollen mother cells be-
tween the telophase of meiosis I and the meta-

WCIC found

4 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

ES 1-5. Anisophyllea. — 1, 2, 4, 5. A. disticha. —3. A. sp. (D. W. Thomas 3494, MO).— 1. Transverse
CN (TS) of a young anther showing the five cell-layered wall structure. Bar — 10 um.— 2. TS of an older
anther with degenerating middle layers (arrow). Bar = 10 um.—3. TS of a developed anther. Its wall consists
of the fibrous Mna epum and the epidermis. Bar = 50 um.—4. Two-celled mature pollen at the time of
shedding. Arrows indicate nuclei of the two cells. Bar = 10 um.— 5. Chromosomes of a pollen mother cell at a
stage between Victus I and metaphase II. n = 7. Bar = 10 um. ep, epidermis; et, endothecium; ml, middle
layer; t, tapetum; mc, microspore mother cell.

1987]

phase of meiosis II were fixed by chance and
included in microtome sections, we were able to
count the chromosome number of Anisophyllea
for the first time: A. disticha has n = 7 (Fig. 5)
Megagametophyte and nucellus. The ovule
is anatropous. A single archesporial cell differ-
entiates beneath the apical dermal layer of the
nucellus (Fig. 6). The archesporial cell divides
periclinally into two: the upper primary parietal
cell and the lower sporogenous cell (Fig. 7). The
primary parietal cell divides periclinally, and its
derivatives further divide anticlinally and peri-
clinally, forming parietal tissue with three to five
layers above the embryo sac. The sporogenous
cell develops into a megaspore mother cell and
undergoes meiosis, giving rise to a linear tetrad
of megaspores. A triad of megaspores may also
be formed by suppression of the second, mitotic
division on the micropylar side. In the mega-
spore tetrad (or triad), the chalazal megaspore
functions (Fig. 8). A functional megaspore de-
velops successively into a 2- (Fig. 9), 4- (Fig. 10),
and 8-nucleate embryo sac (Fig. 11). Thus the
mode of the embryo sac formation is of the Po-
lygonum type. The synergids are slightly hooked

polar nuc
which is positioned near the egg apparatus (Fig.
13). Consequently an organized mature embryo
sac has only five nuclei or cells: an egg cell, two
synergids, and two polar nuclei (as a single cen-
tral nucleus; Figs. 12, 13).
uring megasporogenesis and megagameto-

genesis, apical epidermal cells of the nucellus di-
vide periclinally, and their daughter cells also
repeat periclinal divisions. As a result, a four to
six cell-layered nucellar cap is formed above the
embryo sac (Fig. 14); Karsten (1891) also illus-
trated such a nucellar growth in “Anisophylleia
zeylanica." The nucellar cap and the other nu-
cellar tissue, both of which enclose the embryo
sac, persist into younger stages of fruit devel-
opment (Figs. 11, 14, 17). There is no case in
which the nucellar tissue degenerates before fer-
tilization so that the embryo sac directly borders
on the integument.

Integument. The ovule has a single integu-
ment (Figs. 7, 15), although Karsten (1891) de-
scribed the ovule of * "Anisophylleia zeylanica"

nucellar tissue surrounding the embryo sac as the

TOBE & RAVEN—ANISOPHYLLEACEAE 5

inner integument and the (only) true integument
as the outer integument.

A micropyle is always formed by the integu-
ment, excepting one very unusual case in whic
the integument did not grow beyond the nucellar
apex (Fig. 16). In this respect as well, Karsten
(1891) seems to have erred: he considered the
persistent, lateral nucellar tissue to be the inner
integument, which he concluded did not enclose
the nucellar apex. In fact, Karsten concluded that
a micropyle is not formed in “‘Anisophylleia zey-
lanica.” Referring to Karsten’s drawing of ovules
and descriptions, we also mistakenly character-
ized the ovule of Anisophyllea not only as being
bitegmic but also as having a nucellar beak (which
actually was the well-developed persistent nu-
cellar cap; see Tobe & Raven, 1983)

The integument is about five to seven cells
thick in Anisophyllea disticha (Fig. 15) and about
four to five cells thick in A. sp. (Fig. 17). The
thickness of the integument is not different from
one part of ovule to another, and therefore the
cross section of the ovule is nearly circular (Fig.
17). A raphe bundle ramifies oblique-laterally
toward the chalazal end (Fig. 18). Therefore in
cross section the ovule or fruit has four to five
vascular bundles at the peripheral part of the
integument of testa (Fig. 19).

Throughout the development of the ovule or
fruit, the integument or seed coat is thickened
by secondary multiplication. However, the in-
nermost cell layer never differentiates toward the
so-called endothelium

Fertilization, endosperm, and embryo. De-
spite their multiovular condition in Aniso-
phyllea (usually four and rarely three ovules per
ovary), fruits were always one-seeded. Fertiliza-
tion is porogamous. Endosperm formation is of
the Nuclear type (Fig. 20). Because of incom-
pleteness of our fruit sample, we could not con-
firm whether or not wall formation takes place
in free endosperm nuclei. Hand-sectioned ma-
ture seeds of the two species we studied lacked
endosperm (Fig. 21). Concerning the presence of
endosperm in seeds, Hou (1958) described that
in Anisophyllea disticha seeds consist of a solid

ody, of which the main part is formed by a
thick, hard albumen. Geh and Keng (1974) stated
that in Anisophyllea disticha, the entire undiffer-
entiated embryo is embedded in endosperm;
consequently, they characterized the seed of Ani-
sophylleae (Anisophyllea and Combretocarpus)
as albuminous. Based on results of our obser-
vations, however, it seems that what Hou thought

6 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

FIGURES 6-14. Anisophyllea. —6, 7, 9-13. A. disticha. — 8. A. sp. (D. W. Thomas 3494, MO).—6. Longitudinal
Mice (LS) of an ovular primordium with the 1-celled archesporium. Bar = 10 um.— 7. LS of a young ovule
with the primary parietal cell. Bar — 10 um.— 8. LS of a young ovule with the con megaspore. Arrows

the 2-nucleate embryo sac stage. Bar = 10 um.— 10. LS of an older ovule at the 4-nucleate Vigo sac stage.
One of the two nuclei at the micropylar side appears in the next section. Bar — 10 um.— 11. LS of a nearly

1987]

to be the thick, hard albumen was actually the
embryo itself, and that the seed that Geh and
Keng observed was too young to confirm the
endosperm condition.

Although we did not pursue the whole process
of embryogenesis either, the development of

cotyledons. Geh and Keng (1974), however, re-
ported two protuberances on the apical part of
the embryo in Anisophyllea disticha, which they
interpreted as two cotyledons. We conclude that
the cotyledons of Anisoph dps either develop in-
completely or are essentially absent in Aniso-

phyllea. No hypostase is a after fer-
tilization.

Mature seed and seed coat. The mature seed
is narrowly cylindrical, 13.0-13.5 mm long and
3.8-4.0 mm thick in Anisophyllea disticha,
whereas it is ovoid or elliptical in outline, 13.0—
13.8 mm long and 6.0—6.4 mm thick in A. sp.
(Fig. 21). In the young seed, the seed coat appears
to be constructed of a thick, massive tissue, with

ple inner layer about 25—30 cells thick
(Fig. 23). The cells of the outer epidermis are
thick-walled and cuboid, whereas those of the
underlying multiple inner layer are also thick-
walled but extremely stretched tangentially.

COMBRETOCARPUS HOOK F.

Anther and microspores. The anther is tetra-

ing maturation, the cells of the epidermis are
somewhat enlarged and become tanniferous; the
cells of the endothecium are also enlarged; the
middle layers degenerate (Fig. 25). The tapetum
is glandular, and its cells become 2-nucleate. The

TOBE & RAVEN—ANISOPHYLLEACEAE 7

two nuclei in a tapetal cell later are fused with
each other. Thus the mature anther wall is com-
posed of the persistent but somewhat collapsed
epidermis and the fibrous endothecium (Fig. 26).
By the time of anther dehiscence, the connective
tissue between two microsporangia of each teca
degenerates completely. After dehiscence by lon-
gitudinal slits, the anther wall is remarkably re-
flexed (Fig. 2

Meiosis in ihe microspore mother cells is ac-
companied by simultaneous cytokinesis. The
shape of the resultant tetrads, on the basis of the
examination of 50 selected tetrads, is “usually”
(78%) tetrahedral, **occasionally" (1496) decus-
sate, and “very occasionally" (896) isobilateral.
The pollen ien are two-celled at the time of
shedding (Fig.

Chromosomes. “Pollen mother cells at the
metaphase of meiosis I happened to be fixed and
appeared in microtome sections. On the basis of
those sections, we observed the chromosomes of
Combretocarpus for the first time and deter-
mined n = 8 (Fig. 29). Size differences seem to
be present among those eight chromosomes.

Megagametophyte and nucellus. The ovule
is anatropous and crassinucellate. The archespo-
rium is nearly always 1-celled (Fig. 30). A multi-
cellular archesporium may very rarely differen-
tiate—an ovule or young fruit containing twin
ambs sacs was very ay observed (Fig. 3
Th lly into two:
the upper primary parietal “cell and the lower
sporogenous cell. The primary parietal cell may
or may not divide further periclinally; if it does
so, a two cell-layered parietal tissue is formed.
The sporogenous cell develops into a megaspore
mother cell (Fig. 31). After enlarging in volume,

J

ecome 2-nucleate (Fig. 32).
spore is functional. Then, while the micropylar
megaspore degenerates, the two nuclei in the

—

mature ovule at the 8-nucleate embryo sac stage. Of the eight nuclei, two polar nuclei are fused into a single

central nucleus, and three antipodal cells are degenerating. Bar — 50 um.— 12, 13. Tw
owing the egg apparatus and the central nucleus. Bars = 10 um

mature ovule sho

o serial LSs of a part of
—14. Same as Figure 11,

but at a lower magnification. Note that the nucellar tissue is persistent and that no cell layer of the integument

shows differentiation into an endothelium. Bar = 50
sporogenous cell; fc, functional megaspore; n, nucleus of the em

um. arc, SA cell; p, primary parietal cell; s,

c; eg, egg cell; ega, egg apparatus; cn,

central nucleus; ant, antipodal cell; sy, synergid; nuc, nucellar tissue; pig integument; cp, nucellar cap.

8 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FIGURES 15-23. Anisophyllea. — 15,
MO).— 15. Longitudinal section (LS) of a young ovule. Note that the avuke i is unite

18-20, 22. A. disticha.—16, 17, 21, 23. A. ics (D. W. Thomas 3494,

Bar = 50 um.—16. LS
um. —17. fae ries (TS) of a mature
LS of a mature ovule tangentially cut through a raphe showing the ramification of a
raphe bundle. Bar = 100 um.— 19. TS of a young seed. Note that the thick seed coat contains several vascular
bundles at the peripheral part. Bar = 1 mm.— 20. LS ofa young seed containing a proembryo and free endosperm

of an unusual mature ovule lacking a micropyle. Bar = 50
ovule. Bar = 50 um.— 18.

1987] TOBE & RAVEN—

ANISOPH Y LLEACEAE 9

functional chalazal megaspore separate from each

ther: one moves toward the micropylar end,
while the other moves toward the chalazal end
(Fig. 33). Each nucleus divides successively to
form a 4- and an 8-nucleate sac (Fig. 34). Thus
the embryo sac formation conforms to the bispor-
ic Allium type. The synergids are slightly hooked,
and the antipodals are ephemeral, disappearing
before fertilization. Two polar nuclei do not fuse
with each other until fertilization takes place;
they are positioned near the egg apparatus. A
mature embryo sac just before fertilization is
composed of five nuclei or cells: an egg cell, two
synergids, and two polar nuclei.

Embryo sacs characteristically accumulate an
abundance of starch grains (Fig. 35). The starch
grains, which begin to accumulate from the
2-nucleate embryo sac stage, are most abundant
in the 8-nucleate embryo sac stage but disappear
after fertilization.

During megasporogenesis and megagameto-
genesis, the nucellar tissue does not show any
particular differentiation and persists at least un-
til the earliest fruit stages (Figs. 36, 38). Apical
dermal cells of the nucellus do divide periclinally
(Fig. 37), and their daughter cells also repeat peri-
clinal divisions, thus forming a nucellar cap four
to six cell layers thick above the embryo sac (Fig.
36)

Integument. The ovule is unitegmic (Figs. 31,
36). The growing integument is about four or five
cells thick (Figs. 36; see also Fig. 38). No differ-
ence in thickness exists between the different parts
of the ovule. Therefore, except for the raphe, the
cross section of ovule is nearly circular (Fig. 38).
The integument is not vascularized. Neither sec-
ondary multiplication of the integument nor dif-
ferentiation of the innermost cell layer into a so-
called endothelium occur.

e integument elongates beyond the nucellar

rogamous. After fertilization, the fruit elongates
remarkably (Fig. 39). Endosperm formation is of
the Nuclear type (Fig. 40). In the early stages,
free endosperm nuclei are located around the

proembryo (Fig. 40) and at the peripheral region
of the embryo sac (Fig. 43). Because of incom-
pleteness of our fruit sample, we could not ob-
serve to what degree an amount of endosperm
increases later. The mature seeds lack endosperm
(Figs. 41, 42). We did not investigate embryo-
genesis in detail but can state, on the basis of our
observations of a few microtome-sectioned
proembryos, that it proceeds normally (Fig. 40).

ithin the mature fruit of Combretocarpus,
the embryo is elongate and nearly circular in
cross section (Figs. ). e embryo is
cotyledonous with so small cotyledons ele a
long hypocotyl (Fig. 41

The hypostase is not differentiated even after
fertilization.

Mature seed and seed coat. The mature seed
is linear, 9.5-10.4 mm long and 1.2-1.3 mm
thick; it contains several vascular bundles in the
raphe (Fig. 42), which are derived by ramifica-
tion from a raphe bundle. These bundles are re-
stricted to the raphe, never entering the integu-
ment or testa.

In the young seed, the seed coat is composed
of a tanniferous outer epidermis and a multiple
inner layer, which degenerates (Fig. 43). Even-
tually, in the mature seed, the seed coat com-
prises only the outer epidermis, which is formed
of pigmented, cuboid cells (Fig. 44)

POGA PIERRE

Anther and microspores. The anther is tetra-
sporangiate. The wall prior to maturation com-
prises five cell layers: an epidermis, endotheci-
um, two middle layers, and a tapetum (Fig. 45).
Wall formation conforms to the Basic type. Dur-
ing maturation, cells of the epidermis as well as
of the endothecium enlarge, while the middle
layers degenerate (Fig. 46). The tapetum is glan-
dular, and its cells become 2-nucleate (Fig. 46).
The two nuclei in a tapetal cell are not fused with
each other. The mature anther wall is composed
of the persistent epidermis and the endothecium.
The epidermis is tanniferous, and the endothe-
cium develops fibrous thickenings (Fig. 47). The
anther dehisces by longitudinal slits. By the time
of dehiscence, the connective tissue between two

<

nuclei. Bar = 20 um.— 21. Longitudinal hand-section ofa mature seed. ^
Bar

44 4 14

ar = 5 mm.— 22. LS of a young seed showing a thick seed coat. Bar = 200 um.— 23. LS of a mature seed coat
that is formed by both the multiple inner layer and the conspicuous outer epidermis. Bar — 40 um. in, integument;
nuc, nucellar tissue; rb, raphe bundle; b, vascular bundle; pem, proembryo; fe, free endosperm nucleus; em,

mbryo

; SC, seed coat; epi, epidermis of seed coat; inl, multiple inner layer.

10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

d i
reflexed. Bar = 50 um.— 28. Two-celled mature pollen at pi time of shedding. Bar = 10 um.— romosomes
of pollen mother cell at metaphase I. n — & Bar = 2 um. ep, epidermis; et, endothecium; ml, middle layer; t,
tapetum; mc, microspore mother cell.

1987]

microsporangia of each theca degenerates com-
l

tely.

Meiosis in the microspore mother cell is ac-
companied by simultaneous cytokinesis. The
shape of the resultant tetrads, on the basis of the
examination of 50 selected tetrads are “usually”
(86%) tetrahedral, “occasionally” (12%) decus-
sate, and “rarely” (2%) isobilateral. The pollen
grains are 2-celled at the time of shedding (Fig.
48

Chromosomes. Using serially sectioned pol-
len mother cells that were fixed at the later pro-
phase of meiosis I, we observed the chromo-
somes of Poga for the first time and determined
the chromosome number n = 7 (Figs. 49-51).

Nucellus and integuments. Although female
flowers were not available, we confirmed by us-
ing mature ovules and very young fruits that the
nucellar tissue enclosing the embryo sac persists
at least until the early stages of fruit development
(Figs. 52, 54). No hypostase is differentiated even
after fertilization.

The ovule is bitegmic, i.e., possessing the outer
and the inner integument (Fig. 52). The outer
integument is originally about four or five cells
thick, and the inner integument two cells thick
(Fig. 53). The cells of the outer epidermis of the
outer integument, which later become those of
the outermost layer of the exotesta, are conspic-
uously enlarged into cuboid cells. The raphe bun-
dle ramifies and vascularizes the outer integu-
ment. In a cross section of a young fruit, six to
eight vascular bundles in addition to several raphe
bundles are observed in the testa (Fig. 54).

The lom is formed by both integuments
(Fig. 5

l and embryo. We could not ob-
serve either the mode of endosperm formation
or embryogenesis. But we can say at least that
the mature seed completely lacks endosperm
(Figs. 55, 56) as Vaughan (1970) described, and
that the embryo does not have cotyledons. Con-
cerning the cotyledons, Vaughan (1970) men-
tioned that they are fused. However, judging from
the resemblance in exomorphology of the em-
bryo with Anisophyllea, it seems that Poga also
lacks cotyledons from the beginning.

Mature seed and seed coat. The fruits are al-
ways one-seeded. The mature seed is 20.0-22.5
mm long and 12.0-13.5 mm thick, is ovoid and
slightly suppressed toward the raphe-antiraphe
direction, and has a thick, dark brown seed coat
(Figs. 55, 56).

TOBE & RAVEN—ANISOPHYLLEACEAE 11

In the young seed, the seed coat is composed
only of a thick testa and lacks a tegmen. It seems
that, during the process of seed development, the
inner integument or tegmen is crushed, while the
outer integument or testa increases in thickness
by secondary multiplication. Within the young
testa, a differentiation into a multiple outer layer
and a multiple inner layer can be observed (Fig.
57). The multiple outer layer is about 8 cells thick
and has cells that are more or less enlarged. In
contrast, the multiple inner layer is 7—10 cells
thick, with the cells stretched tangentially (Fig.
57). The structure of the mature seed coat ba-
sically does not differ from that of the young seed
coat. In the mature seed coat, however, the walls
of the constituent cells are thickened, and the
multiple inner layer occupies nearly one-third of
the whole thickness ofthe testa, with the multiple
outer layer occupying the remaining two-thirds
(Fig. 58). Because our microtome sections of the
seed coat were not very good, we could not ex-
amine the details of cell structure. Vaughan
(1970), however, gave a drawing of the anatom-
ical structure of the testa, which consists of an
inner layer that is about 12 cells thick and an
outer layer about six or seven cells thick. Refer-
ring to Vaughan (1970), Corner (1976) described
the outer epidermis of the multiple outer layer
as composed of cuboid cells with slightly thick-
ened, lignified walls, and the other cells of the
multiple outer layer as thin-walle

The mature seed coat structure of Poga, which
is bitegmic, seems comparable with that of An-
isophyllea, which is unitegmic. In both genera,
the mature seed coat contains a similar (probably
identical) multiple inner layer, which is charac-
teristically composed of tangentially stretched
cells with thick walls. The only evident difference
between the mature seed coat structure of Poga
and that of Anisophyllea lies in thickness of the
outer layer, i.e., about six or seven cells thick in
Poga (see Vaughan, 1970) and one cell thick (out-
er epidermis only) in Anisophyllea. In other
words, T seed coat of Anisophyllea, like that of

oga, may also be constructed principally of the
34 or “outer Integument”), which of course
is not differentiated in the single integument of
Anisophyllea. Therefore the seed of unitegmic
Anisophyllea and even of unitegmic Combreto-
carpus (with a mature seed coat consisting only
ofthe outer epidermis) may be regarded as testal,
and the seed of bitegmic Poga can also be defined
in this way.

12 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

FIGURES 30-38. _Combretocarpus rotundatus. — 30. Longitudinal section (LS) of an ovule with the 1-celled
archesporium. Bar = 10 um.—31. LS of a young ovule with the primary parietal cell and the megaspore mother
cell. Note that the ovule has only a single integument. Bar = 10 um.— 32. LS ofa young ovule with the Beers gii
dyad. Note that each megaspore has two nuclei. Bar = 10 um.—3. LS of an ovule at the 2-nucleate embryo sa
stage. Bar — 10 um.— 34. LS of a nearly mature ovule at the 8- nucleate embryo sac stage. Bar = 10 uv idee

1987]

POLYGONANTHUS DUCKE

Anther and microspores. The anther is basi-
cally tetrasporangiate. The microsporogenous
tissue, however, is occasionally transversely di-
vided by a septum composed of tapetal cells (Fig.
59). Although we could not determine the modes
of anther wall formation, the wall prior to mat-
uration comprises five cell layers: an epidermis,
endothecium, two middle layers, and a tapetum.
During maturation, the middle layers degener-
ate, while the cells of both the epidermis and the
endothecium become enlarged (Fig. 59). The ta-
petum is glandular, and its cells become 2-nu-
cleate before degeneration (Fig. 60). The two nu-
clei in a tapetal cell do not fuse with each other.
Eventually the mature anther wall is composed
of a persistent epidermis, whose cells are some-
what collapsed in places, and a fibrous endothe-
cium (Fig. 61). The connective tissue between
two microsporangia of each pius b
completely before the anthers dehi

pollen grains are 2-celled at the time of shedding
(Fig. 62).

Chromosomes. Using serially sectioned mi-
crospore mother cells that were fixed at the late
prophase of meiosis I, the chromosomes of Po-
lygonanthus were observed for the first time.
Throughout, at examination and reconfirmation
in many cells, we determined the chromosome
number of P. amazonicus as n = 7 (Figs. 63-65).

Megagametophyte and nucellus. Although
our observations are fragmentary, we were able
to observe some aspects of the process of mega-
sporogenesis and megagametogenesis in Polygo-
nanthus. The ovule is anatropous and crassinu-
cellate. At least one parietal cell is cut off above
the megaspore mother cell (Figs. 66, 67). Al-
though the mode of embryo sac formation was
not determined, the 2- nucleate a sac of

ct from
‘hit ‘of Combretocarpus rotundatus Gi de-
velops a bisporic A//ium type embryo sac) (Fig.

TOBE & RAVEN —ANISOPHYLLEACEAE 13

68; compare with Fig. 33). The accumulation of
starch grains in the embryo sac, which is char-
acteristic of Combretocarpus, does not occur in
Polygonanthus amazonicus. The antipodal cells
are probably ephemeral, because they are absent
in the organized mature embryo sacs (Fig. 69)

The nucellar tissue enclosing the embryo sac
is persistent until at least the stage of fertilization
(Fig. 69). Periclinal divisions occur in the apical
dermal cells of the nucellus. Therefore the nu-
cellar cap is probably formed by derivatives of
the apical dermal cells.

Integuments. The ovule is bitegmic, i.e., it
has both an outer and an inner integument (Fig.
66). The outer integument is initially about five
cells thick, but later becomes seven to nine or
more cells thick because of secondary multipli-
cation (Fig. 66). The inner integument, in con-
trast, is two cells thick (Fig. 67). In the later stages,

u g t
creases in cn t 69). Although we could
not observe any stages of the development of the
seed coat, it seems very unlikely that the inner
integument or tegmen contributes to its structure
when mature. The raphe bundle ramifies
throughout the outer integument, which is there-
fore vascularized. In cross section, seven or eight
bundles in siii to several raphe bundles are
observed (Fig

he be is ‘fered by both integuments
(Fig. 69).

DISCUSSION

Our own results on the embryology and chro-
mosome numbers of Anisophylleaceae, together
with some data on ovule and seed morphology
published earlier (Karsten, 1891; Vaughan, 1970;
Geh & Keng, 1974), are presented in Table 2.
On this basis, we summarize the embryological
features of Anisophylleaceae as follows

Anther tetrasporangiate, but occasionally
polysporangiate because of insertion of tapetal
septa (Polygonanthus); anther wall with five cell

—

Polarized view of the same as that shown in Figure 34, showing a conspicuous accumulation of starch grains

in the embryo sac. Bar = 10 um.—36. LS ofa

mature ovule with an | organized embryo sac. Note that the nucellar

tissue is persistent. Bar = 50 um

ofay
divisions occurring in apical epidermal cells of the nucellus. Bar = = 10 umn. — —38. Transverse section ofa young

seed with twin embryo sacs. Bar
cell; in, integument; n, nucleus

O um. arc, archesporial cell; p, primary p
of the embryo sac: sy, a m pos eta ja purius a cell; nuc, nucellar

cell; mc, megaspore mother

tissue; pd, periclinal cell division; sc, seed coat; es, embryo

ANNALS OF THE MISSOURI BOTANICAL GARDEN

MEO

v
|

[T Q) £ L hI

IGURES 39—44. deer dud rotundatus. — 3 i
—40. LS of a proembryo with m endosperm nuclei surrounding it. Bar — 50 um. .—41.

fruit. Bar — el
i wath a cotyledonous embryo. Not

e that the mature seed is exalbuminous. Bar = 1 mm.—

of a matur
42 “atuntami section of a mature seed with several vascular bundles at the raphe. Bar = 500 um.—43. LS of

a young seed coat

Bar = 10 um.—44. LS ofa mature seed coat. Bar = 10 um. pem, proembryo; fe, free endosperm

nucellus; cot, cotyledon; em, embryo; sc, seed coat; rb, raphe bundle

layers, its formation of the Basic type; anther
epidermis persistent, consisting of more or less
collapsed cells; endothecium persistent and de-
veloping fibrous thickenings; middle layers

ephemeral; tapetum glandular, its cells 2-nu-

tually fused in Anisophyllea and Combretocar-
pus, but not in Poga and Polygonanthus.

1987] TOBE & RAVEN—ANISOPHYLLEACEAE 15

FIGURES 45-51. Poga oleosa. —45. Transverse section (TS) of a young anther showing the wall structure with
. TS of an older w). Bar = 10

five cell layers. Bar = ofa ther with degenerating middle layers (arro

m.—47. TS of a nearly mature ther wall consists of the fibrous endoth m and the epidermis. Bar =
100 um.—48. Two-celled mature pollen at the time of sheddi ndicate nuclei of the two cells. Bar =
10 e serial sections of pollen mother cell at the late prophas win romosomes of n

um.— 49-51. Thre S
7. Seven chromosomes are numbered | to 7. Bars = 10 um. ep, epidermis; et, endothecium; ml, middle layer;
t, tapetum

Cytokinesis in the microspore mother cells si- Gametic chromosome number n = 8 in Com-
multaneous; microspore tetrads tetrahedral, de- — bretocarpus, n = 7 in Anisophyllea, Poga, and
cussate, or isobilateral; pollen grains 2-celled Po/ygonanthus.

when shed. Ovule anatropous and crassinucellate; arche-

16 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FIGURES 52-58. Pogaoleosa.—52.1 itudinal section (LS) of ote that the ovule is a
and has a persistent nucellar tissue. Bar = 100 um.— 53. Transverse section Mes + a mature ovule showing the
inner integument, with two cell layers, and the eae hg oy with four cell layers. Bar = 10 um. M TS
ofa = 500 um 5. Longitudinal hand-section of the mature

sced. Note that the mature seed is exalbuminous. Bar = 5 mm.— — 56. Transverse hand-section of the mature
seed. Bar = 5 mm.— 57. TS of a young seed coat. Note that there is no tegmen and that the testa is E
into the multiple i inner layer and the multiple outer layer. Bar = 100 um.— 58. LS of a mature seed coat. Bar =

m. ii, inner integument; oi, outer integument; es, embryo sac; nuc, nucellar tissue; rb, eee bundle: b,
vascular bundle; sc, seed coat; em, embryo; inl, multiple i inner layer; oul, multiple outer layer

1987] TOBE & RAVEN—ANISOPHYLLEACEAE 17

Ficures 59-65. Polygonanthus amazonicus. — 59. Longitudinal section is a developed anther. Arrow indi-
cates the a septum dividing a microsporangium. Bar = 50 um.—60. Transverse section of a young Tax š
Bar = 10 um.— 61. TS of a mature anther at the time of dehiscence. ‘Its wall consi of the fibrous endothec

and the ME Bar = 50 um.— 62. Two-celled mature pollen at the time of shedding. Arrows indicate nuai
of the two cells. Bar = 10 um.—63-65. Three serial sections of pollen mother cell at E. prophase I showing
chromosomes of n = 7. Seven chromosomes are numbered 1 to 7. Bars = 10 um. t, tapetum; ep, epidermis; et,
endothecium.

18 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FiGURES 66-70. Polygonanthus amazonicus.—66. Lon gitudinal section (LS) of a young phon Note that the

ovule is bitegmic. Bar = 50 um.— 67.

stage. Bar — 10 um. —

ol, Outer integument; p, pri imary parietal cell; mc, geie

Same as that shown in Figure 66,
that the inner integument is two cells thick. Bar = 20 4
9. LS of a mature ovule. Aies in: the nucellar tissue is persistent.

outer integument has vascular

but shown at higher magnification. Note
at the 2- ien ons embryo sac
Bar = 50 um.— 70.
bundles. ii, inner integument,
r cell; n, nucleus of the embryo sac; es,

m.—68. LS of an ovule

ore mother

mbryo sac; nuc, nucellar tissue; rb, raphe bundle; b, vascular bundle.

ari l sus ine: off a primary parietal

mbryo formation of the Polygonum
oe » nisoph opk or the Allium type (Combre-
tocarpus), synergids slightly hooked; antipodals
ephemeral; polar nuclei fused before fertilization
(Anisophyllea) or not fused (Combretocarpus).
Nucellar tissue not degenerating at least until the
early stages of seed development; apical dermal

nucellar cells dividing periclinally, forming a nu-
cellar cap, chalaza without a hypostase.

Ovule unitegmic (Anisophyllea and Combre-
tocarpus) or bitegmic (Poga and Polygonanthus);
in bitegmic ovules, the inner integument two cells
thick and the outer integument thicker; outer
integument vascularized due to ramification of
raphe bundles, but not vascularized in Combre-

1987]

tocarpus, micropyle formed by either the one

uclear type; seed exalbuminous;
mode of embryogenesis not determined; embryo
(potentially) dicotyledonous with a long hypo-
cotyl, having either small cotyledons (Combre-
tocarpus) or rudimentary and/or no cotyledons
(Anisophyllea and Poga). Seed coat testal (Poga)
or logically testal (Anisophyllea and Combreto-
carpus); mature seed coat formed by the outer
epidermis alone (Combretocarpus), both the out-
er epidermis and the multiple inner layer (Ani-
sophyllea), or both the multiple outer layer and
the multiple inner layer (Poga).

RELATIONSHIPS WITH RHIZOPHORACEAE

Although, as shown in Table 2, some of the
embryological features of Anisophylleaceae are
diverse, the family is consistent enough in most
such characteristics to allow a more critical com-
parison with Rhizophoraceae than has hitherto
been possible. In summary, these two families
share only a few embryological features. They do
agree, for example, in having a crassinucellate,
bitegmic ovule and the Nuclear type of endo-
sperm formation; but a combination of these and
other shared features is widespread among many
other unrelated families of the angiosperms as
well.

In contrast, psi Sana ipie from Rhi-
zophoraceae in som portant embryological
features. First of all, in i yaya the nu-

ceae the nucellar tissue is ephemeral, disappear-
ing completely by the time of fertilization (see
Karsten, 1891, for Rhizophora, Ceriops, Bru-
guiera, and Carallia; Cook, 1907, for Rhizopho-
ra; Carey, 1934, for Rhizophora; Mauritzon,
1939, for Gynotroches; Juncosa, 1984a, 1984b,
for Bruguiera and Cassipourea). Therefore in
Rhizophoraceae the Midi sac borders directly
on the inner integum

Secondly, in ee NR (Poga and Po-
lygonanthus) the inner integument is character-
istically two cells thick, whereas in Rhizophora-
ceae it is much thicker. Indeed, an inner
integument with four to eight layers has been
illustrated by Karsten (1891) for Bruguiera, Ceri-
ops, and Carallia, by Carey (1934) for Rhizopho-
ra, and by Mauritzon (1939) for Bruguiera and

TOBE & RAVEN — ANISOPHYLLEACEAE 19

Gynotroches. Juncosa (1984a, 1984b) described
the inner integument of Bruguiera exaristata as
initially being about 10 cells thick and that of
Cassipourea elliptica as being about five to eight
cells thick. In addition, a specialization of the
innermost cell layer of the inner integument into
an endothelium has been reported in some in-
land genera of Rhizophoraceae, including Car-
allia (Karsten, 1891), Gynotroches (Mauritzon,
1939), and Cassipourea (Juncosa, 1984a). An en-
dothelium is never formed in Anisophylleaceae.

Thirdly, the mature seed is exalbuminous in
Anisophylleaceae, but albuminous in Rhizopho-
raceae. The presence of abundant endosperm in
mature seeds has been reported for Rhizophora
(Cook, 1907; Carey, 1934; Juncosa, 1982), Ceri-
ops (Carey, 1934), and Cassipourea (Juncosa,
1984a

Some critical differences in embryo and seed
coat morphology might also be added It distin-
guishing /
(see Comer, 1976). 2 evel, studies on those
ll too limited
to allow this. Further studies on the embryology
of Rhizophoraceae, including embryo and seed
coat morphology, are needed to clarify the dif-
ferences between this family and Anisophylle-
aceae

To sum up, despite insufficient information on
the embryology of Rhizophoraceae, the available
data indicate that Anisophylleaceae differ sig-
nificantly from them. If Anisophylleaceae were
included as a tribe or subfamily, Rhizophoraceae
sensu lato would be defined very broadly. With
the support of exclusive occurrence of the nature
of aluminum accumulation (Chenery, 1948; Ku-
kachka & Miller, 1980); alternate, exstipulate
leaves; three or four free styles (Geh & Keng,

ai
Anisophylleaceae and Rhiz
closely related and warrants regarding Aniso-
phylleaceae as a distinct family.

SYSTEMATIC POSITION OF ANISOPHYLLEACEAE

Cronquist (1981, 1983) has proposed assigning
Within Rosales,

(Hydrangeaceae, Columelliaceae, Grossulari-
aceae, Greyiaceae, Bruniaceae, and Alseuosmi-
aceae). Of these, only Grossulariaceae have been

20

ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE 2. Embryological and chromosomal data of Anisophylleaceae.

[VoL. 74

Number of integu-
ments

Thickness of integ-
uments when bi-
tegmic

i.i. 2 cell-layered;
0.i. 4—5 cell-lay-
ered

Character Anisophyllea Combretocarpus Poga Polygonanthus
Anther and microspores:
Number of sporan- 4 4 4, sporangium occa-
gia sionally divided
Anther wall devel- Basic type Basic type Basic type
opment
Anther epidermis Persistent Persistent a Persistent
Endothecium Fibrous Fibrous Fibro ibrous
Tapetum Glandular Glandular Cur Glandular
Number of tapetal 2 2 2 2
nuclei
Tapetal nuclear fu- Occur Occur Not occur Not occur
sion
Cytokinesis in Simultaneous Simultaneous Simultaneous Simultaneous
meiosis
Shape of micro- Usually tetrahe- Usually tetrahe- Usually tetrahe- ?
spore tetrad dral, very occa- dral, oo dral, occasional-
sionally decus- ly dec ly decussate,
: very aie rarely isobilater-
isobilateral isobilateral a
Mature pollen 2-celled lled 2-celled 2-celled
Chromosomes:
Base number x27 x=8 x=7 x27
Megagametophyte and nucellus:
Ovule curvature Ana Anatropous Anatropous Anatropous
Nature of nucellus Crassinucellate Crassinucellate Crassinucellate Crassinucellate
Archesporium 1-celled arly al ? ?
1-celled, very
rarely 2-celled
Thickness of pari- 3-5 cell-layered 1-2 cell-layered ? ?
etal tissue
Shape of mega- Linear Linear ? ?
spore tetrad
Functional mega- Chalazal cell Chalazal cell ? ?
spore
Pattern of embryo Polygonum type Allium type T ?
sac formation
Synergids Slightly hooked Slightly hooked ? ?
Antipodal cells Ephemeral Ephemeral ? Probably ephemeral
r of nuclei 5 5 ? 5
or 5 cells in ma-
e embryo sac
Accumulation of Not occur Occur ? Not occur
starch grains in
embryo sac
Nucellar tissue Persistent Persistent Persistent Persistent
Nucellar cap orme Formed ? Probably formed
Hypostase Not formed Not formed Not formed Not formed
Integuments:
l l 2 2

1.1. 2 cell-layered;
o.i. about 5 cell-
layered

1987]

TABLE 2. Continued.

TOBE & RAVEN—ANISOPHYLLEACEAE 21

Character Anisophyllea Combretocarpus Poga Polygonanthus
Vasculature Present Absent Present Present
Micropyle forma- By the only integu- By the only integu- By both integu- By both integu-

tion ment ment ments ments

Differentiation of Not occur Not occur Not occur Not occur

endothelium
Fertilization, endosperm, and embryo:

Path of pollen tube Porogamous Porogamous ? ?

Tem forma- Nuclear type Nuclear type ? ?

ius in ma- Absent Absent Absent ?
ture see

Embryogenesis ? ? ? ?

Embryo in mature e Or Cotyledonous Not cotyledonous ?

seed ot cotyledonous

Size of cotyledons un small (rudi- Small — ?

when present mentary)
Mature seed and seed coat:

Shape of seed Narrow-cylindrical Linear Ovoid and slightly ?
(A. disticha); suppressed to-
ovoid or ellipti- ward raphe-an-
cal (A. sp.) tiraphe direction

Size of seed 13.0-13.6 mm 9.5-10.4 mm long 20.5-22.5 mm
ii. = 3.8-4.0 and 1.2-1.3 mm long, 12.0-13.5

m. (A. diam. mm wide
pus 13.0-
13.8 mm long
and 6.0-6.4 mm
diam (A. sp.)
Tegmen — — Ephemeral ?
Whole thickness of 26-31 cell-layered 1 cell-layer 17-20 cell-layered ?
see C
Thickness of inner 25-30 cell-layered — 7-10 cell-layered 7
layer o

Thickness of outer l cell-layer 1 cell-layer About lO cell-lay- ?

layer of SC ered

i.i., inner integument; o.i., outer integument.

relatively well studied embryologically, whereas
the others have been studied little or not at all

Grossularine h exalbuminous seeds
(Cronquist, 1). On the other hand, so-
phylleaceae resemble Grossulariaceae (princi-

pally Ribes, from which most data are available)
in nearly all features of anther and microspore
development; in their anatropous, bitegmic, and
crassinucellate ovule; Polygonum-type embryo
sac; ephemeral antipodal cells; inner integument
with two cell layers (see Davis, 1966; Corner,
1976; Cronquist, 1981, for data on Grossulari-
aceae). However, Anisophylleaceae differ from
Grossulariaceae in several embryological fea-
tures. For example, the tapetal cell is basically

2-nucleate in Anisophylleaceae, but multinu-
cleate in Grossulariaceae; the integument is vas-
cularized in Anisophylleaceae, but not vascular-
ized in Grossulariaceae; endosperm formation is
of the Nuclear type in Anisophylleaceae, but of
the Cellular or the Helobial type in Grossulari-
aceae; the tegmen is ephemeral in Anisophylle-
aceae (Poga), but persists in Grossulariaceae; the
seeds are non-arillate in Anisophylleaceae, but
arillate in Grossulariaceae (see 3j que 1926;
Davis, 1966; Corner, 1976, for data on Gros-

sulariaceae). Therefore it seems that available

R
In contrast, Dahlgren (1983) pum Aniso-

22 ANNALS OF THE MISSOURI BOTANICAL GARDEN

phylleaceae in the Cornales, which comprise 27
families including Hydrangeaceae (and five fam-
ilies whose position is uncertain; see also Dahl-
gren & Thorne, 1984). Of 27 families, nine have
either not been studied embryologically, or have
been studied only to a very limited degree. Of
the 18 remaining families, nearly all share a uni-
tegmic ovule, ephemeral nucellar tissue, endo-
thelium, Cellular type of endosperm formation,
and albuminous seed. The Cornales thus seem
to be very well defined by a combination of those
shared embryological features. Anisophylle-
aceae, which lack any of those characteristic em-
bryological features of the Cornales (almost cer-
tainly unitegmic ovule in Anisophyllea and
Combretocarpus), seem clearly distinct from the
Cornalean families and do not belong in that
order.

We would rather suggest Myrtalean affinities
for Anisophylleaceae. Embryologically, Aniso-
phylleaceae agree almost completely with Myr-
tales, and in fact share the eight ordinal embry-
ological features (see Tobe & Raven, 1983, 1984):
1) anther tapetum glandular, 2) ovule crassinu-
cellate, 3) inner integument two cells thick, 4)
micropyle formed by both integuments, 5) an-
tipodal cells ephemeral, 6) endosperm forma-
tion— Nuclear type, 7) seed exalbuminous, and
8) mature pollen 2-celled. One might point out
a fusion of tapetal nuclei (in Combretocarpus and
Anisophyllea), formation of the nucellar cap, and
testal seed as features distinguishing Anisophyl-
leaceae from Myrtales. However, nuclear fusion
in the tapetal cells is undoubtedly a secondary
characteristsic that evolved in two genera of An-
isophylleaceae. Indeed Poga and Polygonanthus,
both of which have many primitive features, as
will be discussed later, have unfused tapetal nu-
clei. The nucellar cap, which is formed by deriv-
atives of the apical nucellar dermal cells, is com-
monly observed in Combretaceae (Myrtales;
Venkateswarlu & Rao, 1972). A seed coat lacking
a tegmen is frequent in Melastomataceae (Myr-
tales; Corner, 1976). Anisophylleaceae may dif-
fer from Myrtales in having embryos with re-
duced or rudimentary cotyledons and a long
hypocotyl. Such an embryo morphology seems
to result in hypogeal germination, which is re-
ported in at least Anisophyllea disticha (Geh &
Keng, 1974). The peculiar embryo morphology
and germination habit may suggest a specialized
position of Anisophylleaceae. However, embryos
devoid of cotyledons are recorded in many un-
related families, a majority of them growing in

[VoL. 74

ecologically specialized habitats (Natesh & Rau,
1984, review). Study of embryogenesis and or-
ganogenesis in seeds through germination seems
to be needed for the elucidation of the ecological
significance of such specialized embryos in An-
isophylleaceae. Except for the difference in em-
bryo morphology, there seems to be essentially
a perfect correspondence in embryological fea-
tures between Anisophylleaceae and Myrtales.
Viewing other reproductive and vegetative
character states, Anisophylleaceae lack both the
intraxylary phloem and the vestured pits, which
are regarded as characteristic features defining
the Myrtales (Van Vliet, 1976; Van Vliet & Baas,
1984; Dahlgren & Thorne, 1984). However, the
occurrence of S-type sieve-element plastids in
Anisophyllea and Combretocarpus, in contrast
with the P-type plastids in Rhizopt agrees
with Myrtales (Behnke, 1982, 1984). Tricolpor-
ate pollen morphology in Anisophylleaceae (as
well as in Rhizophoraceae) is of the basic type
found in the Myrtales (Erdtman, 1966; see also
Dahlgren & Thorne, 1984). Aluminum accu-
mulation characteristic of Anisophylleaceae (un-
known in Rhizophoraceae) is known to occur in
Crypteroniaceae and Melastomataceae of Myr-
tales (Chenery, 1948; Kukachka & Miller, 1980).
Thus, considering a considerable number of
coincidences (in reproductive anatomy) in con-
trast with a limited number of differences (in
wood anatomy), in conjunction with support by
sieve-element plastid type, palynology, and alu-
minum accumulation, it seems that Anisophyl-
leaceae are closely related to Myrtales. Depend-
ing on how we interpret the lack of intraxylary
phloem and vestured pits in Anisophylleaceae,
it might even be justifiable to place Anisophyl-
leaceae in the Myrtales. According to Van Vliet
and Baas (1984), the combined occurrence of
intraxylary phloem and vestured pits is very re-
stricted in the dicotyledons; in fact, except for
the Myrtales, this combination is found only in
part of the Gentianales, Thymelaeales, Poly-
galales, and Polygonales. Outside these orders,
either one of these features (but not both) spo-
radically occurs in many different groups of di-
cotyledons (see Van Vliet & Baas, 1985: 784, fig.
1). Only a few orders are characterized by con-
sistent possession of one or both of those two
wood anatomical features. Therefore it does not
seem that the lack of intraxylary phloem and
vestured pits in Anisophylleaceae necessarily
precludes a possibility of close affinity with Myr-
tales. Based on total evidence now available, we

1987]

would suggest that Anisophylleaceae be placed
near Myrtales. Perhaps Anisophylleaceae rep-
resent one of the groups that diverged from a
common ancestral stock with Myrtales and then
spread widely. The position of Anisophylleaceae
will be evaluated better as the Rosiflorae or the
Rosales, which are considered phylogenetically
basic in position with respect to Myrtales, are
understood better embryologically.

INTERRELATIONSHIPS AND EVOLUTION
OF THE GENERA

Because of many shared embryological fea-
tures, as shown in Table 2, as well as of shared
vegetative and some other shared reproductive
features (see also Geh & Keng, 1974; Van Vliet,
1976), there is no doubt that Aniopibiled. Com-
bretocarpus, Poga, and Polygonanthus are mono-
phyletic. Despite the lack of data about several
features in Poga and Polygonanthus, the avail-
able embryological data are now enough to allow
us to compare all four genera.

Of these, Combretocarpus is the most distinct.
It has a gametic chromosome number of n = 8,
Allium type embryo sac, nonvascularized integ-
ument, starch grains in the embryo sac, cotyle-
donous embryo, and thin mature seed coat one
cell layer thick. In contrast, Anisophyllea, Poga,
and Polygonanthus have a chromosome number
of x = 7, Polygonum type embryo sac (unknown
in Poga and Polygonanthus), vascularized integ-
ument, no starch grains in the embryo sac, non-
cotyledonous embryo (unknown in Polygonan-
thus), and thick mature seed coat (unknown in
Polygonanthus). Combretocarpus agrees with
Anisophyllea only in having fused tapetal nuclei
and a unitegmic ovule. On the contrary, Aniso-
phyllea differs from Poga and Polygonanthus in
sharing neither bitegmic ovules nor distinct ta-
petal nuclei as well as in not sharing a multiple
outer layer in the mature seed coat (though un-
certain in Polygonanthus). Polygonanthus differs
from Poga only in its occasionally divided mi-
crosporogenous tissue. Except for this, there is
no essential difference between Poga and Poly-
gonanthus, as far as the data available are con-
cerned.

In order to clarify phylogenetic interrelation-
ships ofthe genera, it seems necessary to evaluate
each of the characters showing differences be-

ua mbryologi
phylleacca following Eldredge and Cracraft
(19

° + 4 o

TOBE & RAVEN—ANISOPHYLLEACEAE 23

of character state similarities (i.e., synapomor-
phies or symplesiomorphies). Of the embryolog-
ical features, the Polygonum type embryo sac
formation (Anisophyllea) that is characteristic of
a majority of the dicotyledons (Davis, 1966) is
undoubtedly primitive to the Allium type man-
ner (Combretocarpus), and also bitegmy (Poga
and Polygonanthus) is primitive to the unitegmy
(Anisophyllea and Combretocarpus, Bouman,
1984).

Concerning the vasculature of the integument,
there 1s no consensus regarding whether or not
the vascularized integument represents an ar-
chaic condition. Bouman (1984) suggested that
there seems to be a general relation between the
size of ovules or seeds and the degree of vascu-
larization. As far as Anisophylleaceae are con-
cerned, the vascularized integument or testa (Ani-
sephyllet, Poga, and Polygonanthus) i is baee
he
ii vaseclaeieed one Fe aR E
bretocarpus has multiple vascular pieds in vin
raphe of the mature seed (see Fig. 42). This vas-
cular condition in Combretocarpus is probably
derived from the condition seen in the three oth-
er genera by suppression of vascular extension
into the integument, because Combretocarpus has
the thin integument that eventually becomes the
one cell-layered seed coat at maturity. In this
connection, the thick mature seed coat or testa
is probably primitive (symplesiomorphous) to
the thin, one cell-layered mature seed coat. Com-
pared with Poga, Anisophyllea lacks a hypo-

oga;
Innerlayerand the hypodermal tissue ofthe mul-
tiple outer layer. Corner (1976, vol. 1: 57) has
considered the limitation of a multiple mechan-
ical tissue (probably like that of Poga) into one
cell-layered as one of specialization trends of seed
coat. Following Corner, we may be able to pos-
tulate that the successive or simultaneous re-
duction of the multiple inner layer and the hypo-
dermal tissue of the multiple outer layer had
occurred in the seed coat evolution of Aniso-
phylleaceae so that only the epidermis was per-
sistent, as in Combretocarpus. Although we did
not observe the anatomy of the testa of Polygo-
nanthus, it was confirmed that the (outer) integ-
ument shows a seconda
dition that is cle

ombretocarpus. Therefore it seems very likely
that Polygonanthus would form a mature seed
with as thick a testa as in Poga.

24 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE 3. Evolutionary trend of karyological and
some embryological characters in Anisophylleaceae.

Plesio-
Characters morphy Apomorphy

1. pei of integu- 2 l

2. Pio nuclei Not fused Fused

3. “Outer layer" of the Thick Thin, 1 cell
“thick” seed coat thick
(thickness)

4. Chromosome number =7 n=8

5. Pattern of embryo sac Polygonum Allium type
formation type

6. “Inner layer" of the Present Absent
mature seed coat

7. Accumulation of Absent Present
starch grains in de-
veloping embryo

8. Present Absent

sac
Vasculature of integu-
ments

The accumulation of starch grains in the de-
veloping embryo sac (Combretocarpus) is known
to occur in many unrelated families of dicoty-
ledons (see Davis, 1966). Even within a family,
however, their occurrence is in general restricted

y by particular groups in many unrelated
families, probably because of the necessity of dif-
ferent metabolic activity during megagameto-
genesis.

Embryos with moderately developed cotyle-
dons are almost universal among dicotyledons
and therefore must be primary. On the contrary,
because of their restricted occurrence (see Natesh
& Ram, 1984), embryos lacking cotyledons seem
to be secondary in the evolutionary trend. In this
respect, it might be interpreted that an embryo
with small cotyledons (Combretocarpus) seems
less specialized than that which lacks cotyledons,

or at the most has rudimentary ones (Aniso-
phyllea and Poga). Such differences in the degree
of size reduction of cotyledons, however, may be

share a conspicuous hypocotyl, a truly significant
and unusual feature

It may be difficult to determine whether the
chromosome base number of Anisophylleaceae
is n = 7 or 8. Noticeably n = 8 occurs only in

[Vor. 74

Combretocarpus, a genus that is furnished with
many advanced and fewer primitive character
states as discussed above. In contrast, n — 7 is
shared by Anisophyllea, Poga, and Polygonan-
thus, all of which — particularly the latter two—
retain a combination of primitive character states.
Thus it seems likely that n = 7 is the base number
of the family, and n = 8 the derived.

The fusion of tapetal nuclei (Anisophyllea and

t O

— (Poga and Polygonanthus), as discussed
arlier. These nuclei also remain distinct in most

Made, (Tobe & Raven, 1983).

The results of our evaluation of these character
states are summarized in Table 3. On this basis,
we constructed a cladogram illustrating the evo-
lutionary interrelationships of the genera (Fig.
71). The cladogram indicates that the proto-Ani-
sophylleaceae, a hypothetical ancestor of the
family, had nearly all of the embryological fea-
tures that are presently retained by Poga and

tually uncertain in Poga an
tegmy, vascularized integument, thick seed coat
consisting ofa multiple inner layer and a multiple
outer layer, no starch grain accumulation during
megagametogenesis, and non-fused tapetal nu-
clei. An tral luti y li toh

diverged into two main branches: one leading to
Combretocarpus and Anisophyllea, and the other
leading to Poga and Polygonanthus. In the for-
mer branch, the ovule became unitegmic; tapetal
nuclei fusion has been generalized, and the thick-
ness of the multiple outer layer was reduced into
one cell layer (i.e., the outer epidermis); all three
characters are synapomorphies common to
Combretocarpus and Anisophyllea. This branch
further diverged into two branchlets: one leading
to Combretocarpus, and the other leading to Ani-
sophyllea. In the branchlet leading to Combre-
tocarpus, chromosome base number changed to
n = 8; the Allium type embryo sac and unitegmy
were derived; complete reduction of seed coat
tissue except for the outer epidermis (i.e., of both
a multiple inner layer and the hypodermal tissue
of the original thick seed coat) and reduction of
mentary vasculature occurred nearly si-

e.
B

ge
B
O

ri gam
respect to embryolog
ing change has pique in the other branchlet

1987]

e
° SY
S o n

` C

„O

"2
S)

d

o

[— 5
[ . 1- pe 4
e 3
=C 2
F 1
FiGuRE 71. A cladogram illustrating postulated

evolutionary interrelationships of the genera of Aniso-
phylleaceae. Corresponding characters numbered 1 to
8 and their evolutionary states are shown in Table 3.
Shaded rectangles indicate apomorphies. *Character
state uncertain in Polygonanthus; "Character state un-
certain in Poga.

leading to Anisophyllea, or in the other main
evolutionary line leading to Poga and Polygo-
nanthus.

Consequently, Combretocarpus differs sub-
stantially from the three other genera and is ap-
parently the most specialized member of the
family. In contrast, most plesiomorphies are re-
ferred to Poga and Polygonanthus, which both
retain a combination of primitive embryological
character states common to the ancestor of Ani-
sophylleaceae. Anisophyllea stands in a more or
less intermediate position between Combreto-
carpus, on the one hand, and the group com-
prising Poga and Polygonanthus, on the other.

TOBE & RAVEN—ANISOPHYLLEACEAE 25

Indeed Anisophyllea shares most of its archaic
features with Poga and Polygonanthus but shares
its apomorphies, including unitegmy and tapetal
nuclear fusion, with Combretocarpus.
Phylogenetic interpretations of the infrafamil-
ial relationships as discussed above are primar-
ily based on embryological character state eval-
uation and chromosome number. The cladogram
shown in Figure 71 is to be regarded as limited
in this respect and provisional. The evolutionary
trend in certain characters (such as chromosome
number and seed coat structure) might be the
opposite of what we have proposed, and certain
characters (such as tapetal nuclear condition)
might be of much less fundamental significance
than others. Earlier studies on Anisophylleaceae
he

whether or not Polygonanthus was closely related
to the other three genera (for instance, Pires &
Rodrigues, 1971) or whether or not Anisophyl-
leaceae (*Anisophylleae") should be excluded
from Rhizophoraceae (for instance, Van Vliet,
1976). We now regard both of these questions as
definitively solved and hope that our phyloge-
netic diagram will stimulate further research on
the family from various other points of view and
will thus be improved as a result of these inves-
tigations.

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en
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TRAPLINERS IN THE TREES: HUMMINGBIRD
POLLINATION OF ERYTHRINA SECT. ERYTHRINA
(LEGUMINOSAE: PAPILIONOIDEAE)!

DaviD A. NEILL?

ABSTRACT

Erythrina sect. Erythrina Mdh re) specie
tributed principally in Mesoam
nators— were observed at 17 ciun P 13 spe

pollinators were all "high- reward traplining" ada

behavior, including in arduus
short- € d

es t
defense by a hummingbird of a ui tree is piri eui The traplining hummii

n flo ral visit tors—includin

s of hummingbird- viae i -= and shrubs, dis-
g nectar -

sins southern Mexi xico an pos Rica

e high-reward

traplining systems involving hermit hummingbirds and understory plants such as Heliconia (Musa-
ae).

Erythrina L. (Leguminosae: Papilionoideae)
comprises about 112 species distributed
throughout the tropical regions of the world and
extending into warm temperate areas in both the
northern and southern hemispheres (Krukoff &
Barneby, 1974; Neill, in press). Most species are
trees or shrubs, but about 10 species occurring
in climates with pronounced dry and/or cool sea-
sons are perennial herbs with large, woody root-
stocks. Erythrina species occur in a very wide
variety of habitats, from lowland tropical rain
forests to very arid subtropical deserts to high-
land coniferous forests above 2,500 m in ele-
vation.

Erythrina species have red or orange flowers
and copious nectar and are adapted to pollina-
tion by nectarivorous birds. Two distinct syn-
dromes of ornithophily are evident. All 42 of the
World species and 15 of the 70 New World

ine birds cannot hover efficiently or for any length

of time, and the inflorescences of passerine-pol-
linated Erythrina are oriented in such a way that
the birds can perch while feeding on nectar from
the flowers. The corolla standard is usually broad

and the flowers are open, with exposed repro-
PER S parts. Pollen is deposited on the feeding
bird’s breast (Cruden & Toledo, 1977). In con-
trast, 55 of the New World species of Erythrina
(nearly half the genus) are pollinated by hum-
mingbirds (Trochilidae), which occur only in the

“pseudotube,” concealing the wing and keel pet-
als as well as the reproductive parts. The flower
resembles the tubular corollas of many gamopet-
alous hummingbird-pollinated plants, but in Er-
ythrina the pseudotube is not sealed on the ven-
tral side where the margins of the corolla standard
meet. The inflorescence axis of the humming-

' Discussions with Peter Feinsinger, Peter Raven, and Héctor Hernandez were helpful in the planning and
execution of this research and in the interpretation of the results. Frederick Neill’s Tae and companionship

all during the fieldwork in Mexico was invaluable. Francisco Guerrero and Rom

Velásquez, biologists at El

Sumidero National Park, helped with the fieldwork there. Alina Chacón, Mary Merello, Gloria Hoch, and Donna

Krausz assisted with the preparation of the manuscript

and figu

and Missouri Botanical Garden. The research was funded by grants from the National Science Foundation (DEB
81-20386), the Jesse Noyes Smith Foundation and the Organization for Tropical Studies for fieldwork in Costa

Rica, and Elizabeth Neill

? Missouri Botanical Garden, St. Louis, Missouri 63166.

ANN. Missouni Bor. GARD. 74: 27-41. 1987.

28 ANNALS OF THE MISSOURI BOTANICAL GARDEN

bird-pollinated species is erect and the flowers
are oriented outward, providing no perch for the
hovering hummingbirds.
The American |
are included in six different sections of subg. Er-
ythrina; these I believe to have been derived from
passerine-pollinated groups by convergent evo-
lution in several independent lineages. The larg-
est by far of the hummingbird-pollinated groups
is sect. Erythrina (36 species) with its center of
diversity in northern Central ; outh-
ern Mexico (Neill, in press). Most species a sect.
Erythrina are canopy or subcanopy trees, ranging
in height from 5 m in semiarid scrub to 25 m or
more in lowland rain forest or cloud forest. A
few species inhabit the understory and light gaps
of wet forests. Many have restricted ranges and
are edaphic specialists; an example is E. tux-
tlana, which grows only on limestone outcrops
in lowland wet forests in southeastern Mexico.
A number of field studies of pollination in Er-
ythrina have been conducted in recent years, in-
cluding observations of passerine- pira as
well as ł
1974; Toledo & ] Hernández, 1979; ED à
Toledo, 1979, 1982; Cruden & Toledo, 1977;
Feinsinger et al., 1979; Morton, 1979; Steiner,
1979; Guillarmod et al., 1979). An extensive sur-
vey of hummingbird pollination of Erythrina tree
pecies, however, was lacking prior to the re-
search reported here.

5 Ree | 11; 4 2

9

HUMMINGBIRD FORAGING BEHAVIOR:
A REVIEW

Hummingbird species differ in size, bill mor-
phology, and foraging behavior as do the floral
morphology and flowering patterns of the plant
species they visit (Feinsinger & Colwell, 1978).
The two principal behavioral types are territo-
rialists, which feed at large patches of flowers,
defen ing the flowers against usurpers; and trap-

relatively small, short- billed birds, and “hig

reward" trapliners, birds with relatively i.
bodies, high energetic requirements, and long or
curved bills. High-reward trapliners are the most
"specialized" of hummingbirds in the sense that
they have the highest fidelity to particular plant
species. These are the only birds capable of pol-

[VoL. 74

linating plants with iene pula flowers, and they

ceto plant
species with low population densities. The most
well-studied of the high-reward trapliners are the
hermit hummingbirds (Phaethorninae), which
forage principally in lowland and mid-elevation
wet forest understories on widely spaced, nectar-
rich herbs and shrubs.

Stiles (1978, 1981) noted that very few hum-
mingbird-pollinated plants are canopy trees; in
contrast, many ornithophilous plants of the Old
World, pollinated by passerine birds, are trees.
Stiles reasoned that this difference is related to
the social systems of the two bird groups. Pas-

lly forage in large flocks,
whereas hummingbirds are virtually always sol-
itary. A large concentration of flowers on a large
tree would be parceled up into feeding territories

pollination. A large flock of pass
contrast, could quickly exhaust the resources of
even a large tree, and the flock would be com-
pelled to move on to the next tree, thus effecting
cross-pollination.

Erythrina is an exception to the general paucity
of hummingbird-pollinated canopy or subcan-
opy trees. It should be instructive to determine
by observations in the field whether Stiles’s pre-
diction holds true for hummingbird-pollinated
Erythrina. If hummingbirds parcel the crown of
a tree into several feeding territories, they may
reduce intertree pollen flow; alternatively, hum-
mingbird-pollinated Erythrina trees may possess
adaptations that reduce territorial behavior and
promote intertree movement of the humming-
bird pollen vectors.

In the present study of hummingbird polli-
nation in natural populations of trees of sect.
Erythrina, several questions were addressed: Do
the hummingbirds that pollinate Erythrina be-
have as territorialists or as trapliners? How spe-
cialized are Erythrina pollinators? Another goal
was to assess the potential for pollen transfer
between different species of Erythrina. Experi-
mental hybridization studies (Neill, in press) in-
dicate that species of sect. Erythrina are highly
interfertile, but unless the hummingbird pollen
vectors carry pollen from one species to another,
hybridization will not take place under natural
conditions. This assessment required informa-
tion about the flight and foraging patterns of the
birds, and whether the Erythrina species shared
the same pollinators or had different, host-spe-
cific pollinators.

1987]

MATERIALS AND METHODS
FLORAL BIOLOGY AND BREEDING SYSTEMS

Observations on phenology and other details
of Erythrina floral biology, as well as experi-
mental studies of genetic self-incompatibility,
were made on wild populations in Mesoamerica
and on cultivated trees in Hawaii. The experi-
mental methods for the breeding system studies
are described by Neill (in press).

OBSERVATIONS OF FLORAL VISITORS

Floral visitors to 17 populations of 13 species
of hummingbird-pollinated Erythrina trees were
observed in Mesoamerica. These included four
wet season-flowering species in Costa Rica, July—
September 1981, and nine dry season-flowering
species in southern Mexico, January—April 1983.
All but one of the species is in sect. Erythrina;
the exception is E. gibbosa from Costa Rica, in
the monotypic sect. Gibbosae. Flowering and
pollination patterns in this species are very sim-
ilar to those in sect. Erythrina.

For the 13 species a total of 195 person-hours
of observation was conducted. For a population
the number of observation hours ranged from
2.5 to 45.5. Most of the observations were made
between dawn and 12 P.M., because floral visitor
activity is usually greater in the early morning
and drops substantially by noon. Some obser-
vations were conducted in the late afternoon Saen
avian floral
after the midday lul

At the beginning y each observation day I
counted the number of open flowers on each tree.
For each bird visit to a tree crown, I recorded
the time, duration of visit, number of flowers
probed, and direction of arrival and departure.
I judged qualitatively the frequency of the visi-
tor's contact with the reproductive parts of the
flower, as a measure of potential pollination ef-
ficacy. Many of the non-pollinating passerine bird
visitors actually destroyed or removed a consid-
erable number of flowers daily, reducing the
number available to subsequent visitors to the
tree. This activity was recorded and entered into
the daily flower censuses.

NECTAR PRODUCTION

I sampled daily nectar production in four
species in sect. Erythrina: E. globocalyx and E.

NEILL— ERYTHRINA POLLINATION 29

costaricensis inhabiting wet forests in Costa Rica,
and E. chiapasana and E. goldmanii in dry for-
ests of Chiapas, Mexico: Because the inflores-
cences in the tree crowns were difficult to reach
from the ground, repeated sampling of nectar
secretion of individual flowers over the course
of a day was not possible; only a day’s total pro-
duction was sampled.

In the late afternoon, flowers due to open the
following day were bagged with mosquito net-
ting. These *'first-day" flowers were removed at
about 4 P.M. the following day, and nectar vol-
ume from each flower was measured by repeated
probing with a 10- or 25-ml calibrated micro-
pipette.

Ns production for *second-day" flowers of

. goldmanii and E. chiapasana was estimated
in the following manner: inflorescences were
bagged with mosquito netting in the late after-
noon as for first-day flowers, and nectar was al-
lowed to accumulate in the isolated flowers for
two days. Total nectar accumulation was mea-
sured at 4 P.M. on the second day. The mean
first-day accumulation of nectar subtracted from
the mean 2-day accumulation yields the esti-
mated second-day nectar production. (This
method assumes that nectar removal does not
influence secretion.) All bagged flowers fell off by
the morning of the third day.

Sugar concentration expressed as percent su-
crose equivalence (Bolten et al., 1980) was mea-
sured for each flower or for the pooled nectar of
several flowers using an American Optical model
10431 temperature-compensated hand refrac-
tometer. With the figures for nectar volume and
sugar content, the mean caloric value of the nec-
tar per flower was calculated for each population
sampled.

RESULTS
FLORAL BIOLOGY AND BREEDING SYSTEMS

In most passerine-pollinated species of Ery-
thrina the stamens and stigma are well separated
at anthesis and the flowers are homogamous, i.e.,
pollen is released from the anthers and the stigma

close proximity, tightly enclosed in the floral
pseudotube, and the flowers are protandrous.
On the first day of anthesis the staminal fila-

30

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

TABLE 1. Observations of avian floral visitors to Erythrina tree species.
Nectar Thieves & Robbers
Humming-
Species Locality' Trees? Hours? Pollinators birds Passerines
1. E. ameri- Mexico: Oaxaca 4 9 Heliomaster — Icterus galbula
cana Coatlán constantii
2. E. berte- Mexico: Chiapas 5 3.5 Anthraco- Anthraco- Icterus macu-
roana x alenque thorax pre- thorax pre- lialatus
E. folkersii tii vostii Pheucticus lu-
dovicianus
3. E. bere- Mexico: Veracruz 2 2 Eugenes ful. | — —
nices Tequila gens
4. E. chiapa- Mexico: Chiapas 8 15.5 Heliomaster — Icterus wagleri
sana El Sumidero constantii
5. E. chiapa- | Mexico: Chiapas 6 18 Eugenes ful- Hylocharis Diglossa barit-
sana Teopisca gens leucotis ula
Icterus galbula
6. E. coch- Costa Rica: Heredia 6 20 Heliomaster Chalybura —
leata La Selva longirostris urochrysia
7. E. costari- Costa Rica: Puntarenas 1 5 Heliomaster Phaeochroa —
censis San Vito de Java longirostris cuvierii
8. E. gibbosa Costa Rica: Alajuela 2 11 Phaethornis Lampornis —
Monteverde guy hemileucus
9. E. globo- Costa Rica: San Jose 5 8.5 Eugenes ful- — —
calyx Las Nubes gens
Campylop-
terus hemi-
leucurus
10. E. gold- Mexico: Chiapas 12 45.5 Heliomaster — Icterus gularis
manii El Sumidero constantii I. pectoralis
I. wagleri
11. E. folkersii Mexico: Veracruz 2 3.5 Phaethornis — —
Los Tuxtlas supercilio-
Sus
Campylop-
terus hemi-
C. curvipennis
12. E. folkersii Mexico: Chiapas 2 45 Phaethornis Amazilia tza- Icterus galbula
Palenque supercilio- I. prostheme-
SUS las
13. E. lanata Mexico: Oaxaca 2 4.5 Heliomaster — —
erto Escondido constantii
14. E. lanata Mexico: Jalisco 2 7.0 | Heliomaster — Cassiculus
Chamela constantii melanicte-
rus
15. E. pudica Mexico: Chiapas 8 16.0 Heliomaster — Icterus gularis
El Aguacero constantii
16. E. tux- Mexico: Chiapas 3 15.0 Heliomaster Amazilia Coereba flave-
tlana Malpaso constantii cyanoceph-
Anthraco- ala Cyanerpes lu-
thorax pre- A. tzacatl cidus
vostii

Icterus gradu-
acauda

1987] NEILL—ERYTHRINA POLLINATION 31
TABLE 1. Continued.
Nectar Thieves & Robbers
Humming-
Species Locality! Trees Hours? Pollinators irds Passerines
Eugenes ful- Pheucticus lu-
gens dovicianus
17. E. tux- Mexico: Veracruz l 5.5 | Campylop- Eupherusa —
tlana Uxpanapa terus curvi- imi
pennis

l asa we locality and voucher data for each observation are listed in Appendix.
2 Number of indi rved.

vidual trees obse
2 Paeon Tua of observation

ments are fully grown and pollen is released from
the stamens situated near the apex of the corolla
within the pseudotube. At this stage the style is
shorter than the stamens and the stigma is not
receptive. The style and ovary continue to elon-
gate during the night after the first day of flow-
ering. By the second day, when most ofthe pollen
has been removed by floral visitors, the stigma,
now receptive and with a sticky exudate on its
surface, is held a few mm beyond the anthers,
just inside the mouth of the pseudotube at the
apex of the corolla. Each flower, then, is func-
tionally male on the first day and functionally
female on the second. Unpollinated flowers usu-
ally abort after the second day but sometimes
remain for a third or fourth day before aborting;
the stigma appears to remain receptive during
this time.

The inflorescence of sect. Erythrina is a pseu-
doraceme with fascicles of three flowers each ar-

cence, and usually one or two fascicles of
flowers open each day. An individual inflores-
cence, with 30-50 fascicles, blooms for two or
three weeks. An inflorescence in “full bloom" is
composed of three to nine functionally female
flowers at the bottom (in one to three fascicles),
three to six functionally male flowers in the fas-
cicles just above the females, and above them
floral buds at progressively younger develop-
mental stages.

Inflorescence development in an individual
Erythrina tree is staggered, so a tree often blooms
for two to three months or more. Blooming among
trees in a population is also staggered, so a pop-
ulation is often in bloom for four to five months
annually or even longer. Some species remain in
bloom for a shorter period, one to two months

within a population. Detailed, multi-year phe-
nological data for any particular site, however,
is not available.

Most species of sect. Erythrina flower during
the dry season, from January to May in Me-
soamerica, and are leafless when in flower. Leaves
flush, in general, after flowering has ceased and
while fruits are developing, prior to or just after
the onset of the rainy season in May or June.
Some species flower during the rainy season, June
to October. These species also usually shed all
their leaves before flowering and flush a new set
of leaves as flowering ceases. This behavior is
unusual: few other tree taxa in the Neotropics,
especially in very wet forests, shed leaves during
the rainier portion of ai year and retain them
during the drier portio

The data presently E nsi: suggest that all
Erythrina species are genetically self-compatible
(Neill, in press). The fitness of progeny resulting
from self-pollination is significantly lower than
that of progeny resulting from outcrossing; out-
crossing appears to be predominant in the breed-
ing systems of Erythrina. Some seed set from
geitonogamous pollinations and even from oc-
casional autogamy, which probably occurs in
natural populations.

FLORAL VISITORS

The avian visitors to the flowers of most ob-
served Erythrina por ions include d h
mingbird and passerine “‘illegi

on, gi m
linators (Table 1). Pollination records for each
Erythrina species are summarized in Table 2,
which encompasses prior reports on humming-
bird-pollinated tree species: Erythrina lanceo-

32 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74
TABLE 2. Species of hummingbirds observed as legitimate pollinators on 15 species of Erythrina. Abbre-
viations in parentheses refer to the countries in which species were observed (M = Mexico, C = Costa Rica,
and T = Trinidad).
Hubiniasbirds Erythrina Species
_ " G
S 6 š € 2 “< _ = < 6
3 4 = 3 RE š s S ç B š Bo Q
S a uw S S S 8 š gs 8 3 S BB
E Š 8 S & ÈA &8 S S S E 8 § Š
S S š 8 3S E $S B E S E i $ š 3
& & S S $ s Š l š & g 3S BES Š
uj nj bJ bj ud us yy ye ye ye S SQ OR og
Heliomaster constantii + + + + + +
H. longirostri + + + +
Eugenes fulgens + + + + +
pylopte
hemileucurus + + +
C. curvipennis + +
Anthracothorax prevostii + + +
Phaethornis guy + +
Ph. superciliosus +
Glaucis hirsuta +

lata in Costa Rica (P. Feinsinger, pers. comm.)
and E. pallida in Trinidad and Tobago (Fein-
singer et al., 1979)

For this discussion I have adopted a modified
version of Inouye’s (1980) terminology for floral
larceny: “nectar robbers” make a hole or oth-
erwise damage floral tissue to gain access to the
nectar, while

cludes pollination. The distinction between nec-
tar thieves and robbers is important because rob-
bers may damage the ovary or stylar tissue and
often destroy or remove the entire flower, and
thus may have a much greater effect in reducing
the reproductive potential of the plant than do
nectar thieves.
Billl bt

4 : :1
-

from Ridgway,
1911) of the hummingbird species observed as
pollinators and illegitimate visitors to Erythrina
trees are compared in Figure 1. Legitimate pol-
linators all have bills longer than 28 mm, where-
as nectar thieves and robbers, with one excep-
tion, have bills shorter than 22 mm

POLLINATORS

Nine species of hummingbird were observed
as legitimate pollinators of the 15 species of Er-

ythrina trees. All ofthe hummingbird pollinators

may be characterized as long-billed, high-reward
trapliners, although some behaved as territori-
alists or even nectar thieves on certain occasions,
as detailed below.

I observed two species of hermit humming-
birds (Phaethorninae) pollinating Erythrina:
Phaethornis guy and P. superciliosus. A third
hermit species, G/aucis hirsuta, was reported as
a pollinator of E. pallida in Trinidad by Fein-
singer et al. (1979). The hermits visited princi-
pally the smaller understory species of Erythrina
but rarely were in the forest canopy. Their for-
aging behavior at understory Erythrina is similar
to that documented for understory herbs such as
Heliconia (Stiles, 1975) or shrubs such as Aphe-
landra (McDade, 1984).

The remaining six Erythrina pollinators are
non-hermits (Trochilinae); all are long-billed,
high-reward trapliners which forage like hermits,
but usually in the forest canopy or open areas
rather than in the understory. Heliomaster con-
stantii is the principal or sole known pollinator
of at least six Erythrina species in the dry forests

on both the Pacific and Caribbean slopes of Me-

NEILL— ERYTHRINA POLLINATION

ERYTHRINA FLORAL VISITORS: HUMMINGBIRDS

s

NS NN.

\\ \ \ x
30_N NN NY w
ail

S A N AAAA
s2 \ \ \ \ \ YX
all

AN
- \\\\\ \\V
all

NN N N N N \
N NN N N N \
y>FFFOOQOA

Nectar Thieves & Robbers

FIGURE 1. Bill lengths of Bop éd. visitors to Erythrina trees, including pollinators and legitimate

visitors. Ph gu = Phaethornis guy; Ph s
Heliomaster longirostris; Gl hi =

= Phaethornis superciliosus, He co =
Glaucis hirsuta; Ca he = Campylopterus hemileucurus; Eu fu =

Heliomaster constantii; He lo
E genes fulgens:

Ca cu = grey pif curvipennis, An pr = Anthrocothorax prevostii, Ph cu = Phaeochroa cuvieri

Amazilia tzacatl, Am
ba = Heliothryx baro

soamerica, and from E. pallida in Trinidad
(Feinsinger et al., 1979). Eugenes fulgens is found
mostly in the highlands of Mesoamerica above
1,500 m. It is the principal pollinator of at least
four highland species of Erythrina, and I also
observed it below 1,000 m pollinating E. tux-
tlana in southern Mexico. Campylopterus hemi-
leucurus 1s a bird of wet forests from near sea
level to 1,800 m and has been observed polli-
nating three Erythrina species in such habitats.
Campylopterus curvipennis was observed as a
pollinator of two Erythrina species in low- to
mid-elevation wet forests of southern Mexico.
Anthracothorax prevostii was the least consis-
tent Erythrina pollinator. This bird of low- to

= Amazilia cyanocephala; Hy le = Hylocharis leucotis; Eu ex = [eredi eximia; He

mid-elevation humid fi

as a legitimate pollinator and contacted the re-
productive parts of Erythrina like the other long-
billed trapliners. More frequently, however, An-
thracothorax was a nectar thief, as described
below.

The species of hummingbirds that pollinate
sect. Erythrina all behave in a similar manner
when feeding at the flowers. The hummingbird
first approaches the inflorescence and hovers to
align its bill precisely with the axis of the first
flower it is to visit (Fig. 2). It inserts its bill at
the apex of the flower; at full penetration the
reproductive parts of the flower always contact
the bird’s throat, upper chest, or the base of the

34 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FIGURES 2-5.
liomaster constantii a self to feed at flowers of e lanata; dry forest near the Pacific coast, Pochutla,

Oaxaca, Mexico. Bill i
pine-oak forest, Teopisca. Chia
upper throat. — 4-5.

inserted at apex of corolla. — 3.
pas, Mexico. At full insertion of the bill, pollen is deposited at base of bill or on
Hylocharis wasps a short-billed nectar thief, approaching and fora

Hummingbird pollinators and nectar thieves of Erythrina sect. Erythrina.—2. Pollinator He-

Eugenes fulgens pollinating E. chiapasana; highland

ging at flowers of E

chiapasana; same 2 sss as in Figure 3. The bill is inserted into the mouth of the calyx or the slit of the corolla
**pseudotube" to obtain nectar; reproductive parts of the flower are not contacted.

bill (Fig. 3). The bird remains hovering at this
position for up to five seconds, then withdraws
and moves to another flower in the inflorescence
or to another inflorescence.

The pollination records summarized in Table
2 indicate that there is no species-specific, one-
to-one relationship of Erythrina species and
pecies. The ecological and geo-

birds have been observed at several species of

Erythrina at different localities in the birds’ range,

and more than one pollinator has been recorded
for many of the Erythrina species. The bird
species are quite similar to one another behav-
iorally and morphologically, and the tree species
are also quite similar to one another in terms of
floral morphology, flowering behavior, and nec-
tar rewards (the last is discussed below). Ery-
thrina sect. Erythrina and related groups are ev-
idently adapted to pollination by the high-reward
trapliner guild of hummingbirds as a whole. This,
however, is a small subset of all hummingbirds.

1987]

In Mesoamerica, there are very few additional
hummingbird species with the appropriate mor-
phology and behavior for Erythrina tree polli-
nation, other than the nine species listed in Table
2. (Doryfera ludoviciae in the highlands of Costa
Rica and Panama may be the only high-reward
trapliner in the region not reported as an Ery-
thrina pollinator [P. Feinsinger, pers. comm.].)

the local community level, often only a
single high-reward traplining hummingbird is
present so that many populations of Erythrina
are pollinated by a single bird species. In the
lowland dry forests of Pacific Mesoamerica, He-
liomaster constantii is the only appropriate hum-
mingbird present, so it is undoubtedly the sole
pollinator of the Erythrina tree species restricted
to Pacific dry forests.

The hummingbirds are not strict specialists on
Erythrina. Well-studied species such as Eugenes
fulgens have been reported visiting a number of
other plant species. The two Heliomaster species
may be more specialized as Erythrina foragers
than are the other hummingbird genera. During
the course of his extensive community-level
studies of hummingbirds, P. Feinsinger (pers.
comm.) observed Heliomaster longirostris on
Trinidad to feed only at Erythrina pallida and
at the apocynaceous vine Mandevilla hirsuta;
while at Monteverde, Costa Rica, Heliomaster
constantii visited exclusively different species of
the same two plant genera.

NECTAR THIEVES

The nectar thieves of Erythrina sect. Erythrina

chrysia (Tab le 1). The short bills of these birds
preclude them from reaching the floral nectar by
inserting their bills at the apex of the corolla.
Nectar thieves take advantage ofthe incomplete-
ly sealed tube of the Erythrina corolla. They ap-
proach the flower from below (Fig. 4) and, often
with some struggle and manipulation of their
bills and bodies, slip their bills into the proximal
end of the ventral slit of the pseudotube formed
by the corolla standard, or into the mouth of the
calyx (Fig. 5). They are thus able to gain access

damaging
tivity is concentrated at the base of the corolla,
and thieves were never observed to contact the

NEILL— ERYTHRINA POLLINATION 35

reproductive parts of the flowers at the apex of
the corolla. Unlike the pollinators, thieves fre-
quently grasp the flowers with their feet when
feeding and clamber over the inflorescence to
reach adjacent flowers. They may damage the
surface of the corolla somewhat, but they do not
appear to damage the ovary itself. Small nectar
thieves may lack the power to pierce the thick
Erythrina perianth as do the larger-bodied nectar
robbers.

Most of the hummingbird nectar thieves are
small species with bills under 21 mm long and
bodies weighing less than 6 g. An exception is
Anthracothorax prevostii, whose bill length of 28
mm is within the low end of the range of the
legitimate pollinators and whose body size of 10
g is equivalent to that of the pollinators. A. pre-
vostii was occasionally seen visiting Erythrina
flowers in the manner of a true pollinator, but
more often it behaved as a nectar thief in a man-
ner similar to the smaller opportunistic birds.

NECTAR ROBBERS: HUMMINGBIRDS

Nectar-robbing hummingbirds have shorter
bills than the pollinators but are larger in body
size and more powerful than the nectar thieves.
The robbers pierce the calyx or base ofthe corolla
with their needle-like bills to gain access to Er-
ythrina nectar.

I observed the large (12 g) hummingbird
Phaeochroa cuvierii repeatedly robbing flowers
of a roadside Erythrina costaricensis near San
Vito de Java in southern Costa Rica. Hovering
beneath or beside the inflorescence, the bird
placed the tip of its bill against the fleshy calyx
and with three or four sharp thrusts punctured
through the calyx to plunder the nectar. Usually
the robber hovered while piercing the flower, but
sometimes it perched on the inflorescence. Flow-
ers strewn on the ground below the tree had up
to six puncture holes through the underside of
the calyx, indicating that Phaeochroa returned to
a single flower several times to drain it of nectar.
Most pierced flowers were soon aborted, and
some had signs of damage to the ovary caused
by the robber's bill. On two successive mornings,
a Phaeochroa repeatedly robbed an E. costari-
censis tree that was also visited at intervals by a
legitimate pollinator, Heliomaster longirostris.
When on occasion the two birds arrived to fee
at the tree simultaneously, a territorial fight en-
ued.

In the same region of southern Costa Rica

o

36 ANNALS OF THE MISSOURI BOTANICAL GARDEN

where I made the foregoing observations of E.
costaricensis, Skutch (1971) reported that Heli-
omaster longirostris pollinated the flowers of Er-
ythrina berteroana while Phaeochroa cuvierii and
another short-billed hummer, Heliothryx bar-

american rainy season, August-November, while
E. berteroana flowers during the dry season, late
December to March. The similarity of visitation
patterns reported for these two species with dif-
ferent flowering phenologies in the same region
(they are not strictly sympatric) suggests that to-
gether they support the same pollinators and il-
legitimate visitors in succession for a consider-
able portion of the year

NECTAR ROBBERS: PASSERINE BIRDS AND PARROTS

All of the | non. -hummingbird visitors to d
thrina sect. E
erally FROM De reproductive potential of the
flowers they visit. These robbers include passer-
ine birds in the families Icteridae, Coerebidae,
and Fringillidae, and the non-passerine parrot
family Psittacidae. Icterids and coerebids are le-
gitimate pollinators of some Erythrina species in
the Neotropics, but on sect. Erythrina they are
strictly parasitic.

I observed seven species of orioles (Icterus spp.)
robbing eight species of Erythrina in Mexico and
the icterid e eines Neige. one
Erythrina speci rioles were the most fre-
quently ED p all robbers a Expl the
most complex behavior to obtain the nectar.
Typically, an oriole would pluck a flower with
its bill, then hold it against a branch with one
foot and jab its bill into the mouth of the calyx
to reach the nectar. After plundering the flower,
the oriole would drop it and pluck another. The

sumed daily by the orioles. Sometimes an oriole
would impale a flower on a thorn of an Erythrina
tree branch to hold it in place while the oriole
imbibed the nectar; the impaled flowers were left
hanging on the branch

Oriole species vary in their degree of special-
ization as nectar feeders (Stiles, 1981). Some ev-
idently obtain a high proportion of their caloric
requirements from floral nectar, at least during
certain seasons of the year. Orioles are important

[Vor. 74

legitimate pollinators of some Erythrina species
such as the widespread E. fusca (sect. Duchas-
saingia) (Morton, 1979) and the central Mexican
endemic E. oliviae (sect. Olivianae) (Toledo &
Hernandez, 1979). These passerine-pollinated
Erythrina species are presumed to represent an
ancestral condition with respect to the hum-
mingbird-pollinated groups including sect. Ery-
thrina. Orioles and other icterids are known to
behave as nectar robbers, in a similar manner to
their behavior at Erythrina, at other plant species
in the Neotropics, including banana (Musa par-
adisiaca), which was introduced from the Old
World tropics (Skutch, 1954). In the case of Er-
ythrina, the evolutionary relationship of the ori-
oles’ nectar-robbing behavior of the humming-
bird-pollinated species to their legitimate
Senay of the putatively is pean
pollin n. Did
nm switch to Sud c bbs after having
evolved nectar-feeding behavior as legitimate
pollinators, or was the order reversed?

everal species of the honeycreeper family
(Coerebidae) are nectar robbers of Erythrina
flowers. I observed the flower-piercer Diglossa
baritula robbing E. chiapasana in the highlands
of southern Mexico by holding the corolla with
its specialized hooked upper mandible and pierc-
ing it with its lower mandible to extract the nec-
tar. Hernández and Toledo (1979) observed sim-
ilar behavior by the same bird species at Erythrina
leptorhiza, an herbaceous species of highland
central Mexico.

Two coerebid bird species, the shining hon-
eycreeper Cyanerpes lucidus and the bananaquit
Coereba flaveola, were nectar robbers of Ery-
thrina tuxtlana. These birds sometimes pierced
calyces in the manner of Diglossa, and at other
times slipped their bills into the calyx mouth
without puncturing it, in the manner ofthe short-
billed hummingbird nectar thieves. Like the ori-
oles, these two honeycreepers that behave as rob-
bers of hummingbird-pollinated Erythrina
species are also important legitimate pollinators
of passerine-pollinated Erythrina including E.
poeppigiana in Trinidad (Feinsinger et al., 1979)
and E. megistophylla in Ecuador (Steiner, 1979).

migrant rose-breasted grosbeak Pheucti-
cus ludovicianus (Fringillidae) nectar-robbed a
living fencepost row of hybrid Erythrina berte-
roana x E. folkersii and natural populations of
E. folkersii and E. tuxtlana, all on the Atlantic
slope of Chiapas, Mexico. Unlike the other pas-

L

1987] NEILL—ERYTHRINA POLLINATION 37

TABLE 3. Daily nectar production in flowers of Erythrina sect. Erythrina.

X Sucrose
Species Nectar Volume Equivalence X Calories
(Locality) X ul + s.d. Wt/Vol per Flower N
A. First-day flowers
E. costaricensis (San Vito de Java) 36.4 + 14.7 29% 43.8 10
E. globocalyx (Las Nubes) 31.4 + 19.5 22.8% 29.3 10
E. chiapasana (El Sumidero) 29.6 + 12.0 27.3% + 3.9% 33.3 12
E. goldmanii (El Sumidero) 31.9 + 14.6 28.9% + 2.2% 38.2 10
B. Two-day accumulation of nectar
E. chiapasana (El Sumidero) 49.8 + 26.1 27.3 56.0 6
Estimated caloric production of
econd- flower (B-A): 22.7
E. goldmanii (El Sumidero) 52.3 + 31.1 68.1 9
"wn m caloric production of
second-day flower (B-A): 29.9

serine robbers, grosbeaks actually consumed flo- varied relatively little within or among popula-
ral tissue as well as nectar. Usually they plucked tions (2396-2996). The calculated mean caloric
the flower and either crushed the calyx with their value of the nectar per flower ranged from 29 to
bills and dropped the flower or consumed the 43 cal among the sampled populations, with an
entire flower. At times grosbeaks merely bit off overall mean of 36 cal.
the end of the corolla (and pistil), leaving the Nectar continued to accumulate on the second
flower attached with the calyx and corolla stump. day of flowering in the bagged flowers of E. chia-
On several occasions I observed the short-billed pasana and E. goldmanii. The estimated caloric
hummingbird Amazilia tzacatl follow a foraging production of the second-day flowers (overall
grosbeak and insert its bill into the decapitated mean = 27 cal) was somewhat less than in the
Erythrina corolla tube to extract the remaining first-day flowers, but this may have been due to
nectar. inhibition of production by the accumulation of
On a number of occasions I observed parrots large nectar volumes in the protected flowers, in
(Psittacidae) consume immature seeds of Ery- the absence of removal by nectarivores.
thrina trees, but never the flowers. Skutch (1971), The results of the nectar sampling from the
however, reported the orange-chinned parakeet different species were similar enough to allow a
Brotogeris jugularis to be an important nectar rough estimate of the daily caloric production
robber of Erythrina berter insouthern Costa per flower for hummingbird-pollinated sect. Er-
Rica. The parakeets plucked the flowers with their — ytArina in general. For purposes of the discussion
bill or feet, bit through the calyx to extract the below, an average caloric production of 35 cal
nectar, and dropped the flowers without consum- per flower per day is assumed as an approxi-

ing floral tissue. mation. This is somewhat less than the values
reported by Stiles (1975) for hermit-pollinated
NECTAR PRODUCTION AND CALORIC VALUE Heliconia species (48—141 cal) but is an order of

magnitude or greater than the production typical
of plants pollinated by short-billed generalist
hummingbirds (Feinsinger, 1978).

Daily nectar production per flower in the sam-
pled populations of Erythrina chiapasana, E.
costaricensis, E. globocalyx and E. goldmanii is
shown in Table 3. Within populations, the vari-
ance in nectar production per flower was high
(for example, the range in E. globocalyx was 10— Macmillen and Carpenter (1977) derived a
67 ul). The mean nectar volume for each of the regression equation for the 24-hour energy costs
four populations (30-36 ul) was quite similar, of nectar-feeding birds, based on empirical data
however. The sugar concentration of the nectar on basal metabolic rates and energetic costs of

HUMMINGBIRD ENERGETICS AND NECTAR REWARDS

38 ANNALS OF THE MISSOURI BOTANICAL GARDEN

flight for hummingbirds, Hawaiian honeycreep-
ers, and African sunbirds. Using this equation,
the daily energetic cost for a Heliomaster weigh-
ing 8.0 g is calculated to be 10.9 kcal. This is the
equivalent of 311 Erythrina flowers at 35 cal/
flower. (Hummingbirds do gain some nourish-
ment by consuming arthropods, so the actual
daily nectar consumption of an Erythrina pol-
linator may be somewhat less.)

Erythrina species in sect. Erythrina — even large
canopy trees at the peak of their blooming pe-
riod — do not generally produce as many as 300
flowers per day. Therefore an individual is prob-
ably not “worth” defending as a feeding territory
by a hummingbird; several trees must be visited
daily to satisfy the bird's energetic requirements.

Intertree movement of the foraging birds, and
consequent pollen flow between trees, is evi-
dently promoted by the limited number of flow-
ers produced on an individual tree. In contrast,
me extended blooming period of Erythrina and

£41

of the nectar resource promotes
the high fidelity of visitation exhibited by the
traplining pollinators. This syndrome is exem-
plified by my data on flowering behavior and
pollination observations of populations of sev-
eral different species of Erythrina, discussed be-

w.

Erythrina cochleata. Erythrina cochleata is a
25-m tall canopy tree at La Selva Biological Sta-
tion, a tropical wet forest site in Costa Rica. I
attempted to locate every reproductively mature
individual of this species in an area of about 2
km? at La Selva and found a total of 10 trees.
This species is confined to the alluvial terraces
of the rivers and major streams, so on a large
scale the trees were clumped, but no individual
was less than 50 m from its nearest conspecific
neighbor. Erythrina cochleata flowers during the
wet season, and the population was in flower
continuously at least from May through Septem-
ber 1981

I observed flower production and floral visi-
tors for six days at four different individual trees.
The trees averaged 112.5 (range 84-181) open
flowers per day. The pattern of floral visitation
each day was very consistent. The only pollinator
and regular visitor was Heliomaster longirostris.
Each morning between 7:00 A.M. and 8:30 A.M.
a solitary Heliomaster would arrive, visit as few
as four to as many as 45 Erythrina flowers, and
depart. These visits were repeated at sporadic
intervals during the morning. (Birds were not

[Vor. 74

tagged, so subsequent visits may have been made
by different individual birds.)

On only two occasions was more than one
hummingbird seen at a time in an Erythrina
cochleata crown, and both times the interloper
(once another Heliomaster, once a nectar-thiev-
ing Chalybura urochrysia) was chased away by
the Heliomaster.

Heliomaster longirostris is considered an **un-
common" bird at La Selva (Slud, 1960), yet I
saw this species every time I looked for it at
flowering Erythrina trees. Evidently the small,
scattered population of Erythrina cochleata sup-
ports the nutritional requirements of, and re-
ceives consistent pollination service from, a small
population of Heliomaster for a period of several
months each year. What the Heliomaster hum-
mingbirds do when the trees cease flowering is

own. They may migrate to populations of
other Erythrina species on the Atlantic slope of
Costa Rica, such as E. steyermarkii, that flower
during the dry season, or they may forage at other
canopy flowers such as Mandevilla spp. vines
(besides Erythrina, there are no other hum-
mingbird-pollinated canopy-level trees in the re-

ion).

Erythrina goldmanii. The habitat and pop-
ulation structure of Erythrina goldmanii at Can-
on del Sumidero National Park in Chiapas, Mex-
ico, where I observed this species, are very
different from those of E. cochleata at La Selva,
but the pollination systems of the two species are
quite similar.

Erythrina goldmanii is a dry forest species and
is rather scrubby, rarely attaining a height of over
6 m. At El Sumidero, on a slope above the semi-
arid basin ofthe Rio Grijalva, E. goldmanii forms
dense populations of small trees in disturbed sec-
ondary forest. I made observations in a 2 ha plot
containing 54 plants. Most trees had only one or
two inflorescences in bloom, with less than 10
flowers per tree; the largest tree had 36 flowers.
In all there were 475 flowers in the 2 ha plot at
peak flowering.

Heliomaster constantii was the only pollinator
and the only hummingbird visitor seen at Ery-
thrina goldmanii in over 45 hours of observa-
tion. At least three Heliomaster individuals were

beyond the boundaries of the plot. Their fidelity
to Erythrina was very high: only twice did I see

1987]

a Heliomaster visit any other plant, and then only
for single floral probes
Orioles were Beguen visitors to Erythrina
goldmanii, and they destroyed an estimated 21%
of the flowers daily in the manner described pre-
viously. There may have been competition for
nectar resources between the orioles and the
liomasters, but I never observed any aggressive
interactions between orioles and hummingbirds.
pecies of Erythrina sect. Erythrina are usually
allopatric, being separated by elevation and hab-
itat; but at El Sumidero two species come into
contact. Erythrina goldmanii inhabits the dry
lower slopes at about 800 m, and E. chiapasana
occurs in the moister forest on top of the plateau
at 1,100 m. Heliomaster constantii visited Ery-
thrina chiapasana just as it did E. goldmanii less

in the intermediate zone, on the upper
slopes of the El Sumidero escarpment (Neill, in
press). The birds evidently do not discriminate
among Erythrina species when the species occur
together. Heliomaster hummingbirds are cer-
tainly the pollen vectors implicated in interspe-
cific gene flow between Erythrina species at El
Sumidero.

Erythrina tuxtlana. One final observation
indicates that traplining hummingbirds will
sometimes behave as facultative territorialists if
they are given the opportunity. I observed floral
visitors to a 20 m tall Erythrina tuxtlana in mid-
elevation wet forest near Malpaso, Chiapas. The
tree had a broad-spreading crown with 1,400 open
flowers. According to the estimates of nectar pro-
duction in other species, this should have been

ofthe nee into bading tert ories and maintained
thet hen not
feeding, each bird generally perched within its
territory, and many aggressive interactions en-
sued when one bird crossed into another’s ter-
ritory. This was the only instance of consistent
within-tree territoriality I observed in any Ery-
thrina population.

CONCLUSIONS: Is IT COEVOLUTION?
The flowers of Erythrina sect. Erythrina pro-
vide a rich nectar resource that is fed upon by
many species of birds besides the legitimate pol-

NEILL—ERYTHRINA POLLINATION 39

padi The pollinators, however, constitute a
mall guild of “high-reward traplining" hum-
ied about eight species in Mesoamerica.
hese are mostly non-hermits of the subfamily
Trochilinae. Predominant among these is the ge-
nus Heliomaster. The pollinators are highly
faithful visitors to Erythrina, which provides
them with a consistent nectar resource for long
periods. The limited caloric value of nectar pro-
duced per tree per day usually precludes the
maintenance of permanent feeding territories at
a single tree by the hummingbird visitors. The
consequent nomadic or **traplining" behavior of
the hummingbirds and the dispersal patterns of
the pollen they transport among the scattered
individual E rythrina trees may be a critical factor

summary, is a canopy-level analogue of the high-
reward traplining pollination systems of Heli-
conia and similar understory plants. In this sense
the pollination system of sect. Erythrina, togeth-
er with the other hummingbird-pollinated sec-
tions of Erythrina trees (sects. Stenotropis, Pseu-
do-edules, Gibbosae, and Corallodendra; cf. Neill,

in press) may be unique. Hummingbird-polli-
nated canopy and subcanopy trees are in them-
selves uncommon (Stiles, 1978), and I know of
no other genus of canopy trees besides Erythrina
that is adapted to pollination by the traplining
guild of hummingbirds.

To what extent have species of sect. Erythrina
and their hummingbird pollinators coevolved?
To what extent is this a specialized mutualism?
Feinsinger (1983) indicated that a highly

Occurrence; although te may oer ona
particular plant ost hermit
species forage on a number of different plants.
Similarly, most hermit-pollinated plants are ser-
viced by several species of hermits, although
shorter-billed birds are excluded as pollen vec-
tors. If one's definition of coevolution requires
a high degree of one-to-one species specificity in
such mutualistic interactions, then hermits and
hermit-pollinated plants cannot be considered
very *'tightly coevolved." Feinsinger (1983) con-
pem Em *most hermits, hermit- like hum-

4 1 4 CJ GNE EL

on
LLUD
coevolution between two divae - proud of
species."

40 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Species of sect. Erythrina vary in the degree
of specificity of their association with the hum-
mingbird pollinators. Erythrina species of the
dry forests of the Pacific slope are pollinated ex-
clusively by Heliomaster constantii, so the plant’s
fitness is directly dependent on the behavior and
morphology of a single bird species. The oppor-
tunity for the plant to evolve adaptations to spe-
cific traits of the bird is clear. Heliomaster con-
stantii, however, feeds upon and pollinates a
number of Eryt t the bird’s
geographic range, andi it also feeds ı upon and pol-
linates at least one other plant genus (Mandevilla;
Feinsinger, pers. comm.). Although there is un-

oubtedly a temporary sort of exclusivity in the
Erythrina-Heliomaster association in certain
ecological communities at certain seasons of the
year, the association cannot really be considered
an obligate mutualism

The plant-pollinator association is less specific
for Erythrina species of highland and wet-forest
communities, where several species of traplining
hummingbirds often visit and pollinate an in-
dividual tree on a single day. Hummingbirds such
as Eugenes and Campylopterus visit a rather wide
variety of plants besides Erythrina. Even in these
cases, however, such plant-pollinator associa-
tions involving high-reward traplining hum-
mingbirds are much more exclusive than those
involving short-billed generalist hummingbirds
and een E plants.

g the species of sect. Erythrina, there is
little difitreptianon in floral morphology, flow-

patterns,

the pollinators—the pollination system of all
species is quite similar. Rather, species are dif-
ferentiated by their restriction to distinct climatic

edaphic conditions. This large group of
species, as a whole, has evolved a particular set
of adaptations to the guild of high-reward trap-
lining hummingbirds.

LITERATURE CITED

BOLTEN, A. B., P. FEINSINGER, I. BAKER & H. G. B
1980. On the calculation of sugar ease
in flower nectar. Oecologia 41: 301-304.

CRUDEN, W. R. & V. M. TOLE 1977. Oriole pol-
lination of Erythrina breviflora bari
evidence for a polytypic v Pla
Syst. Evol. 126: 393-4 03.

FEINSINGER, P. 1978. Ecological interactions between
plants and birds in a successional tropical com-
munity. Ecol. Monogr. 48: 269-287.

—. 1983. Coevolution and pollination. Pp. 282-

[Vor. 74

310 in D. Futuyma and M. Slatkin (editors), Co-

evolution. Sinauer Assoc., Sunderland, Massachu-

setts.

& R. K. CoLweELL. 1978. Community orga-

nization among pongo a S edi birds.

Amer. Zool. Ls -795.

. B. LiN ian A. SWARM & J. A. W
1979. Visio e the ecilination biology of res
Erythrina species on Trinidad 2 Tobago. Ann.
Missouri Bot. Gard. 66: 451-47

GUILLARMOD, A. J., R. A. JUBB & C. 1 SKEAD. 1979.
Field studies m six southern African species of
Erythrina. Fo Missouri Bot. Gard. 66: 521-527.

HERNÁNDEZ, H. 1982. Female sterility in Ery-
thrina M. Allertonia 3(1): 72-76.

. TOLEDO. 1979. The role of nectar

robbers and pollinators in the reproduction of Er-

ythrina leptorhiza. Ann. Missouri Bot. Gard. 66:

512-520.

———— & ————. 1982. Floral biology of Erythrina
Ma d p the evolution of pollination sys-
n American species of the genus. Allertonia

—84.

(1:

Inouye, D. W. 1980. The vr iid of floral lar-
ceny. E 61: 1251-1253.

KRUKOFF, B. A. & R. C. Sr 1974. A con-
spectus a the genus Erythrina. Lloydia 37: 332-

LINHART, Y. B. 1973. Ecological and behavioral de-
terminants of pollen dispersal in hummingbird-
pollinated Heliconia. Amer. Naturalist 107: 511-

McD DADE, L. A. 1984. Systematics and reproductive

MORTON, E. S. 1979. Effective pollination of Ery-

coev ived behavioral manipulation? Ann. Mis-
ouri Bot. Gard. 66: 482-489.

Mina. D. A. In press. j studies on species
relationships in Erythrina (Leguminosae: Papilio-
noideae). Monogr. Syst. Bot. Missouri Bot. Gard.

1911. The birds of North and Middle

I part

. 1954. Life Histories of Central Amer-

ican Birds. Cooper Ornithological Club
1971. A Naturalist in Costa Rica. Univ. of

Florida Press, Gainesville.

SLup, P. 1960. The birds of Finca “La Selva," Costa
Rica: a tropical wet forest locality. Bull. Am. Mus
Nat. Hist. 121: 49-148.

STEINER, K. E. 1979. Passerine pollination of Ery-
thrina megistophylla Diels (Fabaceae). Ann. Mis-
souri Bot. Gard. 66: 490—502.

. 1975. Ecology, or phenology a

i e Costa Rica

y 5

9 Ecological pet volutionaty implica-
tions of bird pollination. Amer. Zool. 18: 603-
615.

1987]

1981. one aspects of bird-flower co-
evolution, with particular reference to Central
ouri Bot. Gard. 68: 323-351.

974. Observations on the relation-

p S led 487.

ERNÁNDEZ. 19 gud iie oliviae:
a new case n oriole pollination i r Mexico. An
Missouri Bot. Gard. 66: 503-51

APPENDIX

Locality and Voucher Data for Erythrina
Populations Used in Observations of Floral Visitors
(Numbers correspond to those listed in Table 1

1. E. americana Miller. Mexico: Oaxaca, 5 km E of
San Pablo Coatlan. 16?12'N; 96?47'W, Elev. 1,450 m.
Disturbed gallery = with Taxodium, and cultivated
fields. Tree to along intermittent stream. 9-10
Feb. 1983. Neill 5421. 5424.

2. E. berteroana Urban x E. folkersii Krukoff &
Moldenke. Mexico: ERAS 3 km S of Palenque. Elev.
100 m. 17?28'N; 92?00'W. Fencepost bordering field.
Tree to 8 m. 18 March 1983. = ill 5533.

. E. berenices Krukoff & Barneby. Mexico: Vera-
cruz, Tequila. 19245'N; 97°03’ W. Elev. 1,650 m. Coffee
plantation; Premontane Wet Forest. Tree to 12 m.
Jan. — 9 Neill 5381.

4. chiapasana Krukoff. erg: Chiapas, El
Su al National Park, Km 14-16. Elev. 1,100 m.
16?47'N; 93°06'W. Disturbed L. Dry Forest,
transition to moist mixed Quercus forest. Tree 15 m
Mar ch 1983. Neill 5455, 5458, 5465.
Krukoff. Mexico: Chiapas, 13 km

6. E. cochleata “naran Costa Rica: ‘Heredia, La
Selva Biological Station. Elev.
84°00'W. Tropical Wet Forest. Tree 25 m. 15-18 Aug.,
18-21 Sept. 1981. Neill 5015, 5101.

. E. costaricensis Micheli. Costa Rica: Puntarenas,
San Vito de Java, Las Cruces Botanical Garden. Elev.

NEILL— ERYTHRINA POLLINATION 41

1,200 m. 8*45'N; 82*55'W. Premontane Rain Forest,
roadside. Tree 6 m. 11-12 Sept. 1981. Neill 5099.

8. E. gibbosa Cufodontis. Costa Rica: Alajuela, up-
per Penas Blancas Valley, below Monteverde Reserve.
Elev. 1,400 m. 10?20'N; 84?45'W. Premontane Rain
Forest; edge of pasture. Tree to 4 m. 4—6 Sept. 1981.

Ne x 5057.

. E. globocalyx Porsch & Cufodontis. Costa Rica:
"s Jose, Las Nubes. Elev. 1,700 m. 9°53'N; Z vials
Fencepost row, border of pasture. Tree to 8 m; sporadic
wo p 14 Aug., 25 Sept. 1981. Neill 5035. 5142.

E. goldmanii Standley. Mexico: Chiapas, El
Sumidero National Park, Km 7. Elev. 900 m. gni N;
93°06'W. Tropical Dry Forest; secondary, disturbed
scrub. Tree 8 m. 25 Feb.-1 March 1983. Neill, MAT

E folkersii Krukoff & Moldenke. Mexico: Ve-

ux ur tree to 8 m. 28 Jan
12. E. folkersii koff
pas, Palenque Archaeological Site. 17?29'N; 92?01'W.
Tropical Wet Forest; forest edge. Tree 5 m. 19 March
1983. Neill clk
13. E. lana ae Mexico: Oaxaca, 37 km W of
Puerto Escon dido. ev. m. 15?90'N; 97?20'W.
Tropical Dry sid. scrub. Tree 6 m. 13 Feb. 1983.
"t 430.
a Rose. Mexico: Jalisco, Chamela Bio-
local Station. Elev. 250 m. — N; 105?03'W. Tree
13 Jan. 1983. Neill 532
E. pudica Krukoff & M Mexico: Chiapas,
Rio de la Venta, Cascada El Aguacero. Elev. 750 m.
16°46'N; 93°33'W. Tropical Dry Forest, scrub. Tree 6
m. 27, 31 March 1983. Neill 5512.

16. E. tuxtlana Krukoff & Barneby. Mexico: Chia-
pas, 25 km N of Ocozocuautla. Elev. 700 m. 16?48'N;
93?25'W. Premontane Wet Forest; karst limestone. Tree
to 20 m. 28 March, 9 April 1983. Neill 5486, 5621.
Barneby. Mexico: Ve-
ra 90 m. 17?11'N; 94?39'W.
Tropical Wet Forest; kars limestone. Tree 15 m. 16—
17 April 1983. Neill 564

A COMPARISON OF THE DIVERSITY, DENSITY, AND
FORAGING BEHAVIOR OF BEES AND WASPS
ON AUSTRALIAN ACACIA!

PETER BERNHARDT?

ABSTRACT

Twenty-seven bee taxa and 24 wasp taxa were collected on the open inflorescences and/or extra-

floral nectaries of eight Ac

acia species in Victoria,
taxonomic diversity, bees outnumbered wasp foragers by 88% of

Australia. Despite this superficial similarity in
the combined catch of winged

d, with the short-tongued Halicti-

dae and Colletidae comprising the largest unit of native Apoidea o
lossum (Halictidae) and Leioproctus (Colletidae) comprised
i Th

the genera Lasioglos.
o short-ton

foraging female bees o
% of the combined catch of the tw

Acacia species. No wasps, however, were collected on var. retinodes o f A. retinodes

e families e number of bee

n the Acacia species studied. Pollen

taxa collected on
er through late autumn. Polylectic foraging bee
o

% of the

on extra-floral nectar before foraging on nectarless inflorescences. The density and taxonomic rediere d
of wasps remained highest on the Acacia species that offered the greatest volume of sucrose- “rich, extra

floral nectar (i.e., A terminalis). B

Australia than are wasps. The direct influence of wasps on polyad oan appears to be nominal
except in those Acacia species bearing functional extra-floral nectaries

Winged Hymenoptera (bees and wasps) have
been observed to forage frequently on inflores-
cences of Australian Acacia. In contrast to the
Psyllidae and some Coleoptera, bees and non-
parasitic wasps are not destructive to the small
flowers that comprise an Acacia head or spike
(Bernhardt, 1982). Instead evidence suggests that
Hymenoptera are often pollinators of some Aca-
cia species in southeastern Australia (Bernhardt,
1982; Bernhardt et al., 1984; Knox et al., 1985).

The flowers of all Australian Acacia examined
thus far are nectarless (Bernhardt, 1982; Bern-
hardt et al., 1984; Bernhardt & Walker, 1984,
1985; J. Kenrick & G. Beresford, pers. comm.).

ar

When the anthers Pen is eight polyads pres-
ent in each anther are extruded, or partially ex-
truded, from their respective sacs (Kenrick &

Knox, 1979; Knox & Kenrick, 1982). Conse-
quently, bees, wasps, and certain flies easily col-
lect polyads from the synchronously opening flo-
rets in an inflorescence.

Female bees are known to collect Acacia poly-
ads to feed to their larvae. Foraging bees remove
polyads from the anthers via thoracic vibration
of whole inflorescences (Buchmann, 1983) or by
scraping anthers directly with their forelegs (Vo-
gel, 1978), or both (Bernhardt & Walker, 1984,
1985). Acacia species are usually self-incompat-
ible. Seed set tends to occur only when pollina-
tors move between genotypes belonging to the
same species (Knox & Kenrick, 1982; Bernhardt
et al., 1984; Kenrick et al., 1984b).

Capture records of insects foraging on Acacia
in southeastern Australia and analyses of their
pollen loads suggest that Hymenoptera are often
more important as polyad vectors than are either

! Research was conducted at the Plant Cell Biology Research Centre of the School of Botany, University of

Melbourne under the supervision of R. B. Knox.

unding was provided by the Australian Research Grants

Scheme and the Australian Department of Education (CPPER). I thank A. Heisler and the rangers of the National
Parks Service, Victoria (Brisbane Ranges, Cape Schanck) for their cooperation. This study would not have been

possible without the timely D of J. Kenrick, G. Beresford, R. Ma
m of Victoria identified Hymenoptera a

Walker of the National Mus

rginson, P. O'Neal, and T. Hough. J.
nd sent pedet wasps to other
Knox and D. M.

Australian authorities. I am nu grateful for the continuing interest and mPp port R. B.

Calder. C. D.

Michener provided a most valuable critique of the o

riginal man

2 Department of Biology, Saint Louis University, 3507 Laclede, St. Louis, Missouri 63103.

ANN. MIssouRI Bor. GARD. 74: 42-50. 1987.

1987]

beetle or fly taxa. Calliphorid and syrphid flies
transport polyads from Acacia inflorescence to
inflorescence without damage, but they may oc-
cur on Acacia inflorescences at lower density and
diversity than Hymenoptera (Bernhardt et al.,
1984; Knox et al., 1985).

Excluding the rare documentation of pollina-
tion by birds or marsupials (see review by Turner,
1982; Knox et al., 1985) the flowering behavior,
floral presentation, and polyad presentation of
most Acacia species would be expected to favor
a system of generalist entomophily (Bernhardt,
1982; Bernhardt et al., 1984). That is, all insects
that forage for polyads have immediate access
to the inflorescences of Acacia and could effect
deposition of polyads on respective stigmas.

Therefore the purpose of this study was to de-
termine which groups within the Hymenoptera
were major vectors of Acacia polyads with suf-
ficient fidelity to regularly effect seed set. To ac-
complish this end the density and taxonomic di-
versity of polyad foragers were compared to their
respective activities on Acacia inflorescences.

MATERIALS AND METHODS

Acacia species and study sites. Eight Acacia
were selected to determine interspecific and in-
traspecific foraging preferences of Hymenoptera.
The species of Acacia may be found in flower
throughout the year (Kenrick et al. 1984a, 1984b;
Bernhardt, 1982) with the majority flowering
from August through October (Costermans,
1983). Therefore the eight selected species rep-
resented the 12-month flowering season of the
genus but emphasized the period of intensively
overlapping floral phenology from the last month
of winter (August) until the second month of
spring (October). The periods of fieldwork, study
sites, and habitats of each Acacia species are list-
ed below. Descriptions of floristic alliances fol-
low Specht and colleagues (1979).

1) Acacia longifolia Willd. 31/vi11/84—28/1x/84.
Langwarren Reserve: Tall shrubland with dis-
rupted epacrid heath (Bernhardt, 1986).

2) A. mearnsii De. Wild. 8/xi/84—2 1/xi/84. Cor-
anderrk Reserve: Moist sclerophyll wood-
land/shrubland (see Bernhardt & Calder,
1981).

3) A. mitchelii Benth. 16/vii/82-21/1/83. Bris-
bane Ranges National Park: Dry sclerophyll
woodland/shrubland (see Bernhardt & Walk-
er, 1984).

4) A. myrtifolia Willd. 19/viii/82-7/x/82. Bris-

BERNHARDT — AUSTRALIAN ACACIA 43

bane Ranges National Park: Dry sclerophyll
woodland/shrubland (see Bernhardt & Walk-
er, 1984).

5) A. paradoxa DC. (syn. A. armata R. Br.). 12/
1x/84-31/x/84. Brisbane Ranges National
Park: Dry sclerophyll woodland/shrubland
(see Bernhardt and Walker, 1984).

6) A. pycnantha Benth. 11/viii/82-16/ix/82.
Brisbane Ranges National Park: Dry sclero-
phyll woodland/shrubland (see Bernhardt &
Walker, 1984).

7) A. retinodes var. retinodes Schdl. 16/i/82—17/
1/82. Grampians National Park: Montane and
valley dry sclerophyll forest/shrubland with
adjacent epacrid heaths (see Bernhardt &
Walker, 1985)

8) A. retinodes var. uncifolia J. Black. 15/xi/81—
5/11/82. Cape Schanck National Park: Coast-
al calcareous dune flora consisting of tall
shrubland and invasive, naturalized shrubs
and herbs (see Bernhardt et al., 1984).

9) A. terminalis Macbr. 18/iii/83-28/iv/83. Er-
ica-Moe (south Gippsland) and Boolah Boo-
lah State Forest. Moist sclerophyll woodland/
forest with a rain forest element (see Knox et
al., 1985).

Analysis of Hymenoptera. The foraging be-
havior of Hymenoptera was observed and re-
corded over the respective periods of fieldwork
every day or every other day. Insects were col-
lected selectively from 8 A.M. until 2 P.M. as for-
aging behavior becomes negligible by mid after-
noon.
Insects were collected only if they were ob-
served foraging on the open inflorescences of
Acacia and/or taking nectar from extra-floral
nectaries on the leaves or phyllodes of A. /on-
gifolia, A. myrtifolia, A. pycnantha (Bernhardt &
Walker, 1984), and A. terminalis (Knox et al.,
1985). Foraging is defined here as the active re-
moval of polyads from anthers or the probing of
flowers and extra-floral nectaries with mouth-
parts (Bernhardt et al., 1984). Terminology for
bouts of foraging by bee taxa follows Michener
(1979).

Insects were killed communally in jars con-

species carried by insects, each insect was placed
on a clean glass slide and “bathed” in a couple
of drops of 100% ethanol. When ethanol had
evaporated, the white residue remaining on the

44 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

TABLE 1. Bee taxa collected on the inflorescences of Acacia.

Acacia Species on Which Bee Taxon Was Captured*

No.
Bee Taxon LO ME MI MY PA PY RR RU TE Bees
Anthophoridae:
Exoneura (Brevineura) spp. 0 4 0 0 0 0 0 0 0 4
Exoneura (Exoneura) spp. 0 18 0 0 0 0 2 0 11 31
35
Apidae:
Apis mellifera” A l l 5 4 A A A A
Colletidae:
Amphylae 0 0 0 0 0 0 0 0 1 l
Calli om 0 0 0 0 0 0 0 0 4 4
Callomelitta spp. 0 0 l 2 0 0 l 0 l 5
Euhesma spp 0 0 0 0 0 3 8 0 0 11
E a al spp. l 0 0 0 2 0 0 0 0 3
Hylaeus sp. 0 0 0 0 0 0 l 0 0 l
Leionrocs metallescens 0 0 0 0 0 0 0 14 0 14
. plumo 0 0 0 0 0 0 0 l 0 l
ene s (E uryglossidia) spp. 0 0 0 0 42 0 2 0 0 44
Leioproctus (Leioproctus) spp. 3 0 2 12 7 23 19 0 2 79
Trichocolletes sp. 0 0 0 0 l 0 0 0 0 | 1
164
Halictidae:
Homalictus brisbanensis 0 2 0 0 0 0 0 10 0 12
H. demissus 0 0 0 0 0 0 l 0 0 l
H. dixon 0 0 0 0 0 0 0 0 5 5
H. holochorus l 0 0 0 0 0 0 0 0 l
H. megastigmus 0 0 2 0 7 0 l 0 0 9
H. oxoniellus 0 0 0 0 0 0 0 4 0 4
H. punctatus 0 0 0 l 0 0 1 0 0 2
Lasioglossum (Australictus) spp. 0 12 0 0 0 0 0 0 0 12
(Austrevylaeus) spp 0 0 0 0 0 0 0 0 l l
La. (Chilalictus) spp. 0 9 12 5 2 2 6 l 4 41
La. (Parasphecodes) spp. 2 l 9 9 22 6 16 71 22 158
Nomia spp. 0 5 5 0 0 0 0 0 0 _10
256
Megachilidae:
Megachile sp. 0 0 0 0 0 0 0 2 0 2
2
Grand Total 7 51 31 29 83 34 58 103 51 457
a LO = A. -e ME = A. mearnsii, MI = A. mitchelii, MY = A. myrtifolia, PA = A. paradoxa; PY = A.
pycnantha: RR = etin mie var. retinodes; RU - A. ret. var. uncifolia; TE — A. terminalis.

— Abundant oe more than five bees caught but far more than that number were identified on sight on
this Acacia species).

slide was mounted in one or two drops of Cal- dividual unless 25 polyads could be counted in
berla's fluid (Ogden et al., 1974). Identification that single sample. Taxa producing monad pol-
of pollen was made under light microscopy. len were not counted as present in a sample un-
However, since insects were killed in the same less 25 individual grains were identified on a
jar, contamination of pollen species was possible. single slide (Bernhardt et al., 1984). Insects
Therefore an Acacia species or species of Epa- washed of pollen were air dried, numbered, place
cridaceae was not counted as present on an in- in individual glassine bags, and identified.

1987]

BERNHARDT — AUSTRALIAN ACACIA 45

TABLE 2. Characteristics of the secretion of extra-floral nectar in four Acacia species.*

Characteristics of the Nectary or Nectar LO MY PY TE
Nectary secretes throughout the flowering seasons of + + +
Nectary secretes > 1 ml every morning? = = = +
Nectar hexose ric ? = + —
Nectar sucrose rich or sucrose dominant ? + = +

? Derive

> Observations and collections of nectar made bet

from Bernhardt (1982), Bernhardt and Walker AR and Knox et al. (1985).

tween 7-9 A

LO = A. longifolia, MY = A. myrtifolia; PY = A. pycnantha; TE = A. terminalis.

RESULTS
BEES

Bees foraged for polyads on each of the eight
Acacia species (Table 1). A total of 457 bees were
caught on the eight Acacia species. The greatest
numbers of bees collected belonged to the short-
tongued family of Halictidae. Of all five families
the Halictidae offered the greatest number of gen-
era collected. Lasioglossum (subgenus Parasphe-
codes; Halictidae) species were collected in great-
er numbers on Acacia compared to all other

locally abundant on half of the Acacia species
studied and could be identified effortlessly on
sight (Table 1). This bee was the dominant for-
ager on A. /ongifolia.

Five bee families were identified on eight Aca-
cia species. However, seven out of the 27 bee
taxa represented single captures. The long-
tongued families Apidae and Megachilidae were
represented by only one taxon each. The third
long-tongued family, Anthophoridae, was rep-
resented by two subgenera of Exoneura (Table
1). All remaining bee taxa belonged to the short-
tongued families Halictidae and Colletidae.
Eighty-four percent of the Colletidae collected
belonged to the genus Leioproctus s.l. Eighty-
three percent of the Halictidae belonged to the
genus Lasioglossum s.l. (Table 1). Lasioglossum
(subgenus Parasphecodes) species and A. mellif-
era were the only bee taxa collected on each of
the eight Acacia species (Table 1). Leioproctus
(subgenus Leioproctus) Lud were caught on
seven out of eight Acacia speci

Representatives from the Ë wiis Antho-
phoridae, Apidae, ae i and Halictidae were
recorded as foraging on the extra-floral nectaries
of A. longifolia, A. myrtifolia, A. pycnantha, and
A. terminalis (Tables 1, 2). There was no evi-
dence that Acacia inflorescences growing on

shoots bearing extra-floral nectar received great-
er numbers of bees (or mare bee taxa) compared
to those A
(Table 1).

There was no obvious correlation between the
number of bee taxa collected on an Acacia species
and the habitat in which the species was studied
(Table 3). At the intraspecific level, though, the
dry sclerophyll woodlands and heaths of the
Grampians National Park offered a wider fauna
for A. retinodes var. retinodes than did the coastal
dunes of Cape Schanck (Table 3). The number of
bee taxa below the family level collected on four

Acacia species sympatric through the Brisbane

Ranges National Park varied from five to eight.
No more than four families of bees were collected
on any Acacia species (Tables 1, 3).

Variation in the number of bee taxa on an

Acacia species did correlate positively with the

flowering seasons of the plants and increased from
mid winter (July) until the end of autumn (May;
see Table 4). The number of bee taxa collected

terminalis (Table 4). The greatest number of bee
taxa was collected on the summer-flowering A.
retinodes var. uncifolia (Table 4).

The majority of bee taxa foraging on the eight
Acacia species visited a wide range of plants for
pollen and/or nectar (Tables 4, 5). Technically
these bees must be classified as polylectic for-
agers (Armstrong, 1979; Michener, 1979). From
mid winter until late spring, and then from au-
tumn to early winter, 21-55% of the bees cap-
tured on any Acacia species carried the polyads
of that species mixed with the pollen of one or
more sympatric species. Furthermore 63-91% of
the bees captured on Acacia from late spring to
late summer carried the pollen of one or more
sympatric plants mixed with Acacia polyads (Ta-
ble 4). Bees collected on A. pycnantha showed

46 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE 3. Variation in the number of bee taxa col-
lected at different sites.

No. Bee
Taxa
Acacia (No.

Site? Taxon Families)
Brisbane Ranges MI 7 (3)
MY 6 (3)
PA 8 (3)
PY 5 (3)
Cape Schanck RU 8 (4)
Coranderrk Res ME 8 (3)
Erica-Moe; Bub on TE 10 (4)
Grampians RR 12 (4)
Langwarren Reserve LO 5 (3)

? See study sites for description of habita
A. ae. PA =
t.

A, retinodes var. ` retinodes. LO = A. longifolia.

the lowest level of polylectic foragers, whereas
bees collected on A. mearnsii showed the highest
(Table 4

Sixty-three percent of the bees bearing loads
of Acacia polyads that were mixed with the pol-
len of other plants carried monads of one or more
genera of Myrtaceae (Table 5). Ninety-two per-
cent of the bees carrying mixed pollen loads car-
ried the pollen of plants that bear floral nectaries

Table 5)

WASPS

Wasps were collected on all Acacia species ex-
cluding A. retinodes var. retinodes. Only 58 wasps
were collected on the Acacia species representing
seven wasp families (Table 6). Sixty-four percent
of the wasps were divided equally between the
Sphecidae and Tiphiidae. The greatest numbers
belonged to Cerceris s.l., but these insects were
collected only on A. paradoxa and A. terminalis

Table 6)

A total of 24 wasp taxa below the family level
were collected on Acacia. Eleven of these rep-
resented single captures. No wasp taxon was re-
corded on more than two Acacia species. Acacia
species offering extra-floral nectar appeared to
attract a minimum of twice as many wasp taxa
as those lacking extra-floral nectar (Tables 2, 6).
Wasps observed and collected on A. /ongifolia,
A. myrtifolia, and A. terminalis consistently flew
to the extra-floral nectary before attempting to
forage on the nectarless florets.

[Vor. 74

TABLE 4. Flowering patterns of Acacia species as
correlated with the taxonomic diversity and foraging
behavior of the bees.

No.
Bee No. Bees
axa on Bearing*
Flowering Period Acacia Acacia
and Acacia Species sp. Polyads Ratio’?
Mid Winter-Early Spring
A. longifolia 22 (9) 0.40
A. pycnantha 41 (9) 0.21
Mid Winter-Mid Spring
A. myrtifolia 29 (16) 0.55
Early Spring-Late Spring
A. paradoxa 96 (50) 0.52
Late Spring
A. mearnsii 8 59 (45) 0.91
Early Summer-Mid Summer
A. mitchelii 7 32 (25) 0.78
Early Summer-Late Summer
A. retinodes
var. retinodes 12 67 (56) 0.8
A. retinodes
var. uncifolia 8 121 (77) 0.63
Autumn-Early Winter
A. terminalis 10 50 (16) 0.32
a The first number in the column refers to the total

number of bees caught on the particular Acacia sp. that
carried polyads of Acacia on their bodies. The second
number, in parentheses, refers to the total number of

+L 4 a Te | $ 1 4 1 L 11 y EMT NP

genera,
° The ratio = number of bees that carried Acacia
use plus pollen of other genera divided by total
mber of bees that carried Acacia polyads.

Acacia terminalis offered more extra-floral
nectar per gland on a daily basis than did the
three other nectariferous species (Table 2). Aca-
cia terminalis recelved 33—66% more wasp taxa
than were captured on the other three nectarifer-
ous species.

Wasps foraged selectively on Acacia species
am dunchona ind Pora Hectanes. These

te id
spring and from autumn to early mns (Tables
4, 6). Consequently the density and diversity of
wasps on Acacia was heaviest during the coldest
seasons.

Seventeen wasp taxa carried the polyads of at
least one of the Acacia species on which they
were caught: Antamenes sp., Anthobosca sp.,

47

1987] BERNHARDT—AUSTRALIAN ACACIA
TABLE 5. Comparative frequencies of pollen from other sr phyte famili identified on bees carrying
Acacia polyads.
No. of Bees Bearing Acacia Polyads Mixed wh
of Other Families Captured on each Acacia
Spermatophyte
Family? LO ME MI MY PA PY RR RU TE Total
Compositae + — 0 4 0 0 3 I 6 26 0 40
Dilleniaceae — 0 0 3 0 4 l 0 0 0 8
= semaine + 0 0 0 l 3 l 0 0 3 8
umin
Tanaman PNA l 4 0 0 23 0 0 0 0 28
Liliaceae s.l. + 0 6 0 0 2 0 0 0 0 8
Myrtaceae + 2 47 19 2 26 0 71 43 15 225
inaceae — 9 0 0 0 1 l 0 0 0 11
Pittosporaceae + 0 0 0 0 0 0 2 0 0 2
Proteaceae + 0 0 0 4 0 3 0 0 0 7
Rhamnaceae + 0 0 0 12 2 4 0 0 0 18
* + = bees bearing pollen of genera with nectariferous flowers; — = bees bearing poll fg tł tarl

flowers.

LO = A. longifolia, ME = A. mearnsii, MI =

A. mitchelii MY = A. myrtifolia, PA = A. paradoxa; PY = A.

pycnantha; RR = A. retinodes var. retinodes, RU = A. ret. var. uncifolia, TE = A. terminalis.

Cerceris antipodes, Cerceris sp., Labium sp., Lis-
sopimpla excelsa, Lissopimpla i Lophocheilus
s sp., Phyma re tiles pygi-
. Sp. near nonilicornis, pd us Sp.,
Pseudozethus sp., Rhagigaster PASEAR A Sphex
sp., Tachynomia moerens, and the unidentified
species of Tiphiidae. Mixed loads of Acacia poly-
ads plus the pollen of sympatric plants were con-
fined to those angiosperm families with nectar-
iferous flowers: Compositae, Epacridaceae,
Myrtaceae, Rhamnaceae, and Solanaceae.

DISCUSSION

Wasps versus bees as Acacia pollinators. The

asps, however, comprise only 11%
of the total population of foraging Hymenoptera.
This supports previous reports that bees forage
for Acacia polyads in far greater numbers than
do wasps (Bernhardt, 1982; Bernhardt et al., 1984;
Bernhardt & Walker, 1984, 1985; Knox et al.,
1985). Female bees appear to forage primarily
for polyads and secondarily for extra-floral nec-
tar (Bernhardt & Walker, 1984; Knox et al., 1985).
The results presented in this paper indicate that
wasps tend to forage preferentially for extra-flo-
ral nectar.

Large quantities of extra-floral nectar fail to
assure polyad transport by wasp foragers (Knox
et al., 1985). Almost equal numbers of bees and
wasps foraged for nectar and polyads on valley

populations of A. terminalis, but bees consis-
tently carried polyads of A. terminalis more often
than did wasps (Knox et al., 1985). The low den-
sity of wasp populations on flowering Acacia,
combined with the foraging preferences of these
insects, suggests that wasps are of secondary im-
portance to seed set, at best. Under certain cir-
cumstances, pollen-eating flies are probably su-
perior polyad vectors compared to wasps
(Bernhardt et al., 1984)

This does not mean to suggest that wasps are
always nominal pollinators ofthe Australian flora.
Unfortunately few studies have been done to as-
sess the selective pressure of wasp foraging on
other angiosperms (Armstrong, 1979). Waspsare
important pollinators of nectariferous and nec-
tarless (pseudocopulatory) Orchidaceae in Aus-
tralia (Beardsell & Bernhardt, 1982). Wasps may
also pollinate the nectariferous Epacridaceae
(Bernhardt, pers. obs.; Knox et al., 1985). Of
course, wasps are important pollinators of Pro-
sopis (Simpson et al., 1977), but these mimosoid
shrubs bear floral nectaries.

Bee diversity on Acacia species. Although all
representatives from all five families of Apoidea
found in Australia (Armstrong, 1979) were iden-
tified on Acacia in this study, some families were
more common than others. This is partially ex-
plained by biogeography. The Apidae are poorly
represented in Australia (Michener, 1979), and
the native genus Trigona is uncommon south of
the Tropic of Capricorn (K. Walker, pers. comm.).

48 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

TABLE 6. Wasp taxa collected on the inflorescences of Acacia species.

Acacia spp. on Which Wasp Was Captured

No.
Wasp Taxon LO ME MI MY PA PY RR RU TE Wasps
Braconidae:
Apanteles sp. l 0 0 0 0 0 0 0 0 1
l
Eumenidae:
Antamenes sp. 0 3 0 0 0 0 0 0 0 3
Pseudozethus sp. 0 0 0 0 0 0 0 0 5 5
Unidentified sp. 0 0 l 0 0 0 0 0 1
9
Ichneumonidae:
Biconus sp. 0 0 0 1 0 0 0 0 0 1
Labium spp l 0 0 0 0 0 0 0 3 4
Lissopimpla excelsa 0 0 0 1 0 0 0 0 0 l
Lissopimpla sp 0 0 0 0 0 0 0 0 2 2
8
Pergidae:
Neourys sp. 0 0 0 0 0 l 0 0 9 l
l
Pompilidae:
Chirodamus sp. 0 0 0 0 0 0 0 0 1 l
Pompilius sp. 0 0 0 0 0 0 0 0 1 1
2
Sphecidae:
Cerceris antipodes 0 0 0 0 0 0 0 0 7 Ji
Cerceris spp. 0 0 0 0 1 0 0 0 6 7
n sp. 1 1 0 0 0 0 0 0 0 2
Rhopalum 0 0 0 0 0 0 0 0 l l
Sphex sp. 0 0 0 l 0 0 0 0 0 mt
18
Tiphiidae:
Anthobosca sp. 0 0 0 0 0 0 0 3 5
E anilitatus 0 0 0 l 0 0 0 0 2
Phymatothynnus pygidalis 0 0 0 0 1 0 0 9 l
Phymatothynnus sp. nov.
n. nonilicornis* 0 0 0 0 0 3 0 0 0 3
Phymatothynnus 0 0 0 1 0 0 0 0 0 l
hagigaster comparatus? 0 0 0 0 0 2 0 0 0 2
Tachynomia moerens* 0 0 0 0 0 2 0 0 0 2
Unidentified spp. 2 0 0 0 1 0 0 0 0 3
19
Total No. Nerd
Acacia specie 5 4 1 5 2 10 0 3 28 58
Total No. a Taxa/
Acacia species 4 2 l 5 2 6 0 l 9

a Refers to wasp taxa caught en asa
= A. longifolia, ME = A. mea : MI = A. mitchelii; MY = A. myrtifolia, PA = A. paradoxa; PY = A.
pycnantha; RR = A. retinodes var. wna ri RU = A. ret. var. uncifolia; TE = A. terminalis.

1987]

Trigona may be important in the pollination of
paleotropical Acacia through northeastern Aus-
tralia (Bernhardt, 1982; Armstrong, 1979).
Biogeography alone, however, cannot explain
the comparative dearth of anthophorid and
megachilid bees on Acacia species in southeast-
em Australia, While it is true that neither family
ity of Colletidae
in Australia (Michener, 1979), there are far more
megachilid and anthophorid taxa in Victoria than
were sampled in this study (Michener,

sidered families of long-tongued bees. Although
they may be common throughout the habitats
described in this paper, evidence suggests that
they forage preferentially on flowers offering pol-
len and nectar (Armstrong, 1979). In particular,
Megachilidae often are associated with zygo-
morphic flowers, particularly papilionoid le-
gumes (Michener, 1965; Armstrong, 1979). The
only anthophorid bees relatively common on
Acacia were Exoneura. These bees were largely
confined to the only two arborescent Acacia
species studied bearing bipinnately compound
leaves: A. mearnsii and A. terminalis. This is
probably coincidental, but it would be worth-
while exploring “presumed” foraging preferences
that Exoneura may express towards sections of
Acacia s.l.

Climatic conditions may also influence the di-
versity of bee taxa on Acacia with overlapping
distributions. During the spectacular drought of
1982 and 1983 no specimens of Leioproctus
(subgen. Euryglossidia) were collected on Acacia
in the Brisbane Ranges (Bernhardt & Walker,
1984). The Brisbane Ranges had a cool, wet spring
in 1984, and in that year Leioproctus (subgen.
Euryglossidia) far outnumbered previously
abundant Leioproctus (subgen. Leioproctus).

Considering the broad polylecticism of the bee
taxa on these eight Acacia species, one cannot
refer to any bee taxon as an “acacia bee" as some
entomologists speak of **clover bees,” “orchid
bees,” and “squash and gourd bees” (Michener,
1974, 1979). The very absence of floral and ex-
tra-floral nectar in so many Australian Acacia
species forces bees to forage on other sources.
Oligolecty continues to decline as bees exploit
co-blooming sources of nectar and additional
pollen in competition with Acacia.

Lasioglossum (Halictidae) and Leioproctus
(Colletidae) remain the predominant polyad for-
agers of many Acacia species (Bernhardt, 1982;
Bernhardt et al., 1984; Bernhardt & Walker, 1984,

=
oO

BERNHARDT—AUSTRALIAN ACACIA 49

1985; Knox et al., 1985). Lasioglossum species
may be more important pollinators than Leio-
proctus species. Lasioglossum were caught on all
Acacia species studied and were collected in
greater numbers than were Leioproctus.
Parsphecodes and Chilalictus were the most
commonly collected subgenera on six out of the
eight Acacia species studied. Parasphecodes bees
outnumbered Chilalictus bees. These allied sub-
genera may exert preferential foraging patterns
when nectarless angiosperms have overlapping
flowering periods. From later winter through late
spring Acacia species and some Hibbertia species
(Dilleniaceae) have H unes flowering pe-
riods. Lagio, en. Chilalictus species
forage iq e on yrs (Bernhardt,
1984, 1986), but Parasphecodes forage in greater
ni Des on Acacia (Bernhardt & Walker, 1985).
Polylecticisms and the role of polylectic bees in
Acacia cross-pollination. The Halictidae and
Colletidae are both families of short-tongued bees
that forage for nectar on flowers with shallow

widely distributed family of nectariferous angio-
sperms in Australia (Costermans, 1983). Most
shrubby Myrtaceae flower through the warm
spring-summer months when the greatest num-
ber of bees (both short- and long-tongued) are
active. Out of the four most important nectarif-
erous families that were patronized most fre-
quently by bees in this study, three have flowers
with short-tubular or shallow-bowl perianths
(Faegri & van der Pijl, 1979): Myrtaceae, Com-
positae, and Rhamnaceae. The keeled “flag”
flowers of the papilionoid legumes are also vis-
ited by short-tongued bees probably because the
weight of comparatively large-bodied Leioproc-
tus and Lasioglossum triggers access to the con-
cealed nectaries.

Bees that pollinate nectarless flowers tend to
belong to polylectic genera: Apis (Michener,
1974), Bombus (Buchmann, 1983), Centris
(Frankie et al., 1983), Lasioglossum (Bernhardt,
1984; Bernhardt & Burns-Balogh, 1986). It is not
unusual for an individual bee that belongs to a
taxon in which foragers are not fed by siblings
to visit one to four nectarless but pollen-rich
species and six to eight nectariferous species dur-
ing flowering seasons (Macior, 1968; Bernhardt
& Montalvo, 1979; Bernhardt, 1984; Bernhardt
& Burns-Balogh, 1986).

Acacia species in Australia are, in general, mass-
flowering plants that are also highly self-incom-

50 ANNALS OF THE MISSOURI BOTANICAL GARDEN

patible. Seed set will not occur unless bees move
spontaneously from one shrub to another. The
absence of floral nectar forces polyad foragers to
break their bouts on Acacia species with trips to
nectariferous flowers growing on nonphyloge-
netically related plants (Bernhardt & Walker,
1984, 1985). When the bees are replete with
chemical energy (i.e., nectar) they may return to
Acacia, but in the mosaic distribution of scle-
rophyll shrubs it will probably be to a shrub dif-
ferent from the one abandoned for nectar.

LITERATURE CITED

ARMSTRONG, J. A. 1979. Biotic pollination mecha-
nisms in the Australian flora—a review. New Zea-

. BERNHARDT. 1982. Pollination
biology of Australian terrestrial orchids. de l
183 ¿n E. G. Williams, R. B. Knox, J. H . Gilbert
& P. Bernhardt (editors), Pollination 82. Plant
Cell Biology Research Centre, Univ. of Mel-
rne, Australia.
Maid P. 1982. Insect pollination of Australian
a. Pp. 85-97 in E. G. Williams, E. Knox, J.
ilber & P. Bernhardt (editors), Pollination
'82. Plant Cell rid Research Centre, Univ. of
n Austra
4. The s sete biology of Hibbertia
sri oS PI. Syst. Evol. 147: 267-277.
Bee-pollination in Hibbertia fascicu-
lata Dilniaeae. Pl. nr Evol. 152: 231-241.
1986. Floral mimesis
» Thelma Mene: Br. Pl. Syst. Evol. 151: 187-

D. M. CALDER. 1981. The floral ecology of
pasau populations of Amyema pendulum and
yema T (lanta 2). Bull. Torr.
Club 108: 213-230.

&

. MONTALVO. 1979. The pollination
ecolo ogy of Echeandia macrocarpa (Liliaceae).
Brittonia 31: 64-71.

K. WALKER. 1984. Bee foraging on three
patric species of pueda Acacia. Int. J.
Entomol. 26: 322-330.

85. Insect foraging on Acacia
ee var. . retinodes. Int. J. Entomol. 27: 97-
l

. KENRICK & R. B. KNox. 1984. Pollination
biology ay is breeding system of Acacia reti-
es (Leguminoseae: Mimosoideae). Ann. Mis-

souri Bot. Gard. 71: 17-29.
BUCHMANN, S. L. 1983. Buzz pollination in angio-
rms. Pp. 73-114 in C. E. Jones & R. J. Little
(editors), Handbook of Experimental Pollination
Biology. Van Nostrand wee ias Inc., New York.
COSTERMANS, L. tive Trees and Shrubs of
sm Australia. "Rigby Publishers, Austra-

maa N DER Pur. 1979. Principles of
Pollination Y Ecology, 3rd edition. Pergamon, Ox-
ford/New Y

[VoL. 74

FRANKIE, G. W., W. A. HABER, P. A. OPLER & K. S.
Bawa. 1983. you and organization of

Little (editors), Handbook of Experimental Pol-
lination Biology. Van Nostrand Rheinhold Inc.,
New York.
Kenrick, J. & R. B. KNox. 1979. Pollen develop-
ment and cytochemistry in some Australian species
of rer ore J. Bot. 27: 412-427.

L & R. B. Knox. 1984a. Self-incom-
ud in Acacia. —a pre- or post zygotic mech-
anism? Pp. 146-153 in E. G. Williams & R. B.
Knox (editors), Pollination '84. Plant Cell Biology
Research Centre, School of Botany, Univ. of Mel-
bourne, Australi

ERN

ali
HARDT, ` R. MARGINSON, G. BERESFOR
& R. B. KNOX.

: in E.
Pollination '84. Plant Cell Biology Research Cent
Wigs of Botany, Univ. of Melbourne, Austra lia.

Knox, R. B. & J. KENRICK. 1982. Pol dup —_—

in Suus to the breeding system of 4

411—418 in D. Mulcahy & E. Ottaviano editor

Pollen Biology. oleis Holland Press, Amsterdam

,P HARDT, R. MARGINSON,
BERESFORD, I. BAKER & H. G. BAKER. 1985. Ex-
tra- in nectaries as sr ashi Sd bird polli-
ation t. 72: 1185-

196.

MacioR, L. W. 1968. Bombus (Hymenoptera, Api-
dae) queen foraging in relation to vernal pollina-
tion in Wisconsin. Ecology 49: 20-25

MICHENER, C. D 65. A classification of the bees
of the Australian and South see regions. Bull.
Amer. Mus. Natur. Hist. 130: 62.

. 1974. The Social ake of the Bees. Belk-

nap Press of Harvard Univ., Cambridge, Massa-

chusetts.
1979. Seer dd of the bees. Ann. Mis-

souri Bot. Gard. 16: 277-347.

pling Airborne Pollen. “Hafner, New York.

Simpson, B. B., J. L. NEFF & A. R. MOLDENKE. 1977.
Pp. 84-107 in B. B. Simpson (editor), wa atl
Its Biology in Two Desert Scrub Ecosyste
Dowden, Hutchinson & Ross, Inc., rai ei
Pennsylvania

SPECHT, R. A., E. M. RAE & V. H. BOUGHTON. 1974.
Conservation of major plant communities in Aus-
tralia and Papua New Guinea. Austral. J. Bot.
Suppl. Ser. 7: 1-667.

TuRNER, V. 1982. Non-flying mammal pollination:
an opportunity in appen Pp. 110-121 in E. G.
Williams, R. B. Knox, J. H. Gilbert & P. Bernhardt
(editors), Pollination '82. Plant Cell Biology Re-
search Centre, oe Botany, Univ. of Mel-
bourne, Austra

VoGEL, S. 1978. c T shifts from reward to
deception in pollen flowers. Pp. 89-96 in A. J.
Richards (editor), x Pollination of Flowers
pied Linn. Soc. Symp. No. 6, Academic Press,

FLOWER LONGEVITY AND PROTANDRY IN TWO SPECIES
OF GENTIANA (GENTIANACEAE)!

C. J. WEBB AND JAN LITTLETON2

ABSTRACT

Both Gentiana saxosa and G. serotina are protandrous. When flowers open, pollen is presented

extrorsely around the closed stigma for one to six days. As the stigma opens, t e toward

the corolla lobes. The length of the female phase, and therefore reproductive flower life, is ae hakai

by pollination, although in both species the corolla may remain fresh for longer than one month. Fresh
ale- i i

flowers pollinated on their tenth day of stigma presentation seme no om ipii they appeared
fresh. Senescence of unpollinated flowers differed between species: in G. saxosa the flowers remained
open and gradually deteriorated, but in G. serotina the flowers asus d diei before full senescence.
Pollination-induced flower senescence has been demonstrated for a number of other angiosperms, and
the usual reactions to pollination are corolla abscission, color change, or wilting. In Gentiana, the
closed corolla enfolds the large superior ovary and may serve to protect it from predators as well as
prevent further pollinator visits. Pollination-i
maintenance costs by ensuring that the flower functions no longer than neces sary. One correlate of
this =o in hermaphroditic flowers is protandry, which ensures pollen dispatch before flower

clos

Floral senescence may be either time-depen-
dent (endogenous) or exogenous (usually polli-
nation-induced). However, there have been few
detailed investigations of the factors that deter-
mine floral senescence, and hence, floral longev-
ity (Primack, 1985). Most such studies have con-
ceptrated on corolla tor changes or other

t and

follow p

signal senescence (e.g., Arditti et al., 1973; yee
ditti, 1976; Gottsberger, 1971; Gori, 1983; Strauss
& Arditti, 1984; Casper & La Pine, 1984; Halevy,
1984), and less attention has been afforded struc-
tural changes such as wilting, flower closure, and
corolla abscission (Mayak & Halevy, 1980). Most
of this research is concerned with the proximate
determinants of floral longevity rather than the
evolution of particular responses (but see Stead
& Moore, 1979; Gori, 1983; Casper & La Pine,
1984; Devlin & Stephenson, 1984). The paucity

system and ultimately for the plant’s overall re-
productive strategy. For instance, in hermaph-
rodite flowers pollination-induced flower senes-
cence will limit the duration of pollen and stigma

! We thank I. C. Brown and garden staff at Lincoln,

presentation and so may influence or be influ-
enced by the extent and nature of dichogamy
(Lloyd & Webb, 1986).

New Zealand species of Gentiana (in the
southern group of Philipson, 1972) are protan-
drous (Thomson, 1881; Simpson & Webb, 1980;
Webb, 1984a). Their large, relatively simple

owers make them particularly suitable subjects
for experimental studies of flower function. This
paper describes the response of flowers of two
species to pollination and reports the results of
experiments to determine the functional dura-
tion of male and female phases in terms of pollen
presentation and seed production.

MATERIALS AND METHODS

The two species of Gentiana selected were those
that grew best in cultivation. Gentiana saxosa
Forster f. grows naturally in coastal sites of south-
ern South Island and Stewart Island, New Zea-
land; plants were collected from Curio Bay,
Southland. enti serotina Cockayne occurs
in grassland in h Island; plants
were collected from Lake Lyndon, Canterbury.
The plants were grown in clay pots in an insect-
proof cage in greenhouses at Lincoln, Canter-

Canterbury, for maintaining the plants in cultivation,

J. Miles for spen with pye and P. Brooke for pp aaa 2 = > We are grateful to L. F.
Edgar,

Daph- Lively, nd D. oyd for co

mments on a draft

e manu

2 Botany n Department an E and Industrial Meu y see Bag, Christchurch. New Zealand.

ANN. Missouni Bor. GARD. 74: 51-57. 1987.

52 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE 1. Results of controlled pollinations to de-
termine self-compatibility in Gentiana saxosa and G.
serotina.

%
Seeds
Pro-
u ap- duced
ber of sules per
ow- ro ap-
Treatment ers duced sule
G. saxosa Cross- 10 10 94.2
pollinated
Self- 10 10 94.2
pollinated
Unpollinated 10 0 —
G. serotina Cross- 5 5 57.9
pollinated
Self- 5 5 67.6
pollinated
Unpollinated 5 0 —

bury. Field observations were made at Lake Lyn-

n.

To test for self-incompatibility and autogamy
10 flowers in G. saxosa and five flowers in G.
serotina were assigned to each of three treat-
ments: self- and cross-pollination by hand, and
unpollinated (Table 1). Flowers were tagged in-
dividually and observed daily. When capsules
matured, they were harvested and good seeds
and aborted seeds or undeveloped ovules were
counted.

The duration of male and female phases was
determined for individual tagged flowers. Pollen
and stigma presentation were observed daily un-
til petals withered and turned brown. Of 61 flow-
ers of G. saxosa observed daily, five were cross-
pollinated on each of the first to tenth days of
their female phase, and 11 left unpollinated. Of
34 flowers of G. serotina, eight were pollinated
on each of the first, fifth, and eighth days of their
female phase, five on the tenth day, and five
flowers were left unpollinated. For G. serotina,
three flowers from each of the first, fifth, and
eighth day pollinations were collected two days
after pollination, the stigma dissected out, fixed,
stained with aniline blue, and examined under
fluorescent microscopy to determine the extent
of pollen germination and pollen tube growth.
For the remaining flowers, the responsiveness of
each to pollination was quantified as the number
of days until (a) the flower closed completely, (b)
closed to its greatest extent (for flowers that closed

[Vor. 74

incompletely), or (c) the number of days until
the corolla withered and turned brown (for flow-
ers that never closed). When capsules were ma-
ture they were harvested and seed production
scored.

At Lake Lyndon, 10 fresh, unbagged, female-
phase flowers of G. serotina were cross-pollinat-
ed by hand, and ten were left unpollinated. These
flowers were examined the following day.

RESULTS

Flower form, phenology, and pollina-
tion. Both Gentiana saxosa and G. serotina are
perennial herbs with a central leafy rosette and
stout taproot. The flowers are on annual, leafy,
lateral flowering branches that bear from one to
eight flowers in G. serotina and one to 30 or more
in G. saxosa. The corolla is deeply 5-lobed, white
with translucent stripes on the lobes and upper
tube (Fig. 1), and greenish-yellow toward the base
of the tube. The five stamens are attached to the
corolla tube; the anthers are purple and the pollen
is cream to brownish-yellow. Nectar is produced
between the bases of the filaments near the base
of the tube. The central ovary contains an av-
erage of 29 ovules in G. saxosa and 47 in G.
serotina (Webb, 1984a). The flowers lack a well-
defined style; the stigma is 2-lobed, distinctly
papillate, and dry.

In the field, G. saxosa blooms in summer and
autumn (January to May), and G. serotina from
late summer to autumn (February to May). Un-
der greenl diti G hed peak
flowering in December and G. serotina in March.
The flowers are sweetly scented and visited on
fine days by a range of insects. At Lake Lyndon,
G. serotina was most frequently visited by syr-
phid flies and solitary bees (Lasioglossum sor-
didum). During anthesis, flowers opened com-
pletely only on fine days, and almost completely
closed at night. On wet or cold, dull days few
new flowers opened, and in those that had opened
previously the corolla lobes did not spread com-

etel

=

Both G. saxosa and G. serotina are self-com-
patible (Table 1); however, flowers that were not
pollinated and from which insects had been ex-
cluded failed to produce any seed, so biotic pol-
lination is necessary for seed set.

Protandry. The flowers of both species are
distinctly protandrous. In neither was there any
obvious synchrony of male and female phases
among flowers within a plant or even a flowering
branch, so that within large plants pollen and

1987]

WEBB & LITTLETON—GENTIANA 53

FiGures 1-8.— 1-7. Protandry and reaction to pollination in flowers of Gentiana serotina (bars = 0.5 cm

1. First open, male phase.—2. Fourth
corolla. —
after pollination of fresh Pani phase flower.— 6.
eighth day of female phase.—
(bar = 0.5 cm)

stigmas were presented simultaneously in differ-
ent flowers and geitenogamy could occur.
When a flower first opens, the anthers have
already dehisced to present pollen ext ly near
the center of the flower and the stigmatic lobes
are tightly closed (Fig. 1). Later in the male phase,
the corolla lobes open further, the stigma begins
to open, and the stamens move outwards toward
the corolla (Fig. 2). In G. saxosa, the stigma may
open and stamens recurve late on the first day
of anthesis, or as late as the fifth day. The mean
duration of the male phase is 1.41 days (N = 57).
In G. serotina, the stigma opens and stamens
recurve between the third and seventh days
(mean =4.73 days, N = 30). In G. saxosa, an-
thers have usually moved halfway toward the
corolla when the stigma opens and have reached
the corolla lobes in up to three days after that.
In G. serotina, the anthers reach the petals and
usually wither one or two days after the stigma
opens (Fig. 3), and in some cases they eventually
protrude between the corolla lobes. Although
pollen is no longer presented in a central position
by the time the stigma opens, there may be some
overlap between pollen and stigma presentation
within the flower, especially in G. saxosa, for
which the male phase is shorter. However, under

4. Flower closing, one day after arcade of fresh female-phase flower. —
Flower incompletely closed, five days after pollination on
7. Unpollinated flower, 18 days.—8. Unpollinated flower of G. saxosa, 30 days

day, stigma opening.—3. Seventh day, 1 igma open, anthers near

5. Flower closed, two days

field conditions, little pollen usually remains by
the time the stigma opens.

When the stigma opens the two lobes occupy
the position in which pollen was presented in the
newly opened flower (Figs. 1, 3). The duration
of the female phase is dependent on pollination.

Reaction of the flower to pollination. Both
species showed similar reactions to pollination.
When fresh, first-day, female-phase flowers were
pollinated, they partially closed that night and
failed to reopen the following day (Fig. 4). Two
days following pollination the corolla lobes were
imbricate to form a neat, tight structure similar
to, but slightly larger than a late bud (Fig. 5). The
petals slowly turned brown and withered as the
capsule reached maturity after about four weeks.

When the flower was pollinated on the second
to fifth days of the female phase, the corolla re-
acted as quickly and folded as neatly as it did
following first-day pollination. After the fifth day,

u

„ei
close or closing incomipietely
third of the flowers pollinated after the fifth day
of their female phase never closed completely.
There was some variation among flowers of the

2 4 6 8 10
DAY OF FEMALE PHASE
FIGURE 10. Average percentage of ovules produc-

54 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74
17 100.,
Z
5
; Oso |
|
C5 O
Z p =)
c z C60 .
T / O
= / a
= 10 4 40
OQ
na
ul
S TE
n WY 20 |
D o
o IN
S
O te ae 0 T T T T *—. 1
O
=
(O
>
<
e

o

NEVER R POLLINATED
DAY OF FEMALE PHASE `
E9. Days taken for flowers of Gentiana sax-
osa (solid dots) and G. serotina (open squares) to close
or wither NE pollination on different days of
emale phas

same age, but the trend to decreasing reactivity
with increasing age was clear in both species (Fig.

).
In the field, flowers of G. serotina reacted sim-
ilarly to those in the greenhouse. Ten fresh fe-
male-phase flowers were all closed on the day
following hand pollinations, whereas eight of 10
flowers not pollinated by hand had reopened.
Under fluorescent ai ee all three stig-
mas from flowers of G. serotina that had been
pollinated on the first day of the female phase
had numerous germinating pollen grains and
many well-formed pollen tubes penetrating the
stigma. Of the three stigmas pollinated on their
fifth day, two showed good pollen germination
and the third had only a few germinating grains.
Two of the three stigmas that had not been pol-
linated until their eighth day had no pollen ger-
minating, and the third had good germination.
The proportion of ovules that developed into
good seeds was clearly affected by the age of the

production for flowers pollinated later than the
fourth or fifth days of the female phase (Fig. 10).
In G. saxosa, flowers pollinated on the first to
fourth days of the female phase consistently ma-

ing seed for flowers of Gentiana saxosa (solid dots) and
: í Y fall : 11: 4i am ALS

ferent days of female phase (mean + standard error
for G. saxosa, mean only for G. serotina).

tured a high proportion of their ovules. Those
pollinated on the fifth to seventh days were more
variable in seed production, and seed production
was related to reactivity of individual flowers.
For example, a flower pollinated on the fifth day
failed to close and produced no seed, whereas a
flower pollinated on the seventh day closed al-
most completely and had 25% seed production.
Few flowers pollinated on the eighth day of the
female phase, and none pollinated on the ninth
or tenth days peoduced soea, The panem of seed
production for fl

in G. serotina was similar (Fig. 10), but the small-
er sample size does not allow a detailed analysis.

Thus, although the corolla and stigma may
appear fresh for much longer, the functional fe-
male phase in both species is only four to eight
days. Adding together male and female phases,
one gets a reproductively functional flower life
of five to 12 days for G. saxosa, and six to 14
days for G. serotina.

The reaction of flowers that were never pol-
linated differed between species. In G. serotina,
the corolla lobes continued to diverge as the flow-
er aged, but eventually began to close (Fig. 7),
finally reaching a position similar to that of a
late-pollinated flower (Fig. 6) before turning

recurved position and in all but one of 11 flowers
showed no sign of returning to a closed position
(Fig. 8) and turned brown after an average of 17
days. In both species some flowers had white,

1987]

turgid corollas for longer than a month, although
the stigmas were dull and blackened by this time
and the stamens had altogether withered.

DISCUSSION

Floral longevity of individual flowers may be
defined in terms of reproductive function as the
period over which a flower is able to receive and/
or dispatch pollen. However, outside this period,
flowers may perform functions not directly re-
lated to their own reproductive success, as in the
case of supplemental pollinator attraction pro-
vided by late buds and older flowers. Thus, a
more practical definition of floral longevity in
terms of reproductive success of whole plants
may be the period for which a flower is able to
attract pollinators. For many plants, particularly
tropical species, floral longevity appears to be a
predetermined, endogenous characteristic with
many such species having 1-day flowers (Dob-
kin, 1984; Primack, 1985). In some such species,
non-induced floral changes, which indicate that
the flower is no longer reproductively functional,
may occur before full senescence (indicated by
abscission or wilting of petals). In others, flowers
may be reproductively functional until full se-
nescence (Gori, 1983). The second major pattern
is provided by those species in which flower life
may be curtailed by pollination (Kerner, 1902;
Arditti, 1976; Stead & Moore, 1979; Devlin &
Stephenson, 1984). It is important to note that
for species in which floral changes are pollina-
tion-induced, the potential flower life is still en-
dogenously determined and the pollination-in-
duced changes occur within this outer limit.

Gori (1983) gnized and ized
able data on five basic types of floral change that
follow pollination or indicate the end of repro-
ductive function before full senescence: color
change, termination of odor and/or nectar pro-
duction, change in flower orientation, collapse of
flower parts, and corolla abscission. To this may
be added flower closure as described here for
Gentiana, and reported by Kerner (1902) for
Mammillaria glochidiata. In fact, flower closure
is likely to be the response in many species in
which petals close regularly at night or in dull
weather. Although floral changes may prevent or
deter pollinator visits, the reproductive, func-
tional end of flower life occurs when no viable
pollen is available for dispatch and the stigma
ceases to be receptive. In Gentiana serotina and
G. saxosa, the end of reproductive flower life is
indicated either by flower closure or the loss of

WEBB & LITTLETON — GENTIANA 55

responsiveness of the flower to pollination. Once
this occurs, pollen does not germinate on the
stigma, and no seeds are produced. The begin-
ning of flower life is marked by the corolla lobes
opening to expose the dehisced anthers; all pollen
is normally lost before the beginning of the fe-
male phase, so pollen availability does not de-
termine the end of flower life, although it may
affect flower longevity if the male phase is pro-
longed by a paucity of pollinator visits as in Lo-
belia cardinalis (Devlin & Stephenson, 1985).

When full senescence ofa flower finally occurs,
floral changes as listed above may occur. In ad-
dition, the whole flower may be aborted, even if
pollination has occurred, especially in species that
use flower abortion as a means of maternal reg-
ulation (Lloyd, 1980; Bawa & Webb, 1984).
Within a species, it is possible that the floral
changes that occur following pollination may dif-
fer from those of flowers that are never polli-
nated. In both Gentiana saxosa and G. serotina,
fresh female-phase flowers closed in response to
pollination; in G. saxosa, but not G. serotina, the
reaction of never-pollinated flowers differed in
that flower closure did not occur and petals with-
ered in a recurved position. However, the end
of functional pollen and stigma presentation in
never-pollinated flowers occurred well before the
morphological change of flower closure in G. se-
rotina and corolla wilting in G. saxosa. The per-
sistence of these flowers well beyond their re-
productively functional life may add to the overall
floral display of the plant.

Pollination-induced changes in flowers have
been interpreted as signals that direct pollinators
to unvisited flowers (Allen, 1898; Kerner, 1902;
Arditti, 1976; Stead & Moore, 1979; Casper &
La Pine, 1984) or help to conceal the developing
seeds within pollinated flowers from predators
(Allen, 1898), or simply minimize costs of flower
maintenance by retaining the perianth no longer
than is necessary (Kerner, 1902). Gori (1983)
considered three aspects of the first alternative:
avoidance of pollinator interference within pol-
linated flowers, increasing the pollinator's for-
aging efficiency and so increasing the residence
time on the plant, and increasing ppilinatiog: ef-
ficiency by restricting p
productively fonction) flowers. In ‘the buo
species of Gentiana studied here, corolla closure
clearly signals that the flower is unavailable for
visits; in fact it precludes visits, and the corolla
also tightly enfolds the developing ovary until it
is almost mature, making it less accessible to

56 ANNALS OF THE MISSOURI BOTANICAL GARDEN

predators. Field experiments, as described by
Gori (1983) for Lupinus, would be necessary to
determine the precise function of pollination-in-
duced flower closure in Gentiana. In many in-
sect-pollinated species, the end of flower life is
signalled by corolla abscission and the devel-
oping ovary is either inferior or enclosed within
infolded bracts or calyx lobes as in Malvaviscus
arboreus (Webb, 1984b). Corolla closure, rather
than abscission, may be necessary to protect the
ovary in Gentiana because the large superior
ovary extends well beyond the small calyx lobes.

In terms of natural selection, flower life for a
particular species is likely to be a trade-off be-
tween the cost of maintaining the flower in a
receptive state and the probability that it has
been pollinated. Thus, as originally suggested by
Kerner (1902), flower life is likely to be longer
for species that normally experience unpredict-
able weather conditions allowing fewer suitable
times for pollinator visits, for species with few,
large flowers per plant, and for species that are
obligately outcrossing. These suggestions are
supported by data on flower life that show that
many tropical plants, particularly those flowering
in the lowlands d the dry season, have 1-day
, 1984; Primack, 1985). Under

contrast, plants of higher altitudes and of tem-
perate areas have long-lived flowers, as might be
expected when many days are unsuitable for
flower visits. That many large orchid flowers with
complex outcrossing mechanisms are long-lived
is to be expected because they may have low rates
of flower visitation.

There are three strategies that might to some
extent ameliorate the difficulty of unpredictable
pollination. First, if flower life is pollination-de-

endent, as in the two species of Gentiana de-
ied here, then flower maintenance costs are
minimized. There is, however, probably a cost in
for pol-

lination-induced floral changes (Gori, 1983).
Second, under conditions unfavorable to polli-
nators, autogamy may occur (Kerner, 1902; Fae-
gri & van der Pijl, 1979); the evolution of autog-
amy often involves loss of dichogamy or
herkogamy, and also a reduction of flower life
(Morin, 1983). In some species selfing may be
delayed until an opportunity for outcrossing has
been provided or may even be a direct response
to unfavorable weather conditions. In Gentiana
lineata, the flowers open fully in warm sunny
weather suitable for insect pollination, but on

[Vor. 74

wet or dull days the flowers open only partly so
that the anthers dehisce directly onto the stigma

(1984). T
during rather unfavorable to pollination, or
flowers may open only on suitable days with the
floral parts protected during inclement weather
(Kerner, 1902).

When floral changes are pollination-induced,
flowers are likely to be non-dichogamous or pro-
tandrous because HOWEN: closure, are c
sion, and other t rtail

the time over which pollen can be pene
(Lloyd & Webb, 1986). The pollination-induced
flower closure described here for Gentiana can
be effective only if flowers are protandrous; pro-
togyny would preclude pollen presentation, and
adichogamy would severely limit the male phase.

Flower life may appear to be the simple result
of proximate causes—especially of weather con-
ditions and pollinator availability. However, the
average flower life for a population, the particular
response of flowers to pollination, and the cues
used to determine the time of senescence, must
all be the result of natural selection. Yet, the
selection of these factors has seldom received the
attention of reproductive biologists in spite of
the important part they play in determining the
success of plants as pollen or seed parents.

LITERATURE CITED

ALLEN, G. 1898. Flashlights on Nature. Grant Rich-
ards, London.

ARDITTI, J. 1976. Post-pollination phenomena in or-
chid flowers. The Orchid Review 84: 261-

OGAN & A. V. CHADWICK. 1973. Post-
pollination phenomena in orchid flowers. IV. Ef-
fects of ethylene. Amer. J. Bot. 60: 883-888.

Bawa, K. S. & C. J. WEBB. 1984. Flower, fruit and
seed abortion in tropical forest trees: implications
for the evolution of paternal and maternal repro-
ductive patterns. Amer. J. Bot. 71: 736-751.

Casper, B. B. & T. R. LA Pine. 1984. Changes in
corolla color and other floral characteristics in
Cryptantha humilis (Boraginaceae): cues to dis
courage pollinators? Evolution 38: 128-141.

DevLin, B. & A. G. STEPHENSON. 1984. Factors that
influence the duration of the staminate and pis-
tillate phases of Lobelia cardinalis flowers. Bot.
Gaz. 145: 323-328.

Sex differential floral lon-
gevity, nectar secretion, and pollinator foraging in
a protandrous species 72: 303-310.

D 1984

i

d resident nectarivores
FAEGRI, K. & L. VAN DER PUL.

. Oecologia 64: 245-254.
1979. The Principles

1987]

of Pollination Ecology, 3rd edition. Pergamon
Press, Oxford.

GonRi, D. F. 1983. Post-pollination phenomena and

tive floral changes. In . Jones & R. J.

Little (editors), Handbook of Experimental Pol-

lination Biology. Van Nostrand Reinhold, New

GOTTSBERGER, G. 1971. Colour change of petals in
ow arboreus flowers. Acta Bot. Neerl. 20:
—388.

unre A. H., C. S. WHITEHEAD & A. M. KOFRANEK.
1984. Does pollination induce abscission of Cy-
clamen flowers promoting ethylene produc-
tion? Plant Physiol. 75: 1090-1093.

KERNER VON MARILAUN, A. 1902. The Natural His-
tory of Plants, odes F. W. Oliver (translator).
Blackie & Son, Lon

LrLovp, D. G. 1980. Sexual strategies in plants. I. An
hypothesis of serial adjustment of maternal in-

vestment during one reproductive session. New
Phytol. 86: 69-79.
. J. WEBB. 1986. Avoidance of interfer-

ence between the presentation of pollen and stig-
mas in angiosperms. I. Dichogamy. New Zeal. J.
2

1980. Flower senes-

V. Thimann (editor), Senescence in
Plants. CRC Aes Boca Raton, Florida.

Morin, N. 1983. Ap of Githopsis (Campan-
ulaceae). Syst. Bot. 8: 436-468.

PHILIPSON, W. R. 1972. The generic status of the

WEBB & LITTLETON—GENTIANA 57

southern hemisphere gentians. Adv. Pl. Morph.
1972: 417-422.

PRIMACK, R. B. 1985. Longevity of individual flow-

SCHOEN, D. The selection
of cleistogamy and heteromorphic diaspores. Biol.
J. Linn. . 23: 303-322.

SIMPSON, M. J C. J. WEBB. 1980. Germination
in

udies on flower
longevity in Digitalis. Pollination induced corolla
abscission in Digitalis flowers. Planta 146: 409—
414.

STRAUSS, M. S. & J. ARDITTI. 1984. Postpollination
phenomena in orchid flowers. XII. Effects of pol-
lination, emasculation, and auxin treatment on
flowers of Cattleya porcia *Cannizaro' and the ros-
tellum of Phalaenopsis. Bot. Gaz. 145: 43-49.

THoMsoN, G. M. 1881. On the fertilization, etc., of
New Zealand flowering plants. Trans. Proc. Ne
Zeal. Inst. 13: 241-288.

984a. Constraints on the evolution of

tematics. Academic Press Canada, Don Mills, On-
tario.

1984b. Hummingbird pollination of Mal-
vaviscus arboreus in Costa Rica. New Zeal. J. Bot.
22: 575—581

NOTES ON THE BREEDING SYSTEMS OF
SACOILA LANCEOLATA (AUBLET) GARAY
(ORCHIDACEAE)!

PAUL M. CATLING?

ABSTRACT

To document breeding systems in the Mensa neotropical terrestrial orchid, Sacoila paula Nati
pollination and seed development were studied in the field and in cultivated plants. In the s ern
Florida study area, plants of var. /anceolata were not pollinated ; and hummingbird pollinators were
apparently absent. When plants from ea
were abundant in southern Ontario, huoyalagbini- ñollinstiop was observed on numerous occasions
and the incidence of pollination was approximately 90%. Pollination experiments demonstrated a
reliance on pollen vectors in plants of var. lanceolata from central Guyana, where a variety of hum-
mingbird pollinators are available, but the plants of the same variety from southern Florida were
found to be agamospermic. Examination of serial sections of ovaries in successive developmental

o in portions of the tropical range. Plants of S. ' lanceolata var. paludi cola from southern
Florida were found to self-pollinate. I ponnani indepe in southern Florida
populations of S. lanceolata var. lanceolata and v dia and the apparent absence of races

totally nul upon pollen vectors are aoc With pollinator-paucity

Sacoila lanceolata (Aublet) Garay var. lanceo-
lata is a reddish- to orange-yellow- or occasion-
ally pale greenish-flowered, terrestrial orchid with
a broad neotropical distribution extending from
northern Florida and northern Mexico to north-
ern Uruguay (Luer, 1972). be urpis:
var. paludicola Luer is restricted to extreme
southern Florida [the Siar eines Strand in
Collier Co. (Luer, 1971a), Kendall Hammock in
Dade Co. (pers. obs.), and from one location in
the Everglades (McCartney, 1985)] and the Ca-
ribbean region (based on examination of speci-
mens at AM

varieties are representative of the bird pollina-
tion syndrome (Pijl & Dodson, 1966).

Casual observations of seed development in a
few cultivated plants that were maintained in the
absence of any potential pollen vectors suggested
that both var. /anceolata and var. paludicola may

hk ll; + £ 4 4 +1 3l

southern Florida. Since the breeding systems of
tropical and subtropical terrestrial orchids are
relatively poorly known, a study was undertaken
to document the breeding systems of S. /anceo-
lata.

METHODS

FIELD STUDIES OF POLLINATION IN
SOUTHERN FLORIDA

In order to gather information on natural pol-
lination, populations of S. /anceolata var. lan-
ceolata were observed for an overall total of 20
hours in southern Florida during mid-May 1978
and 1984. The presence of potential pollinators,
including hummingbirds and large bees partic-
ularly, was noted. Large bees are the regular pol-
linators of some related species (Pijl & Dodson,
1966; Catling, 1983)

Since pollination of orchid flowers involves
removal of the pollinarium from the anther, it
is possible to obtain an approximation of the
incidence of pollination by recording pollinaria

! Living plants of Sacoila lanceolata var. paludicola ki provided by C. A. Luer, H. Brown, and G. Matous.

C. A. Luer and V. R

and N. H. Wil
? Agriculture Canada

Brownell assisted with field work
(University of nd C. A. Luer (Missouri Botanical Garde

Useful criticisms were provided by S. C. H. Barrett
n), N. R. Morin (Missouri Botanical Garden),
k.

liams (Univ. of Florida). W. Wojtas kindly anda with the wir pes wor
Research Institute, Centr:

rm, Ottawa, Ontario, Canada

KIA 0C6.

ANN. MISSOURI Bor. GARD. 74: 58-68. 1987.

1987]

FIGURE 1. x. B,
d pen Een

ers, Immokalee, Collier Co., Florida,

CATLING—BREEDING SYSTEMS OF SACOILA 59

1cm

pou

. S. lanceolata var. lanceolata.—A. Frontal view of flowers, ps oisi Co.,
—B. y la spike of cultivated plant ey central Guyana, South Am —D.

ca, 13
May.—C. S. z var. e ned spike,

Side of flow
vated plant from Fahkahatchee Strand, Collier e. Florida, 15 Febru

removal. This is more easily recorded than the
presence of pollen tubes in the deteriorating col-
umn of faded flowers. It is, however, only an
approximation since pollinaria removal is oc-
casionally (although apparently rarely) prevent-

ed by malformation or drying out of the viscid-
ium or by the difficulty of attaching to an existing
excess load of pollinaria. The incidence of pol-
lination in southern Florida, as determined by
the absence of the pollinaria in faded flowers,

60 ANNALS OF THE MISSOURI BOTANICAL GARDEN

was determined by examining 500 flowers from
a total of 100 inflorescences representing eight
locations.

STUDIES OF POLLINATION OF TRANSPLANTS

Plants from the Sarasota and Immokalee areas
of southern Florida were observed for approxi-
mately 33 hours after transplanting into a rural
setting in southeastern Ontario, Canada, in late
May 1984. The presence of potential pollinators
was again recorded. The approximate incidence
of pollination was determined by pollinaria re-
moval (see above) in five groups of five experi-
mental plants each, including a total of 429 flow-

POLLINATION EXPERIMENTS

Pollination experiments were performed on
plants maintained in a glass house in Ottawa in
April and May 1984 from which potential pol-
linators (insects, hummingbirds) were excluded.
Voucher specimens are contained in the Agri-
culture Canada herbarium of the D.
Research Institute in Ottawa, Ontario (DAO
The experiments were performed on ra groups
of plants: 1) 60 plants of S. /anceolata var. lan-
ceolata, four from each of 15 localities in south-
ern Florida distributed in a broad band, ca. 30
km wide, from Sarasota south to Immokalee; 2)
three plants of S. /anceolata var. lanceolata from
near Mahdia, central Guyana, South America;
and 3) six plants of S. /anceolata var. paludicola
including three from the Fahkahatchee Strand,
Collier Co. and three from Kendall Hammock,

ade Co., both in southern Florida

Starting from the base of a spike, each se-

pollinated, the fourth pollinated with pollen from
a different flower on the same plant (geitonoga-
mous pollination), and the fifth was pollinated
with pollen from a flower on a different plant
(cross-pollination). With the exception of the
plants from Guyana, cross-pollination was car-
ried out using plants from different localities.
Each ovary was appropriately tagged to indicate
the treatment of the corresponding flower: The
Hull ber
as the percentage of embryo sacs containing em-
bryos. The latter value was estimated by emp-

well

[Vor. 74

tying the capsule and determining the presence
or absence of embryos in the first 200 embryo
sacs observed. This was followed by a scan of a
few thousand to make certain that the value ob-
tained was representative. If the value was con-
sidered not representative, two more samples of
200 were scored and the average percentage was
then recorded.

ANATOMICAL STUDY

Each of the three groups of plants used in the
pollination experiments was studied anatomi-
cally to dete ermine whether or not peeudogamy
tre equiring
was operating and to determine the method of
agamospermy in plants producing seed without
any pollination. Serial sections of ovaries were
examined in successive developmental stages.
Ovaries fixed and preserved in formalin-acetic
acid-alcohol (FAA) were dehydrated, embedded
in paraffin, sectioned, mounted, and stained us-
ing a safranin-hematoxylin combination (Lillie,
1969; Jensen, 1958). Mature seeds of experi-
mental plants were stained for examination using
the differential stain technique described by Ow-
czarzak (1952). The percentage of polyembryo-
nic seed was determined in the same way as the
percentage of embryo sacs containing embryos
(see above).

(asexual seed

STUDY OF DRIED MATERIAL

An attempt was made to determine the dis-
tribution of breeding systems through the ex-
amination of herbarium material from AMES,
MICH, SEL, FTG, and DAO (acronyms from
Holmgren et al., 1981). This is made possible,
to a degree, by association of various morpho-
logical features with a specific breeding system.
Plants were considered agamospermic if they
possessed any polyembryonic seed. Relatively
small flower size was considered potentially in-
dicative of autogamy (see Ornduff, 1969). Lack
of expansion of some ovaries in fruiting material
was considered suggestive of obligately cross-
pollinated races.

RESULTS

FIELD STUDIES OF POLLINATION IN
SOUTHERN FLORIDA

In southern Florida, insects (including larger

bees) were present but were not observed visiting

the flowers of S. lanceolata var. lanceolata. No

1987]

CATLING — BREEDING SYSTEMS OF SACOILA 61

E2. Ruby-throated yanmar (Archilochus colubris) pollinating Sacoila lanceolata var. lanceolata
ida, 25 May

FIG
fede: som ota Co., Flori

. Female probin

ower directly through the floral tube. Female

withdrawing with a pale yellowish Sdn attached to the tip of the bill. — C. Male probing a dower from the
right side below the lip.—D. Male probing flower directly.

hummingbirds were observed. There was no evi-
dence of pollination, in the form of pollinarium
removal, in any of the eight southern Florida
locations.

STUDIES OF POLLINATION OF TRANSPLANTS

In southeastern Ontario, large insects includ-
ing large bees were common, but they totally
ignored the flowers. Ruby-throated Humming-
birds (Archilochus colubris) were also common.

pollinators. Females were observed visiting in-
florescences on 25 separate occasions and males
on 46. A visit usually involved probing three to
five flowers on an inflorescence and visiting one
to three inflorescences in a group of five. Visits

were 15 minutes to two hours apart and were
most regular in the morning between 9 A.M. and
11:30 A.M. Visits by one or more females (i.e.,
female visits) involved a direct probing of the
flowers (Figs. 2A, 3A), the pollinia becoming at-
tached, by the elongate cushion-type viscidium
that sheaths the rostellum (Fig. 3C, D, E, and
Greenwood, 1982), to the distal portion of the
bill (Figs. 2B, 3B). Male visits involved probing
from the side and below the lip approximately
50% of the time (Fig. 2C, D). Probably as a con-
sequence of this “robbing,” male visits resulted

5 experimental plants, which were
observed being pollinated by the hummingbirds,
the incidence of pollination was 90% (1.e., 38
of 429 flowers).

62 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

oe

| 10 mm |

FIGURE 3.
a flower of Sacoila (alent see var. lanceolata. — A. B
tw

Column viewed from below
viewed from
Co., Florida

POLLINATION EXPERIMENTS

Pollination was clearly necessary for seed de-

in the absence of pollination. Pollination at any
time is followed by fading in two or three days.

In S. /anceolata var. lanceolata from southern
Florida, seed developed regardless of pollinar-
ium removal (Table 1), and these plants were
clearly autonomously agamospermic. The flow-
ers lasted 5-15 days in the absence of pollination,
but pollination resulted in fading within two days.

Seed development occurred in undisturbed
flowers of S. /anceolata var. paludicola, but not
when the pollinarium was removed from the
flower at an early stage (Table 1), indicating

it | the di (darkenedi =

C D E

iagrams of a pM eager (Archilochus colubris) ia aen pollinia from
ird with bill inserted in flower, prior

pward movement
—B. Bird with pollinia attached near tip | of bill. —

afte
below showing viscidium (darkened) and pollinia. All diagrams based on material from Sarasota

autogamy (self-pollination) or pseudogamy (pol-
lination-induced agamospermy) in this variety.
Self-pollination occurred after the flowers had

en open for one or two days. The flowers are
short-lived, lasting for approximately five days.
The mechanism of self-pollination is a simple

lengths of the pollinia and the basal part of the
column (Fig. 4A).

ANATOMICAL STUDY

The Guyana plants of var. lanceolata and plants
of var. paludicola from southern Florida dem-
onstrated a normal sequence of pollination, pol-

1987] CATLING — BREEDING SYSTEMS OF SACOILA 63
TABLE 1. Pollination experiments with Sacoila lanceolata.
No./96 No./%
No. of No. of Ovaries Ovaries % Sacs with
Treatment Plants Ovaries Expanded* with Seed* Embryos*
S. lanceolata var. lanceolata—Guyana
Undisturbed 3 6 0/0 0/0 0
Pollen removed 3 6 0/0 0/0 0
| botes 3 6 6/100 4/66 50-100
Geitonogamous-pollination 3 6 6/100 4/66 60-100
Batic 3 6 6/100 4/66 60-100
S. lanceolata var. lanceolata—South Florida
Undisturbed 60 200 200/100 200/100 90-100
Pollen removed 60 180 180/100 180/100 97-100
Self-pollination 60 180 180/100 180/100 96-100
Geitonogamous-pollination 60 180 180/100 180/100 97-100
Cross-pollination 60 180 180/100 180/100 97-100
S. lanceolata var. paludicola—South Florida
Undisturbed 6 25 25/10 25/10 95-100
ollen removed 6 12 0/0 0/0 0
Self-pollination 6 12 12/100 12/100 95-100
Geitonogamous- eee 6 12 12/100 12/100 95-100
Cross-pollinatio 6 12 12/100 12/100 95-100

* As a percentage of the total ovaries tested.

len tube growth, fertilization, and embryo de-
velopment (see Catling, 1982). In plants of var.
lanceolata from Florida, the gametophyte de-
generated and one or more cells of the inner in-
tegument proliferated to become embryos. The
proliferation of these cells was indicated by the
more deeply staining protoplasts, relatively large
nuclei, and well-developed nucleoli (Fig. 6). Pro-
liferation had initiated in the ovaries of flowers
open for five days and may have initiated earlier
but was not evident in mature ovaries with buds
on the point of opening. Using ovaries of in-
creasing age within an inflorescence, it was pos-
sible to trace the darkly staining and proliferating
micropylar integument to embryos in mature
seeds. The seed resulting from adventitious em-
bryony (Fig. 7C) differs from that resulting from
fertilization (Fig. 7A, B) in having a proportion
of the seeds polyembryonic (range 28—95%, mean
71.3%, standard deviation 17, based on 15 flow-
ers representing 15 different southern Florida
populations).

ive capsules from flowers cross-pollinated on

(range 56-94%, mean 73.5%, standard deviation
17.9; range 30-84%, mean 57.2%, standard de-
viation 23.5, respectively).

STUDY OF DRIED MATERIAL

Many more collections and some supporting
fieldwork would be necessary to allow a reliable
assessment of the distribution of different breed-
ing systems throughout the range of S. lanceo-
lata. However, an appraisal of the extent to which
the breeding systems of the southern Florida

que p ible with the ma-

terial aiid:

Agamospermy, as evidenced by polyembryon-
ic seed, was found in all fruiting plants from
Florida and in some plants from Guatemala, Be-
lize (British Honduras), and Bolivia. In some
cases this was associated with non-opening (i.e
cleistapogamous, see Sheviak, 1982) flowers (for
example, district of Peten, Guatemala, 12 June
1933, C. L. Lundell 3896, MICH). The occur-
rence of agamospermy within the main tropical
range is not clearly associated with relatively high
altitude, a situation that involves some of the
same effects as high latitude, and the sample is
not large enough or sufficiently well documented
to reliably reflect a trend.

Relatively small flowers (lateral sepals 14—17
mm long), suggestive of self-pollination, were
characteristic of plants from a number of Carib-
bean islands (for example, Bahamas, St. Vincent,
based on specimens at AMES and FTG). Limited

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Ry

L 0.5mm |

| 0.5mm |

A. Column of Sacoila lanceolata var. lanceolata viewed diy below (left) and diagrams of cross-
rida. — B.

sections (right) based on material from Sarasota Co., Flo

n of Sacoila t var. paludicola

viewed from below and diagram of cross-section (right) of flowers esse sa from Collier Co., Florida. C,-C,,
cross-sections illustrated on the right-hand side of the figure: P, pollinarium; PG, pollen ebrius S, stigmatic

surface; V, viscidium.

fruit set, suggestive of obligate cross-pollination,
was restricted to Costa Rica and northern South
America.

DISCUSSION

amospermy is uncommon in the orchid
y reported only in

and
wari, 1952; Ca ding: 1982), Prasophyllum (Bates,
1984a), Microtus (Bates, 1984b), Paracaleana

(Jones, 1977), Dactylorhiza (Dressler, 1981),
Zygopetalum (Dressler, 1981), and Pterygodium
(Schelpe, 1970). Autogamy is much more prev-
alent than agamospermy in vascular plants gen-
erally and in the orchid family, where it has been
reported in over 60 genera (Catling, unpubl. data).

In S. lanceolata var. lanceolata, it is not known
whether agamospermy is facultative. Although
there is no evidence of seed resulting from cross-
fertilization in terms ofa lower incidence of poly-

1987]

e
0.5 mm

CATLING — BREEDING SYSTEMS OF SACOILA 65

J

» ^ e

Cross-section of column of S. /anceolata var. paludicola, from Collier Co., Florida. This section

IGU ]
represents the position C,—C, in Figure 4B. AC, anther cap; P, pollinarium; PG, pollen germinating; S, stigmatic
surface.

embryony, this may have been the result of cross-
pollinations using genetically identical individ-
uals with some degree of self-incompatibility. Al-
though the crossings involved plants from dif-
ferent localities in southern Florida, it is
conceivable that the different populations had
the same agamospermic origin and are geneti-
cally identical.

Pollination by hummingbirds in S. lanceolata
is to be expected on the basis of various floral
features such as the tubular, horizontally orient-
ed, reddish flowers without internal marking and
lacking odor (Austin, 1975; Pijl & Dodson, 1966),
but no previous observations of hummingbird
pollination have been reported. The only feature
of characteristically bird-pollinated orchid flow-
ers that is lacking is a dark pollinarium (Dressler,
1971), that of S. /anceolata being pale yellow.
Although the observation of hummingbird-pol-
lination reported here involved transplanted and
cultivated plants, there seems to be no reason to

doubt that these observations are indicative of
the pollination mechanism in various other sit-
uations where one or more species of humming-
birds are available.

The suggestion of a lack of natural pollinators
in southern Florida pineland is of interest insofar
as it may provide an explanation for the polli-
nator-independent breeding systems. Although
self. nollination and iated

with pollinator-paucity and colonization of new
territory (for example, Allard, 1965; Jain, 1976;
Lloyd, 1978) the actual absence of pollinators
has not often been well established (for example,
Kevan, 1972); nor has “newness” been quanti-
fied.

In the southern Florida pinelands in May, Fr-
ythrina herbacea and Ipomea microdactyla are
among the few characteristic hummingbird blos-
soms available, and these species were not ob-
served near colonies of var. /anceolata. Only one
species of hummingbird (A. colubris) is present,

66 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FIGURE 6. A, B. Cross-sections of ovules of S. lanceolata var. lanceolata from Sarasota Co., Florida.—C.
Longitudinal section. g — degenerate gametophyte, ii — proliferating inner integument, oi — outer integument.
Material from Sarasota Co., Florida.

and by April and May it has already migrated

localized (Sprunt, 1954), and there is very little
evidence of breeding in southern Florida (Rob-
ertson, 1974). Thus there is relatively little to
attract hummingbirds to the vicinity of many SS.
lanceolata colonies in southern Florida during
the May blooming period, and there is only one
potential hummingbird pollinator, which is ap-
parently not abundant. The var. paludicola also
exists in the absence of a diverse hummingbird-
pollinated flora in southern Florida, although a
few bromeliads exhibiting the syndrome do flow-
er simultaneously in the hammocks. The situa-
tion over much of the range of var. /anceolata is
quite different since many different humming-
bird species are available and there is a diversity
and continuity of hummingbird-pollinated blos-

soms on a year-round basis (Grant & Grant,
1968).
Populations of five other tropical orchids (£ p-

that promotes self-pollination. This is in contrast
to parts of (or all of) the warmer continental neo-
tropics where these same species have a column
structure adapted to cross-pollination, or where
this latter column structure is at least predomi-
nant (Luer, 1971b, 1972; Pijl & Dodson, 1966;
pers. obs.). Pollinators are not clearly lacking in
the case of these species. It is possible that con-
tinual near extinctions due to frost (Singleton,
1936; Stowers & Le Vasseur, 1983), hurricanes
(Craighead & Gilbert, 1962; Alexander, 1967)
and sea-level fluctuations (Fairbridge, 1974;

1987]

CATLING — BREEDING SYSTEMS OF SACOILA 67

FIGURE 7.
C. Seed of S. lanceolata var. lanceolata from Sarasota Co., Flo

Long, 1974) resulted in a strong selection for
colonizing ability, regardless of pollinator-avail-

nator-paucity is considered to provide an ade-
quate explanation for their pollinator-indepen-
dent breeding systems in southern Florida.

LITERATURE CITED

ALEXANDER, T. R. 1967. Effect of hurricane Betsy on
the southeastern Everglades. Florida Acad. Sci
Quart. J. 30: 10-

ALLARD, R. W. 1965. Genetic systems associated with

OHA

species. Pp. 49- 76 in H. G. Baker & G. C. Stebbins

A, B. Seed of Sacoila lanceolata var. paludicola. — A. Collier Co., Florida. — B. Dade Co., Florida. —
rida.

(editors), The Genetics of Colonizing Species. Ac-
ademic Press, New York.

AUSTIN, D. F. 1975. Bird flowers in the eastern United
States. Florida Scientist 38: 1-12

BATES, R. 1984a. Apomixy in four south Australian
species of Prasophyllum R.Br. J. Native Orchid
Soc. South Austr. 8(4): 38-39

1984b. The genus Microtus R.Br. (Orchida-
ceae): a taxonomic revision with notes on biology.
J. Adelaide Bot. Gard. 7(1): 45-89

CATLING, P. M. 1982. Breeding systems of north-
eastern North American Spiranthes (Orchida-
ceae). ae J. Bot. 60(1 2): ador 3039.

. Pollination of northeastern North
Ame erican uem e (Orchi pres Canad. J. Bot.
3

V.C. GILBERT. 1962. The effects

orida.
DRESSLER, R. L. TL Dark pollinia j in humming-

68 ANNALS OF THE MISSOURI BOTANICAL GARDEN

bird-pollinated orchids, or do hummingbirds suf-
fer from strabismus? Amer. Naturalist 105: 80-
83

———, The Orchids, Natural History and
Classification. Harvard Univ. Press, Cambridge,
Massachusetts.

FAIRBRIDGE, R. W. 1974. The holocene sea-level rec-
ord in south Florida. Pp. 223-232 in P. J. Gleason
(editor), Environments of South Florida: Present
and Past. Miami Geological Society Mem

GRANT, K. A. & V. GRANT. 1968. Hummingbirds
and Their Flowers. Columbia Univ. Press, New

GREENW he = 982. Viscidium types in the
ani ou (Mex.) 8(2): 283-310.
HOLMGREN, P UKEN & E. K. SCHOFIELD.
1981. Index Herbariorum, ~ 2 The Herbaria

of the World. Dr. W. Junk, B

JAIN, S. K. 1976. Theevolution "e NER plants.
Annual Rev. Ecol.

JENSEN, W. A. Bo tanical Mini NM Prin-
ciples and Practice. W. H. Freeman and Co., San
Francisco.

1977. Miscellaneous notes on Australian

to synonymy. The Orchadian 5: 126-1 128

KEvAN, P. G. 1972. Insect eta of high arctic
flowers. J. Ecol. ~~ 83

LirLig, R. D. H. J. Fai S Biological Stains.
The Williams and Wilkins Co., Baltimore

Lrovp, D. G. 78. Demographic factors and self-
fertilization in plants. Pp. 67-88 in O. T. Solbrig
(editor), Demography and the Dynamics of Plant
Populations. Blackwell, Oxford

Lona, R. W Ç
southern Florida. Pp. 28—36 in P. J. Gleason (ed-
itor), Environments of South Florida: Present and
Past. Miami Geological Society Mem. 2

Origin of the vascular flora of

[VoL. 74

d C. A. 1971a. A variety of Spiranthes lanceolata
n Florida. Florida Orchidist 14: 19.

1971b. Abnormal development of the

ther—a report of two cases. Florida Orchidist 14:

26-29.

72. The Native Orchids of Florida. New
York Botanical Garden, Bronx, New York
MAHESHWARI, . Polyembryony in angio-
sperms. Palaeo- botanist (Lucknow) 1: 319-329.
McCartney, C. 1985. Orchids of Florida—the or-
chids of Everglades National Park— 1. Amer. Or-
chid Soc. Bull. 54(3): 265-2
ORNDuFF, R. 1969. Reproductive biology in relation
to ipei Taxon 18: 121-133.
A.

OWCZARZAK, 952. Arapid method for mounting
rige BUDE p Technol. 27: 249-251.
Pu, L. C. 1966. Orchid

Flowers: Their Pollination and Evolution. Univ.
of Miami Press, Coral Gables, Florida.
ROBERTSON, R. 1974. The southern Florida
avifauna. Pp. 414-4 52 in P. J. Gleason (editor),
Environments of South dine: Present and Past.
Miami Geological Soc
SCHELPE, E. A. 1970. Seif pollination, aie ain d

African Orchid Society Journal, March: 9- D

SHEVIAK, C. J. 1982. Biosystematic study of the Spi-
ranthes cernua complex. New York State Museum
(Albany) Bull. 448, 73 pp.

SINGLETON, G. 1936. The freeze of 1934. Florida
Acad. Sc. Proc. 1: 23-

SPRUNT, A., JR. 1954. Florida Bird Life. Coward-
McCann Inc. and National Audubon Society, New
Yor

STOWERS, D M., JR. & M. LEVASSEUR. 1983. The
Florida freeze of 13 January 1981: an impact study
of west-central Florida. Florida Scientist 46: 72-
83

FLOWER AND FRUIT BIOLOGY IN SOUTHERN SPANISH
MEDITERRANEAN SHRUBLANDS!

JAVIER HERRERA?

ABSTRACT

Flower and fruit biology was studied in a coastal, southern Spanish serok community composed of

30 plant taxa. Data on breeding systems; rewards offered to vectors; flower, fruit, a
xa in the community have insect- pollinated hermaphroditic
rphology. Dioecious species are relatively well represented

fruiting intensities are reported. Most ta
flowers that are largely unspecialized in mo

nd seed sizes; and

(27% of the total), as are vertebrate-dispersed species (43%). Bagging experiments demonstrated that
apap were required for maximum fruit production, but the existence of ee rent systems

was not t

indicia was investigated, it was found that sprouting taxa

ted. When the relationship between fruiting intensity and the words perform vegetative

had, on average, lower fruit production

than those that were unable to sprout. Low fruit production is Polen a relation to reproductive

allocation trade-offs.

Mediterranean-type vegetation has been the
subject of research for investigators who wish to
h 1 } in OP(YOTra he

ically distant areas with similar environmental
factors (Specht, 1969; Mooney & Dunn, 1970;
Cody & Mooney, 1978). Also, a great emphasis
has been put upon plant development and the
adaptive features of plants in this hia seasonal
climate oe & Parsons, 1973; oney et
al., 1974; Kummerov, 1983). Cares lit-
tle is kapun. however, about other characteris-
ticsin the biology of mediterranean species, such
as their reproductive biology. The paucity of in-
formation is particularly noticeable with respect
to plants living in the Mediterranean region itself
(but see C. M. Herrera, 1981, 1984; Jordano,
1982, 1984, for plant-frugivorous bird relation-
ships, and J. Herrera, 1985, for nectar secretion
patterns in scrub). Some information is available
for mediterranean areas in America, Australia,
and South Africa (for example, Moldenke, 1975;
Specht et al., 1981; Kruger, 1981). But in spite
of this, our present knowledge of the reproduc-
tive biology of mediterranean shrublands is low
compared to our knowledge of tropical (for ex-
ample, Frankie et al., 1974, 1983; Heithaus, 1974;
Bawa, 1979; Bawa & Opler, 1975; Opler et al.,
1980) and temperate plant communities (for ex-
ample, Mosquin, 1971; Kevan, 1972; Pojar, 1974;
Reader, 1977; Primack, 1983).

This paper presents part of a study designed

to investigate the reproductive biology of a
southern Spanish sclerophyllous scrub commu-
nity. Flower and fruit features in a number of
taxa are used to elucidate reproductive patterns.
The relationship between sprouting behavior (the
production of new stems from established Kd
zomes, lignotubers, or burls; James, 1984) an
pollination—reproduction variables is also inves-
tigated. Pollination relationships at the com-
munity level will be dealt with elsewhere (J. Her-
rera, in prep.).

STUDY AREA AND METHODS

This study was conducted in the Donana Bi-
ological Reserve (Donana National Park, Spain).
The reserve is located on the Atlantic coast of
southwestern Spain, in an area with a mediter-
ranean-type climate where the vegetation is com-
posed mainly of mediterranean sclerophyllous
shrublands with some planted pine woods. An-
nual precipitation averages 537 mm. Mean an-
nual temperature is 16°C, January being the cool-
est month (9.8°C) and July the hottest (24.6°C).
Temperatures rarely descend below zero, and
summer drought covers five months on average
(May through September). The soil is sandy, and
maximum elevation above sea level is 3

Basically the vegetation encompasses two types

t
of the underground water table is relatively near

'This study received financial support from the Spanish Comision Asesora de Investigacion Cientifica y
Técnica (CAICYT), through a grant (82/264) to Salvador Talavera (Departamento de Botanica, Facultad de
Biologia, Universidad de iii i ae Dr. P. E. Gibbs for correcting the English, and two reviewers for

helpful comments on the ma

? Departamento de Bout jm de Biologia, 41080 Sevilla, Spain.

ANN. Missouni Bor. GARD. 74: 69—78. 1987.

70 ANNALS OF THE MISSOURI BOTANICAL GARDEN

soil surface, the scrub becomes dominated by
hygrophytic species (heath). Otherwise, a highly
xerophytic scrub vegetation is found (see Allier
et al., 1974; Rivas-Martinez et al., 1980). The
most representative species, both in hygrophytic
and xerophytic scrub, were selected for study giv-
ing a total of 30 taxa. From December 1982
through March 1984 the study plot was visited
weekly, and data on flowering phenology, pol-
lination relationships, and flower-fruit biology
of the species were gathered. Here I report only
on aspects of flower and fruit biology.
Individual plants or branches were marked be-
fore flowering, and their area estimated from
maximum and minimum diameters. Flowers
were counted every week during the time of study,
nd the maximum numbers registered for the
plants of a species were averaged to give an es-
timate of flower production per unit area.
Fresh samples of flowers and fruits of all taxa
were taken to determine the most outstanding
external features, such as dimensions, color, and
sex. Ten to 20 flowers of each species were mea-
sured (length, maximum and minimum diame-
ters), and 20 to 500 complete flowers without
their pedicels were air dried and weighed. The
maximum dimension of a flower was found to
be directly correlated with its dry weight (r, =
0.806, N = 30, P < 0.001). Therefore dry weight
has been used subsequently as an estimate of
flower size, since it is less dependent on the ar-

termined from external features and visitor cen-
suses (J. Herrera, in prep.). Flower forms were
referred to those of Faegri and van der Pijl (1979).

Pollen production and the number of ovules
in the ovary of each flower were determined for
10 to 15 flowers per plant species. Ovules were
counted under a dissecting microscope. Pollen
production was assessed in the same flowers by
macerating one or two anthers in a known vol-
ume of detergent-safranine solution and count-
ing the number of grains in 10 replicates of 5 ul.
Pollen-ovule ratios of hermaphroditic species
were then compared with those given by Cruden
(1977).

To investigate nectar secretion, flowers were
observed in the field and in the laboratory under
a dissecting microscope. In doubtful cases, the
arrival of insect visitors to flowers was precluded
by bagging branches with white nylon mesh;
flowers were examined after 24 hours and, when
possible, the accumulated nectar was quantified.

[VoL. 74

The volume per flower was measured by the
length of the column, in mm, inside 5 ul micro-
pipettes. Concentration, on a weight : weight ba-
sis, was measured in two ATAGO refractometers
(models N1 and N2) compensated to ambient
temperature. The milligrams of nectar sugar plus
diluted solids (Inouye et al., 1980) were com-
puted by the product of volume x concentration
(Bolten et al., 1979)

Percent fruit production in open pollinated
flowers of the species was estimated from flower
and ripe fruit counts on marked branches. In the
same individual plants that were used to estimate
percent fruiting, some branches were also bagged
with white nylon mesh (or with glassine paper
bags in anemophilous taxa) to determine fruiting
intensities when pollinators were excluded. Bag-
ging experiments were not carried out with dioe-
cious taxa. Samples of fruit were collected and
oven dried to a constant weight, and the number
of seeds, along with the weights ofthe whole fruit,
all seeds, and that of an individual seed were
averaged and recorded. Notes on fruit and flower
predation were also taken and, when possible,
the agents responsible were identified.

The ability to perform vegetative regeneration
after complete destruction of aboveground bio-
mass (sprouting) was assessed for every species.
PAS on ou of these Species w were tns
estudy
area on plants damaged by fire, M or
human disturbance during the years 1982 through
1985

RESULTS
FLOWER BIOLOGY

The names of the species studied, together with
their most distinctive floral features, are sum-
marized in Table 1. Our set of mediterranean
plants consists of 30 taxa in 17 families and 25
genera. Most families contribute one or two
species, with only Cistaceae, Leguminosae, La-
biatae, and Ericaceae being relatively well rep-
resented (5, 4, 3, and 3 species, respectively).

Pollination by insect vectors is dominant in
the community, where entomophilous species
account for more than 80% of the total (Table 1).
Only five species s rely on wind for pollen dis-

dioecious species are relatively well represented
(27%). Thymus tomentosus is the only species
with a different sexual condition (gynodioecy).

HERRERA—FLOWER & FRUIT BIOLOGY

1987]

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72 ANNALS OF THE MISSOURI BOTANICAL GARDEN

|
Oo
S. 2^
a e
m. ° š,
> =
se. "Ka
2
c 2r ° e,
> 4 A
S1 á
u.

7 0

FLOWER DRY wees wm
El. Relationship een the weight of an
individual flower and the mean number of flowers per
square meter in peak blo ien plants. Triangles —
sprouting taxa; circles = nonsprouting taxa.

Flower sizes (as dry weight of the complete flow-
er) in the sample range between 2.2 mg for the
female flowers of Pistacia lentiscus and 65.5 mg
for Cistus salvifolius. More than 5096 of the taxa
have flowers of 5 mg or less in dry weight. Floral
dry weight is not independent of the breeding
system in this set of species. If taxa are segregated
in two categories, less or more than 5 mg (a value
that is near the median in the frequency distri-
bution), dioecious taxa predominate in the lower
class (8 out of 9), whereas hermaphroditic taxa
are evenly distributed among the two classes (10
and 11, respectively). The differences are statis-
tically significant (G = 5.04, df = 1, P < 0.025).

The predominant flower morphology is of the
dishbowl type (Faegri & van der Pijl, 1979), which
is present in the Cistaceae and also in all the
entomophilous dioecious taxa. Species in the La-
biatae have short, tubular corollas; while the le-
gumes present typical flag blossoms. The only
long-tubed flowers are those of Lonicera pericly-
menum. Generally speaking, flowers are small
and/or have corollas that impose little or no re-
striction on floral visitors (except for L. pericly-
menum and the legumes). All species in the Le-
guminosae and three in the Cistaceae have yellow
flowers, which is the most common color (nine
of total); whereas six species have white corollas.
Many of these yellow- or white-flowered taxa
offer pollen as the main reward to pollinators
(Table 1). There are five species with pink flow-
ers, which provide predominantly nectar as the
reward.

[VoL. 74

Among entomophilous taxa, pollen and nectar
are offered by approximately the same number
of species (12 and 13, respectively). However,
since wind-pollinated taxa are also occasionally
used as a pollen source by insects (pers. obs.),
this food material is more readily available in
our community than is nectar, at least in terms
of plant taxa. Furthermore, we have succeeded
in quantifying the secretion in only six out of 13
species in which nectar is the main reward. These
species were Daphne gnidium (0.17 mg sugar/
flower per 24 hr.), Erica ciliaris (0.08 mg), La-
vandula stoechas (0.15 mg), Lonicera pericly-
menum (2.00 mg), Rosmarinus officinalis (0.20
mg), and Rubus ulmifolius (1.20 mg). In the re-
maining seven species the yield of nectar sugar
per flower per 24 hr. was too scarce to be quan-
tified with our method (presumably less than 0.08
mg). Maximum concentration values ranged be-
tween 9.5% (Erica ciliaris) and 60% (Rubus ul-
mifoliusy, volumes per flower on a daily basis
ranged between 0.9 ul (Lavandula stoechas,
Daphne gnidium) and 10 ul (Lonicera pericly-

enum).

Pollen yield of the individual flowers is given
in Table 1. The lowest is that of Armeria velutina
(10? grains) and the highest that of Myrtus com-
munis (599 x 10?), which has insect-pollinated,
nectarless flowers of the brush type common in
the Myrtaceae. Since pollen production is not
independent of flower size, a more correct esti-
mate of staminate effort on a per flower basis
would be the number of pollen grains produced
per milligram of flower dry weight (Relative Sta-
minate Effort, RSE). Defined in this way, RSE is
minimum for entomophilous taxa such as Ar-
meria velutina (500) and vain periclymenum
(524). The former has dimorphic pollen grains
and stigmas, and the latter a the most spe-
cialized (long-tubed, highly nectariferous) flowers
in the sample. Maximum values of RSE are found
in anemophilous species such as Corema album
(2.3 x 10°) and Erica scoparia (120 x 10). Mean
RSEi y higher for
than for entomophilous ones (U — 121, N = 5,
25, P < 0.001).

Values of pollen-ovule ratios for hermaphro-
ditic species are given in Table 1. In most taxa
this ratio is greater than 2,000 and thus referable
to the allogamous class of Cruden (1977). Only
three species have pollen-ovule ratios lower than

,000. Mean number of flowers for plants in peak
bloom are also shown in Table 1. A highly sig-
nificant relationship exists between flower num-

1987]

HERRERA—FLOWER & FRUIT BIOLOGY

TABLE 2. Fruit and seed characteristics for 29 scrub species. Numbers in parentheses are sample sizes.

DE. Mean Dry Weight (mg) Pre-
Species Type! of Seeds Whole Fruit All Seeds Individual Seed dation

Armeria velutina D l 3 (50) 1 l (50) E
Asparagus aphyllus F 1.2 46 (100) 23 19 (122) *
Calluna vulgaris D 3.6 (100) 0.2 0.1 (325) =
Chamaerops humilis F l 1,364 (18) 781 781 (18) +
Cistus libanotis D 22.3 59 (20) 23 1 (445) +
Cistus salvifolius D 51 137 (12) 53 1 (612) +

F 3 57 (30) 35 12 (90) =
Cytisus grandiflorus D 6.9 214 (20) 39 6 (137) =
Daphne gnidium F l 18 (75) 8 8 (75) =
Erica ciliaris D 17.9 5 (80) 0.2 0.01 (1,150) +
Erica scopari D 6.5 1 (60) 0.1 0.02 (1,000) +
Halimium commutatum D 2.6 33 (20) 14 (52) +
Halimium halimifolium D 25.7 59 (20) 15 0.6 (514) =
Helianthemum croceum D 10.3 34 (15) 15 1 (155) +
Helichrysum picardii D l 0.6 (750) 0.6 0.6 (750) on
Lavandula stoechas D 2.3 4 (50) l 0.6 (114) +
Lonicera periclymenum F 2.3 52 (30) 15 7 (30) =
Myrtus communis F 5.7 108 (75) 56 10 (430) =
Osyris alba F l 157 (30) 90 90 (30) +
Osyris quadripart F l 131 (75) 73 73 (75) -
Phillyrea angestfola F l 35 (100) 12 12 (100) —
Pistacia lent F l 22 (80) 13 13 (80) =
Rhamnus oem F 1.6 63 (30) 26 16 (30) =
Rosmarinus officinalis D 2.9 4 (20) 2 0.6 (59) +
Rubus ulmifolius F 40.4 213 (30) 87 2 (30)
Smilax asper. F 1.4 78 (100) 51 36 (140) —
2. i genistoides D 2.5 59 (15) 13 5 (38) +
Ulex m D 2.1 22 (20) 7 3 (35) +
Ulex xn an D 2 47 (10) 12 6 (20) +

' D, dry fruits; F, fleshy, vertebrate-ingested fruits.

ber and flower size (dry weight); this relationship
can be easily appreciated in Figure | (r= —0.826,
= 27, P < 0.001).

T of flowers by insect predators, mostly
noctuid larvae, coleopteran larvae, and oth
unidentified insects, was observed in only four
species (Table 1).

O
+

FRUIT BIOLOGY

Table 2 summarizes various characteristics of
the fruits in our set of mediterranean plant species.
Fruit production in Thymus tomentosus was so
sparse that we were unable to gather a reasonable
sample of fruits and seeds; this species has there-
fore been excluded from the analyses. According
to the way in which their seeds are dispersed our
taxa are easily divided in two groups: those whose
seeds are dispersed by vertebrates (13 taxa) and
those whose seeds are dispersed by the wind or
in a largely passive way (16 taxa). The former

have fleshy fruits, while the latter have dry fruits
(mostly capsules, legumes, or achenes). Seed col-
lecting by ants has been observed in a few taxa,
but data are not conclusive enough to recognize
a third (ant- -dispersed) class. Most vertebrate-

number of seeds per fruit ranges between
l and 51, and the fruit size (fruit dry weight)
between 0.6 mg (Helichrysum picardii) and 1,364
mg (Chamaerops humilis). The lightest seeds are
those of Erica ciliaris (2 x 10? mg) and the
heaviest those of Chamaerops humilis (781 mg).
Predation of fruits, mainly by Curculionidae, lar-
vae of Tortricidae, Noctuidae, and parasitic hy-
menopterans, is far more common than flower
predation (55% and 13% of species pectively).
Damage by predators was observed more fre-
quently on dry than on fleshy fruits (G — 4.02,
df = 1, P < 0.05)

74 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE 3. Results of independence tests performed
between the type of fruit (dry or fleshy) and other vari-
ables of flowers and fruits. Class intervals considered
were weight of fruit, less than 20 mg, 20-40 mg, 40-
60 mg, or more than 60 mg; weight of the paneer
seed, less than 10 mg, or more than 10 mg; num
seeds per fruit, less than 2, 2-4, or more than 4.

Independence of

G df P
i (hermaphrodit-
oec 11.72 1 «0.001
Weight ier 804 3 «0.05
Weight of cu seed 23.32 | «0.001
Seeds per fruit 13.88 2 <0.001

G, value of the statistic in the G test of independence;
df, degrees of freedom.

Results of tests for the independence of fruit
type and a few variables of flowers and fruits are
given in Table 3. There is a significant tendency
for species with hermaphroditic flowers to pro-
duce dry fruits, and for species with unisexual
flowers to produce fleshy fruits. The indepen-
dence hypothesis of fruit type and breeding sys-
tem (Bawa, 1980) is thus rejected (Table 3). The
only gynodioecious taxon in the sample has been
included (conservatively) in the dioecious class
for this analysis. The independence hypothesis
for the type of fruit and the variables weight of
fruit, weight of an individual seed, and seeds per
fruit are all rejected. In fact, fleshy fruits tend to
be, on average, heavier than dry ones (U = 185.5,
N = 16, 13, P < 0.05) and to have fewer and
heavier seeds (U = 203, N = 16, 13, P < 0.001).
Note that we are reporting dry weights, so that
differences are not due to the high water content
of fleshy, vertebrate-ingested fruits.

Fruiting intensities for 28 of the studied species
can be seen in Table 4. Data are not available
for two dioecious species, Corema album and
Rhamnus lycioides. Fruit production after exclu-
sion of pollinators is also reported for hermaph-
roditic taxa. Percent of fruit production in open
pollinated flowers ranged between 2% for Thy-
mus tomentosus and Daphne gnidium, and 9296
for Halimium commutatum. Bagging of flowers
in most cases had a clear effect of decreasing fruit
production—to levels as low as 0-196 in 15
species. There was just one taxon that was un-
affected by exclusion of pollinators (Daphne gni-
dium, 296 fruit production under both treat-
ments). Some fruit was produced (10—3196) in
bagged flowers of a few taxa (Calluna vulgaris,

[Vor. 74

Erica scoparia, Myrtus communis, and Rosma-
rinus officinalis), but even in such cases polli-
nating vectors were needed for fruiting to arrive
at its maximum.

REPRODUCTIVE TRADE-OFFS

Figure 2 shows the relationship between per-
cent fruiting and the quotient of the fruit dry
weight to the flower dry weight (fruit : flower size
ratio hereafter), which indicates about how high
the increase in size is from the first reproductive
unit to the second. This variable will be em-
ployed below to investigate some patterns of re-
productive allocation in our set of species. A
highly significant negative relationship exists be-
tween percent fruit production and fruit : flower
size ratio (r = —0.6780, N = 26, P « 0.001, log
transformed data). Thus species in which there
is a great increase in dry weight during the tran-
sition from flower to fruit are those with the low-
est percent fruiting, whereas species in which this
process involves little gain in weight have rela-
tively greater fruiting success.

Species known to perform vegetative regen-
eration (sprouting) are PEPE, from non-
sprouting ones in Table 4, and their respective
distribution in the relationship defined by per-
cent fruit production and fruit: flower size ratio
is Shown in Figure 2. Sprouting taxa tend to have
low values of fruiting and high values of fruit:
flower size ratio. Mean percent fruit production
is not, however, significantly different for sprout
ing and nonsprouting taxa (U = 109, = 14,
12, P > 0.2); differences in mean dry weight
increase from flower to fruit are weakly signifi-
cant (U = 117.5, P > 0.05). Species in the Eri-
caceae lower the coherence of the sprouting group,
since they are well-known sprouters but have
relatively low fruit: flower size ratios. If the Er-
icaceae are removed, the sprouting group is en-
tirely composed of taxa with few-seeded, verte-
brate-ingested fruits; with the exclusion of
ericaceous taxa, the nonsprouting group overlaps
entirely with the dry fruit group, and the sprout-
ing group does likewise with the fleshy fruit group.
Differences in mean percent fruit production and
fruit: flower size ratio are now significant (U =
101.5, P < 0.05; U= 116.5, P < 0.002, N = 12,
11, respectively).

Figure 1 shows an inverse relationship be-
tween the number of flowers at peak blooming
and UNE dry WORK Both sprouting and non-

enly distributed along this

contia so dat i mean number of flowers at

1987] HERRERA— FLOWER & FRUIT BIOLOGY 75

TABLI Fruit production in open pollinated and bagged flowers of the studied species. Numbers in paren-
theses indicate the number of flowers, and N the number of plants. The sample of the only gynodioecious species
(Thymus tomentosus) includes three hermaphrodites and four female plants. Species known to be capable of

sprouting are marked with an asterisk.

Fruit Production (%)

Species Open Pollinated Bagged N
Armeria velutina 59 (478) 3 (476) 5
Asparagus aphyllus* 24 (804) — 6
Calluna vulgaris* 88 (100) 24 (50) 5
Chamaerops humilis* 4 (4,560) — 7
Cistus libanotis 55 (5,822) 0.2 (538) 4
Cistus salvifolius 50 (3,014) 0 (92) 10
Cytisus grandiflorus 12 (150) 0 (134) 4
Daphne gnidium* 2 (23,749) 2 (2,660) 10
Erica ciliaris* 45 (147) 1 (105) 10
Erica scoparia* 89 (85) 31 (65) 5
Halimium commutatum 92 (100) 0 (100) 5
Halimium halimifolium 41 (70) 1 (1,259) 5
Helianthemum croceum — 0 (171) 3
Helichrysum picardii 68 (1,308) 1 (1,443) 5
Lavandula stoechas 69 (200) 0.4 (1,040) 10
Lonicera periclymenum* 8 (1,537) 4 (211) 4
Myrtus communi. 68 (857) 23 (78) 4
Osyris alba 7 (1,385) — 8
Osyris quadripartita* 5 (2,400) — 5
Phillyrea angustifolia* 14 (1,229) 0 (735) 5
Pistacia lentiscu 16 (1,368) — 5
Rosmarinus Pi daa 31 (256) 10 (797) 5
Rubus le 78 (931) 0 (158) 10
Smilax a 9 (4,053) — 5
ADU genistoides 40 (115) 0.2 (651) 4
Ti us tomentosus 2 (210) 0 (210) 7
Ulex minor 16 (238) 0 (221) 10
Ulex parviflorus 5 (310) 0 (471) 5

peak blooming is not significantly different be-
tween them (U — 101, P 0.1); neither is mean
flower dry weight significantly different (U =
113.5, P > 0.1, N = 14, 13). Both sprouting and
nonsprouting taxa may be many- and small- or
few- and big-flowered.

DISCUSSION

The 30 mediterranean plant species studied
represent a relatively small sample ofthe regional
flora (more than 2,000 taxa for southern Spain,
of which nearly 300 are woody). Furthermore,
shrublands have many different and diverse
species compositions in southern Spain, depend-
ing on elevation, rainfall, edaphic factors, etc.,
so that the results reported here should be ex-
tended only with caution to other scrub com-
munities in the region. Nevertheless many ofthe

studied taxa are widespread, and the community
they form is undoubtedly a clear example of
coastal scrub on sandy soils, which is character-
istic of other areas in southern Spain.

In the studied community a sizeable hetero-
geneity in reproductive traits was likely to occur,
since 30 plant species were distributed among 17
families. However, certain groups (virtually Cis-

did others. Hence a phylogenetic component in
the reproductive patterns recognized should not
be ruled out, in addition to an ecological com-
ponent and to the fact that the plants form a
steady, long-lasting community achieving repro-
duction year after year. Several aspects in the
reproductive patterns are not restricted to our
particular community. The relative dominance
of taxa in which nectar yield is low or even zero,

76 ANNALS OF THE MISSOURI BOTANICAL GARDEN
100r
aa °.
Ë ^
s ° ° a
= °
t SOF p’
— ° .
3 e
u T ô
° A A
Eom, ur A,
2

0 1
FRUIT WEIGHT: FLOWER WEIGHT(log)
URE 2. Relationship between fruit: flower dry

species; black triangles = species in the family Erica-
e.

for example, is general in the region (J. Herrera,
1985). The consequences of this factor upon the
i gena relationships will be dealt with else-

ere (J. Herrera, in prep.). Dispersal by inan-
imate or vertebrate agents creates a dichotomy
that is also found in other mediterranean-type
communities (Bullock, 1978, chaparral; C. M.
Herrera, 1984b; Jordano, 1984, southern Spain).

nities (Reader, 1977; Ruiz-Zapata & Arroy
1978; Bawa, 1979; Bawa & Beach, 1983) un
nators are needed for maximum fruit produc-
tion. Further investigations will determine the
extent to which incompatibility systems occur in
shrublands of southern Spain.

Sprouting is also a common regenerative strat-
egy of woody plants in many and diverse eco-
systems (James, 1984). It is particularly impor-
tant in regions of a mediterranean-type climate
that experience summer droughts and frequent
fires. This aspect has received much attention in

of a continuum) have been
recognized: the “seeder” strategy consists of ob-
ligate seed reproduction and incapacity to form
new stem sprouts after destruction of aerial parts;
the *sprouters" can produce stem or root sprouts
that enable repeated shoot production despite
frequent damage (Malanson & Westman, 1985;
see James, 1984, for an extensive review). It ap-
pears that both strategies also are found among
sclerophyllous species of the Mediterranean Ba-

[Vor. 74

sin. In our community most plants fall clearly
into one category or the other, although species
in the Ericaceae could be easily included in both:
they produce plenty of seeds and are also able to
sprout, which was reported some time ago for
Calluna vulgaris (Kayll & Gimingham, 1965).
It has been hypothesized that pollination-re-
production variables may be associated with each
of the sprouting or nonsprouting strategies. In
seeders, for example, the pollination system must
be efficient enough to assure the release of nu-
merous seeds (which could open a way to self-
compatibility), produce numerous flowers highly
attractive to pollinators, and offer a greater re-
ward per flower than sprouters (Carpenter &
Recher, 1979). In contrast, sprouting taxa are
proposed to have a high rate of outcrossing, al-
though often with a low production of flowers,
which would be relatively low rewarding (Fulton
& Carpenter, 1979; Carpenter & Recher, 1979).
"ee hypotheses are supported in part by our
dat

^ high rate of outcrossing has to be expected
in the sprouting group of species, since many
dioecious taxa are included in that group. There
are no evident signs, however, of a high incidence
of self-pollination in the nonsprouting group. In
contrast, mean percent fruit production was sig-
nificantly higher for nonsprouters than for
sprouting species, which the hypothesis
outlined above. Low pollination efficiency due
to unisexuality can be reasonably proposed to
explain low fruit production in the sprouting
group, but we wish to put forward a comple-
mentary view. Percent of fruit production was
found to be inversely related to fruit : flower size
ratio (i.e., the increase in mass needed to produce
a fruit from a flower). Such a relationship sug-
gests the existence of a trade-off between the en-
ergy that a plant allocates to an individual fruit
or seed and the number it can successfully form.
Low fruit production in vertebrate-dispersed, big-
seeded species may reflect low pollination suc-
cess and/or the impossibility of developing every
fruit with pollinated ovules, due to the relatively
igh costs involved in the ripening process. It
has been demonstrated, for example, that some
species experience low fruit production despite
good levels of pollination, which is apparently
due to intrinsic regulatory mechanisms (Lloyd,
1980; Wyatt, 1981; Casper & Wiens, 1981; Cas-
per, 1983; Bookman, 1983; Bawa & Webb, 1984;
see Stephenson, 1981, for a review). Flower size
and number showed a negative relationship too,

1987]

but the predicted tendency for sprouting taxa to
appear at one end of this continuum has not been
detected by

Wells (1969) pointed out that the capacity to
sprout vegetatively from underground parts is
probably an ancestral trait. James (1984), how-
ever, suggested that sprouting cannot always be
seen as an ancestral trait in woody dicotyledons.
In the present study the capacity to sprout is
associated with other traits, such as the produc-
tion of fleshy, vertebrate-ingested fruits, heavy
seeds, and low fruit production, along with a
relatively high incidence of dioecy (species in the
genera Asparagus, Chamaerops, Lonicera, Osy-
ris, Pistacia, Rhamnus and Smilax, for exam-
ple). This group of taxa has subtropical affinities:
sclerophyllous species in these genera existed well
before the Pleistocene and the evolution of true
mediterranean climatic conditions (Raven, 1973;
Axelrod, 1975; Pons, 1981). In contrast, species
in the genera Armeria, Cistus, Cytisus, Hali-
mium, Lavandula, Rosmarinus, or Thymus are
nonsprouters; and the high number of taxa en-
demic to the Mediterranean Basin makes clear
that species in these genera originated much more
recently (Quezel, 1978, 1981; Pons, 1981). The
capacity to sprout is thus lacking in the typically
mediterranean taxa but is jiu in the more
mediterranean ones. This supports
s (1969) iat pene se is an
ancestral trait, a it may indicate that only those
"tertiary flora" subtropical sclerophyllous taxa
with a capacity to sprout were able to survive
the shift to seasonal dryness associated with the
mediterranean climate.

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REPRODUCTIVE SYSTEMS AND CHROMOSOME RACES OF
OXALIS PES-CAPRAE L. AND THEIR BEARING
ON THE GENESIS OF A NOXIOUS WEED!

ROBERT ORNDUFF?

ABSTRACT

In its native South Africa, Oxalis pes-caprae is represented by diploid and tetraploid races; the
short-styled,
d So uth

ual proportions in natural populations. The diploid and tetraploid races of the species have a well-
developed incompatibility system associated with their floral trimorphism. Outside South Africa, the

is there evi idence of any aggressive superiority of the short- styled morph i in the

South

Africa. W

h Afri
asa eed dn the diploids and tetraploids within South Africa, is unknow

Oxalis pes-caprae L. is a troublesome and
widespread agricultural and garden weed, par-
ticularly in areas of the world with a mediter-
ranean climate such as central Chile, the Medi-
terranean basin, parts of Australia, and California
(Salter, 1944; Young, 1958; Munz, 1959; Mi-
chael, 1964). The species is native to southern
Africa, where it is variable (Salter, 1944), and
where it is distributed from Namibia (South West
Africa) southward to the Cape region and around
the Indian Ocean coast at least as far north as
the Knysna area, sometimes ranging well inland.
There it occurs as a “well-behaved” native of
relatively undisturbed sites as well as a weed,
and it is particularly common in vineyards and
along roadsides. In southern Africa, the species
is tristylous (Fig. 1), but throughout most of its
exotic range it is represented by a short-styled
form, which is pentaploid (25 — 35), and which
reproduces asexually via bulbils. In parts of its
exotic range, tetraploid and presumably sexual
populations also occur, though less commonly
than the sterile pentaploid.

In view of the importance of this species as a
weed, certain features of the reproductive biology

es
rica, but apparent less successful

and chromosome cytology of plants originating
in South Africa were studied in the hopes of elu-
cidating the events that led to the origin of the
aggressive weedy races from the native races.

MATERIALS AND METHODS

Bulbs collected from natural populations of
Oxalis pes-caprae in the Cape Province region
in South Africa in 1970 and 1971 were later grown
at Berkeley for chromosome studies. One col-
lection was provided by Sherwin Carlquist.
Chromosome numbers were determined by ex-
amining microsporogenesis in flower buds of
these cultivated plants preserved in 3: 1 ethanol:
acetic acid and stained in acetocarmine. Bulbs
collected by Peter Goldblatt from two localities
in 1984 provided plants used in an artificial
crossing program to determine the presence and

and a vineyard at Rustenberg, near Stellenbosch,
Cape Province. The two localities are ca. 50 km
from each other. Crosses were made in the spring
of 1985 and 1986 by transferring pollen from

! Supported in part by grants from the National eir Foundation, the Council for Scientific and Industrial
ke

Research of South Africa, and the Committee on R

Carlquist, T. I. Chuang, Rivka cea = Peter Goldblatt
n, and Patricia Watters for assisting this a in various
0.

Kim E. Steiner, David E. Sym

rch, University = lifornia, Berkeley. I thank Sherwin

, R. Harry Koga, Peter W. Mic = se Donald Pfister,

? Department of Botany, unity of California, Berkeley, California 9472

ANN. MIssouRI Bor. GARD. 74: 79-84. 1987.

80 ANNALS OF THE MISSOURI BOTANICAL GARDEN

aie

5 mm

Mid

Long

FIGURE 1.
ignation of each floral form
the text are given to the iw of the figure.

dehisced anthers to stigmas under insect-free
conditions. Seed set was determined by collect-
ing nearly mature capsules and counting the seeds
released by them in seed packets. Pollen size for
the three morphs was determined by mounting
i grains in aniline blue-lactophenol and mea-

ring them with an ocular micrometer; pollen
stalna bility was determined by counting the
number of stained grains in a sample of 100
mounted in this medium

RESULTS

Chromosome numbers. Seven ofthe nine na-
tive populations examined were tetraploid (n =
14); two were diploid (n = 7; Table 1). Previously
published reports, summarized in Table 2, in-
dicate only tetraploidy and pentaploidy for the
species.

Weediness. The two diploid populations and
four of the tetraploid populations were growing
under disturbed conditions such as roadsides,
vineyards, or grainfields and were considered
weedy; the others occurred in undisturbed con-

ed (field data
MIURI Udta

ditions and
are lacking for one tetraploid Bauen Table

1).
Morph ratios. Morph ratios have been re-
ported earlier (Ornduff, 1974). In two popula-

[VoL. 74
long -level
mid-level

l £ short-level
Short

Illustrations d gynoecia and androecia of the three floral forms of Oxalis pes-caprae. The des-
s given below each illustration. The stigma and anther level designations used in

tions, morph ratios of Longs : Mids: Shorts were
1:1: 1, both populations were considered, with
some doubt, to be non-weedy. In four popula-
tions, morph ratios deviated from equality. In
one of these, Longs were deficient; in another,
Mids; in third, Shorts; and in the fourth, Longs
and Mids were greatly outnumbered by Shorts.
Three of these four populations were character-
ized as weedy (two for which chromosome counts
are available were tetraploid).

Pollen size and stainability. Pollen size of
diploids and tetraploids is trimorphic (Table 1).
Pollen from the long-level anthers is largest, that
from mid-level anthers is smaller, and that from
short-level anthers is smallest. Pollen from an-
thers at equivalent levels in different morphs was
generally of di same size duin a population,
but there were i pol-
len size from equivalent anther sets. There was
no correlation between differences in pollen size
and in chromosome number of the diploids and
tetraploids. Pollen from presumed pentaploids
collected in California was extremely variable in
size, even from an individual anther. Pollen
stainability of diploids and tetraploids was vari-

able, but was mostly over 6096. One Short tet-
d (7038, Table 1) had pollen with 1296 and
25% stainability from its two sets of anthers. One

lege Ave., Berkeley

1987] ORNDUFF—ONXALIS PES-CAPRAE
TABLE l. Chromosome numbers, pollen size (um), and pollen stainability in Oxalis pes-caprae.
Origin of Pollen
ollection (authors Chrome: (size; standard deviation; stainability in percent)
or collector's field some Floral Long-level Mid-level Short-level
umb Number Morph thers Anthers Anthers
Cape Province, South Africa:

7035: Worcester, river- n= 14 Long 41.7; 1.83 (—) 35.8; 2.03 (—)
bank, non-weedy Mid 45.7; 2.65 (—) 33.0; 2.05 (—)

7038: Klapmuts, road- n = 14 Short 54.1: 3.94 (12%) 41.2; 3.10 (25%)
side, weedy

7041: Stellenbosch, — Short 49.5; 3.20 (66%) 41.8; 1.83 (69%)

vineyard, weedy

7096: Mamre Road Sta- n=7 Mid 46.4; 1.95 (99%) 33.7; 1.91 (98%)
tion, roadside, weedy

7248: Gouda/Hermon, n] Mid 42.7; 3.94 (—) 31.1; 2.20 (—)
roadside, weedy

7292: KL on yman — Long 41.8; 1.83 (8990) 42.7; 4.84 (88%)

ry, field, non- Mid 50.1; 2.38 (86%) 38.9; 1.83 (7890)
eed Short — 50.4; 4.15 (93%) 41.5; 2.93 (98%)

7301: — n" n = 14 Long 39.0; 1.83 (78%) 30.3; 3.92 (79%)

baan, field, non- Mid 46.2; 1.68 (83%) 31.2; 1.54 (81%)
weedy Short 48.3; 5.73 (81%) 34.4; 2.03 (77%)

8053: Calvinia, grain n=14 Long 42.6; 2.42 (65%) 36.2; 2.52 (60%)

field, weedy id 52.2; 2.67 (64%) 35.0; 1.81 (40%)
Short — 47.2: 2.75 (74%) 38.0; 2.55 (73%)

8055: Clanwilliam, — Long 38.2; 2.49 (75%) 32.6; 2.07 (68%)

veld, non-weedy Mid 45.1; 2.89 (94%) 32.6; 2.30 (86%)
Short 48.3; 2.87 (88%) 38.8; 2.83 (82%)

Goldblatt s.n.: Rustenberg, n = 14 Long (90%) —-;— (84%)
near Stellenbosch, Mid —;— (6696) —;— (62%)
abandoned farmland Short — — (70%) —;— (8196)

Goldblatt s.n.: Noordhoek, n = 14 Long —;— (93%) —;— (88%)
Cape Peninsula Mid —;— (81%) —;— (79%)

Short —;— (88%) -— (7896)

Carlquist s.n.: South- n= 14 Long 40.2; 2.46 (80%) 32.0; 1.93 (46%)
western Cape Prov- Mid 50.8; 4.84 (92%) 34.5; 2.01 (96%)
ince Short 49.7; 4.37 (67%) 41.8; 3.71 (63%)

California, U.S.A.:

Ornduff s.n.: Univ. — Short —;— (62%) —;— (87%)
California campus, — Short —;— (58%) —;— (51%)
Berkeley

R. Dulberger s.n.: as — Mid =; — (49%) —;— (5896)
above

R. Dulberger s.n.: Col- — Mid —;— (32%) —;— (29%)

set ofanthers ofa few other collections (e.g., 8053
Mid, Carlquist s.n. Long) had pollen with low
stainability. The two collections used in the
crossing program had pollen stainabilities ex-
ceeding 62%. Presumed Short pentaploids col-
lected in California had pollen stainabilities

ranging from 29% to 87%. However, these pollen
grains were variable in size, and the total number
per anther was reduced compared with pollen
production of diploids and tetraploids as esti-
mated from the density of the pollen grains on
the prepared slides.

82 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

TABLE 2. Published chromosome counts of Oxalis pes-caprae.

Chromosome Number Locality of Population Examined Reference
2n = 28 Cape Town, South Africa Marks, 1956
India (garden plants) Mathew, 1958
Madeira Borgen, 1974
South Australia Oram in Symon, 1961
Western Australia Michael, 1964
2n = 34 India (Punjab) Bir and Sidhu, 1978, 1979, 1980
Sidhu, 1979
2n = ca. 35 Italy Vignoli, 1935, 1937
2n = 35 Unknown

South Australia
Western Australia

Cape Town, South Africa

Franklin in Michael, 1964
Franklin in Michael, 1964

Compatibility relationships. Table 3 presents

ent plants from Rustenberg and Noordhoek. Le-
gitimate pollinations are those between anthers
and stigmas at equivalent levels; all other pol-
linations are termed illegitimate (Darwin, 1877).
Only 4% of the illegitimately pollinated flowers
produced capsules compared with 87% of the
legitimately pollinated flowers. The average
number of seeds per FEE eine jus
legitimate crosses was 15.9; fi t
it was 0.2. Self-pollinations dud 0-0.2 ied
per pollination. Intermorph, illegitimate polli-
nations produced 0—0.5 seeds per pollination. In-
termorph, legitimate pollinations produced 13.0-
20.9 seeds per pollination. The two populations
used in the crossing program produced similar
results for all classes of crosses. Seed production
of Shorts following legitimate pollinations was
slightly greater than that ofthe other two morphs.
oth populations are tetraploid (Table 1).

DISCUSSION

In its native range in South Africa, Oxalis pes-
caprae is represented by indigenous weedy and
non-weedy races with a conventional tristylous
floral morphology, including pollen-size tri-
morphism, and a fully pia trimorphic in-
compatibility system. viduals are strongly
self-incompatible and oaa cross-polli-
nations produce little or no seed. Legitimate pol-
linations produce considerable quantities of seed.
The pentaploid form has been reported from that
country as well (Michael, 1964), but it is unclear
how common this race is there.

In South Africa, tetraploids are apparently more
common than diploids, but weedy populations

of both are known. Unequal morph ratios may
be more common in weedy South African races
than in non-weedy ones and are likely a conse-
quence of vegetative propagation that is en-
hanced by physical disturbances of the habitat
during agricultural or road-building activities.
Morph ratios were found in which Longs, Mids,
and Shorts, respectively, were deficient in num-
bers. Each unequal morph ratio differed from the
others, with no morph(s) predominating overall.
Thus there is no basis from observation of native
races that accounts for the fact that the aggressive
morph outside South Africa is short-styled.

Over most of its exotic range of distribution,
Oxalis pes-caprae appears to be represented by
a fairly sterile, pentaploid short-styled morph.
As early as 1887, Hildebrand noted the preva-
lence of this short-styled form and its lack of seed
set. Henslow (1891, 1910) also noted these fea-
tures and described in some detail the means of
vegetative reproduction later amplified by Galil
(1968). Where introduced, the species is distrib-
uted by human agents and, in places, by other
animals such as the mole-rat (Galil, 1967) or by
birds (Young, 1958).

Despite the high level of pollen sterility of the
short-styled pentaploid, it apparently reproduces
occasionally by seed. This may result from self-
or geitonogamous pollinations in populations
where Shorts alone are represented. Illegitimate
pollinations of sexual Shorts carried out in the
present study produced small amounts of seed,
which offers support for this suggestion. Vignoli
(1937), in an embryological study of the species,
noted rare sexual reproduction in the pentaploid
but concluded also that apogamy may rarely oc-
cur in this race. Another line of evidence for

1987]

TABLE 3.

ORNDUFF=— OXALIS PES-CAPRAE 83

Results of legitimate and illegitimate pollinations in two populations of Oxalis pes-caprae. The

first figures summarize results for the Rustenberg population; the second figures summarize results for the

Noordhoek populatio

Number of Average
Style Length (9 parent) x Number of Flowers Number
nther Level/Style Length Flowers Producing umber o of See
rent)! Pollinated Capsules Seeds Obtained Pollination
Self-pollinations (all illegitimate):
L x m/L selfed 18; 55 0; 6 0; 34 0; 0.6
L x s/L selfed 20; 12 0; 0 0; 0 0; 0
M x I/M selfed 38; 33 0; 1 0; 5 0; 0.2
M x s/M selfed 23; 35 0; 0 0; 0 0; 0
S x VS selfed 18; 20 1; 0 1; 0 0.1; 0
S x m/S selfed 15; 12 0; 0 0; 0 0; 0
Intermorph, illegitimate pollinations:
L x s/M 26; 20 0; 0 0; 0 0; 0
L x m/S 23; 13 1; 2 10; 7 0.4; 0.5
M x s/L 25; 29 0; 2 0; 8 0; 0.3
M x I/S 15; 26 0; 1 0; 1 0; 0
S x l/M 15; 21 0; 3 0; 9 0; 0.4
S x m/L 25; 30 1;4 17; 6 0.7; 0.2
Intermorph, legitimate pollinations:
x l/M 18; 48 12; 41 279; 738 15.5; 15.4
L x I/S 27; 34 23; 30 430; 583 15.9; 17.2
M x m/L 33; 37 29; 38 413; 533 12.5; 14.4
M x m/S 54; 42 46; 34 797; 546 14.8; 13.0
S x s/L 13; 22 11; 20 222; 460 17.1; 20.9
S x s/M 11; 29 11; 25 255; 588 23.2; 20.1
Summary of all legitimate, illegitimate pollinations:
Legitimate 156; 212 132; 188 2,396; 3,448 15.4; 16.3
Illegitimate 261; 306 3; 19 28; 70 0.1; 0.2

tation is as follows: “L x m/L selfed” means a long- styled flower (L) was self-pollinated with its own

s from the mid-level (m) set of anthers. ‘

means a long-styled flower was pollinated with pollen

from the short-level stamens of a mid-styled flower. Figure l illustrates the three flower types

occasional sexual reproduction of this form comes

rwi e
southern Italy by Vignoli (1935) and in central
California by Dulberger (pers. comm.). It also
has been suggested that in Western Australia,
where the pentaploid sometimes exists in mixed
populations with tetraploids, hybridization be-
tween the two may occur (Michael, 1964). The
variability of the species there would suggest fre-

1961) and

and Madeira (Mathew, 1958; Borgen, 1974), but

it is unknown whether these are composed of
more than one morph.

Although the occurrence of tetraploids and

pentaploids in Australia as a consequence of in-
dependent introductions is documented (Mi-
chael, 1964), it is possible to explain the origin
of tetraploids within pentaploid populations by
another mechanism. The pentaploids proque
some viable pollen (Table i Vignoli,
& Sidhu, 1980; Michael, 1964). The illustrations
and discussion of microsporogenesis in penta-
ploids by Vignoli (1935, 1937) indicate that pol-
len grains with n = 14 can be produced by such
plants. If megasporogenesis also results in eggs
with 71 — 14, fertilization of these eggs by diploid
sperm would result in a tetraploid zygote.

The place and mode of origin of the weedy

84 ANNALS OF THE MISSOURI BOTANICAL GARDEN

pentaploid race are uncertain. Lower (1963) sug-
gested that it may have originated outside South
Africa. A 5x chromosome count reported by Mi-
chael (1964) indicates that this race is present in
South Africa, but it is apparently not common
(although this remains to be documented). The
origin of the weedy pentaploid race can only ten-
tatively be reconstructed, but I believe that it
likely occurred in South Africa. The simplest ex-
planation for the origin of pentaploidy is that it
resulted from the union of an unreduced gamete
of a tetraploid plant with a haploid gamete from
a diploid plant. Since diploids are unknown out-
side South Africa, this event must be assumed
to have occurred in South Africa. Although dip-
loids and tetraploids are not known to be sym-
patric in South Africa, they have been collected
very near each other, so sympatry may occur.
Weediness clearly preceded the occurrence of
pentaploidy since sexual diploids and tetraploids
are frequently aggressively weedy: they com-
monly occur in cultivated fields and along road-
sides. Pentaploidy itself has not conferred weedi-
ness on the species. Likewise, prodigious means
of vegetative reproduction occur in diploids and
tetraploids with a fully developed sexual appa-
ratus, so that the almost exclusively vegetative
propagation characteristic of the pentaploid is a
condition that likewise preceded the origin of
pentaploidy.

Present evidence, although scanty, suggests that
the largely sterile pentaploid race is less success-
ful as a weed in South Africa than are the sexual
diploids and tetraploids. Outside South Africa,
however, the pentaploid seems to prevail, pos-
sibly as a consequence of its greater competitive
success Over sexual races under exotic conditions
(as, for example, seems to be the case i in Austra-
lia) or
of introduction that led to this race being more
abundant than its apparent diploid and tetra-
ploid precursors. There is a possibility that there
are competitive differences among the chromo-
somal races of Oxalis pes-caprae, or among its
morphs (as suggested ae io ns Tue re-

11

suggests the possibility That: in these, the sexual
apparatus may be impaired and that asexual
mechanisms are more important in their repro-
ductive mode. Both suggestions merit study.
Nevertheless, the sequence of events leading to
the origin of pentaploidy and the routes of hu-
man-aided migration and introduction of this
species to other continents will probably never
be fully reconstructed.

[VoL. 74

LITERATURE CITED

Bir, S. S. & M. SipHu. 1978. In IOPB chromosome
number reports LX. Taxon 27: 223-231.

1979. Cytological observations in weed flora
of orchards of Patiala TA Punjab. Recent Res.
Pl. Sci. (New gs li) 7 1-271.

jon aul studies
on weed flora k cultivable lands of Patiala district,
Punjab. J. Palynology 16: 85-

BORGEN, L. romosome nu mbers of Maca-
ronesian flowering plants II. Norwegian J. Bot. 21:
195-210.
DARWIN, C. The Different Forms of Flowers
on Plants of the Same Species. John Murray, Lon-
on.
Gaui, J. 1967. On the dispersal of the bulbs of

Oxalis cernua Thunb. by rare (Spalax eh-

bii Nehring). J. Ecol. 55: 787-7

968. Miren dispersal in Oxalis cernua.
m J. Bot. 55: 68-7

HrENsLOW, G. 1891. On the northern distribution of
Oxalis cernua Thunb. Proc. Linnean Soc. Lond.
1890-1891: 31-36.

1910. Remarkable instances of plant disper-

2-361.

eh oy. Hort. Soc
Mes BRAN Experimente über die ge-
aie ake Fortpflanzungsweise der Oxalisar-
ten. Bot. Zeit —6; 33-40.
in H. F. rt on Oxalis pes-caprae L.

963.
n South Africa. Unpublished document supplied
by Waite Institute, Glen Osmond, South Austra-

lia.

Marks, G. E. 1956. Chromosome numbers in the
genus Oxalis. New Phytol. 55: 120-129.

MATHEW, P. M. 1958. Cytology of Oxalidaceae. Cy-
tologia 23: 200-

MICHAEL, P. W. 196 e identity and origin of

ifornia Press, Berkeley and Los Angeles.
ORNDUFF, R. 1974. Heterostyly in South African
plants: a conspectus. J. South African Bot. 40: 169-
187.

ios J. R. 1973. Soursob (Oxalis pes-caprae L.)
n We stern Australia: its life history, distribution,

Distributional and Cyt tological
Flora of Cultivable Fields of

. E. The species of Oxalis established
n South Australia. Trans. Royal Soc. S. Aus
-71.

TN L. 1935. Ricerche preliminari di citologia
sull'Oxalis cernua Thunb. Nuovo Giorn. Bot. Ital.

42: 668-669.

— P. 1937. Fenomeni reproduttivi di Oxalis cer-
nua Thunb. Lavori Inst. Bot. Palermo 8: 5-30.

Y AMASHITA, K. 1935. Zytologische Studien an Oxalis

I. Jap. J. Genet. 11: 36.

Young, D. P. 1958. Oxalis in the British Isles. Wat-
sonia 4: 51—69.

FLORA OF THE VENEZUELAN GUAYANA — II

JULIAN A. STEYERMARK!

ABSTRACT

A total of 18 species, four subspecies, and one variety are newly described from the Venezuelan
a

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phologically closely d but —— ape yos subspecies isolated on different sandstone
descri

table mountains. A key to the subspecies

provided, as well as a ie to the BE acted species s of Bonnet

BROMELIACEAE

Brocchinia oliva-estevae Steyerm. & Lyman B.
Smith, sp. nov. TYPE: VENEZUELA. Bolivar:
summit of Auyan-tepui, extreme north end
above Angel Falls, Dec. 1984, Francisco
Oliva Esteva s.n. (holotype, VEN). Figure 1.

lanta parva caulescens florifera 3.7 dm alta, caule

s, nervis paullo
obscuris; scapo folioso, scapi bracteis lanceolato-ligu-
latuis 6-8 cm longis, 1.4—1.7 cm latis; VENE ag
erecta laxe bipinnatim Mr 2.1 dm longa tenui,
arsim brunneo-furfuracea xem. axibus
c

pedicellis 1—
2: pro ngis minute furfuraceis; petalis albidis lanceo-
latis pesce al Med res tuse € acutis haud Baguicutatis 3

m longis, longis
0.5 mm in vario inferiore aban oeda
3 mm longo; ovulis aes e ia appendicibus cau-
datis

Caulescent, small, herbaceous plant, flowering
3.7 dm tall; stem erect, 13 cm tall. Leaves as-
cending, pale dull green both sides with about
10 parallel, slightly darker longitudinal lines be-
coming bronzy where entering sheath, submem-
branous, flexible, concave above, convex below,

ibed Sauvagesia marahuacensis is
tia.

not contracted at base, the larger ones ligulate-
lanceolate, acute, mucronate, 10 cm long, 1.7-
1.8 cm wide, lower leaves ovate, acuminate, 1.5—
2.8 cm long, 1-1.4 cm wide, the upper ones lon-
ger than the internodes, minutely pale lepidote.
Scape foliose, scape bracts lanceolate-ligulate, 6—
8 cm long, 1.4-1.7 cm wide. Inflorescence erect,
laxly bipinnately paniculate, 2.1 dm long; rachis
slender, covered with a scattered brown furfura-
ceous indument, with 5 short, racemose, lateral
axes 3.5-6 cm long, each axis 15-17-flowered,
the uppermost part ofthe inflorescence elongated
to 10 cm; primary bracts subtending the lower
three axes ascending, lanceolate, acute, 3 cm long,
0.8 cm wide, entire; floral bracts oe ah acute
to acuminate, 3—3.5 mm long, 0.7 wide at
the base. Flowers shortly pedicellate, E l-
2 mm long, Medie Eco = pale
een, lanceolate, acute, 3—4. ong, | mm
wide, dorsally die -DTE in lower half.

nguiculate,

Anthers suborbicular-oblong, 0.5 mm long, ba-
sally bilobed; filaments white, 1.2 mm long.

vary inferior, pale green, subclavate-cylindric,
3 mm long, 1 mm wide at summit, 0.7 mm wide
at base, pale brown furfuraceous; ovules caudate-
appendaged at both ends.

This species of Brocchinia is characterized by
its bipinnate inflorescence with lepidote, simple,

' Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299,

ANN. MissouRi Bor. GARD. 74: 85-116. 1987.

FIGURE |.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Brocchinia oliva-estevae.— A. Habit.—B. Flower with bract.

[VoL. 74

1987]

racemose axes, primary bracts extending from !⁄4
to 4 the length of the rachis, shortly pedicellate
flowers, non-unguiculate petals, and narrow sub-
membranous, slightly nerved, short leaves only
10 cm long by 1.8 cm wide, which are not con-
tracted toward the base. It is most closely related
to B. cowanii L. B. Smith of Cerro Moriche, Terr.
Fed. Amazonas, Venezuela, in having a bipin-
nate inflorescence with non-unguiculate petals,
but differs from that taxon in the scape bracts
shorter than the internodes and in the shorter
sepals and petals.

It is a pleasure to name this interesting species
for Mr. Francisco Oliva Esteva, Venezuelan
landscape architect, an avid student of Brome-
liaceae, and author of several books on orna-
mental plants of Venezuela.

PIPERACEAE

Peperomia marahuacensis Steyerm., sp. nov.
TYPE: VENEZUELA. Territorio Federal Ama-

65°23'W, 2,520-2,620 m, 26-27 Feb. 1985,
Julian A. Steyermark & Bruce Holst
130742-A (holotype, MO; isotype, VEN).
Figure 2C

Herba effusa; caulibus elongatis 8 dm longis, 2.5-3
scentes ramorum

spicuis nervis duobus lateralibus inconspicuis, supra
nervis principalibus pilosulis, demum glabris, subtus
nervis sparsim pilosulis pilis patentibus 0.2-0.5 mm
longis ceterum glabris vel glabratis, marginibus parte
suprema '4-!6 ciliolatis ceterum glabris; petiolis 5-20
mm longis, laminis 2—3-plo brevioribus pilosis.

Sprawling herbaceous plant with stems elon-
gated to 8 dm, 2-3 mm diam., sparsely puber-
ulous except at the densely pubescent younger
tips. Leaves alternate, orbicular- to triangular-
ovate, acuminate at apex, truncate or broadly
rounded at base, larger blades 3-4.5 cm

them conspicuous and 2 lateat nerves Tant, Lal
brouson theu

the main nerves, de surface glabrous or da
brate, but the nerves sparsely pilosulous with
spreading hairs 0.2-0.5 mm long; leaf margins
ciliolate in the uppermost '4-!5, elsewhere gla-
brous; petioles 5-20 mm long, '4—'2 length of leaf
blade, pilose.

STEYERMARK-— VENEZUELAN GUAYANA 87

From the related P. foveolata Steyerm. of Ce-

cent along the nerves of the upper and lower
surfaces, and in the shorter, pubescent petioles,
which are proportionately shorter in relation to
the length of the leaf blade. Compared with P.
peltoidea Kunth, it differs in the non-peltate, non-

ace, but mainly glabrous below on the
leaf egi itself, with leaf margins ciliolate only
in the uppermost !4—!5 , and in the petioles short-
er than the leaf blades.

Peperomia gentryi Steyerm., sp. nov. TYPE:
VENEZUELA. Territorio Federal Amazonas:
Cerro de la Neblina, Camp V, valley north
base of Pico pus pee 0?49'N, 66°0’W, 1,250
m, 21-24 M 1984, Ronald Liesner &
Brian opie: 16901 (holotype, MO; iso-
type, VEN).

Herba repens; foliis alternis late ovatis vel subrhom-

infimis 5-7-plinerviis praeter margines superiores cili-
atos glabris; spicis geminis 2-3.5 cm longis; drupis el-
lipsoideo-ovoideis rostratis.

Repent herb with elongated glabrous stems 2
mm diam.; internodes 4—9 cm long. Leaves al-
ternate, broadly ovate to subrhombic-ovate, acute
to subacute at apex, truncate to broadly obtuse
at base, 5-7-plinerved, the innermost lateral
nerves forking off the midrib within the lower-
most 5-11 mm, 4.5-9 x 3-5.5 cm, moderately
pilose-ciliate on the upper margins; petiole 3.5—
7 cm long. Inflorescence with paired spikes; pri-
mary peduncle 2.8-5 cm long, minutely puber-
ulent; secondary peduncle subtending the spike
.5-2 cm long, sparsely puberulent 2 or
glabrous; spikes 2-3.5 cm long, 1.5-2 m iam.
Fruit basally attached, p spin ovoid, rostrate,
the body 0.5 x 0.3 mm, the beak 0.2-0.3 mm
long; stigma anterior near base of beak.

—

aratype. Same locality as type, 12 Apr. 1984, Gen-
try & Stein 46542 (MO, VEN). Figure 2A, B

This species is related to both P. distachya and
P. schwackei. From P. distachya (L.) A. Dietr.
this species differs in the 5—7-plinerved broadly
ovate to subrhombic-ovate leaves, which are
acute to subacute instead of acuminate at the
apex and truncate to broadly obtuse at the base.

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

13mm
i
> R
£ E
A ant
i | "d
5 w S
E ;
X š z
NÁ
` € |
k |
\=
N / <
=——
A
S

| M

FIGURE 2. Peperomia. A, B. P. gentryi. — A. Habit. — B. Beaked fruit. —C. P. marahuacensis. — Habit

STEYERMARK

1987]

— VENEZUELAN GUAYANA 89

From P. schwackei C. DC. of southern Brazil, it
differs in the 5-7-plinerved, acute to subacute
leaves and longer secondary peduncle, which
supports the spike.

Piper gentryi Steyerm., sp. nov. TYPE: VENEZUELA.
erritorio Federal Amazonas: Cerro Ne-
blina, trail due south from Base Camp,
stunted swampy forest, alt. 140 m, 23 Apr.
1984, Alwyn Gentry & Bruce Stein 46887
(holotype, MO).

Frutex 1.5-metralis, internodiis 2-3.5 mm diam. gla-
bris; foliis grosse rugosis lanceolato-ellipticis vel ob-
longo-ellipticis apice acutis vel acuminatis basi subae-
quilateraliter acutis vel subacutis 1 7-22 cm longis, 5.5—
7.5 cm latis supra glabris subtus ad nervos hirsutulis
ceterum glabris, nervis lateralibus utroque latere 11-
15 totis pinnatim venosis; petiolis 3 12 mm longis
breviter hirsutulis; spicis 2 cm longis, 6-7 mm latis;
pedunculis 3 mm longis glabris; E floriferis fim-
briatis.

Shrub 1.5 m tall, internodes 2-3.5 mm diam.,
glabrous. Leaves lance-elliptic or oblong-elliptic,
acute to acuminate at apex, subequally acute or
subacute at base, 17-22 cm long, 5.5-7.5 cm
wide, broadest at the middle, pinnately nerved
throughout to apex, glabrous above, punctate,
glabrous below on leaf surface but hirsutulous
with short, spreading hairs on the midrib and
lateral nerves, grossly rugose both sides, venation
sulcate above, elevated below, the areoles formed
by the tertiary veins averaging 5-7 mm diam .;
lateral nerves 11—15 each side, spreading at an
angle of 15—20°, sulcate above, subelevated be-
low, anastomosing at 5-10 mm from the margin;
petioles 5-12 mm long, shortly hirsutulous, vagi-
nate to the base of the blade. Spikes erect-as-
cending, 2 cm long, 6-7 mm wide; peduncle 3

above, depressed at summit, 2 mm diam., gla-

species is related to P. holtii Trel. &
daa from it differs in the glabrous
stems and nae more numerous lateral
nerves, and floral bracts conspicuously fimbriate.

V OCHYSIACEAE

A REEVALUATION OF THE GENUS
EUPHRONIA (VOCHYSIACEAE)

In his publication on a revision and taxonomic

position of the genus Euphronia, Lleras (1976)

showed that the generic name Lightia must be
replaced by the earlier Euphronia. Furthermore,
he transferred the genus from the Trigoniaceae,
where it had been placed previously, to the
chysiaceae, the family he considered as showing
the closest affinity.

Although two species had previously been rec-
ognized under Lightia in Warming's treatment
of the genus (1875), Lleras considered these taxa
as constituting one variable taxon, which he des-
ignated E. hirtelloides Martius ex Martius & Zucc.
His decision in considering this a polymorphic
species was based on what he judged as extreme
variability in leaf morphology and lack of geo-
graphical or other *'consistency."

While preparing a taxonomic treatment for the
flora of the Venezuelan Guayana, I have con-
cluded that three distinct taxa are involved wor-
thy of specific recognition. Consistent differences
have been found in characters such as type of
inflorescence, length of pedicels, apex of calyx
lobes, and quality of pubescence present on the
exterior of the calyx lobes and pedicels. More-
over, the three taxa demonstrate distinct geo-
graphical areas to which they are limited.

KEY TO THE SPECIES OF EUPHRONIA

la. Calyx lobes caudate-acuminate, the exterior
portion pes by a dense, uniform, ap-
ed, lanuginose tomentum; inflorescence

x, covered with
an appressed lanuginose tomentum ..........
ee
Ib. Calyx lobes pee 1 pe tuse, or subro
date at apex, the rior portion with he se,
ascending to spreading hairs throughout or at
least on midrib; flowers 3—23, racemose, the
pe e arising y Ped an oard rachis;
pedicels 1.5-5 mm long, shorter than the ca-
lyx, rr loosely ascending to spreading pu-
bescence

2a. FI

am

rimus paie ml the. epre
branch, mainly (7-)15-23-flowered, 4-

mi
— with shortly appressed tomen-
tu E. hirtelloides
š Florife erous portion of rachis si shortly

N
[^n

(-4.8) cm
long, mainly ovate to ovate-elliptic, gen-

FiGURE3. Euphronia.—A. E. hirtelloides. — A. Habit of edu branch. — AA. Calyx. — B. E. guianensis. —
B. Habit of flowering branch. — BB. Calyx.— C. E. acuminatissima. —C. Habit of Busen: branch. — CC. Calyx

1987]

< °

8
~~

STEYERMARK— VENEZUELAN GUAYANA

zuphronia guianensis o
suphronia hirtelloides A

suphronia acuminatissima W

l
n? AV nz

FIGURE 4. Map showing distribution of Euphronia.

KC | es?

erally rounded to obtuse at apex, 1.75-
2 times longer than broad; inner calyx

white borders; exterior of calyx lobes with
dense, loosely ascending to spreading
hairs over the entire surface .... E. guianensis

Euphronia acuminatissima Steyerm., sp. nov.
TYPE: VENEZUELA. Territorio Federal Ama-
zonas: Rio Orinoco, Sabana Cumare on right
bank of Caño Cumare, Río Atabapo, 20 km

g.l
pie 43762 (holotype, MO;
NY). Figures 3C, CC, 4

Frutex vel arbuscula 0.2—5-metralis; foliis oblongo-
ellipticis vel lanceolato-ellipticis apice rotundatis vel
obtusis Ap cuspidatis basi acutis vel subacutis 2-
4.5 cm longis, 1-2 cm latis; inflorescentia subfascicu-
lata plerumque (2-)3-6-flora pedunculata, pedunculo

mm longo; pedicellis 7- 8r mm longis, indumento

ena

lanceolatis apice caudato- acuminati S | 7-9 mm longis,

indumento lanuginoso pilis adpressis munitis.
Subshrub to small tree 0.2-5 m tall; stems

gray-white lanuginose, becoming glabrescent or

glabrous on older or lower portions. Leaves co-
riaceous, dark green above, white below, oblong-
elliptic to lance-elliptic, rounded to obtuse at the
abruptly cuspidate apex, acute to subacute at base,
2-4.5 cm long, 1—2 cm wide, 2.6-3 times longer
than broad, gray-floccose above on young leaves,
becoming glabrous on older leaves, white-pan-
nose below; petioles 2-6 mm long, gray-white
lanulose. Inflorescence terminal or axillary, sub-
fasciculate, sessile or shortly pedunculate, (2-)
3—6-flowered, the flowers crowded at or near the
summit of the abbreviated rachis; peduncle 0.5—
1 cm long, together with the rachis gray-white
lanuginose. Bracts subtending pedicels subulate,
caudate-acuminate, 3.5-4 mm long, 0.4-1 mm
wide, lanuginose, more persistent than in other
species. Pedicels 7-8 mm long, nearly equaling
the calyx, densely gray-white lanuginose with ap-
pressed, more or less uniform indument. Calyx
10-12 mm long, densely gray-white lanuginose
without, the hairs E appressed; n cam-
panulate-turbinate, 2.5—4 mm long, 3-5 mm wide
at summit; lobes ' rowls lanceolate, maine
to a long caudate acuminate apex, 7-9 mm long,

92 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1.5-2.5 mm wide, gray-white lanuginose with-
out, minutely sericeous within. Petals lavender
or lilac, spatulate, rounded at 2 d
(15-2)18-23 mm long, (4-)6—9 wide near
summit, 1.5 mm wide at or aa, middle, loose-
ly pilose within along middle, glabrous else-
where, sericeous dis bearing shorter hairs

upper port glabrous elsewhere. Anthers
oblong, a chavs bilobed basally, 2.5 mm
long; filaments 9-11 mm long, the sterile one
retrorsely pilose, the others glabrous. Style 10
mm long, antrorsely pilosulous. Fruit cylindric,
13-14 mm long, gray-white lanuginose.

Distribution. Mainly on white sand savannas
of the Territorio Federal Amazonas, Venezuela,
at altitudes of 100-150 meters.

Paratypes. VENEZUELA. TERRITORIO FED
AMAZONAS: Rio Guainia, Sabana El Venado, left bank
l m abo

NY); same locality, Maguire & Wurdack ches (NY):
base of Cerro Yapacana, 125-130 m, Maguire, Cowan
& Wurdack 30479-A, 30509 (NY); Bajo Rio Cha
alrededores de Canaripó, a unos 20 km al E de la con-
fluencia con el Río Orinoco, 4?03'N, 66?49'W, 98 m,
Huber 1070, 1877 (NY, VEN); Rio Ventuari, frente al
caserío de Canaripó, 4?09'N, 66°50’W, 100 m, Huber
2441 (NY, VEN); Río Orinoco, poco más río abajo de
Santa Barbara, 4°02'N, 67°15'W, 100 m, Huber 2471
(NY, VEN); E del Cafio Perro de Agua, a unos 30 km
al SE de la confluencia Orinoco-Ventuari, 3?47'N,
67*00'W, 100 m, Huber & Tillett 2824 (NY, VEN); 10
km al S del Río Autana, 15 km al SW del Cerro ie
4*44'N, 67?33'W, 100 m, Huber 4063 (NY, VEN); Ca

San Miguel between Limoncito and Caño Ikebeme
(about 70 km from river mouth), 100-140 m, Wurdack
& Adderley 43249 (NY); middle Caño Yagua, NE and

base of Cerro Cucurito, 3°36'N, 66°34'W, Huber &

Tillett 2925 (VEN); 20 km NW of Yavita, headwaters
of Caño Pimichín, 3?1'N, 67°33’W, 120 m, Huber &
Medina 5947 (VEN); 4 km west of Serranía He Cuao,
4°59" N, 67°32'W, Huber & Tillett 5293 (VEN); Rio

2021 (MO, TFA V). BOLIVAR: T a 133282

This species differs markedly from both E. hir-
telloides and E. guianensisin the longer, caudate-
acuminate calyx lobes and fewer-flowered,
subfasciculate inflorescence with longer pedicels.
The pubescence, moreover, is quite different from
either of the other species, with both pedicels
and the exterior of the calyx lobes covered by a
short, appressed, finely lanuginose indument, the
other two taxa having a pubescence of longer,
loose, spreading to ascending hairs on the ped-

[Vor. 74

icels and on a part or whole of the outer surface
of the calyx lobes.

Euphronia guianensis (R. H. Schomb.) H. Hal-
ier, Meded. Herb. Leid. 35: 13. 1918, in
obs.: Lightia guianensis. Lightia guianensis

Schomb. in Linnaea 20: 757. 1847; Warm
in Martius’s Flora Brasiliensis 13(2): 122.
1875. Figures 3B, BB, 4

Shrub or tree (0.2—)2—10 meters tall, branches
gray-lanuginose, becoming glabrous below or in
age. Leaves coriaceous, gray to silvery white be-
low, shortly petiolate to subsessile, mainly ovate
to subovate, generally rounded or obtuse at a
minutely mucronate apex, less frequently sub-
acute, chiefly rounded to obtuse at base, (0.6-)
1.5-3 (4.8) cm long, (0.5—)1—2(-2.5) cm wide
(rarely sterile shoots of juvenile branches elliptic-
oblong, 7 x 3-3.5 cm), revolute, densely white
tomentose below with elevated tomentose mid-
rib, young leaves gray-white floccose above, old-
er ones mainly glabrous above with narrowly
sulcate midrib, sometimes white-tomentose along
upper midrib; petioles 1-4 mm long, densely gray
tomentose. Inflorescence terminal or axillary,
mainly simple and short racemose or occasion-
ally the axis with a branch at base, 2-6 cm long
including the peduncle, the flowering portion 1-
4 cm long, 1-2 cm wide, 3-8-flowered; peduncle
0.6—2.5(-4) cm long, together with the rachis
densely tomentose. Bracts subulate, 4-5 mm long,
tomentose, caducous. Pedicels 1-5 mm long,
shorter than the calyx, densely tomentose with
subspreading to loosely ascending hairs. Calyx
6—9 cm long, tube shallowly campanulate, 1.5—
3 mm long, 3-4 mm wide at summit, densely
gray tomentose with short spreading-ascending
hairs; calyx lobes slightly unequal, broadly lan-
ceolate to oblong-lanceolate, acute, the outer lobes
dull gray nearly throughout with narrow paler
margins, the inner lobes with conspicuous broad
white margins and a narrow gray-green median
zone, the longer inner lobes 6.5-7 mm long, 3
mm wide, the shorter outer lobes 4.5-5.5 mm
long, 2.5-3 mm wide. Petals bluish, purplish, or
rose (varying according to different collectors),
spatulate, rounded at apex, unguiculate, 10-15
mm long, m wide, shortly and inconspic-
uously appressed-pubescent without to glabres-
cent, conspicuously long pilose within except near
apex. Anthers narrowly oblong, 2-2.5 mm long;
filaments 10-11 mm long, the sterile one re-
trorsely pilose, the others glabrous. Style 8-10

1987]

mm long, antrorsely pilose. Fruit oblong-cylin-
dric, 9-20 mm long, gray short-tomentose.

me. “‘Curataquilla’’; *'sacarai-
y savannas and open
sandstone exposures of the eastern portion of
Bolivar in the Venezuelan Guayana from the re-
gion of the Gran Sabana at Cerro Roraima west-
ward to the summit of Cerro Guaiquinima, and
in the adjacent Pakaraima Mountains of Guy-
ana, at altitudes of 300—-1,200 m, ascending to
1,600 m on the slopes of Ptari-tepui. [Cardona
2726 from the summit of Auyan-tepui at an al-
titude of 2,500 meters was cited as Euphronia
hirtelloides by Lleras (1976). It is actually Myrtus
alternifolia Gleason, as cited in Flora of Auyan-
tepui (Steyermark, 1967).]

Specimens examined. GUYANA. Mt. Ayanganna,
Pakaraima Mountains, between Chinowieng and Chi-
Chi Landing, 1,000 m, Maguire, Bagshaw & C. K
Maguire 40647 (NY); Kamarang Station, Pakaraima
Mountains, 500 m, Maguire & Fanshawe 32614 (NY).

VENEZUELA. BOLIVAR: Gran Sabana, El Dorado-Sta.
Elena road, 2.5 km before turnoff at San Rafael, 1,030
m, Luteyn, Lebrón-Luteyn & Steyermark 6294 (MO,
NY); km 146 along El Dorado-Sta. Elena road, 1,280
m, Luteyn, Lebrón-Luteyn & Steyermark 6291 (MO,
NY); between Santa Teresita de Kavanayen and base
of Ptari-tepui, 1,220 m, Steyermark 60307 (F, NY);
between Ptari-tepui and Sororopán-tepui, 1,615 m,
Steyermark. 60274 (F, MO); Cerro Manacauaray, head-

1,100 m, Cardona 2609 (NY); alrededores de Sta. Elena
de Uairén, Lasser 1273 (NY, VEN); Gran Sabana, 27
km N of Kama-Merü, carretera El Dorado-Sta. Elena
road, 5?30'N, 61?20'W, 1,300 m, Holst, Steyermark &
Manara 2222 (MO); 5 km east of Kavanayen, 1,200
m Maguire 33717 (NY); Uarupata, Maguire 33283
NY); Kamarang head, Gran Sabana, 800—950 m, Ma-
guire 33293 (NY); región de los rios Icabaru, Hacha y
qe sin nombre, 45
NY ue re del E

an

3°41" g"W. "Sieve

E. Dunsterville 113455, 113119-A (NY
: :

175 south of El Dorado, 1,200 m, Steyermark 111 296
(NY, VEN); Uriman, 300 m, Steyermark 75330 (F,
MO, NY); Río Caroní below Urimán, 393 m, Stey-
ermark & Wurdack 3 (F, MO, NY); región of Canaima,
6°15'N, 62°47'W, 200—500 m, Agostini 258 (NY, VEN);
Hacha ponas Canaima, Prance 16550 (NY, US): be-
tween Luepa and Kavanayén, 1,317-1,375 m, Badillo
& Forsa 6255 (MY); km 135-1 37 south of El Do-
rado, Badillo & Holmquist 6196 (MY); 148 km south
of El Dorado, 1,350-1,400 m, Steyermark & Dunster-
ville 104162 (MY, VEN); Cerro Akurimá, Sta. Elena,

STEYERMARK —VENEZUELAN GUAYANA 93

Tamayo 2699 (US); between Parupa and Kavanayén,
eek 792 (VEN); 7.5 km NE of Santa Elena, 4?40'N,
61?4'W, 880 m, Steyermark & Liesner 127592 (MO,
UB

This species is characteristic ofthe eastern por-
tion of the Venezuelan Guayana in the state of
Bolívar, where it is a common ied on rocky
open sandstone outcrops and savanna

Warming (1875) gave only a brief jm
of Lightia guianensis, mentioning a few salient
characters such as “foliis minoribus, ovatis v.
obovatis, basi rotundato-cuneatis; racemis bre-
vivissimis 4-6-floris" in differentiating this species
onia hirtel-

e

glabris" has been found to be incorrect. Careful
examination of specimens from eastern Vene-
zuela shows the filaments of the fifth sterile sta-
men to be retrorsely pilose and the four fertile
stamens as glabrous, characteristic of the genus.
Although united by Lleras with E. hirtel/loides
(loc. cit.), E. guianensis is amply distinct not only
in its shorter, fewer-flowered racemes and small-
er, differently shaped leaves, as mentioned by
Warming, but also in the hairs of the outer part
of the calyx lobes and pedicels uniformly loosely
ascending to spreading. Moreover, dried speci-
mens manifest broad white marginal zones on
the inner calyx lobes with only a relatively nar-
rower gray-green central midrib. This latter char-
acter is in contrast to the more uniformly gray-
green calyx lobes of E. hirtelloides with only the
central median portion with a loose, dense, as-
cending sericeous pubescence.
e specimens of Prance 16550 and Agostini
258 from the Canaima region need some com-
ment. The material of Prance is sterile with the
leaves larger than usual, and that of Agostini has
inflorescences (up to 7 cm long) and peduncles
(3.5—4 cm long) longer than usual, thus attaining
dimensions similar to those of E. hirtelloides.
However, the short flowering portion (2-3 cm)
is like that of E. guianensis.

Euphronia hirtelloides Martius, Nov. Gen. et Sp.
: 121,4, 73. 1825. Lightia licanioides Spruce
Warm. in Mart rasiliensis

Lightia licanioides. Figures 3A, AA,
Shrub or tree mainly 2-10 m tall, branches
gray lanuginose, becoming glabrous below.
Leaves coriaceous, gray-green above, white be-

94 ANNALS OF THE MISSOURI BOTANICAL GARDEN

low, shortly petiolate, lance- to oblong-elliptic or
ovate, shortly acute at apex, acute to obtuse at
base, 4.5-7.5 cm long, 1.5—4 cm wide, densely
and closely white tomentose below with elon-
gated midrib, g i
tomentose in the sulcate groove; petioles 5-7 mm
long. Inflorescence terminal or axillary, simply
racemose, or with 1 or 2 lateral axes branching
from the base, 5-11.5 cm long including the pe-
duncle, flowering portion 4-10 cm long, 2-2.5
cm wide, (7—)15—23-flowered; peduncle 1-4 cm
long, together with the rachis gray-lanulose. Bracts
lanceolate, acute, 3.5—4 mm long, 0.8-1 mm wide,
sericeous without, caducous. Pedicels 1.5-5 mm
long, mainly shorter than the calyx, with dense,
loosely ascending to spreading hairs. Calyx 6-9
mm long, tube shallowly campanulate, 2 mm
long, 2-4 mm wide at summit, gray-tomentel-
lose, lobes unequal, inner ones oblanceolate-ob-
ovate, shortly acute, broader than the outer and
broadened above the middle, outer ones broadly
iid more narrowed above the middle,

2.5-)5-8 mm long, 2-3 mm broad, sericeous
both sides, more abundantly long-sericeous with
longer, looser hairs on outer midrib, rather uni-
formly gray-sericeous elsewhere, externally with
shorter appressed hairs. Petals rose, purplish, or
lavender, the limb spotted with violet (fide Clark
and Maquirino), spatulate, rounded at base, un-
guiculate, short-sericeous or sometimes glabres-
cent, long pilose within at base and median por-

V11IIVillllwo YY hite

ile one retrorsely pilose, the others glabrous. Style
11-12 mm long, antrorsely pilose. Fruit cylin-
dric, 15-23 mm long, 5-6 mm wide, subobtusely
trigonous, gray lanate.

Distribution. Amazonian Brazil in Estado
Amazonas and Territorio do Roraima, Ama-
zonian Colombia in Vaupés and Caquetá, and
Territorio Federal Amazonas of southern Ven-
ezuela, at altitudes of 100—150 meters.

Specimens examined. | VENEZUELA. TERRITORIO FE-
DERAL AMAZONAS: ad flumina Casiquiari, Vasiva et Pa-
cimoni, 25-26 Feb. 1854, Spruce 3413-x (type of Ligh-
tia licanioides; isotype, NY; photo ;

Carlos de Rio Negro, 1°56’N, 67°03'W, 119 m,
Clark & koa Oe 7338, 7363 (NY); NW base of
rro Yapacana, epee a 34525 (MO,
NY), 34546 (NY); Sa avann rro Yapacana,
m p & Wurdack 305 94 EN

Coro . AMAZONAS: Río Caquetá, Araracuara sa-
vannas, rpe C. K. Maguire & Permie 44153

; Vaupés: lower Río Parana-pichuna, at Mitú,

x
9

[Vor. 74

RAZIL. AMAZONAS: Rio Curicuriary, affluent Rio
Negro, Ducke 337 (GH, MO, NY, US); same locality,
Ducke 159-A, 23869 (NY, US); Manaus-Caracarai Road
(BR-174), km 115, campina adjacent to Igarapé Lajes,
Zarucchi, Almeida & rae eee 2544 (NY); Rio Negro,
Preto, Fróes 22753 (M i iuxi
200 km above mouth, 2 of Rio Negro, Prance et
al. 15502 (NY, US); Rio Negro to 608 (NY).
Territorio Federal Rora nco, Sáo José de
51

(MO, NY, US); Caracarahy, Rio Branco, Ducke ] 407
(GH, NY, US); Rio Negro, Rio Tea 40 km above mouth,
village Bacuri, Kubitzki et al. 79. 240 (US).

In his description of Lightia licanioides (=Eu-
phronia hirtelloides), Warming (1875) incorrect-
ly described all five filaments of the stamens as
retrorsely pilose, and plate 22 (1875) depicts three
of the stamens with retrorse pubescence. How-
ever, dissection of specimens pertaining to this
taxon reveals the retrorse pilosity present only
on the fifth sterile filament, while the other four
fertile ones are glabrous. As indicated in the pres-
ent key to the species and in comments under E.
guianensis, E. hirtelloides is distinct in having a
more elongated, many-flowered inflorescence;
larger, more acutely tipped leaves; and the pu-
bescence of pedicel and outer surface of calyx
lobes of a different type. It should be noted here
that the calyx of Euphronia is gamophyllous
[Lleras (1976) describes the calyx as consisting
of five sepals], consisting of a calyx tube and five
lobes, as originally defined by Martius and Zuc-
carini (1826), Robert Schomburgk (1847, as
Lightia), and Warming (1875, as Lightia).

AQUIFOLIACEAE

Ilex liesneri Steyerm., sp. nov. TYPE: VENEZUELA.

Territorio Federal Amazonas: Depto. Ata-
erro Marahuaca, 1-2 km N of
3?43'N, 65°31'W, 1,100 m,
8-9 Mar. 1985, Ronald Liesner 18469 (ho-
lotype, MO; isotype, VE

rutex 1-2 m altus glaber; foliis anguste elliptico-
lanceolatis ad apicem obtuse attenuate eque angustatis
4.5-9 cm lon-
=f 0.5-1.5 cm latis subtus sparsim punctulatis, ad
margines crenato-serrulatis, utroque latere 5-13-cre

nulato; inflorescentiis axillaribus et lateralibus solitae

inflorescentiae pedicellis filiformibus sub anthesi 4—
mm n longis sub fructu 4-8 mm longis; floribus 4- meris;

f. calucic

Zarucchi 1986 (GH).

mm longis, 1.5 mm latis; floribus 9: calycis lobis 0.5

1987] STEYERMARK — VENEZUELAN GUAYANA 95

FIGURE 5. Ilex liesneri. — A. Habit of fl ill fl —B. Habit of flowering

[o wasa showing terminal inflorescence. — C. Habit of deos branch. —D. Detail of portion of ins leaf "Pies
showing punctation, revolute margin, and crenulations. — E. Pyrenes, lateral view, left; dorsal view, right. —
Pistillate flower. — G. Staminate flower

mm longis, 1.2 mm latis; petalis suborbicularibus ro- Shrub 1-2 m tall with slender branches. Leaves
tundatis 1.5 mm longis, 2 mm latis; fructu subgloboso — narrowly elliptic-lanceolate, narrowed above to
vel ovoideo-subgloboso 7 mm longo, 7 mm lato; pyre-
nis 4-5 trigonis 5 mm longis, 3.5-5 mm latis, dorso à long obtuse apex, narrowed below to a long
3—5-costatis. acute base, 4.5-9 cm long, 0.5-1.5 cm wide,

96 ANNALS OF THE MISSOURI BOTANICAL GARDEN

sparsely punctate below, midrib narrow, sulcate
above, slightly elevated below, lower surface
enervate, upper surface 7-8-nerved on each side
of midrib, subhorizontally spreading, anasto-
mosing 1-2 mm from margin, remotely crenu-
late-serrulate with 5-13 depressed crenulations
on each margin. Petioles 5-15 mm long. Inflo-
rescence axillary and lateral, solitary, trichoto-
mously cymose, pedunculate; cymes simple or
compound, few-flowered (usually 7) when simple
with one flower on the central axis and three each
on the two lateral axes, when compound each of
the axes 2-3-flowered, the axes divaricately
spreading, 1-3 mm long; pedicels 4-8 mm lon
Bracts subtending pedicels spreading, 0.1 mm
long. Peduncle slender, 0.8-1.5 mm long. Flow-
ers 4-merous. St te flowers: calyx lobes sub-
orbicular, rounded, 1.8 mm long, 1.5 mm wide;
anthers suborbicular, 0.8 x 0.8 mm; filaments
0.8 mm long; pistil rudiment ovoid, 1 mm long.
Pistillate flowers: calyx lobes 0.5 mm long, 1.2
mm wide; petals suborbicular, rounded, 1.5 mm
long, 2 mm wide; ovary ovoid or subglobose, 1—
1.5 mm long, constricted slightly at summit into
a short style 0.2-0.3 mm long; stigma prominent,
ovoid-capitate. Fruit subglobose or ovoid-sub-
globose, 7 x 7 mm; pyrenes 4-5, trigonous, 5
mm long, 3.5-5 mm wide, dorsally 3—5-costate.

Paratypes. VENEZUELA. TERRITORIO FEDERAL
AMAZONAS: Depto. Atabapo, Cerro Marahuaca, Ri
Yameduaka arriba, 3°38'N, 65?28'W, 1,225 m, Liesner
17604 (MO, VEN); “Sima” Camp, wiegen pors
tion along eastern branch of Cerro Negro,
ahuaca, 3°43'N, 65°31'W, 1,140 m, Stevermark & Holst
130575, 130422, 130424 (MO, VEN). Figure 5

This species is distinguished by the narrowly
lance-elliptic, crenulate, punctate leaves, which
are long attenuate at each end, the 3—5-dorsally
ribbed pyrenes, the slender, solitary peduncles
from both axillary and lateral buds, and the fili-
form pedicels on trichotomous cymose inflores-
cences.

OCHNACEAE
Tyleria apiculata Sastre (Fig. 6A-G)

This species was recently described by Sastre
(Phytologia 59: 313-314. 1986) with only a brief
description based upon a single specimen, which
I collected. A later expedition in 1985 to Cerro
Marahuaca by Bruce Holst, Ronald Liesner, and

[VoL. 74

me resulted in the collection of more ample ma-
terial. In view of these additional collections, the
following more detailed description can be fur-
nished.

Shrub or small tree 2-3 m tall. Leaves erect,
crowded, those on sterile branches clustered at
the apex, scattered below the inflorescence on
fertile branches, subobtuse at apex, sometimes
with a minute mucro 2 mm long, the apically
clustered leaves on sterile branches 7-8 cm long,
5-10 mm wide, those on fertile branches often
shorter and 6-13 mm wide, gradually narrowed
to the sessile base, glabrous on both surfaces,
margins finely serrulate-ciliate with closely as-
cending purplish hairs 0.5-0.7 mm long, lateral
nerves finely parallel from base to apex, strongly
ascending to apex. Stipules ovate-triangular or
lanceolate, obtuse to acute, 4-10 mm long, 2 mm
wide, finely parallel-veined. Inflorescence panic-
ulate, terminal, many-flowered, 12-14 cm long,
5—7.5 cm wide in the basal half, 3.5- 4 cm wide
in upper half; flowers pedicellate, pedicels fili-
form, 10-13 mm long, articulate 1-2 mm above
base, dilated below apex, spreading in fruit. Se-
pals membranous, ovate or elliptic-ovate, ob-
tuse, 8 mm wide, 5 mm long. Petals pink, ob-
ovate, rounded at apex, narrowed to the base, 15
mm long, 11 mm wide. Anthers linear-oblong,
4 mm long, acutely apiculate, subsessile; fila-
ments 0.3 mm long. Staminodes spatulate,
rounded at apex, 7 mm long, 2.5 mm wide above,
adnate 1 mm above the base with dimorphic
lateral appendages, the longer appendage deeply
cut into an elongated subulate, simple, distal seg-
ment 5.5-6 mm long attached to a ligulate mul-
tifimbriate portion, and an inner proximal short-
er appendage 5 mm long with lateral fimbrillate
segments. Pistil 9 mm long; style subulate, 5 mm
ong; ovary narrowly conic, 4 mm long, | mm
wide. Capsule oblong-conic, 10-11 mm long.

—

Specimens examined. VENEZUELA. TERRITORIO FE-
L AMAZONAS: Depto. Atabapo, Cerro Marahuaca,
below Salto Los Monos on tributary of headwaters of
Rio Iguapo, 3°35'N, 65?23'W, 1,500-1,600 m, 11 Mar.
1985, Liesner 18511 (MO, VEN); Rio Yameduaka ar-
riba, 3°38’N, 65?28'W, 1,225 m, 17-18 Feb. 1985,
Liesner 17624 (MO, VEN); “Sima Camp,” southcen-
tral portion of forested slopes along eastern branch of
Caño Negro, 3?43'N, 65?31'W, 21-22, 24 Feb. 1985,
Steyermark & Holst 130565 (MO, VEN); below Salto
Los Monos on tributary of headwaters of Río Iguapo,
3°35'N, 65?23'W, 1,500 m, 13-14 Oct. 1983, Steyer-
mark 129649 (P, holotype; MO, VEN, isotypes).
Sastre notes that the species is well marked by its

1987] STEYERMARK— VENEZUELAN GUAYANA 97

FiGure 6. Tyleria sewers — A. Habit of flowering branch.—B. Sterile leafy branch showing terminal
clusters of leaves. — C. Detail of leaf s lower surface. — D. Anther. — E. Anther and staminode, ventral view.—
F. Staminode, dorsal view. nih Cap

apiculate anthers and contrasts it with T. spectabilis species in the piis pedicels in terminal panier
Mag. & Wurd. and T. floribunda Gl. because of its broader and lon sho
shorter hon It also resembles T. linearis Gl. of ad- er petals, and d larger fistibriate lateral appendages of is
jacent Cerro Duida in leaf shape but differs from that — staminodes

98 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Sauvagesia marahuacensis Steyerm., sp. nov.
TYPE: VENEZUELA. Territorio Federal Ama-
zonas: Depto. Atabapo, Cerro Marahuaca,
Rio Yameduaka arriba, 3?38'N, 65?28'W,
1,225 m, Ronald Liesner 17677-A (holo-
type, MO; isotype, VEN). Figure 7.

Suffrutex 0.5-metralis, caulibus virgatis; foliis sub-
sessilibus anguste linearibus vel subulatis apice cus-

nervis elevatis striatis; floribus ad apices ramulorum
pedicellis 3-3.5 mm
oe sepalis lanceolatis acutis 5- 5. 5 mm m longis, = a
m latis;
saree era 7-7.5 mm longis, supra mediu
5 mm latis; uri 5 coronae squamulis X HABI
corona l-seriata basi cum filamentis in columnam
brevem 0304) mm latam coalita, squamulis quinque
1.2-1.5 mm longis, laminis subulato-spathulatis apice
rotundatis m 7-0. 9 mm longis i in stipitem 0. 5m m lon-
gum

2.7 mm ORR 0.2-0.25 mm latis.

Virgate subshrub 0.5 m tall. Stems dichoto-
mously branched or 3—6-verticillate, densely fo-

liose toward the apex, leafless for most of the
length below, 2-3 mm diam. Stipules densely
crowded, overlapping, ge toe ei lanceolate,

2.5-3 mm long, 1 mm wide, dorsally carinate,
conspicuously pectinate from base to apex with
15-18 subulate appendages on each side, each
one tipped by a filiform, white cilium 1 mm long,
lower appendages often deciduous. Leaves sub-

15 times longer than broad, cuspidate at apex,
inconspicuously narrowed at base, margins revo-
lute, minutely glandular serrulate, each margin
with 10-11 minute, appressed-ascending teeth
0.1 mm long, upper surface marked with small
transverse depressions. Flowers numerous near
the apex, conspicuously pedicellate; pedicels 3—
3.5 mm long. Sepals lanceolate, acute, 5-5.5 mm
long, 1.7 mm wide. Petals white, obovate, round-
ed above, narrowed to a subcuneiform base, 7-
7.5 mm long, 4-5 mm wide. Stamens 5; anthers
linear, 2.1-2.7 mm long, 0.2-0.25 mm wide; fil-
aments 0.6 mm long. Staminodes 5, subulate-
spatulate, rounded at apex, 1.2-1. m long,

arrowed basally, the claw 0.5—0.7 mm long, 0.2
mm wide, the laminar portion 0.7—0.9 mm long,
the summit of the laminar portion attaining one-
third to one-quarter length of anther. Ovary
ovoid, 1 mm long; style subulate, 2.9-3 mm long.

[Vor. 74

This species differs from S. guianensis (Eichl.)
Sastre and varieties in the narrower linear-subu-
late leaves 10-15 times longer than broad with
depressed areas on the upper surface and in the
long-pedicellate, more numerous flowers con-
spicuous at the ends of the leafy stems as con-
trasted with the solitary, sessile or barely pedi-
cellate flowers largely hidden amongst the leaves
of S. guianensis.

SAUVAGESIA GUIANENSIS AND VARIATIONS

A study of Sauvagesia guianensis (Eichl.) Sastre
(1970) reveals much variation not only in details
of leaf morphology but in those of staminodial
form as well. This taxon (sensu lato) is distrib-
uted in the region of the Guayana Shield, with
its greatest concentration in the Venezuelan

uayana, but with outliers in adjacent Guyana
and Colombia (Sastre, 1970).

It was originally described by Eichler as Leit-
gebia guianensis (op. cit.), based upon a collec-
tion by Richard Schomburgk from the savannas
of Guyana. Gleason (1931) referred collections
of Tate from the summit of Cerro Duida to this
taxon. In 1946 Lasser described a Steyermark
collection, also from the summit of Duida, as
Leitgebia gleasoniana. At this time Lasser (1946)
associate helps collection from Cerro Pa-
raque (Sipapo) as conspecific with the Steyer-
mark type.

Sastre (1970) eventually transferred Leitgebia
guianensis to Sauvagesia and referred later col-
lections originating from various parts of the
Venezuelan Guayana as one taxon. The vari-
ability of this taxon becomes obvious upon more
detailed study. It is noted that specimens origi-
nating from p Gran Sabana of eastern Vene-
zuela and adjacent Guyana near the type locality
have leaves ain fewer times longer than
broad, relatively shorter and with shorter-point-
ed apices than those from the summits of various
table mountains westward in Venezuela. More-
over, the staminodes of specimens from the east-
ern sector of the range have shorter laminar por-
tions. Additionally, a surprising degree of
variation is shown between plants collected from
various sandstone table mountains. For exam-
ple, plants from the summit of Cerro Guaiqui-
nima in Bolivar have leaves with only 2-4 glands
on sach margin that appear in the upper three-

other parts
of the range ‘manifest 6-14 glands on each mar-
gin, which are distributed from one-fourth to

1987]

STEYERMARK— VENEZUELAN GUAYANA

2.6mm

Le

=
11mm
>

NM j N
V Y
KU

SE >
A =

Z

z
Z

AU

MU
nd

[LL
—

Du
o dA

E
=
16mm
2.9mm

13mm

2.7mm

11.5mm

9.5mm

2.7mm

7.5mm

5.5mm

FIGURE7. A-F. Sauvagesia marahuacensis. — A. Habit. — B. Leaf, lower side. — C. Leaf, upper side. — D. Detail
of upper apical portion of lower side of leaf. —E. Petal.—F. Po rion of stamens and staminodes attached to
membran eek baal Sauv ] j sp. ensis:—A!'. Leaf, lower side. we Apical portion of

rtion of ig pee showing anther fer staminode. — B! t. . gui
aeni subsp. gleasoniana:—B. p f, lower side. — B?. Leaf, upper side. — B?. Apical conta of leaf. — ^. Mar-
i C'-C*. S

sipapoensis: — C'. Leaf, lower side. — C". Marginal leaf glands. — C^. Apic
droecium showing anther and staminode.— D'-D*. S. guianensis subsp. guaiquinimensis: —D'. Lea per

, Up
side. — D?. Leaf, lower side. — D?. Apical portion of leaf. — D+. Marginal leaf glands. — D5. Portion of androecium
showing anther and staminode. — D*. Petal.

100

halfway up the leaf margin to the apex. Further-
more, petals of the Cerro Guaiquinima plants
have acute to subacute apices, whereas those on
specimens elsewhere are rounded. Moreover, in
contrast to the usually encountered elongated,
linear-ligulate, narrowly spatulate, or narrowly
elliptic laminar portion of the staminode, plants
from the summit of Cerro Sipapo (Paraque) have
developed a suborbicular type about as broad as

ng.
Results obtained from a study of the available
material in NY and VEN herbaria indicate that
Sauvagesia guianensis has undergone differen-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

tiation in various portions of its geographical
range resulting in populations showing a diver-
gence in various characters. This geographical
isolation on several of the sandstone table moun-
tains is recognized in the present study as cor-
related with taxonomic characters sufficiently
distinct as to be considered of subspecific sig-
nificance.

The following key encompasses the principal
differences by which these variations may be rec-
ognized within S. guianensis as well as distin-
guishing them from the related S. marahuacen-
Sis.

KEY TO SAUVAGESIA GUIANENSIS AND SUBSPECIES, AND RELATED SAUVAGESIA MARAHUACENSIS

la. Flowers on pedicels 3—3.5 mm long, numerous and conspicuous near apices of the leafy stems; leaves
linear-subulate, 0.8-1 mm wide, 10-15 times longer than broad; upper leaf surface with v l
depressed areas S. marahuacensis

essile

=

wers subs, or on pedicels to 2 mm long pedicellate, few and mainly concealed amidst the
leaves; leaves linear-oblanceolate, 1-3 mm wide, 3-10 times longer than broad; upper leaf surface

ere A depressed areas
s 3-3.5 m

m long, 1-1.5 mm wide; Colombia ......
2b. Saino des 1-1.9 mm long, 0.2-0.5 mm wide; Venezuela 2 Gu

E oe subsp. araracuarensis
3

3a. = liar glands 2-4 on en margin, the lowest starting at 4 ne E below the apex; petals

acute to acute at a

ianensis subsp. guaiquinimensis

w
e

ands 6-14 on e margin, the lowest starting from He the distance below the apex;

minar p

of staminode suborbicular, about as broad as long, 0.5 x 0.4-0.5 mm;
upper aieh of leaf blade with a conspicuous puncticulate appearance

c. ianensis subsp. sipapoensis

4b. Laminar portion of staminode elongated, longer ie broad, linear-ligulate, elliptic-spat-
0.2-0.3 m

ulate, or narrowly Dm 0.8-1.9 x

m; upper surface of leaf blade lacking

a puncticulate appear:

5a. Foliar — 6-1 “Ë on amam ina ma oblanceolate or narrowly oblanceolate, 8-

11 mm

ong, 1.5-3 mm wide, 3—4
to bluntly pointed; ane ee pr stamino 8 mm a ong

mes longer than broad; leaf apex shortly acute
de 0. Fi

is subsp. guianensis

io
o

mm long, 1-2 mm wi

S. gu
. Foliar glands 11-14 on Mrd margin; leaves linear or Uns ane n 10-18(-22
, 7-10 times To than broad; leaf apex prolonged, acu-
minate; laminar portion a staminode 1.2-1 m m

. S. guianensis subsp. gleasoniana

la. Sauvagesia guianensis (Eichl.) Sastre subsp.
guianensis. Sauvagesia guianensis (Eichl.)
Sastre, Caldasia 10: 570. 1970. Leitgebia
guianensis Eichl. in Martius's Flora Brasi-
liensis 13(1): 413. pl. 83, fig. 2. 1871. TYPE:
Guyana. Rich. Schomburgk 1553. Figure

A'—A*

Leaves oblanceolate, acute to bluntly pointed
at apex, 6-11 mm long, 1.5-3 mm wide, 3.5-4
times longer than broad, foliar glands (6—)7-11

spicuous among the leaves; subsessile to 2 mm

long pedicellate. Petals obovate, rounded at apex,
6.8-7.5 mm long, 5 mm wide. Anthers 2.5-3 mm
long. Staminode lamina narrowly to broadly el-
liptic-spatulate, rounded to subacute, 0.8-0.9 mm
long, 0.3 mm wide, the narrower lower portion
about equaling the lamina in length.

Distribution. Guyana and adjacent south-
eastern Venezuela, Bolívar.

Additional E gee) examined. VENEZUELA.
BOLÍVAR: km 1 fEl Dorado, ade of Rio Sacaica,
1,200 m, Soda Md VEN); 52 km N
o ma-meru, carre Do ts Sta. Elena, 5?40'N,
61?25'W, 1,300 m, P9» Steyermark & Manara 2201
(MO, VEN).

1987]

lb. Sauvagesia guianensis subsp. gleasoniana
asser) Steyerm., comb. nov. Leitgebia
gleasoniana Lasser, Bol. Acad. Ci. Venez. 9:
246. 1946. TYPE: VENEZUELA. b ades
Federal Amazonas: Cerro Dui
Savanna Hills, Aug. 1944, Cane 2
(holotype, VEN; isotype, F). Figure 7B'!—B>.

Leaves linear or linear-lanceolate, aristate long-
pointed at apex, 10—18(—22) mm long, 1-2 mm
wide, 7-10 times longer than wide; foliar glands
11-14 each margin, ascending outwardly from
margin, elongated, 0.3-0.5 mm long; upper sur-
face usually marked with iUe transverse stria-
tions. Flowers solitary, few en and incon-
spicuous among the leaves, iban to 1-2 mm
pedicellate. Petals obovate, rounded at apex, 8

mm long, 4.5 mm wide. Anthers 2.3-2.9 mm
long. Staminode 1.2-1.9 mm long, the laminar
portion linear-ligulate or narrowly spatulate, ob-
tuse or rounded, 0.3-1.2 mm long, 0.2 mm wide,
much longer than wide, the lower stipitate por-
tion 0.5-1.3 mm long.

Distribution. Summits of Cerro Duida and
uachamacari, Territorio Federal Amazonas,
Venezuela.

Additional specimens examined. | VENEZUELA.
TERRITORIO FEDERAL AMAZONAS: Cerro Duida, 3?40'N,
65°45'W, 1,500 m, Steyermark, Brewer- Carias & Lies-
ner 124569 (MO, VEN); plateau of Huachamacari,
3?50'N, 65?43'W, 1,720 m, 1 Mar. 1985, Liesner 18057
(MO, VEN); Savanna Hills, Cerro Duida, Steyermark
58248 (F, VEN).

lc. Sauvagesia guianensis subsp. sipapoensis
Steyerm., subsp. nov. TYPE: VENEZUELA.
TERRITORIO FEDERAL AMAZONAS: Serrania
Sipapo, cumbre, sección speed 5?N,
Vli 1,500 m, Ste ark, Bre
Carias & Liesner ee. eae Mk
iniit VEN). Figure 7C!—C4.

A subsp. guianense squamulorum laminis suborbic-
ularibus apice late rotundatis 0.5 mm longis, 0.4—
0.5 mm latis, foliis supra conspicue cellulosis re-
cedit

Leaves linear-lanceolate, ending in a long cus-
pidate apex, 10-13 mm long, 2-2.5 mm wide,
5-6 times longer than broad; foliar glands short,
7-10 each margin, incurved-appressed, the low-
est ones beginning * the distance below the apex;
upper surface with a conspicuous cellular ap-
pearance. Flowers solitary, few, subsessile, in-
conspicuous among the leaves. Petals obovate,
rounded at apex, 5.5 mm long, 4 mm wide. An-

STEYERMARK — VENEZUELAN GUAYANA

101

thers 2-2.5 mm long. Staminode with the lam-
inar portion suborbicular, broadly rounded, 0.3—
0.5 mm long, 0.4—0.5 mm wide, stipitate lower
portion 0.7-0.8 mm long.

Distribution. Summit of Cerro Sipapo, Ter-
ritorio Federal Amazonas, Venezuela

Id. Sauvagesia guianensis subsp. guaiquinimen-
sis Steyerm v E

quinima, cumbre, sector occidental, cerca
del borde cubierto con bosque bordeando
una sabana, 5?45'N, 63?43'W, 1,540 m, 27
May 1978, Steyermark, Berry, G. C. K. &
E. Dunsterville 117498 (holotype, MO; iso-
type, VEN).

A subsp. guianense atque ceteris petalis apice acutis
vel subacutis

e latere 2- 4 secus longitudinem s superiorem

3⁄4 Lx recedit.

Leaves linear-oblanceolate, acute at apex, 6—
9.5 mm long, 1.5-2 mm wide, 4—6 times longer
than wide; foliar glands 2-4 each margin, in-
serted along the upper 34 of the margin, ap-
pressed, short; upper surface with few or no
transverse lines but with a cellular appearance.
Flowers solitary, few, hidden and inconspicuous
among the leaves, subsessile. Petals obovate,
acute at apex, 5.5 mm long, 2 mm wide above
middle. Anthers 2.7 mm long. Staminode 1.6
mm long, the laminar portion ligulate-oblong or
ligulate-obovate, rounded or subobtuse at apex,
1.1 mm long, 0.3-0.4 mm wide, narrowed to a
stipitate portion 0.5 mm long.

Distribution. Known only from the summit
of Cerro Guaiquinima, Bolivar, Venezuela.

Paratypes. VRNDACHEA, Cerro Tag e cumbre,
sector suroeste , $38 N, 63?45'W, 1,650 m, Steyer-
mark, Berry, G. C. K. & E. ud 117434 (MO,
VEN); Cerro ar er cumbre, sector suroeste-
central, 5?45'N, 63?35'W, 950 m, Steyermark, Berry,
G. C. K. & E. Dunsterville 117474 (MO, VEN). Figure

D'-D*.
le. Sauvagesia g b

(Sastre) Steyerm., stat. nov. o mt gui-
anensis var. araracuarensis Sastre, Bull. Mus.
Nat. Hist. Paris 35: 1978. TYPE:
COLOMBIA. Com. Amazonas: Rio Caqueta,
Araracuara, camino a La Sabana, segunda
sabana de Tibeyes, 500—600 m, 5 Jan. 1977,
Sastre & Reichel 5139 (holotype, P; iso-
types, COL, G).

102

Staminodes larger than in the other subspecies,
3-3.5 mm long, 1-1.5 mm wide.

Distribution. Known only from savannas over
sandstone soils at the type locality.

THEACEAE (BONNETIACEAE)

Since the publication of “Flora of the Vene-
zuelan Guayana — I” (Steyermark, 1984), the fol-
lowing additional new taxa are described in Bon-
netia.

iin bolivarensis Steyerm., sp. nov. TYPE:

EZUELA. Bolivar: Ptari-tepui, cumbre,

sai N, 61?47'W, 2,400 m, 19 Nov. 1984,

Otto Huber 9818 (holotype, VEN; isotype,
MO)

Frutex l- metralis; foliis dense tonan Ep
oblongo obtu: e )
2.5-5 x 0.8-1.7 cm; sepalis 11- 12.5 x pecs mm; petalis
21-22 x 14-16 mm; stylis tribus 3-3.5 mm lotes fere
usque ad basem divisis.

Leaves crowded at summit of branches, ob-
long-lanceolate, subsessile, narrowed to an ob-
tusely acute apex, obtuse at base, (2-)2.5-5 x
0.8-1.7 cm, faintly impressed-nerved on both
sides or the lateral nerves not evident, midrib

flower oblong-lanceolate, acute to obtuse, 11-12
x 4.5 mm, dorsally carinate, setulose marginally
with dark setae 1 mm long; sepals lance-oblong,
subacute, 11-12.5 x 4-5 mm, obtusely dorsally
keeled basally and apically; petals white, sub-
cuneately obovate, subtruncate apically with un-
equally rounded sides, narrowed to the base, 21-
22 mm long, 14-16 mm wide at summit, 4 mm
wide at base. Stamens numerous, multiseriate;
filaments 5 mm or less long; anthers 0.5-0.8 x
0.6 mm; pistil 9 mm long; S 3, 3-3.5 mm
long, divided about 73 way dow

The larger flowers and larger, prt leaves
differentiate this taxon from B. chimantensis
Steyerm., B. tepuiensis Kobuski & Steyerm., and
B. toronoensis Steyerm. In its deeply 3-parted
style it differs from B. tepuiensis.

Bonnetia guaiquinimae Steyerm., sp. nov. TYPE:

m, 26 May 1978, Julian A. Steyerm
Berry & G. C. K. & E. sche 117421
(holotype, VEN; isotype, MO).

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

12: =k ` la: hl

non lan

rutex 1.5-metralis; f:
ceolatis Ma oblanceolatis apice acutis basi subobtusis
vel obtus 6 x 1.3-1.6 cm subtus enerv

petalis 20
longis; stylo subulato apice leviter 3-lobato

Leaves coriaceous, entire, oblong-lanceolate to
oblanceolate, acute at apex, gradually narrowed
to a subacute or subobtuse base, 5-6.5 x 1.3-
1.6 cm, enervate below, the midrib subimpressed
below, lateral nerves elevated above; petiole 1—
2 mm long. Sepals coriaceous, suborbicular-ob-
ovate, rounded at a shortly cuspidate apex, 10—
12 mm long, 6-8 mm wide above the middle,
3-4 mm wide at base; petals white, obovate, nar-
rowed to a subunguiculate base, 20 mm long,
15-20 mm wide at the summit, 2-3 mm wide at
base; filaments distinct, 3.5-7 mm long. Anthers
0.7 mm; pistil 9 mm long; style merely
3-lobed at apex.

This taxon is characterized by the shallowly
3-lobed style, entire, oblong-lanceolate, acute
leaves, which are enervate beneath.
distinguished from B. chimantensis Steyerm. by
the larger petals, shallowly 3-lobed style and larg-
er leaves enervate below. From B. toronoensis
Steyerm. it differs in the larger sepals and petals
and larger, entire leaves enervate below, while
from B. tepuiensis and subsp. minor Steyerm. it
is separated by the larger, minutely mucronate
sepals, longer filaments, and entire leaves.

—

Bonnetia cane up Sp. nov. TYPE:
VENEZUELA. var: Ptari-tepui, cumbre,
5°45'N, 61°45’ w, 2,360-2,420 m, 23 Feb.
1978, Steyermark, Carrefio, McDiarmid &
Brewer-Carias 115645 (holotype, VEN; iso-
type, MO).

Frutex 2.5 m; foliis sessilibus seer apice acutis
majoribus 3.5-4.5 x 1-1.5 cm minute obscureque ser-
rulatis; floribus solitariis breviter eR pedun-
culis teretibus vel subteretibus 4—6 mm longis; sepalis
lanceolatis vel suboblanceolatis acutis 12413 x -5
mm; petalis luteis obovatis apice rotundatis 16-18 x
12 mm; antheris 0.8 x 0.7 mm; stylis tribus 6 mm
longis fere usque ad basem divisis.

Leaves coriaceous, lanceolate, acute at apex,
slightly narrowed to the base, the larger ones 3.5-
4.5 x 1-1.5 cm, obsoletely pinnately nerved,
midrib slightly elevated below, microscopically
and obscurely serrulate. Flowers solitary, short-
pedunculate; peduncle terete or subterete, 4—6
mm long; bracts immediately subtending flower
narrowly oblanceolate, acute, 11-12 x 3-4 mm;

1987]

sepals subcoriaceous, lanceolate or suboblanceo-
late, acute, 12-13 mm, 10-striate, mi-
nutely ciliolate, obtusely carinate; petals yellow,
obovate, rounded above, 16-18 mm long (pre-
anthesis), 12 mm wide near apex, 2 mm wide at
base. Filaments 1.5-3.5 mm long (pre-anthesis);
anthers 0.8 x 0.7 mm; pistil 10 mm long; 3 styles
6 mm long, free nearly to the base.

From the yellow-flowered B. wurdackii Ma-
guire, this species differs in the larger, lanceolate,
and acute leaves, longer sepals, larger petals (even
in bud), longer filaments, and longer style
branches. The leaves, furthermore, do not man-
ifest the pale punctate stomata that are clearly
shown in B. wurdackii. From B. tristyla Gleason
it is easily distinguished by the short pedicels,
smaller floral parts and smaller leaves; while from
B. huberiana Steyerm. it is well separated by the
larger, lanceolate leaves, shorter pedicels, and
larger floral parts.

Bonnetia tristyla Gleason subsp. nervosa Stey-
erm., subsp. nov. TYPE: VENEZUELA. Terri-
torio Federal Amazonas: Cerro Yapacana,
below summit, 3?45'N, 66?45'W, 825 m, 5
May 1970, Julian A. Steyermark & George
Bunting 103153 (holotype, VEN).

Frutex vel arbor 1.5—4-metralis; foliis supra costa
media necnon nervis lateralibus nonnullis conspicue
manifesteque elevatis, nervis lateralibus supra inae-
qualiter prominentibus; petalis 25-38 x 15-22 mm.

Leaves suboblong, oblong-oblanceolate, or ob-
long-obovate, obtuse to rounded at apex, nar-

Petiole absent or 1-2 mm long. Peduncle 3-4.3
(—6.5) cm long. Sepals 12-20 mm long, the outer
12-15 mm long, the inner 15-20 mm long. Petals
25-38 x 15-22 mm.
Mee TERRITORIO FEDERAL
o Yapacana, summit, 1,000-1,200 m,
5 May 1970, Steyermark £ Bunting 103103 (VEN);
Cerro ieee 2 Jan. | , Maguire, Cowan & Hat
dack 30632 (N B. Cerro Yapacana, 1,200
Maguire, Cowan & Wurdack 30665 (NY. VEN); Gene

Paratypes.
AMAZONAS: Cerr

STEYERMARK— VENEZUELAN GUAYANA

103

bie 30 km al SSW de Ocamo, 2°31'N, 65?23'W,
440-600 m, 1-2 Mar. 1984, Steyermark, Berry & De-
lascio 130405 (MO, VEN); Cerro de la Neblina, Caño
Grande SSW of Cumbre Camp, 1,050-1,100 m, 25
Dec. 1957, Maguire, Wurdack & C. K. Maguire 42498
(NY, VEN); Cerro de la Neblina, summit, Canon
Grande slopes E of Cumbre Camp, 1,200-1,300
Maguire, Wurdack & C. K. Maguire 42235, 42179
(NY, VEN); Cerro Avispa, Río Siapa, summit, od N,
65°51'W, 1,510 m, G. C. K. & E. Dunsterville s.n. (VEN);
same locality and data, Cardona 3098 (VEN); Serrania
de Vinilla, 20 km SW of Ocamo, 2°20'N, 65°22' 6
m, Huber 6168 (VEN); E side of Río Siapa,
65°41'W, 600 m, Huber 6006 (VEN).

ue

This taxon differs from typical B. tristyla Glea-
son in having smaller petals, upper surface of the
leaf blade with the midrib prominently elevated,
and unequally prominent pairs of lateral nerves,
with 6—8 of the pairs prominently elevated al-
ternating with finer, lightly impressed interme-
diate ones. In B. tristyla subsp. tristyla the large
petals 35-42 x 27-30 mm contrast with the
smaller ones of subsp. nervosa. Moreover, in
subsp. tristyla the upper midrib is shallowly de-
pressed, not elevated, and all the lateral nerves
of the upper surface of the leaf are equally in-
conspicuous and lightly impressed. The leaf
blades in typical B. tristyla tend to be larger (5-)
8-15 x 2.5-5.0 cm as contrasted with 4-8 x
1.5-3.5 cm in subsp. nervosa. The base of the
leaf blade in B. tristyla subsp. tristyla is cuneately

rowed, whereas that of subsp. nervosa is usu-
ally slightly obtusely curved or rounded above
the junction with the petiole.

The geographical ranges of the two subspecies
are distinct. Typical B. tristyla, described origi-
nally from Cerro Duida, occupies the northeast-
ern sector of the range, from Cerro Duida an
Marahuaca east to the Meseta de Jaua and Sari-
sarinama in Bolivar. Bonnetia tristyla subsp. ner-

vosa, in contrast, is found in the more western
sector, ranging from Cerro Yapacana south to
Cerro Vinilla, Avispa, Aracamuni, and Neblina.

The only other species of Bonnetia having large
yellow petals, B. steyermarkii Kobuski, is easily
distinguished from B. tristyla by the larger acute
sepals subtended by 4—6 large sepaloid bracts and
the broader, acutely angled ancipital peduncle.

The following key incorporates the newly de-
scribed taxa of Bonnetia with those previously
described by the author in 1984.

KEY TO THE SPECIES AND SUBSPECIES OF BONNETIA

la. e phan

eral nerves of upper leaf surface elevated, impressed on lower surface; leaf blades mainly 2-4
ns 2.5 cm B.t

tepuiensis subsp. tepuiensis

104

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

2b. Lateral nerves of rpg leaf surface faintly impressed, mostly not evident on lower surface; |
.8-1.4 cm

ades 1-2 x

eaf
“epp subsp. minor

lb. Style divided into 3 anas nearly or all the way to the base, or shallowly 3- lobed at the a

a. Style shallowly 3-lobed

B. pip

3b. Style divided into 3 branches, parted halfway or more to the base.

Petals white or pin

5a. Petals 21-22 mm n long; mg subacute, 1 1-12.5 mm long; leaves obtusely acute at apex,

the larger 4-5 x 1.2-1.7 cm ...

B. bolivarensis

cA
i-a

subacute, 1.2-3.1 x 0.5-1.3 cm

. Petals 9-12 mm long; ra obtuse or rounded at apex, 8-9 mm long; Tee) acute to

6a. mihi in a terminal rosette; upper leaf surface enervate or nerves faint; petals

B. chimantensis

m broad
6b. pes imbricately extending on the branch below its tip; upper leaf surface im-

pressed-nerved; petals 9-10 mm broad

4b. Petals yellow.

toronoensis

7a. Agro elongate, 3-9 cm long, often surpassing the leaves, ebracteat
8

ds

elev
8b. Petals iiir 2.5-3.8 x

7-3 cm; lateral nerves of upper leaf surface ails impressed but
ted; upper midrib shallowly depressed B. tri

a subsp. tristyla
ll

sty
1.5-2.2 cm; lateral nerves of upper leaf eiie unequally
impressed, 6-8 pairs iba elevated; upper midrib —

B. tristyla ges nervosa

N
c

er.

. Peduncle lacking or at most 1.6 cm long, usually hidden among the leaves or mu

9a. socii linear-oblanceolate, 2.5-7 mm wide; petals 8 mm long; peduncle 8-16 m

Sea

9b. Leaves broadly ME Vasin, or oblong-oblanceolate, 8-15 mm wide venas

n vegetative shoots som

wider); peduncles 3-6 mm long.

102. Leaves broadly ons ioe acute, only slightly narrowed at the base,
len

nearly the same widt

or most of

ngth, 3.5-4.5 x 1-1.5 cm; leaves finely

teed. -nerved mnn not pale punctate beneath, stomata not manifest;

sepals 12-13 mm

B. ptariensis

o
=

manifest; sepals 9-10 mm Al

LISSOCARPACEAE

Lissocarpa "m Steyerm., sp. nov. TYPE:
NEZUELA. Territorio Federal Amazonas:
Depto. MD Cerro Marahuaca, riverine
forest upstream from “Sima” Camp, along
branch of Cano Negro, southcentral portion,
3?43'N, 65?31'W, 28 Feb.-1 Mar. 1985,
1,140 m, Steyermark & Holst 130880 (ho-
lotype, MO; isotype, VEN).

r 10-metralis glabra; foliis subcoriaceis oblon-

Sc cm latis, nervis lateralibus numerosis cena

13 mm renee floribus solitariis. bibracteatis supra ax-
illari pedicellis 3-3.5 mm
velit bins ovatis apice rotundatis 2.5 mm longis,
2-2. zm m latis; corollis cylindricis peru maturo)
m longis, basi 1.5 mm latis, medio m latis,
lobis. 4 ligulato-oblongis rotundatis 4 x 2 mm, coronae
segmentis lanceolatis acutis 1.5-2 mm longis; calyce
ig og 6 mm longo, calyce 5 mm Feud l. =
m lato, lobis 4 suborbicularibus rotun 2
es is, 2 mm latis, minute glandulari-ciliolatis; ruc
anguste elliptico-oblongo in prominentiam obtu

ong
; Leaves ias or obovate, obtuse, rounded, or subacute at apex, con-
o the m broadest above the middle, 1

1-3.5 x 0.4-1 (-
e beneath, pale punctate beneath, the stomata
B. wurdackii

panua T abrupte angustato basi rotundato 3-3.3
m longo, 1.2-1.7 cm lato, 2.2-2.5 plo longiore quam
ue

Tree 10 m tall, glabrous throughout. Leaves
subcoriaceous, dark green above, paler below,
oblong-elliptic to elliptic-ovate, obtusely acu-
minate at apex with acumen 0.8-1.2 cm long,
obtuse to subacute at base, 7.5—13 cm long, (2-)
3-5.5 cm wide, glabrous both sides; midrib el-
evated above, impressed or less elevated below;
lateral nerves numerous, faint, about equally
prominulous on both sides as the loosely tertiary
reticulate venation; petiole 5-13 mm long. Flow-
ers solitary, superaxillary or axillary on new or
old branches, pedicellate, bibracteate at base;
pedicels 3-3.5 mm long; bracts ovate, rounded
at apex, 2.5 mm long, 2-2.2 mm wide. Corolla

S

constricted basal part 2 mm long; 4 corolla lobes
ligulate-oblong, rounded, 4 mm long, 2 mm wide;
8 coronal divisions lanceolate, acute, 1.5-2 mm
long. Calyx and hypanthium 6 mm long; calyx

STEYERMARK

1987]

—VENEZUELAN GUAYANA

105

5 mm long, 1.5-2 mm wide, the cylindric tube
4 mm long, 1.5 mm wide; 4 lobes suborbicular,
rounded at apex, 2 mm long, 2 mm wide, mi-
E glandular-ciliolate on margins. Style 1.5
m long, upwardly thickened to a subtruncate-

L. stigma. Fruit yellow-green, narrowly el-

liptic-oblong, abruptly narrowed to a bluntly tri-
angular protuberance at apex, rounded at base,
3-3.3 cm long, 1.2-1.7 cm wide, the subtending
bracts more or less persistent.

This species differs from L. benthamii in the
differently shaped fruit which is 2.2-2.5 times
longer than broad and narrowly elliptic-oblong,
and in the solitary flower instead of the few- to
several-flowered, subracemose inflorescence.
Additionally, its occurrence in the montane for-
est on the slopes of Marahuaca at an elevation
of 1,140 m is in contrast to the lower altitudes
where L. benthamii has been collecte

The differences suggested by Gleason (1926)
to differentiate L. guianensis Gl. from L. ben-
thamii cannot be maintained with respect to the
more prominent upper midrib and conspicu-
ously reticulate veinlets on both surfaces, sup-
posedly characteristic of L. guianensis. Exami-
nation of material collected by Liesner from San
Carlos de Rio Negro, Venezuela, type locality for
L. benthamii, indicates variation in these char-
acters, some specimens showing prominently
raised upper midribs but with only obscure ter-
tiary venation on the ape tolar Space (Liesner
8692; Liesner & Clark 9
1117) have more jeticulain veinlets
on the lower surface. Similarly, specimens de-
termined by White as L. guianensis (Maguire
34618, 34907) show only obscure venation on
the lower surface.

The leaves of the Marahuaca specimens are
prominently reticulate-veined on both surfaces

with the upper midrib manifestly elevated. The
leaves are relatively small in size as use
with either L. guianensis or L. bentham

Although L. benthamii and L. guianensis can-
not be well separated on vegetative characters,
the larger flowers of L. guianensis may be the
best difference in distinguishing the two taxa.
Pending further collections of flowering material,
the two taxa may at present be considered as
separate species.

S

RUBIACEAE

Chomelia stergiosii Steyerm., sp. nov. TYPE:
VENEZUELA. Bolívar: Anacoco, Río Cuyuní,

entre puesto de la Guardia Nacional Acara-
bisi and Anacoco, 2 Aug. 1981, Basil Ster-
gios & Gerardo Aymard 2804 (holotype,
VEN; isotype, PORT).

Arbor 8-metralis, ramulis glabris; foliis elliptico-
ovatis vel lanceolato- ellipticis apice e» tuse acutis basi
bri e

pedunculo 2.5 cm longo glabro; floribus sessilibus; ca-

tulis vel reflexis oblongo- spathulatis apice rotundatis
3-5 mm longis (1—)1.7-1.8 mm latis, intus dimidio
inferiore sparsim pilosulis; hypanthio extus sparsim
a corolla hypocrateriformi, tubo 20 mm longo,

m lato ubique glabro, lobis lineari-ligulatis 4
mm loe 1.2-1.5 mm latis.

Tree 8 m tall, branches slender, glabrous, spines
axillary, 18-19 mm long. Leaves membranous,
elliptic-ovate to lance-elliptic, obtusely acute at
apex, acute to subobtuse at base, glabrous above,
glabrous below except barbellate in the leaf axils
and sometimes sparsely pilosulous along some
of the lateral nerves, 4.5-7 cm long, 2-4 cm wide,
with minute dark dots moderately scattered be-
neath; lateral nerves 4—5 each side, slender,
slightly sulcate above, faintly impressed below
ascending, faintly anastomosing near margin;
petiole 4-5 mm long, ciliate on upper margins,
elsewhere glabrous, canaliculate above. Stipules
triangular-ovate, acute, appressed-pubescent
without, ciliolate at apex, 2-3 mm long. Inflo-
rescence terminal, long-pedunculate, congested-
cymose with 3-6 flowers, ebracteate; peduncle
2.5 cm long, filiform, glabrous; flowers sessile,
the central one solitary and sessile, the others on
short lateral axes 0.5 mm long. Calyx lobes 4,
unequal, the two larger ones oblong-spatulate,
rounded at apex, narrowed in the basal third, 3—
5 mm long, (1-)1.7-1.8 mm wide in upper part,
| mm wide in lower third, spreading to reflexed,
sparsely minutely pilosulous within in lower half,
glabrous without, two smaller lobes ligulate, ob-
tuse, 2 mm long, 0.8 mm wide, spreading to
reflexed; hypanthium clavate-turbinate, 2 mm
ong, | mm wide above middle, sparsely pilo-
sulous with loosely spreading hairs. Corolla sal-
verform, tube 20 mm long, | mm wide just below
summit, 0.7 mm wide at base, glabrous within
and without, lobes linear-ligulate, rounded at
apex, 4 mm long, 1.2-1. wi
within, strigillose without mainly above m
Anthers slightly exserted, linear, 2 mm long, 2
mm wide, glabrous. Style 13.5 mm long, gla-
brous.

106

This species is sympatric with C. delascoi Stey-
erm. but is distinguished from that taxon in the
much longer corolla, foliaceous, manifestly un-
equal, and spreading calyx lobes, which are
sparsely pilosulous on the interior surface of the
larger pair, and larger leaves. It differs from C.
polyantha Blake in the glabrous exterior of the
corolla tube, the longer corollas, and the longer,
spreading calyx lo

I take pleasure in naming this species for Dr.
Basil Stergios, director of the herbarium of PORT,
who has. activated a well- organized. collecting

leagues, has established a
in the Venezuelan Llanos of Edo. Portuguesa.

Coccocypselum croatii Steyerm., sp. nov. TYPE:
VENEZUELA. Bolivar: vicinity of Icabaru,
4°19'N, 61?44"W, 600 m, 25 Jul. 1982,
Thomas B. Croat 54112 (holotype, MO).

, Herba radicans, caulibus repentibus 2 mm | diam.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

out. Fruit blue, 4 mm long, 7 mm wide, glabrous.

This species is related to C. condalia R. & P.
of Peru, from which it differs in the larger ovate
leaves, shorter petioles, more elongate setose cil-
la on the leaf margins, and more numerous, ar-
cuately curved lateral nerves.

Pagameopsis Steyerm.

Recent collections of Pagameopsis have ne-
cessitated a reevaluation of the specific and sub-
specific elements within the genus.

Pagameopsis maguirei Steyerm. subsp. pusillus
Steyerm., subsp. nov. TYPE: VENEZUELA.
Territorio Federal Amazonas: Depto. Rio
Negro, Cerro de la Neblina, Camp III, NW
Plateau, 13.5 km ENE of Base Camp, 0°54'N,
66°4'W, 1,750-1,850 m, 16-18 Feb. 1984,
Ronald Liesner 16013 (holotype, MO; iso-
type, VEN). Figure 8.

Suffrutex 0. 1-0. 2- metralis; foliis ad apices! ramorum

bris, subulatis elongatis 8-10 mm longis glabris, foliis
petiolatis, petiolis 3-4 mm longis, marginibus superi-
oribus setoso-ciliatis ceterum glabris; laminis obtusis
vel rotundatis 4—4.5 cm longis, 1.5-3 cm latis supra
costam mediam pilis setosis munitis atque marginibus
dense adpresso-setoso-ciliatis, ciliis 0.5-0.7 mm longis
ceterum m glabris, nervis lateralibus Wtfoque latere 7-9
arcuato nase ie
longipedunculatis ara floris, pedunculo 2. in 3. cm

pos E
anthesim 1-1.5 mm longo; calycis lobis lineari-lanceo-
latis acutis 2.5-3 mm longis, 0.7—0.9 mm latis omnino
glabris; = (immatura) coerulea 4 mm longa, 7 m

lata glab

Creeping herbaceous plant with rooting gla-
brous stems 2 mm diam.; stipular sheath gla-
brous except densely ciliate on the summit, pro-
longed into a glabrous, elongated, subulate
appendage 8-10 mm long. Leaves petiolate; pet-
ioles 3-4 mm long, upper margins setose-ciliate,
otherwise glabrous; leaf blades ovate, shortly
acute at apex, obtuse or rounded at base, 4—4.5
cm long, 1.5-3 cm wide, upper midrib setose-
ciliate, margins densely idi s setose-ciliate,
the incurved hairs 0.5-0.7 m
glabrous; lateral nerves 7-9 each
ascending, elevated below. Inflorescence long pe-
dunculate, 5-7-flowered; peduncle 2.5-3 cm.
Flowers subsessile: aa thium and calyx a
brous; hypanthium in post-anthesis 1-1.5 m
long; calyx lobes linear-lanceolate, acute, 2.5.3
mm long, 0.7—0.9 mm wide, glabrous through-

elliptici vel lan-

ceolatis apice obtusis vel subobtusis ‘die Ibis l-

2.5 cm longis, 4-7 mm latis, supra dense adpresso-

hirsutulis subtus marginibus dense hirsutulis, nervis
is d

tis |. m
hemisphericis 7-30 mm lo

dunculatis; pedunculo erecto 2: 5-4. 5 cm longo la is A
mm crasso,
ces 4-5 ramorum bre

vium inflorescentiae, quoque
glomerulo 34. floribus; genes 4—5-meris 4 mm lon-
gis, tubo lobisque extus praeter basin glabram dense
strigosis, lobis 4—5 Melee uus subobtusis 2 mm
longis intus dense lanulosis; bg den lineari-
lanceolatis acutis extus den trigoso- “hirsutis intus
glabris in sinubus una sra ie mun

Dwarf ligneous plant 0.1—0.2 m tall. Leaves
densely crowded in an apical rosette, narrowly
lance-elliptic or lanceolate, narrowed to an ob-
tuse or subobtuse apex, narrowed to a subsessile
base, 1-2.5 cm long, 4-7 mm wide, densely ap-
pressed-hirsutulous above, densely hirsutulous
below and on margins; midrib elevated below,
not evident above; lateral nerves not manifest;
petiole 2-3 mm long, scarcely distinguishable
from the leaf base. Stipular sheaths closely

4—5 short axes, the lowest axes 8-15 mm long,

107

STEYERMARK— VENEZUELAN GUAYANA

1987]

EY

3
te
a tte
wok Pees Se ae Mee

ig tror mE Y Ll
ris,
=,”

` SZ,
x?
> Á

X

> “

x, 17

I»

es

yea ore
ee

tes!

FIGURE 8. Pagameopsis maguirei subsp. pusillus. — A. Habit.—B. Flower, with subtending bracts.—C. Co-

rolla, interior view.—D. Calyx and hypanthium, with pistil detached from calyx, interior view.

108

densely hirsute; rachis with 1-3 pairs of hori-
zontally spreading bracts 3 mm long, 1 mm wide,
hirsutulous without, glabrous, the inflorescence
bibracteate at base with the bracts foliose, linear
to oblong-lanceolate, subobtuse, 8-15 mm long,
2-3 mm wide, densely pubescent above, sparsely
or moderately so below. Peduncle 2.5-4.5 cm
long, 1-1.5 mm thick. Flowers in dense nearly
sessile glomerules of 3-4 at the ends of short axes;
2 bracts subtending the base of the calyx navic-
ular, oblong-lanceolate, 1.5-1.8 mm long, 0.8-1
mm wide, glabrous and glandular within. Calyx
lobes slightly unequal, , linear-lanceolate,
acute, 1.5-2 mm long, 0.5-0.6 mm wide, dense
strigose-hirsute without and on margins, gla-
brous within with 1 gland between the sinuses
of the lobes. Corolla 4 mm long, densely strigose
without except where glabrous at base; tube 2
mm long; lobes 4-5, ligulate-lanceolate, subob-
tuse at apex, 2 mm long, densely lanulose within.

This taxon differs from the other known taxa
of Pagameopsis in the very small, narrow, dense-
ly hirsute leaves and dwarf habit. It is closely
related to P. maguirei Steyerm. subsp. neblinen-
sis in having apical leaf clusters, but differs in the
smaller alone m with 3-5 axes, and in the
dense pubescen

abrous im of subsp. pusillus has been

found on nearby Cerro Avispa of Venezuela.

Pagameopsis maguirei subsp. pusillus var. gla-
brus Steyerm., var. nov. A subsp. pusillus
var. pusillus foliis glabris recedit. TYPE:
VENEZUELA. Territorio Federal Amazonas:
summit of Cerro Avispa, Río Siapa, 1?30'N,
65°51'W, 1,510 m, 5 Dec. 1972, Cardona
3096 (holotype, NY; isotype, VEN)

Another variation of Pagameopsis maguirei
from adjacent northern Brazil is here described.

KEY TO THE SPECIES, SUBSPECIES,

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Pagameopsis maguirei subsp. neblinensis Stey-
erm. var. pirapucuensis Steyerm., var. nov.
TYPE: BRAZIL. Serra Pirapucü, 1,300 m, 26
Jan. 1966, Nilo T. Silva & Umbelino Brazao
60880 (holotype, NY).

Frutex; foliis caulinis dispersis haud omnino api-
calibus, longitudinem caulis 2.5-4 cm oc

occupantibus,
oblanceolatis apice subacutis 8-8. 5 cm longis, 1.3-1.5
cm vel saltem 1 parte
costa media

inferiore costae mediae pilosis, subtus
c

-10mm
tis, moderatim sir pud strigosis, pee
dense hirsutis.

Shrub; leaves arranged along the uppermost
length of the stem, not wholly apical, oblanceo-
late, subacute at apex, 8-8.5 cm long, 1.3-1.5 cm
wide, appressed-pubescent above or at least pi-
lose on the lower part of the midrib, abundantly
substrigose or pilose below, otherwise sparsely
puberulous; petioles abundantly strigose or sub-
strigose throughout. Stipular sheaths elongated,
7-10 mm long, wide, moderately to
densely strigose, the summit densely hirsute.

This taxon differs from the other varieties of
subsp. neblinensis in the more densely strigose
stipular sheaths, which are more densely hirsute
at the summit. From var. neblinensis it differs in
having the leaves dispersed along the stem length
or 4 cm, and in the pilose petioles and lower
surface. From subsp. neblinensis var. angustifoli-
us it differs in the leaves pubescent below and
above at least along the midrib or surface. From
subsp. maguirei var. maguirei it may be differ-
entiated by the pubescent upper midrib and low-
er leaf surface.

In order to accommodate the above newly de-
scribed taxa, the following revised key to Pa-
gameopsis is offered.

AND VARIETIES OF PAGAMEOPSIS

la. Verte: aiia spaced along the upper part of n for 2.5-11 cm; stipular sheath elongated, usually

longer than broad or as DR as broad, POD
2a. aa ‘of leaf margins

EE.

haire di 4:

very e
0.5—1 mm long; inflorescence with ner 3-5 densely flowered eee on i2 pairs of lateral
l.

axes, the whole inflorescence 1.5-4 cm

ong, 2-4 cm

wide; leaves mainly 175-3!4 times longer
P. ee

N
og

. Cilia of leaf m
long; inflorescence branche
the

numerous

margins papi 1 or deniers to ascending, less than 0.5 m
glom

mall rules borne on 7 or more branched axes,

whole inflorescence 5— TE. cm long, 3-6 cm wide; pen mainly (2%-)3—6 times longer than

broad
3a. Lower d surface glabrous Or mainly so, sometimes pie upper leaf surface glabrous 4

a. Ud5 »511U1

ti lobe nd mm 1 long, i 1. 1.25 mm wide .

lobe S GCNSC Milivilvuss

NS P. naue subsp. maguirei var. maguirei

4b. Petiolar base Meade above, gl

to sparsely pubescent below; interior of calyx lobes

1987]

STEYERMARK — VENEZUELAN GUAYANA

109

with short strigose pubescence, the exterior with pies hirtellous pubescence; corolla

2.2-3 mm long, 0.7-1 mm wide

lo
3b. Lower leaf surface pubescent; upper |

magui rei su € neblinensis var. Mise E
labrescent, the midrib pubesc

maguirei ake sp. neblinensis var. pirapucuensis

lb. Leaves apically ee = the be AR 0.5- d 5 cm length of stem; stipular sheath c

tracted, m than lon

s broad as long, 1-6 mm lon

5a. Leaves 2-10 cm ae 0. 7-1. 8 cm wide; bracts subtending base of inflorescence ve "d mm long,
4T mm wide; inflorescence branching into numerous sap glomerules borne in 7 or more
branched axes, the whole inflorescence 5-11 cm long, 3 m wide; shru

cA
Ss

long, 2-3(-5) mm wide; inflorescence with o

pairs of lateral axes, the whole inflorescence 0. T d cm

er
. Leaves 1-2.5(-3) cm long, 0.4—0.7 cm wide; Pra ts subtendin g base

y 3-5 yaspa i glomerules borne on 2
long, 2-3 cm wide; d ucl ligneous plant

6a. Upper and lower leaf surfaces densely hirsute „u P. maguirei subsp. pusillus var. pusillus

6b. Upper and lower leaf surfaces glabrous

P. maguirei subsp. pusillus var. glabrus

Psychotria guanchezii Steyerm., sp. nov. TYPE:

Guanchez 3644 (holotype, VEN;
TFAV). Figure 9.

isotype,

Suffrutex 1.5-metralis, ramis glabris; stipulae vagina
uobus

16-20 utroque latere divuehis pate ibus; inflores-
centia terminali pedunculata, pedunculo gracili erecto
3 cm longo, 1 mm lato minute sparsimque puberu-

puberul entibus; bracteis sub fasciculis florium ligulatis
vel lanceolatis 2.5-3 mm longis utrinque minute den-
seque puberulentibus; calyce hypanthioque 1.5-2 mm
opan; calycis lobis deltoideis obtusis 0. 3 mm longis,
m latis; i 5—6 mm lon-

a extus praeter basim glabram ipsa dense puberulenti
pilis patentibus 0.1-0.2 mm longis munita "i dimi-

a parte superiore puberula ceterum glabr:

Slender subshrub 1.5 meters tall, branches gla-
brous. Stipular sheath 2—2.5 mm long, 1.8-2 mm
wide, glabrous, terminating on each side in 2
subulate minutely ciliolate teeth 3 mm long. Leaf

base, glabrous both sides, 4—9.5 cm long, 0.8-2
ide, midrib elevated ni impressed
bove; lateral nerves 16—20 side, divari-

cately spreading at an angle of 5— ioe faintly anas-
tomosing at 1-2 mm from margin; petiole 3—9
mm long, glabrous. Inflorescence small, cy-

mosely umbellate, terminal, 1.2-1.8 cm long, 1.3-

wide, on erect, slender peduncle 4—4.3
cm long, 1 mm wide, minutely and sparsely pu-
berulent; axes of inflorescence 4, three lateral ones
spreading, 4-6 mm long, 0.5-1 mm wide, one
central axis erect and longer, 7-8 mm long, 1
mm wide, all moderately to abundantly puber-
ulent with spreading unequal puberulence, the
longest hairs 0.1 mm long; 3 lateral axes 5—6-
flowered, 4th axis 7-8-flowered. Bracts subtend-
ing base of inflorescence divaricately spreading,
subulate, 2.5-2.7 mm long, 0.3 mm wide, mi-
nutely and moderately puberulent; bracts sub-
tending flower clusters ligulate to lanceolate, acute
to obtuse, 2.5-3 mm long, 0.7 mm wide, mi-
nutely densely puberulent on both sides. Calyx
and hypanthium 1.5-2 mm long; hypanthium
shallowly campanulate, l x 1 mm; ca m

bt wide. Corolla
Infundibuliform, 5—6 mm long; aie 2.2-3.5 mm
ng, 0.8 mm wide at base, 1.3 mm wide at ori-
fice, dency puberulent, except at very base, with
short spreading hairs 0.1—0.2 mm long, pubes-
cent in upper half within from staminal insertion
to orifice, elsewhere within glabrous; lobes 4, lan-
ceolate, subacute, 1.7-2.5 mm long, 1.3 mm wide,
densely puberulous without. Anthers linear, 1.1—
1.2 mm long, included in uppermost part of co-
rolla tube; filaments 0.8 mm long, inserted half-
way up corolla Tube Style 4.2 mm long, aah w ü
d, dilated, exserted, 0.8 m
This species is related to P. capitata ^"; & P.
but is reduced in the size of all parts. Moreover,
the densely puberulous calyx and minute, puber-
ulous corollas, small inflorescence with slender
peduncle, and narrow leaves are noteworthy dis-
tinguishing characters of the taxon. From P. pi-
resii Steyerm. of Amapá, Brazil, it is differen-

d =]

110 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

4mm

yi!

À Ade dad j
key "tul ,^ i
UE L. fed

C

FiGURE9. Psychotria guanchezii. — A. Habit. — B. Corolla. — C. Calyx and hypanthium, with position of disk
indicated. — D. Corolla, pre-anthesis.

1987]

tiated by the smaller, 5-merous corollas, shorter
stipular teeth, and smaller inflorescence wit
fewer axes.

Psychotria ronaldii Steyerm., sp. nov. TYPE:

VENEZUELA. Territorio Federal Amazonas:

Depto. Atabapo, Cerro Huachamacari, for-

ested slope, 3?39'N, 65?42'W, 600-750 m,

4 Mar. 1985, Ronald Liesner 18214 (holo-
type, MO; isotype, VEN). Figure 10.

Frutex 2-metralis, caulibus glabris; stipularum va-

lateralibus 4—6 patent

briato- PRA pep pe edun culo erecto 1.8—2.5
l-

longo, mm crasso glabro; floribus sessilibus
cient s" videtur n ullis; h mm
longo glabro, calyce breviter 5- lobato, dentibus late

deltoideis vel fere truncatis; corolla subinfundibulifor-
.5 mm longa extus glabra, intus infra insertionem
“Yuhu ways pilosulo ceterum glabro, lobis 5.

Shrub 2 m tall; stems glabrous. Stipular sheaths
broadly semicircular, rounded or slightly acute
at apex, buff-pubescent within and on the apical
margins, 2.5 mm high, 5 mm wide. Leaves
broadly elliptic-obovate, caudate-acuminate at
apex with acumen 2 cm long, acutely narrowed
to a decurrent base, 22-30 cm long, 8-10.5 cm
wide, glabrous both sides; lateral nerves 14-17
each side, widely spreading at an angle of 15°
20°, elevated below, tertiary venation beneath
finely grossly reticulate. Petioles 1—1.5 cm long,
glabrous. Inflorescence terminal, paniculate,
subhemispheric, slightly broader than long, 2 cm
long, 2.5-3 cm wide at base; lateral axes divar-
icately spreading, the lower ones 7-8 mm long,
the upper 2-4 mm long; base of axes with fim-
briate-pubescent scars; axes terminating in 7—10-
flowered clusters. Peduncle erect, 1.8—2.5 cm long,
1-1.5 mm thick, glabrous. Flowers in groups of
7-10. Calyx and hypanthium 2.1 mm long, gla-
brous; hypanthium | mm long, | mm wide; calyx
| mm long, 1.2 mm wide, the border shallowly
toothed; calyx teeth broadly deltoid to nearly
truncate. Corolla cream-colored, subinfundibu-
liform, 4.5 mm long, 2 mm wide above, | mm
wide at base, glabrous without, glabrous within
except for a pilosulous zone at the antheriferous

STEYERMARK— VENEZUELAN GUAYANA 111

level; lobes 5, a lanceolate, subacute, 1.5
mm long, | mm w

The immediate <n of this taxon with
other species of the genus is not apparent. The
species 1s characterized by the small corollas, gla-
brous leaves, inflorescence, and stems, and fim-
brillate pubescence of the scars on the inflores-
cence axes.

The species is named for Ronald Liesner, who
made important collections on Cerros Huacha-
macari and Marahuaca during the expedition in
1985 sponsored by the Terramar Foundation.

Remijia marahuacensis Steyerm., sp. nov. TYPE:
VENEZUELA. Territorio Federal Amazonas:
Depto. Atabapo, Cerro Marahuaca, “Sima”
Camp, southcentral portion of forested slopes
along eastern branch of Cano Negro, 3?43'N,
65°31'W, 1,140 m, 21-22, 24 Feb. 1985,
Julian A. Steyermark & Bruce Holst 130440
(holotype, MO; isotype, VEN). Figure 11.

Frutex 2 m altus, ramis dense adpresso- uq
foliis anguste lanceolatis vel lineari-lanceolatis apice
d brem acuti Is vel subac uti IS 11-18 cm
longis, 2 ded. m lat t

cdia s lat alibu usque adpresso- pubescentibus
axillibus barbellabs. ceterum glabris, nervis lateralibus
r petiolis 0.5-1.5 cm longis ad-
: Binforeetenta axillari sub an-
thesi capitata 2 cm longa, 1.5 cm lata bracteis se ses
oblongo-ovatis obtusis vel subobtusis l. 7-2c m longi

8

.8—1.1 cm latis extus adpre ent

ta; infructescencia breviter racemosa 2—4 cm lon

cm lata elongata; rhachidi dense hirsutula; pedun bulo

6- 15 cm longo, 2 mm lato, dense adpresso- hirsutulo;
pedi-

cellis 1.5 mm longis dense hirsutulis, sub fructu 5-7
mm hara nee hypanthioque 4-5 mm longo, hy-
panthio 3-4 mm longo; calyce apice fere truncato extus
sericeo- sirigoso. i rag glabro-eglanduloso; corolla 15-
2l m lon nga, o 10-15 mm | longo, 2. 5 mm lato,

tied

to, lobis 5 lanceolatis subobtusis 5-6 mm longis extus
3

tis ligulato-oblongis extremitatibus rotundatis 5 mm
longis, corpusculo ovali 1.7-2 mm longo, 1.5 mm lato,
alis late oblongis rotundatis 1.5—2.5 mm longis, 1.5
mm latis.

Shrub 2 m tall, young stems densely appressed-
hirsutulous. Stipules ligulate-lanceolate, sub-
acute, 1.5-3 cm long, 4-7 mm wide, appressed
pubescent without, midrib densely pubescent.

ceolate, long acuminate at apex, acute to sub-

112

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

4.5mm

Tr Mn o Aon nts

2.1mm

B

C D

IGURE 10. Psychotria ronaldii. — A. Habit.—B. Corolla. —C. Corolla, interior view. — D. Calyx and hypan-

F
thium, with position of disk indicated.

acute at base, 11-18 cm long, 2.5-5 cm wide,
glabrous above on mature leaves, lower surface
appressed-pubescent on midrib and lateral
nerves, barbellate in axils of nerves beneath, low-
er surface otherwise mainly glabrous; lateral
nerves 9-11 each side, sulcate above, elevated
below, ascending at an angle of 45—50?, curving
into the margin, tertiary venation finely reticu-
late below. Inflorescence axillary, capitate in an-

thesis, 2 cm long, 1.5 cm wide, 6-10-flowered,
subtended by 2 foliose, oblong-ovate, obtuse or
subobtuse bracts 1.7—2 cm long, 0.8-1.1 cm wide,
appressed-pubescent without. Infructescence ra-
cemose, 2-4 cm long, 3 cm wide, rachis densely
hirsutulous, up to 1.5-2 cm long in fruit. Pe-
duncle 6-15 cm long, 2 mm thick, densely ap-
pressed-hirsutulous. Flowers heterostylous, short-
pedicellate, in anthesis pedicels 1.5 mm long, in

113

STEYERMARK — VENEZUELAN GUAYANA

1987]

E E
E iD
ki 15mm
| F
FIGURE ll. Bite marahuacensis. — A. Habit. — B. Calyx and hypanthium, external view. —C. Calyx and
hypant nthium with disk, internal view.— D. Corolla. — E. Short-styled corolla. — F. Long-styled corolla. — G. Fruit-
ing capsule dus dehiscence. — H. Seed.

114

fruit 5-7 mm long, densely hirsutulous. Calyx
and hypanthium short-cylindric, 2.5 mm wide,
densely lanuginose without with cream-colored,
erect-ascending hairs; calyx 1-1.3 mm long, 2.5
mm wide, truncate or shallowly cupulate, stri-
gose-sericeous without, ciliate on margins, gla-
brous and eglandular within. Corolla pink, sub-
hypocrateriform, densely bufflanuginose without,
15-21 mm long, tube 10-15 mm long, 2.5 mm
wide, glabrous within; lobes 5, lanceolate, subob-
tuse, 5-6 mm long, 1.2-2 mm wide. Anther
linear, 5.3 mm long, in short-styled flowers in-
serted in the upper half of the corolla tube, in
long-styled flowers inserted 2-3 mm above base
of corolla tube. Style 5.5-9 mm long. Mature
capsules cylindric, 1.5-2.3 cm long, 5-6 mm wide,
appressed pubescent. Seeds ligulate-oblong,
winged, 5 mm long, the body pale brown, oval,
1.7-2 mm long, 1.5 mm wide, wings broadly
oblong, rounded, 1.5-2.5 mm long, 1.5 mm wide.

o

Distribut tion. Forested slopes of Cerro Mara-
uaca, Territorio Federal Amazonas, Venezuela,
at an altitude i 1,140-1,220 meters.

atype. Cerro Marahuaca, Upper Rio Yame
danke. 3°38'N, 65?28'W, 1,225 m, 22 Feb. T nes
ner 17818 (MO, VEN).

This species is related to R. maguirei of Cerro
de la Neblina, from which it differs in the smaller
seeds, much shorter corolla, eglandular interior
of calyx tube, glabrous petioles, lower leaf sur-
face, and densely pubescent branches.

Of the 20 species of Remijia presently known
from the Guayana, six are endemic to different
sandstone table mountains (R. pilosinervia from

himantá, R. maguirei from Neblina, R. argen-
tea from Yapacana and Moriche, R. steyermarkii
from Duida, and R. roraimae from Roraima and
Chimantá). Remijia densiflora is widespread on
several table mountains and surrounding areas
of Bolívar and Territorio Federal Amazonas. Re-
mijia marahuacensis becomes the seventh species
to add to the list of endemic taxa of this genus.

Schradera marahuacensis Steyerm., sp. nov.
TYPE: VENEZUELA. Territorio Federal Ama-
zonas: Depto. Atabapo, Cerro Marahuaca,
“Sima” Camp, southcentral portion of for-
ested slopes along eastern branch of Cano
Negro, 3?43'N, 65?31'W, 1,140 m, 25 Feb.
1985, Julian A. Steyermark & Bruce Holst
130721 (holotype, MO; isotype, VEN).

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Suffrutex epiphyticus qms stipulis non visis; fo-
liis petiolatis 5-16 mm s; laminis coriaceis ob-
longo-ellipticis apice one Ad subobtusis basi acutis
9.5-13 cm longis, 2.5-5.5 cm latis, costa media subtus
elevata supra paullo anguste sulcata, nervis li:
utroque latere 11-12 adscendentibus subtus subpro-
minulis supra obsoletis; capitulis hemisphaericis 10-
20-floris 2.5-3 cm diam., 1.5-2 cm

solitario terminali 3.5-5.5 cm

o

latis; calyce hypanthioque 10 mm longis, 4-5 mm

calyce cylindrico- -campanulato breviter lateque ee
12m

ideo-dentato

longa extus glabra intus pilosula, lobis 5 at
subacutis 5 mm longis, m latis; staminibus
prope basin tubi corollae insertis.

Woody glabrous epiphyte. Leaf blades coria-
ceous, oblong-elliptic, obtuse to subobtuse at
apex, acute at base, 9.5-13 cm long, 2.5-5.5 cm
wide; midrib elevated below, slightly narrowly
sulcate above, lateral nerves ascending, 11-12
each side, faint or at most slightly prominulous
below, obsolete above. Petioles 5-16 mm long.
"PER RM hemispheric, 10—20-flowered, 2.5-
3 cm diam., 1.5-2 cm high. Peduncle solitary,
terminal, ery 5.5 cm long, 2-2.5 mm wide,
slightly dilated below summit. Involucre 7-8 mm
high, lobed, lobes suborbicular, 8 mm long, 10
mm wide. Calyx and hypanthium 10 mm long,
4-5 mm wide, calyx cylindric-campanulate,
shortly and broadly deltoid-toothed or subtrun-
cate. Corolla fleshy, cylindric, 12 mm long; tube

mm long, pilosulous internally; lobes 5, lan-
ceolate, subacute, 5 mm long, 1.8-2 mm wide.
Stamens 5, inserted near base of corolla tube;
anthers linear, 3.5 mm long, 0.7 mm wide. Pistil
6.5 mm long; stigmas 2, linear.

Distribution. Known only from forested
slopes of Cerro Marahuaca, 1,140-1,225 m, Ter-
ritorio Federal Amazonas, Venezuela.

Paratype. VENEZUELA. TERRITORIO FEDERAL
AMAZONAS. Cerro Marahuaca, upper Rio Yameduaka
3°38'N, 65?28'W, 1,225 m, 22 Feb. 1985, Liesner 17818
(MO, VEN

Sipanea setacea Steyerm., sp. nov. TYPE:
VENEZUELA. Bolívar: Dtto. Piar, Cerro El
Venado, 20 km E of Canaima, 6?17'N,
62°41'W, 1,300 m, 31 Aug. 1983, Otto Hu-
ber, G. T. Prance & C. Alarcon 8246 (ho-
lotype, NY).
rba repens, caulibus tenuibus dense pilosulis pilis

H
patentibus 0.2 mm longis praeditis; stipulae appendice
setiformi 2-3 mm longa; foliorum laminis ovatis vel

1987]

elliptico-ovatis m acutis basi obtusis vel subacutis
.2-2 x 0.9-1.2 cm utrinque dense subadpresso-pi-
losis, deca 3.5-4. 5 mm longis; inflorescentia pedun-
culata triflora, pedunculo terminali 1-1.7 cm longo
delice Be pilis patentibus praedito; floribus sub-
sessilibus, pedicellis ad 0.5 mm longis; corolla pur-
purea 7.5 mm longa, tubo 4 mm longo, lobis 3.5 mm
longis extus pilosulis.

Herbaceous plant; stems creeping, slender, 0.5
mm diam., densely pilosulous with spreading or
loosely ascending hairs 0.2 mm long; internodes
1-1.8 cm long. Stipules setaceous, 2-3 mm long,
0.1 mm wide, densely pilosulous. Petioles 3.5—
4.5 mm long, 0.5-0.7 mm wide, densely pilo-
sulous with spreading hairs. Leaf blades ovate to
elliptic-ovate, acute at apex, obtuse to subacute
at base, 1.2-2 cm long, 0.9-1.2 cm wide, densely
subappressed-pilose both sides with hairs 0.2—

5 ong. Inflorescence terminal, peduncu-
late, 3-flowered, the peduncle slender, 1-1.7 cm
long, densely pilosulous with spreading hairs.
Flowers subsessile; pedicels to 0.5 mm long.
Bracts subtending inflorescence 2, subulate, acu-
minate, 2.2 cm long, 0.1 cm wide, glabrous with-
in, setose-ciliate with hairs 0.3-0.5 mm long. Ca-
lyx lobes subulate or subsetaceous, 2.2-2.5 mm
long, 0.3-0.4 mm wide, densely pilosulous with
hairs 0.4 mm long; hypanthium pyriform, dense-
ly villosulous with spreading white, non-tuber-
culate hairs 0.4-0.5 mm long. Corolla “purple”
(fide Huber et al.), subsalverform, 7.5 mm long,
the tube 4 mm long, shortly pilosulous with hairs
within near the base of the stamens; lobes nar-
rowly oblong, obtuse, 3.5 mm long, 1.2 mm wide,
moderately pilose without. Anthers linear-ob-
long, 1.3 mm long, 0.3 mm wide. One setiform
squamella 0.15—0.2 mm long situated in each
sinus of the calyx lobes.

n its setaceous stipules and pubescent exterior
ofthe corolla lobes, this species differs from other
members of sect. Virecta subsect. Cryptotricha,
but resembles them in habit, slender stems, and
small leaves. From S. ovalifolia Bremek. var.
ovalifolia and var. villosissima Steyerm. it differs
in the shorter peduncle and petioles, longer co-
rolla and corolla lobes, 3-flowered inflorescence
on a shorter peduncle, shorter, non-tuberculate
hairs of the hypanthium, narrower leaves, and
purple instead of white corollas. From S. mi-
crantha Sandw. it may be distinguished by the
spreading pubescence of stems and peduncles,
broader ovate leaves with pubescent upper sur-
face, loosely pubescent calyx lobes, and 3-flow-
ered inflorescence. From S. gleasonii Steyerm. it

STEYERMARK — VENEZUELAN GUAYANA

115

is differentiated by the subsessile flowers, shorter
petiole, calyx lobes, hypanthium, and corolla
tube, smaller leaf blades, shorter foliar pubes-
cence, and non-tuberculate hairs of the hypan-
thium. Finally, from S. cowanii Steyerm. it dif-
fers in the densely spreading hairs of the
hypanthium, the shorter corolla and calyx lobes,
and the 3-flowered inflorescence.

COMPOSITAE

Gongylolepis terramarae Steyerm., sp. nov. TYPE:
VENEZUELA. Territorio Federal Amazonas:
Depto. Atabapo, Cerro Marahuaca, forested
steep sandstone SE-facing slopes and bluffs,
above branch of Caño Negro, southcentral
portion of meseta, downstream from “Sima”
Camp, 3°43'N, 65°31'W, 1,220-1,350 m, 23-
24 Feb. 1985, Julian A. Steyermark & Bruce
Holst 130629 (holotype, MO; isotype, VEN).

Arbor 3-4-metralis, caulibus vegetativis ad apices
lanulosis, internodiis infra pet Th os atque ad nodos mi-
nus lanulosis, caulibus floriferi
sim pilosulis vel glabris; foliis findet, ellipticis Mis
itis au apice obtusis basi d ad petio

nge decurrentibus 9-24 cm longis, 3-6 atis, cione
x media laxe pilosa vel ient. panes costa
media adpresso-pilosa pilis elongatis munita ceterum
utrin nque sparsim adpresso- pilosis vel glabris, venulis

pedunculis corymbosis 1.5-3 cm longis bracteatis
breviter pubescentibus; capitulis aliquot campanulatis
maturis 3-3.5 cm longis, 2-2.5 cm latis 21-floris; in-
volucro 3 cm longo 5-6- seriato, phyllariis glabris ap-
li bracteis ova-
tis apice acutis 2 5- 4 mm longis dorsaliter puberulis
vel glabrescentibus; corollis bilabiatis, tubo 11 mm
longo, 3 mm lato, lob
3.5 mm la to, lobo
bus; achaeniis co linearibus nigris 9.5 mm longis,
1.4 mm latis costat

Tree 3-4 m tall; tips of vegetative stems lan-
ulose, the internodes below the petioles and at
the junction of the petioles sparsely lanulose, the
flowering and older stems sparsely pilosulous to
glabrous. Leaves coriaceous, dark green above,
pale green below, lance-elliptic to oblanceolate,
obtuse at apex, narrowed to an acute long-de-
current base, 9-24 cm long, 3—6 cm wide, midrib
above loosely pilose to glabrescent, below usually
appressed pilose with elongate hairs, elsewhere
sparsely pilosulous to glabrous on both surfaces;
lateral nerves at an angle of 35—45°, faint on both
sides for 75—À distance to the margin, becoming
obsolescent; tertiary venation beneath scarcely

116

or faint only, above slightly or more evidently
reticulate. Petiole on flowering shoots not de-
veloped, up to 10 cm long on vegetative shoots,
sparsely puberulous to lanulose at the base. In-
florescence corymbosely branched, the pedun-
cles 1.5—3 cm long, bracteate, shortly pubescent.
Heads several, campanulate, mature ones 3-3.5
cm long, 2-2.5 cm wide, 21-flowered, on brac-
teolate pubescent pedicels 1.5-4 cm long; brac-
teoles ovate-lanceolate, acute, 3-3.5 mm long,
dorsally e Bode P: 3 cm long,
5—6-seriate, outerm e, 4-5 mm long,
2.5-3 mm wide, dE ones a -oblong,
rounded at apex, 12-13 mm long, 5-6 mm wide,
innermost linear-ligulate, rounded at apex, 23-
30 mm long, 4 mm wide. Bracts of peduncle
ovate, subacute at apex, 2.5-4 mm long, dorsally
more or less appressed-puberulous to glabres-
cent. Corollas white, tube 11 mm long, 3 mm
wide, glabrous, external lobe ligulate, 14 mm long,
3.5 mm wide, minutely 3-lobed at apex, lobes
rounded; inner lobe 12-13 mm long, 2-parted
with linear subacute segments 1 mm wide. An-
ther 11-12 mm long. Pappus buff, 15-17 mm
long, setae numerously barbellate. Achene black,
narrowly linear, 9.5 mm long, 1.4 mm wide, cos-
tate.

Paratype. VENEZUELA. Cerro Marahuaca, slope be-
Upper Río Yameduaka and base of cliff, 3°38'N,
65°28'W, 1,225-1,400 m, Liesner 17751 (MO, VEN).

This species resembles G. huachamacari Ma-
guire & Wurd. of Cerro Huachamacari, from
which it differs chiefly in the smaller flowers,
greatly reduced peduncular bracts, and larger

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

leaves with pubescent midribs. From G. pedun-
culata Mag. & Wurd. of cerros Part and Huacha-
macari it differs in the smaller, less numerously
flowered heads, corymbose inflorescence, more
reduced size of the peduncular bracts, pubescent
midrib, and narrower involucral bracts; while
from G. yapacana Maguire & Wurd. of Cerro
Yapacana, it differs especially in the absence of
large areolate, strongly reticulate venation, and
shorter peduncles.

LITERATURE CITED

EICHLER, A. . ns In Martius (editor),
Flora Brasiliensis 13(1): 4

GLEASON, H. 1926. Studies on M flora of northern
South America — IX. Bull. Torr. Bot. Club 53: 296.
1931. Botanical Sle a the Tyler-Duida

expedition. Bull. Torr 8: 389.

Lasser, T. 1946. Leitgebia — Bol. Acad.
Ci. Venez. 9: 246.

1976. Revision and taxonomic position

Martius, C. F. P. &
Genera et Species Plantarum. Flora 7( 2.
SASTRE, C. 1970. Recherches sur les Ochnacées. Cal-

dasia 10: 570.
1986. Deux Ochnacées nouvelles du Vene-
uela. Phytologia 59: 313-314.
See R.H. 1847. Beischreibung dreir neuen
pflanzen aus dem flussgebeite des Carimani oder
rh mango, p zuflussen des Mazaruni. Lin-
aea 20: 751-760.
E J. Eb jos del Auyan-tepui. Acta
Bot. Venez. 2(5-8): 2
1984. Flora of Td "Venezuelan Guayana —I.

Ann. Missouri Bot. Gard. 71(1): 326-327.
WARMING, E. 1875. Trigoniaceae. 3 Piana (edi-
tor), Flora Brasiliensis 13(2): 118-

A NEW SPECIES OF JATROPHA (EUPHORBIACEAE)
FROM NICARAGUA!

GRADY L. WEBSTER?

ABSTRACT

A new species, Jatropha stevensii, is described from collections in Dept. Boaco in — ge It may

be assigned to subg. Curcas and ap
Mexico. Jatropha stevensii represents yet an

pears related to Jatro
other extension of the Mexican sri ee element into

pha fremontioides, a species from southern

Central America, and the second record for Jatropha subg. Curcas

When Jatropha costaricensis Webster & Po-
veda was described from Guanacaste, Costa Rica,
some years ago (Webster & Poveda, 1978), it was
predicted that other xeric relict taxa might be
discovered elsewhere in Central America. Recent
botanical exploration in Nicaragua has disclosed
not only a number of new populations of Jatro-
pha podagrica Hook. in rocky areas in Dept. Es-
teli, but also on basaltic ridges in Dept. Boaco a
species that appears to be undescribed. Unlike
J. podagrica, which belongs to subg. Jatropha
(Dehgan & Webster, 1979), the plant from Boaco
clearly is referable to subg. Curcas.

Jatropha stevensii Webster, sp. nov. TYPE: Nic-
aragua, Dept. Boaco, 1.6 km SW of Santa
Cruz, low ridge of basaltic lava, 140-160 m,
12?24'N, 85?50'W, 7 June 1984, W. D. Ste-
vens 22902 (MO, holotype; DAV, isotype;
additional isotypes to be distributed). It
should be noted that the type collection in-
cludes both glabrous and pubescent speci-
mens and was apparently gathered from sev-
eral individuals.

Ab J. olivacea differt dichasio menor, minore stipulis
obsoletis, foliis integris marginibus non glanduligeris;
ab J. alamanii foliis elobatis, sepalis minoribus, petalis
tantum !^ longitudinis coalitis; ab J. fremontioide foliis
acuminatis, dichasio non capituliforme.

Deciduous shrub to 3 m high; twigs terete,
greyish- or reddish-brown, glabrous or hispidu-

minate tip, cordate at base, loosely tomentose to
glabrous beneath, 2-4 cm long, 1—3 cm broad,
palmately 5-veined at base, brochidodromous,

beneath midrib and secondaries raised, veinlets
prominent; margins entire, eglandular. Monoe-
cious; dichasia terminal, often paired, with pe-
duncle 1-2.5 cm long, branches hispidulous;
bracts oblong-lanceolate to oblanceolate, gla-
brous or tomentulose, lower ones 4-5 mm long
and 1.2-1.3 mm broad; pistillate flowers 0—2 per
dichasium, at lower nodes; staminate flowers 10—
15, at upper dichotomies. Staminate flower: ped-
icel 0.5-0.8 mm long, articulated at base; sepals
5, lanceolate, obtuse or subacute, entire, glabrous
or hirsutulous, 2.5-4.8 mm long, 1.2-2 mm
broad; petals 5, pale green, oblong, glabrous
abaxially, copiously hirsutulous adaxially in low-
er half, 5.5-6 mm long, coherent into a tube in
lower !^, midrib with several sharply ascending
laterals that dichotomize distally; disk glands 5,
cubical, glabrous, 0.5—0.6 mm high; stamens 10,
biverticellate, monadelphous below into a slen-
der column 3-4 mm high, outer stamens on fil-
aments ca. | mm long. Pistillate flower: pedicel
glabrous or tomentulose, becoming 3-1 1 mm long
in fruit; sepals 5, elliptic or lanceolate, obtuse or
subacute, becoming in fruit 5.5-6.5 mm long,
2.5-3.8 mm broad; petals 5, ovate at base ta-
pering to oblong obtuse tips, pale green, ca. 5.8—
6 mm long, 4—4.2 mm broad, hirsutulous adax-
ially, coherent in lower !^; disk dissected into 5
lobes; ovary smooth, glabrous, sharply carinate,
3-locular; styles ca. 3 mm long, connate in lower
l^, erect, scarcely bifid; stigmas dilated, elliptic,
0.7 mm long, 0.3 mm broad. Capsule smooth,
3-lobed, prominently 3-carinate on back of cocci,
1.5-1.6 mm in diameter; seeds ellipsoidal, ca. 9
mm long, smooth; caruncle brownish, deeply fla-
bellately lobed, 2.5 mm long, 4 mm broad.

Additional collections examined. NICARAGUA
BOACO:1 km E of Santa Cruz, 200 m, Moreno 22467B

! I wish to thank Ms. Lynn Gillespie for the photograph of the type collection.
? Department of Botany, University of California, Davis, California 95616

ANN. MISSOURI Bor. GARD. 74: 117—120. 1987.

118

1 mm|

D E

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

F

FicugE |. Flowers and leaf of Jatropha stevensii (Stevens 22902). — A-C. Staminate flower. — A. Sepal. — B.
Petal.—C. Androecium and disk-segment. — D—F. Pistillate flower. — D. Sepal.—E. Petal.—F. Gynoecium.—G.

Leaf.

(MO). Matagalpa: San Juanillo, 8 km SE of Ciudad
Darío, 500 m, Grijalva 2599 (DAV, MO).

Jatropha stevensii, named in honor of Warren
Douglas Stevens, the leading student and collec-
tor of the Nicaraguan flora, belongs to subg. Cur-
cas (Adans.) Griseb. by virtue of its flowers with
entire imbricate sepals, coherent petals, and an-
droecium of 10 monadelphous stamens (Dehgan
& Webster, 1979). Within subg. Curcas, it is re-
ferable to sect. Platyphyllae Dehgan & Webster
because of its 3-locular carinate fruits and bise-
riate anthers. However, it is divergent from most
species of sect. Platyphyllae in having elongated
seeds with a prominent caruncle. As noted b
McVaugh (1945), J. fremontioides Standl. from
Oaxaca differs from other species of subg. Curcas
in its prominently carunculate seeds. Further-
more, Dehgan (1980) has indicated that J. fre-
montioides is distinguished by anisocytic sto-
mata, a feature unique in the genus.

<

The prominently carunculate seeds, monoe-
cious flower production, and small entire cordate
leaves link J. stevensii with J. fremontioides and
indicate that the latter is the closest relative of
the new species. The two species form a subgroup
of sect. Platyphyllae differing from more typical
species such as J. platyphylla, J. alamanii, and
J. ciliata, which have larger more-or-less lobed
leaves, flowers produced dioeciously, and seeds
with small caruncles. Nevertheless, the stami-
nate and pistillate flowers of such species as J.
ciliata are quite similar overall to those of J.
stevensii and J. fremontioides, and the prepon-
derance of evidence does not seem to compel
any modifications in the circumscription of sect.
Platyphyllae at this time.

The discovery of J. stevensii in Nicaragua brings
to three the number of endemic Central Amer-
ican species of Jatropha: J. podagrica Hook.
(eastern Guatemala to Nicaragua), J. stevensii

1987]

WEBSTER —JATROPHA

119

IGURE 2. Photograph of type collection of Jatropha stevensii (Stevens 22902); left-hand glabrous branch
with capsule and staminate flowers; right-hand pubescent branch with pistillate flowers.

(Nicaragua), and J. costaricensis Webster & Po-
veda (Guanacaste, Costa Rica). These species be-
long to two different subgenera and sections, J.
podagrica in subg. Jatropha sect. Peltatae and J.
stevensii and J. costaricensis in subg. Curcas sect.

Platyphyllae. However, as noted by Webster and
Poveda (1978), J. costaricensis is most closely
related to J. alamanii Muell. Arg. of southern
Mexico (Oaxaca) and therefore not to J. stevensii.

The Central American species of Jatropha ap-

120

ear to represent a depauperate extension of the
large assemblage of endemic species in southern
Mexico, as remarked by Webster and Poveda
(1978). It is interesting that the floristic break for
Jatropha comes not at the Isthmus of Tehuan-
tepec but in Guatemala; however, there is clearly
a decline in diversity east of the Isthmus.

In contrast to studies such as that of Savage
(1982) on vertebrate distributions in Mesoamer-
ica, there is clearly no ancient Central American
center for xeric Euphorbiaceae such as Cnidosco-
lus, Jatropha, and Manihot. Instead, in Meso-
american Jatropha there has been an invasion of
one species of subg. Jatropha from the south (J.
podagrica) and two species of subg. Curcas from
= north (J. costaricensis and J. stevensii). Gen-

ry (1982) that xeric M taxa
are ultimately of southern origin. However, for
Jatropha subg. Curcas and other distinctive
Mexican xerophyte taxa such as the Fouquieri-
aceae, this migration from the south must have
been so ancient that these taxa may be regarded
as autochthonous to Mexico in terms of their
evolutionary diversification. In contrast, the in-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

vasion of plants ancestral to J. podagrica must
be more recent, and indeed perhaps subsequent
to the closing of the Panamanian gap.

LITERATURE CITED
DEHGAN, B. 1980. Application of epidermal mor-
phology to taxonomic delimitations in the genus
Jatropha L. (Euphorbiaceae). Bot. J. Linn. Soc. 80:
8.

L. WEBSTER. 1979. Morphology and
infrageneric relationships of the genus Jatropha
pay pis Univ. Calif. Publ. Bot. 74: 1-73.
sd 2. Neotropical floristic diversity:
phyiogeosraphical connections between Central
uth America, Pleistocene climatic fluctua-
mon or an accident of the Andean orogeny? Ann.
Missouri Bot. Gard. 69: 557-593.
9

MCcVAUGRB, R. . Thegenus Jatropha in America:
principal intrageneric groups. Bull. Torrey Bot.
Club 72: 271-294.

SAVAGE, J. M. 1982. The enigma of the Central Amer-
ican herpetofauna: dispersals or vicariance? Ann.
Missouri Bot. Gard. 69: 464—547.

WEBSTER, G. L. & L. J. Povepa. 1978. A phytogeo-
graphically significant new species of Jatropha
(Euphorbiaceae) from Costa Rica. Brittonia 30:

0.

REUNION OF PHAEOSPHAERION AND COMMELINOPSIS
WITH COMMELINA (COMMELINACEAE)

ROBERT B. FADEN! AND D. R. HUNT?

ABSTRACT

The genera Phaeosphaerion and Commelinopsis are reunited with Commelina because they we
ruits. Sh

onand Commelinopsis but dehiscent ar
and Mada
are mad

The handful of neotropical species referred to
Phaeosphaerion Hassk. and Commelinopsis Pi-
chon differ from the cosmopolitan genus Com-
melina L. in having indehiscent, conspicuous
fruits — fleshy and blue to black in Phaeosphae-
rion; crustaceous and white (with the seeds ad-
hering to the septa) in Commelinopsis. Although
other characters may yet be found to separate
these segregates from Commelina, the present,
admittedly incomplete evidence suggests other-
wise. To date the fruit characters are all that can
be used to justify having three genera rather than
one.

We formerly have recognized Phaeosphaerion
and Commelinopsis as distinct from Commelina
(e.g., Hunt, 1981, 1983). New evidence has con-
vinced us, however, that these genera should no
longer be maintained. First, the principal species
of Phaeosphaerion and Commelinopsis are so
strikingly similar to species of Commelina that
identifying non-fruiting specimens is sometimes
extraordinarily difficult, Indeed, the resemblance
of P. } th.) Hassk. ex
C. B. Clarke to C ommelina enia Matuda
(?= C. pallida Willd.) is so close, at least in her-
barium specimens, that one is tempted to wonder
whether the genetic basis of the fruit difference
could be a relatively p pe one. ape Com-
melinopsis glabrata D. R. nt (= C. persica-
riifolium sensu Pichon, non a persi-
cariifolia Delile) bears a very strong resemblance
to Commelina obliqua Vahl (synonym C. robus-
ta Kunth), although the two can usually be sep-
arated by flower color and leaf pubescence—
flowers white and adaxial leaf surface smooth in
Commelinopsis glabrata; flowers blue and adax-
ial leaf surface scabrous in Commelina obliqua —
when fruits are lacking.

20560, U

owy fruits similar to those of Plu
i i

The second line of evidence is the recent dis-
covery of two undescribed species of Commelina
that bridge the narrow gap in fruit morphology
between that genus on the one hand and Phaeo-
sphaerion and Commelinopsis on the other. The
s of these species is appe from ede collec-

s from the vicinity of Guayaquil, Ecuador

[Gilmartin 762 (US); ii 16628 md Asplund

645 (S)]. The fruits of this species, which re-
AMA. Phaeosphaerion leiocarpum in habit, are
reddish or dark blue and exserted from the spathes
at maturity. Unlike the fruits of Phaeosphaerion,
however, those of the Ecuadorian species are de-
hiscent when fully mature.

The fruits of the second Commelina species,
represented by Bosser 17832 (P) from Madagas-
car, are also exserted from the spathes at ma-
turity; but they more closely resemble the fruits
of Commelinopsis than those of Phaeosphaerion,
being crustaceous and whitish. Unlike Comme-
linopsis fruits, d of Bosser 17832 seem to be
dehiscent when mature.

e conspicuous Sis of Phaeosphaerion and
Commelinopsis and the two undescribed Com-

ommelinopsis
have closer affinities with distantly related species
of Commelina than with each other. They clearly
represent separate and parallel evolutionary
derivatives from Commelina. The Ecuadorian
Commelina is apparently related to Phaeo-
sphaerion leiocarpum, but technically it belongs
to Commelina because of its dehiscent fruit. The
Madagascan Commelina is unrelated to any of
the other fruited species , all of which
are neotropical. It represents yet another evo-
lutionary lineage.

The already weak boundaries between Com-

! Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C.
S.A.

? Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, England.

ANN. MissouRi Bor. GARD. 74: 121-122. 1987.

122

melina and Phaeosphaerion and Commelinopsis,
respectively, are further eroded by these new

ommelina species. It is evident that Comme-
lina has produced species with berry-like fruits
in several evolutionary lines. Either each of these
lines must be recognized as a distinct genus or
all of them must be retained within Commelina.

In the course of our discussions on the treat-
ment of the Commelinaceae for the Flora Me-
soamericana, we have concluded that Phaeo-
sphaerion and Commelinopsis can no longer be
upheld. No new combination in Commelina is
needed for Phaeosphaerion leiocarpum or Com-
melinopsis rufipes because their basionyms are

ommelina leiocarpa Benth. and Commelina
rufipes Seubert, respectively. Commelinopsis
glabrata appears to be conspecific with Com-
melina rufipes and will be treated in the Flora as
a variety:

Commelina rufipes Seubert var. glabrata (D. R.
Hunt) Faden & D. R. Hunt, comb. et stat.
nov. Basionym: Commelinopsis glabrata D.
R. Hunt in Kew Bull. 36: 199. 1981

caring oed nn on (Kuntze)
Steyerm. (basion
sperma Kuntze) i: Isa synonym of Commelina ru-
fipes var. glabrata.

The status of the other specific and varietal
names in Phaeosphaerion and Commelinopsis
needs to be considered. Phaeosphaerion efoveo-
latum C. B. Clarke from Venezuela is so similar
to Commelina leiocarpa that it is probably con-
specific. We are uncertain about the importance
and consistency of the seed character used by
Clarke (1881) to separate these taxa, so we de-
cline to transfer P. efoveolatum to Commelina at
this time.

Commelina scabrata Seubert is the basionym
for Phaeosphaerion persicariifolium var. scabra-
ta (Seubert) C. B. Clarke. Seubert’s species, how-
ever, is a synonym of Commelina obliqua Vahl,
thus it is not a synonym of any taxon of Phaeo-
sphaerion or Commelinopsis.

We are less certain about Phaeosphaerion
mathewsii C. B. Clarke from Peru, which is known
definitely only from the type (Mathews 148—K).
(The original spelling of the specific name, with
two ‘t’s is clearly a typographical error.) Although
it would appear to belong to Commelina rufipes,
it does not exactly match collections of either
variety. It is perhaps closer to some specimens
of C. obliqua, especially Davidse & González

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

19296 (US) from Venezuela, but it does not match
them perfectly either. The type of P. mathewsii
lacks fruits, and therefore its inclusion in Phaeo-
sphaerion by Clarke (1881) is questionable. Be-
cause we cannot place this specimen in any named
species of Commelina with certainty, we are
maintaining its status as a species and are trans-
ferring it to the genus:

Commelina mathewsii (C. B. Clarke) Faden & D.
R. Hunt, comb. nov. Basionym: Phaeo-
sphaerion mathewsii C. B. Clarke in DC.,
Monogr. Phan. 3: 138. 1881.

A comment may be made here about the ge-
neric name Athyrocarpus, which has sometimes
been used interchangeably with Phaeosphaerion.
Athyrocarpus was first mentioned by Schlechten-
dal (1855) as a possible genus, but it was not
validly published until Hasskarl (1866) included
it in his key to the genera of Commelinaceae.
Hasskarl also described Phaeosphaerion in the
same paper, so the priority between the two names
must be determined by the earliest publication
in which they are combined. Clarke (1881) ap-
pears to be the first worker to combine them,
placing i d in synonymy under
Phaeosphae

Finally, it irme be noted that under Art. 10
of ICBN (Sydney edition, 1983), the type of
Commelinopsisis the same as that of Commelina
persicariifolia Delile, which is probably referable
to C. virginica L. or C. paludosa Blume (Hunt,
1981). To retain NON s generic concept, it
would be necessary to conserve Commelinopsis
under Art. 10.3 with a specimen that Pichon had
examined, or else to choose a new name for the
genus.

LITERATURE CITED

CLARKE, C. B. 1881. Commelinaceae. Jn A. & C. De
lle. M g phi DL g 3: 113-

ji J. K. 1866. Ueber die Commelinaceen.
Flor: d 209-216.

HUNT, D. n det it notes on Commeli-

naceae 2*5 ora o nidad and Tobago.

American ie x. Kew Bull. 36: 195—

1983. Commelinaceae. Pp. 255-275 in Flora
of Trinidad and Tobago 3, part 3. Government
Printer, Port-of-Spain, Trinidad.

F. L. vo

ee D. : deuda
rvationum in plantas hortenses Halae
onum anno MDCCCLIV e

et jam prius a in-
stitutarum. Linnaea 26: 452—488.

CHROMOSOME NUMBERS OF MADAGASCAR PLANTS!

ELISABETH RABAKONANDRIANINA? AND GERALD D. CARR?

ABSTRACT

romosome numbers are reported for 19 species in eight families of flowering plants of Madagascar.

n = 7) is reported for the genus Vernonia, and the

possible allopolyploid origin of New World taxa of the genus based on x = 17 is discussed.

Although the flora of Madagascar contains
many interesting and poorly known genera and
families, chromosome counts are available for
relatively few species. The present paper inau-
gurates what is intended to be a sustained effort
to record the chromosome numbers of Mada-
gascar plants so they will be available to assist
in interpretation of phytogeographic and evo-
lutionary relationships.

MATERIALS AND METHODS

With two exceptions, chromosome numbers
reported here were determined from microspo-
rocytes undergoing meiotic divisions. The two
counts in the genus A/oe were made from mi-
crocytes undergoing the first mitotic division of
microgametogenesis. All floral bud materials used
for chromosomal determinations were preserved
in a modified Carnoy’s fixative (6 chloroform : 3
absolute ethanol: | glacial acetic acid). These were
stored in the fixative under refrigeration until
they were transported to the University of Ha-
wali where acetocarmine slide preparations were
made according to a modification of Beeks’
method (Beeks, 1955). Observations were made
with a Zeiss Photoscope III equipped with phase
contrast optics. Voucher specimens of all cyto-
logical determinations have been deposited in
the Herbarium ofthe Service de Botanique, Uni-
versité de Madagascar, Tananarive (TAN).

RESULTS

The results are listed in Table 1.

DISCUSSION

Clusiaceae. Thecount of n= 10 for Haronga
madagascariensis (Table 1) is the first count for

this monotypic genus of tropical Africa and Mad-
agascar. The same chromosome number char-
acterizes the related genera Hypericum and Vis-
mia (Moore, 1973, 1977; Goldblatt, 1983).

Ericaceae. First reports here of n = 12 for
Vaccinium emirnense and V. secundiflorum agree
with many other reports for the genus (Fedorov,
1974; Moore, 1973; Goldblatt, 1981, 1983).

Crassulaceae. The report of n = 18 given here
for Kalanchoe beharensis (Table 1) agrees with
two previous counts for this species (Baldwin,
1938; Friedmann, 1971

Thymelaeaceae. The report here for the
Madagascar endemic, Gnidia bakeri, agrees with
an earlier report of n = 9 for Gnidia carinata
Thunb. (Venkateswarlu, 1946). This number is
also characteristic of at least five other genera of
the family (Moore, 1977; Goldblatt, 1981, 1983).

Cornaceae. Thechromosome number of Ka-
liphora madagascariensis, an endemic monotyp-
ic genus, is here determined to be n = 16. Aucuba
appears to be the only other genus in the family
known to possess this number (Fedorov, 1974;
Moore, 1973; Goldblatt, 1983)

Compositae. The Madagascar endemic, An-
isopappus anemonifolius, is here reported to have
n= 7. The only other report for the genus appears
to be 2n = 28 for A. africanus (Hook. f.) Oliv.
& Hiern (Auquier & Renard, 1975). Apparently
on the basis of this latter count, Merxmiller et
al. (1977) proposed the base number of x = 7 for
the genus. Our determination for A. anemoni-

folius supports their proposal. The first report

here for the endemic Conyza garnieri is the same
as that found in many other species of this genus
previously investigated (Fedorov, 1974; Moore,
77; Goldblatt, 1981,

rt
citrina (Table 1) is a number that (along with n =

! This work was facilitated by a Fulbright award to the senior author.
? Université de Madagascar, Etablissement d'enseignement Supérieur des Sciences, Service de Botanique, B.P.

906, Tananarive, Madagascar

? Department of Botany, University of Hawaii, 3190 Maile Way, Honolulu, Hawaii 96822.

ANN. Missouri Bor. GARD. 74: 123-125. 1987.

124 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

TABLE 1. Chromosome numbers of Madagascar plants.

Taxon n Collection Data

Clusiaceae

Haronga madagascariensis Choisy 10 142-84! Ambohitantely
Ericaceae

Vaccinium emirnense Hook. 12 145-84 Ambohitantely

Vaccinium secundiflorum Hook. 12 148-84 Ambohitantely
Crassulaceae

Kalanchoë beharensis Drake 18 Cultivated, Université de Madagascar
Thymelaeaceae

Gnidia bakeri Gilg 9 152-84 Ambohitantely
Cornaceae

Kaliphora madagascariensis Hook. 16 120-84 Ambohitantely
Compositae

Anisopappus anemonifolius (DC.) G. Taylor 7 Ankatso

Conyza garnieri Klatt 9 Tsimbazaza

Helichrysum s 7 139-84 Ambohitantely

Emilia Ead DC. 15 Ankatso

Emilia citrina DC. 5 Ankatso

Mikania ey se Willd. 19 143-84 Ambohitantely

Vernonia appendiculata Less. 7 Tananarive

Vernonia diversifolia Bojer 9 101-84 Ambohitantely

Vernonia garnieriana Klatt 30 114-84 Ambohitantely

Vernonia pectoralis Baker 10 Ankatso
Aloeaceae

Aloe deltoideodonta Baker 7 Station Manambaro?

Aloe divaricata Berger 7 Fort Dauphin?
Orchidaceae

Calanthe silvatica Lindley 20 133-84 Ambohitantely

! All collection numbers are those of the senior author.

? Collected by Lydia Rason.

10) is well established in the genus (Fedorov,
1974; Moore, 1973, 1974, 1977; Goldblatt, 1981,
1983). However, the number n = 15 reported
here for E. adscendens is apparently otherwise
known only in E. sonchifolia (L.) DC. (Torres &
Liogier, 1970). The report here for Mikania
scandens is in agreement with earlier reports for
the species (Fedorov, 1974; Moore, 1973; Gold-
blatt, 1981). The counts here of n = 9 for Ver-
nonia diversifolia and n = 10 for V. pectoralis
appear to represent first reports for these endem-
ic species. These numbers agree with previous
reports and are found to be frequent in Old World
taxa of the genus (Jones, 1979). Vernonia gar-
nieriana, another endemic reported for the first
time, has n = 30 (Table 1). This number appears
to have been reported previously for only two
species of Vernonia, i.e., V. glabra (Steetz) Vatke
(Jones, 1979) and V. travancorica Hook. f. (Na-

rayana, 1979). That these are hexaploids based
on x = 10 is suggested by the report of both n =
10 and n = 30 in V. glabra by Jones (1979). The
first report here of n = 7 for the endemic Ver-
nonia appendiculata represents a number that is
otherwise unknown in the genus. Moreover, this
number represents the lowest haploid number
known in Vernonia and allows speculation of a
new, low base number of x = 7 for the genus.
Jones (1979) has suggested that New World taxa
based on x = 17 may have originated through
aneuploidy from posia based on x = 9.
owever, the discovery o = 7 in Vernonia
RE offers an lade explanation.
World taxa based on x = ould have
arisen by way of allopolyploidy involving ta taxa
with x = 7 and x = 10. In this connection, 1
be mentioned that Humbert (1960) eens ge
96 species listed for Madagascar into six groups,

1987]

according to apparent affinities. The group that
includes V. appendiculata contains 38 other en-
demic species. It would be tempting to speculate
that at least some of those also exhibit x = 7. As
Jones (1979) pointed out, much cytological work
remains to be done in the Vernonieae.

Aloeaceae. The endemic species Aloe deltoi-
deodonta and A. divaricata are reported here to
have n = 7, a number that agrees with previous
reports for these and many other species of the
genus (Amano et al., 1972).

Orchidaceae. The first report here of n = 20
for Calanthe silvatica agrees with previous re-
ports for the genus (Fedorov, 1974; Moore, 1973,
1974, 1977; Goldblatt, 1981, 1983).

LITERATURE CITED
AMANO, M., Y. TAKAHASHI & N. KoNpo. 1972. A
comparative karyotype study in Madagascan A/-
oes. Rep. Inst. Breed. Res. Tokyo Univ. Agr. 3:
7-12.

AUQUIER, P. & R. RENARD. 1975. stats $ ed

da, Burundi et Kivu (Zaire) — L Bull. ry ‘Te.
Nat. Belg. 45: 421-

BALDWIN, J. T. 1938. po» the genus and its
chromosomes. Amer. J. Bot. 25: 572-579.

Beeks, R. M. 1955. ovements in the squash
technique for plant chromosomes. Aliso 3: 131-

FEDOROV, A. (editor). 1974. Chromosome Numbers
of Flowering Plants. Otto Koeltz Science Publish-
ers, Koinigstein.

FRIEDMANN, F. 1971. Sur de nouveaux nombres

RABAKONANDRIANINA & CARR—

MADAGASCAR PLANTS 125

chromosomiques dans le genre — E

6: 103-10

T to plant a

e numbers 1975-1978. Monogr. Syst. Bot.

Missouri Bot. Gard., Volume 5.

š 83. Index to plant chromosome numbers
1979-1981. — Syst. Bot. Missouri Bot.

ard., Volum

pe H. 1960. Flore de "x Cowon et des Co-

s. Composées. Tom 71 in Ver-

nia. Mision national d ee d nüitursfis (Pha-

ri

JONES, S. B. 1979. Ciwomasime numbers of Ver-

nonieae (Compositae). Bull. Torrey Bot. Club 106:
9-84.

MERXMULLER, H., D. LriNs & H. ROESSLER. 1977.
Inuleae—systematic review. Pp. 577—602 in V.H.
Heywood, J. B. Harborne & B. L. Turner (editors),
The Biology and Chemistry of the Compositae,
Volume I. Academic Press, London

Moore, R. J. (editor). 1973. Index to plant chro-
mosome numbers 1967-1971. Regnum Veg. 90:
—

—— ———. 1974. Index to plant a numbers
for a Regnum Veg. 91: 1-108.

1977. Index to plant kampus numbers
for 1973/74. Regnum : 1-
— B. M. 1979. tological investigations
ribe Vernonieae el Current Sci.
004.

. H. LioGIER. 1970. Chromosome
numbers of Dominican Compositae. Brittonia 22:
VENKATESWARLU, J. 1946. Chromosome numbers of
o members of Thymelaeaceae. Current Sci. 15:

CYTOTAXONOMIC STUDIES IN THE GENUS URGINEA
STEIN IN WEST AFRICA. II. KARYOTYPE EVOLUTION
IN URGINEA ALTISSIMA (L.) BAKER!

S. O. OYEWOLE?

ABSTRACT

Mitotic and meiotic studies were carried out on Urginea altissima with 2n — 22. At pachynema-
diakinesis pollen mother cells had 9 (1. 61%), — (93. 9%), or 11 (5. 0%) chromosome bodies. Pollen
viability was 93.56%. S between the proportion of PMCs
with (a) 9 bodies and those with anaphase brides (b) 10 bodies and those with normal anaphase
movements, and pollen viability, and (c) 11 bodies and those with excluded chromosomes. The
eleventh homologue was associated with the dioses largest homologue in the PMCs with 10 bodies.
This scindit is specific and ensures the successful transmission of the eleventh homologue to the
spores. Failure of th the breakdown of normal meiotic
behavior. In PRU the eleventh pair behaved normally. Hence the correct diploid number is 20 +

2 homologous fragments.

Urginea Stein is a genus of bulbous geophytes
in the Liliaceae. It is represented by the basic
chromosome numbers of 5 and 7 (Darlington &
Wylie, 1955; Jones & Smith, 1967). The known
tropical African members have a somatic com-
plement of 14 (a species endemic to Madagascar),
20 or 20 + 0-6 B chromosomes (Jones & Smith,
1967), or 22 (Oyewole, 1975b). The two West
African species with a somatic chromosome
number of 22, viz., U. altissima (L.) Baker sensu
stricto and U. gigantea (Jacq.), represent a de-
parture from the established basic chromosome
numbers. They therefore offer a novel opportu-
nity for investigating and understanding the evo-
lution of genetic systems in the genus, possibly
opening up avenues for understanding the mode
of speciation in a family known to contain groups
of morphologically similar and closely related
taxa. Hence the behavior of the chromosomes of
U. altissima sensu stricto has been investigated
in this wor

MATERIALS AND METHODS

Sample collections from natural populations
of U. altissima sensu stricto were cultivated in
experimental gardens at Ibadan and later at Ilo-
rin. These were investigated cytologically, using

root tip squashes for mitosis and flower bud
(anther) squashes for meiosis, both pretreated for
one hour in sat. aq. solution of p-dichloroben-
zene. Twenty-five plant stands from various pop-
ulation locations in western Nigeria were inves-
tigated. Conventional methods (Darlington & La
Cour, 1942; Marenah & Holden, 1967) of squash
preparation were employed. Two percent acetic
orcein was used. Viability of pollen from unde-
hisced anthers extracted from open flowers was
estimated by observing the ability of grains to
stain in 196 acetocarmine within two to three
minutes.

RESULTS

All 22 chromosomes of the somatic comple-
ment behave normally at mitosis (Fig. 1). None
shows differential staining except at the centro-
meric points. They all move normally at ana-
phase; the number and form, from one cell gen-
eration to another (and indeed in different tissues),
remain consistent (Oyewole, 1975b).

The pairing behavior at meiosis, as well as
pollen viability estimates (Table 1), shows that
the taxon has a stable chromosomal system, with
a high average pollen viability of 93.56%. How-
ever, meiotic formations of 9, 10 or 11 chro-

! The University of Ilorin Senate Research Grant No. 8.184.22, 1978/79, from which the final phase of this

work was funded, is gratefully acknowledged. I also thank the various member
University of Nairobi for their technical assistance — most especially their secretary, Mrs.
. The moral support of the University of Nairobi, , Kenya, and the timely financial

u
work at the AETFAT 1982 Congress wou
? De

rs of the Department of Botany,
Ruth Watulo, who

uld not have been possible, are -5 gratefully acknowledged.

partment of Biological Sciences, University of Ilorin, Ilorin, Nigeri

ANN. Missounmi Bor. GARD. 74: 126-130. 1987.

1987]

OYEWOLE— URGINEA ALTISSIMA

127

1-9.—1
which _ nine chromosome bodies. The hexavalent is arrowed.
4.P at Metaphase I. Arrow indicates the PMC showing 11 bivalents.—5. A PMC with Anaphase I s md
persisting into MII.—6. A PMC at Anaphase I with normal movement. There are an “u paqaq, per grou

FIGURES

. Somatic complement of 22 chromosomes.— 2. PMCs at Metaphase I-Anaphase I, one of

— 3. PMCs at Metaphase I showing 10 bodies. —

g. 1).— 7. PMC me ea

* AL with a chromosome bridge (dicentric).
excluded chromosomes (arrowed). — ar (

mosome bodies per pollen mother cell (PMC)
during the first prophase to metaphase were ob-
served (Figs. 2-4). The nine bodies consist of
one hexavalent and eight bivalents; the 10 bodies
consist of nine bivalents and one quadrivalent;
the 1 1 bodies are all bivalents. These formations

B. Persistent AI and AII s —9. ism I PMCs. One has
) represents 10 um

occur at of 1.61%, 93.39%, and 5.0%,
respectively. Figures 5-7 demonstrate first ana-
phases: anaphase bridges (Fig. 5), normal move-
ment (Fig. 6), and lagging and an excluded pair
of chromosomes (Fig. 7). These occur at fre-
quencies of 1.23%, 93.51%, and 5.44% respec-

128

Meiotic behavior of U. altissima Baker sensu stricto.

TABLE 1.

Examined at

AII-TII

AI-TI

Pollen Viability

Total PMCs
Diplonema-MI

Total

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Pollen

Aborted Exam-

Normal

Total

Chromosome Bodies Formed

ined

Pollen
322

Pollen
4,678

PMCs

somes

mation somes

ment

11

579

Number

6.44

93.51 1.23 5.44 100
Telophase I; AII = Anaphase II; TII

100
metaphase I; AI

5.0

Telophase II.

Anaphase I; TI

PMCS - pollen mother cells; MI

[Vor. 74

TABLE 2. Results of x? to test the null hypothesis
that:

% pollen mother cells (PMCs) showing 10 chro-

mosome bodies, normal anaphase Velas

and normal pollen are equal (ratio 1 : 21);

(2) %PMCs with 9 chromosome bodies B x PMCs
with Anaphase I bridge formation are equal (ratio
1:1);

(3) 96 PMCs with 11 chromosome bodies and %
PMCs with Telophase I excluded chromosomes
are equal (ratio 1: 1); an

(4) Total % PMCs with 9 and 11 chromosome bod-

ies, % PMCs with AI bridge formation and TI

excluded chromosomes, % PMCs with AII bridge
formation and TII excluded chromosomes, and

96 aborted pollen grains are equal.

~
—
—

x? Value Probability Value
(1) 0.000605 P » 9996
(2) 0.0508 90% > P > 80%
(3) 0.01854 90% > P > 80%
(4) 0.0096 98% > P > 95%

Level of significance is set at 0.05. The null hypoth-
esis is proved in each

tively. Figures 8 and 9 show Anaphase II (AII)
diagonal bridge and Telophase II (TII) excluded
chromosomes, which also occur at a total fre-
quency of 6.36% while normal AII movement
occurs at a frequency of 93.64%. dere anal-
yses of these data (Table 2) show that pollen
viability percentage and the frequencies of AI
and AII normal movements and the formation
of 10 chromosome bodies per PMC during the
first prophase to metaphase (93.56%, 93.64%,
93.51%, and 93.39%, respectively) are not sig-
nificantly different; that the frequency of for-
mation of nine bodies per PMC is not signifi-
cantly different from the frequency of occurrence
of laggards and excluded chromosomes at TI;
and that the total frequencies of formation of
nine and 11 bodies, AI bridge formation and TI
chromosome exclusion, AII diagonal bridge for-
mation and TII chromosome exclusion, and per-
centage pollen abortion are all not significantly
different.

In the formation of nine bodies, the hexavalent

second largest pair (cf. chromosome bodies in
Figs. 2-4). When this association occurs, the first

1987]

and second large pairs become about equal in
length at MI (see Fig. 4; see also mitotic chro-
mosome lengths of the two large and eleventh

ofits own (in 11 separate bivalents), or associates
with any homologues other than the second larg-
est, meiotic behavior becomes erratic and meiot-
ic products are inviable.

DISCUSSION

Some representatives of Urginea in other areas
have been shown to have B chromosomes (De
Wet, 1957; Jones & Smith, 1967), but accessory
chromosomes have not been reported in any of
the west tropical African materials. Where they
have been reported, these accessory chromo-
somes are known to be different from the auto-
somal members of their complement. They are
heterochromatic, their numbers vary within
populations, their transmission at mitosis is not
regular, and at meiosis there has not been any
mechanism of transmission comparable to that
shown in U. altissima, where association of the
extra pair with a specific pair of autosomes is the
mechanism for successful transmission of the ex-
tra chromosomes into the spores and thereby
into the next generation of the plant. Hence the
extra pair of chromosomes in U. altissima can-
not be seen as essentially inert as in the cases of
the B chromosomes reported in other species.

The formation of a hexavalent involving the
extra pair and two other pairs may indicate some
genetic affinities between the three pairs involved
(Battaglia, 1964), or show the probable origin of
the extra pair (Wedberg et al., 1968). However,
the formation of an association between the extra
, the second large
pair, with a much higher NN than in the
hexavalent formation, and leading to successful
meiotic behavior, may identify the true origin of
the extra pair. This extra pair cannot be regarded
as accessory because it is indistinguishable from
the other pairs in mitotic behavior and structure
(cf. chromosomes in Clarkia unguiculata, and
C. williamsonii in Mooring, 1960, and Wedberg
et al., 1968, respectively). The extra pair behaves
normally at mitosis as in the large B chromo-
somes of Brachychome lineariloba (Carter &
Smith-White, 1972), but its successful transmis-
sion from one generation of the plant to another
depends upon a meiotic mechanism of its as-
sociation with a specific pair of autosomes in the

OYEWOLE— URGINEA ALTISSIMA

129

complement. Hence, it differs from the B chro-
mosomes of B. lineariloba.

The origin of ie extra pair is undoubtedly
betrayed in its association with the second large
pair — only such association leads to viable spore
formation. The mode of origin of the extra pair
is probably by fragmentation at a heterochro-
matic region along the second (short) arm of the
parent (second large) autosomal pair. This is why
the point of breakage on the otherwise acentric
fragment is able to exercise secondarily a spindle-
fiber organizing function at mitosis for effective
polar movement—and, hence, normal autoso-
mal behavior— but which is incidentally too weak
to effectively organize the movement of the frag-
ments at meiosis whenever they form a bivalent
of their own. They then remain excluded on the
equatorial plane after polar movement of the
other chromosomes.

Again, in the formation of 11 normal biva-
lents, incomplete pairing or total asynapsis in the
extra pair, which would lead to early repulsion
ofits members prior to anaphase movement, was
not observed at all. That is, the extra pair shows
synapsis (and probably chiasma formation). This
means that (1) the extra pair contains essential

1

ing, (2) the extra pair came from homologous
segments of a homologous pair of parent chro-
mosomes, and (3) the extra pair carries genic
matter essential to the survival and success of
the plant. This is why its transmission is effected
by a genetic mechanism. The failure of this
mechanism ultimately leads to the formation of
inviable meiotic products.

The behavior of this extra pair during cell di-
vision is of interest. The incidence of fragments
within chromosome complements has been re-
ported, both in nature and in experiments, by

tution, or rejoin with broken ends of other chro-
mosomes (Lea, 1946; Hair, 1952; Giles, 1954).
If restitution is to occur, it occurs immediately
after the breakage and is dependent on oxidative
enzyme metabolism (Wolff & Luippold, 1955).
The extra pair of chromosomes (fragments) in
U. altissima has healed broken ends in somatic
cells and hence behaves as a normal autosomal
pair during mitosis. During meiosis, however,
the ends either remain healed and the pair forms
a bivalent of its own, which becomes excluded

130

during polar movement, or the broken ends be-
come reactivated and thereby the pair associates
with other chromosome pairs. Such a behavior
has not been reported earlier. Rees (1958), how-
ever, working on Scilla, reported the behavior
of chromosome fragments in pollen mother cells.
In Scilla, the fragments become attached either
to their parent autosomes or to any other chro-
mosomes, depending on the relative distance of
the fragments to one or the other. The attach-

1

ment may | yn-
chronous attachment must take place within a
sticky matrix. As no sticky matrix was found in
U. altissima, there was no evidence to suggest
that the association of the extra pair with the
second large autosomal pair was anything but
synchronous. Hence the situation in U. altissima
differs from that of Scilla in that it shows clear
evidence of a genetically controlled mechanism
of association, thereby stabilizing the resultant
genetic system to ensure a high percentage fer-
tility of sexual reproduction. This is in keeping
with the suggestions of Blackwood (1956) and
Rutishauser (1956).

It will be interesting to find out what initiates
the mechanism of healing and reactivation of the
broken ends of the fragments and the second
large autosomal pair during mitosis and meiosis.
It is likely that the stage of development at which
the dissociation occurs is the interphase between
the end of spore formation and the beginning of
spore development, while the reactivation of the
broken ends would take place at the onset of
meiosis in t
ual reproduction are = generally controlled hor-
monally, these events are likely to be part of the
effects of the hormonal control, which in itself is
genetic. This allows repetition of this event
whenever necessary.

In this case, the extra pair of chromosomes
should be regarded as fragments that are geni-
cally essential to the survival and success of the
genetic system of the plant species. By this frag-
mentation, an otherwise somatic s of2n —
20 has become 2n = 22. This — 22 should,
however, be seen as 2n = 20 + 2 ff.

he evolution of this new number may be con-
nected with the morphological differentiation and
differential ecological preference that have led to

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

the recognition of U. a/tissima sensu stricto from
its relatives U. gigantea and U. viridula Baker
in the U. altissima complex dcus 19752).

LITERATURE CITED

BATTAGLIA, E. 19 Cytogenetics of 6-chromo-
somes. Cytologia 17: 245-299,
BLACKWOOD, E. 1 The inheritance of 8-chro-

CARTER, C. R
Brachychome lineariloba. 3. Accessory chromo-
omes. Chromosoma (Berl.) 39: 361-379
Du C. D. & L. F. LA Cour. 1942. The
Handling of Chromosomes. George Allen & Un-
win, London.
& A. P. WYLIE. 1955. Chromosome Atlas of
Flowering Plants. George Allen & Unwin, Lon-
don

DE WET, J. M.J. 1957. Chromosome numbers in the
Scilleae. Cytologia 22: 145-159.

GILes, N. H. 54. Radiation- oua chromosome
aberrations in Tradescantia. 713-762 in A.
Hollaender (editor), cee Biology, Volume
1. McGraw Hill, New York

Hair, J. B. 1952. The origin of new chromosomes
in Agropyron. Heredity 6, Vn een on
Chromosome Breakage]: 215-2

a 1967. i i TUE of

the Liliaceae: . The karyotypes of twenty-five
tropical mid ‘Kew Bull. 21: 31-38.

LEA, D. E. . Actions of Radiations on Living
Cells, 2nd edition. Cambridge Univ. Press, Cam-
bridge

ge.

MARENAH, L. J. & J. H. W. HOLDEN. 1967. Karyotype
studies in Avena: I. The karyotype of Avena i
cultivar Condor. Chromosoma 22: 456-464

1960. Cytogenetic studies of Clarkia

unguiculata: II. merary chromosomes.

OYEWOLE, S. O. 1975a. Taxonomic treatment of the
Urginea altissima (L.) Baker oo in West Af-
rica. Bol. Soc. Brot. 49: 163-172.

1975b. Cytotaxonomic studies in the genus
Urginea Stein in West Africa: k Karyotype anal-
ysis in U. altissima Baker, U. gigantea ire
Oyewole m U. viridula Biker (emend. ). Bo
Brot. 49: 2

Rees, H. 1958. pes rential behavior of chromosomes

a9

taneous chromosome breakage in Trillium
: 367

Translocation heterozygotes a nd supern ary
ch osom populations of Clarkia wil-
pment Evolution 22: 93-107.

Worrr, S. & H POLD. 1955. Metabolism and
AE A E Science 122: 231-232.

CYTOTAXONOMIC STUDIES IN THE GENUS URGINEA
STEIN IN WEST AFRICA. III. THE CASE OF
URGINEA INDICA (ROXB.) KUNTH IN NIGERIA!

S. O. OYEWOLE?

ABSTRACT

Urginea indica (Roxb.) Kunth is a morphologically variable species of wide distribution in the Old

World. In West Tropical Africa, it has
The large form

been considered to consist of a large form and a dwarf

rm was investigated morphologically in as field and in

form.
the laboratory, as well as karyo-
ue in the field. E differ in both vegetative
wed differences in ecological

an orp ere fo
preferences. Biometrical analysis of vegetative and fora peer showed that the four variants
relim

hat they are similar but not identical in karyomorphology. Art

inary karyotype analysis showed
Sek crosses between them failed to

suc viable seeds. It is concluded that U. indica is an incipient polyspecies or species complex.

Urginea indica (Roxb.) Kunth is one of the
four species of Urginea recognized in the latest
revision of Liliaceae in Flora of West Tropical
Africa (Hepper, 1968). It is distinguished from
the other species by its globose capsule. It is wide-
spread in the Old World tropics, inhabiting sa-
vanna vegetation of tropical and subtropical areas
of Africa, southern India and further east (Thi-
selton-Dyer, 1898). In Nigeria, it is widespread
in the central segment of the country, occurring
between latitudes 7°N and 10°N. It occurs in open,
heavy soil with a top layer of humus or in clay
soil of seasonally flooded plains.

It occurs in different ecological niches in var-

(1968) observed this morphological variation and
contended that there are at least two different
forms of this variable species: a large form and
a dwarf form. These two are easily distinguish-
able in the field. The large form has light green
leaves whose undersurface, immediately out of
the bulb, is pinkish; the reproductive shoot is
also pinkish, 45-150 cm tall, with not less than
15 flowers in the lax, racemose inflorescence. The
tepal is also pinkish with a greenish keel. Leaves
and flowers are never borne together. The dwarf
form, on the other hand, has dark green leaves;
the reproductive shoot is green, generally less
than 40 cm tall, with flowers usually ranging be-
tween one and 12; the tepals are yellowish green
with a green keel. Leaves and flowers are never

borne together. Field studies of this taxon during
vegetative growth and flowering revealed that
Hepper’s contention was a rather conservative
estimate. The present paper therefore aims at
establishing the taxonomic status of U. indica
through morphological studies, starting with the
large form

MATERIALS AND METHODS

Natural populations of the large form were
studied morphologically during several field trips.
Representative samples were collected and
brought into cultivation in nurseries first at the
University of Ibadan (southwestern Nigeria), then
at Ahmadu Bello University, Zaria (northcentral
Nigeria) and later at the University of Ilorin
(westcentral Nigeria). Each asec site was
visited at least twice—duri the vegetative
growth period (May to Mic and during the
flowering and fruiting period ewe to
March). Altogether about 200 bulbs were brought
into cultivation. Four morphological varias
were identified from differen

collection. Titled A-D, they were grown on ad-
jacent nursery beds. No intermediate forms were
encountered, even where the variants grew to-
gether or in contiguous sites. Records of mor-
phological features were kept and carefully fol-
lowed in order to identify any environmentally
induced features. These collections have been in
cultivation since 1972/73.

Each bulb is normally not more than 3—4 inches

! The financial support from the "pn edad à iS Ilorin Senate Research Grant No. 8.184.22, especially for the
ed.

final phase of this work, is gratefully acknow

? Department of Biological Sciences, oe of Ilorin, Ilorin, Nigeria.

ANN. Missounmi Bor. GARD. 74: 131-136. 1987.

132

deep in the soil. Soil samples were usually taken
along with each bulb, and both texture and com-
position of the soil were determined later in the
laboratory. The type of vegetation in which the
population was found was also note
M hological cl teristi divided into

qualitative and quantitative features. The qual-
itative features are bulb shape, shoot color, in-
florescence color, leaf form, perianth color, ovary
shape, ovary color, filament, style, and anther
color. These were assessed visually. The quan-
titative features are shoot and inflorescence
height, number of flowers in the inflorescence,
pedicel length, tepal length and width, length of
ovary, style and stigma, anther, filament, and
leaf, and leaf width. The leaf length-to-width ra-
tio, called leaf ratio (l/w), was calculated for each

meters or with millimeter graph paper. The shoot
was measured from its base at bulb surface to its

hile th ight was taken from
the base of the first flower to the tip of the shoot.
Both measurements were taken after the last
(youngest and apical) flower had either opened
or withered to ensure that the reproductive shoot
had stopped elongating. Floral parts from newly
opened flowers were dissected out for measure-
ments.

T

43 g£ + 1

CIV HIANY
in order to investigate leaf cuticular surface pat-
tern, leaf margin, number of veins, structure of
veins, stomatal structure, and the general pattern
of tissue distribution in the lea

Chromosome number and karyomorphology
were studied using root tip squashes. Root tips
were harvested at about 8 A.M., pretreated for
one hour in saturated aqueous solution of p-dich-
lorobenzene, fixed in fresh 1:3 acetic alcohol
(glacial acetic acid and absolute ethanol) and
stored for at least 30 minutes at about — 4°C
before hydrolysis. Hydrolyzed root tips were
squashed in 2% acetic orcein.

RESULTS

The major collection areas are indicated in
Figure 1. Except for Groups B and D, which were
found growing together in a wide expanse of land,
different groups were found in different popu-
lations and in different ecological niches. All the
populations encountered were found growin
either in large numbers or as a few individuals
dispersed in open savanna, with very light grass
and forb cover and a few short, scattered trees

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

and shrubs. The soil was heavy clay with or with-
d/or pebbles. The top layer was dark
humus. All the populations were found in flood
plains with little organic topsoil during the rains.
In the dry season, the soil was hot, dry, hard and
cakey.

Figure 2 illustrates the vegetative morphology
of the four groups, while Table 1 contains a sum-
mary of all the morphological features investi-

ated. Leaves are produced from the onset of the
rains, and the plants remain vegetative for most
of the rainy season. The leaves dry up towards
the end of the rains. The early annual savanna
fire of October to November burns the dry leaves,
and reproductive shoots may be produced any
time from two weeks after the burning. The ma-
ture flower opens into a bell shape; the pedicel
curves back to carry the flower face downwards
at anthesis and until after pollination, after which
the pedicel straightens up. The flower withers if
not fertilized. When fertilization occurs, how-
ever, the young fruit enlarges while the fading
tepals close over it and shrivel into a little cap
on top of the fruit by the time of maturation.

Groups B and D are most similar, being dis-
tinguished by shoot and inflorescence height, leaf
length and form, and number of flowers per in-
florescence only. D is distinguished from C by
the lengths of tepal, style and stigma, filament
and anther, filament color, bulb shape, shoot and
inflorescence color, and ovary shape and color.
B differs from A in shoot height and color, in-
florescence height, lengths of pedicel, style and
stigma, filament, anther, ovary shape, and fila-
ment color. A and C are distinguished by shoot
height, inflorescence height and color, number
of flowers, lengths of pedicel, tepal, anther and
leaf, leaf width, and ovary shape. A and D differ
in shoot and inflorescence height, number of
flowers, lengths of tepal, style and stigma, fila-
ment and leaf, filament color, leaf form and width,
ovary shape, and bulb shape.

Figure 3 illustrates variations in leaf surface
patterns, leaf margi

out st

tinctly different from the others in each of the
features exhibited

The leaf surface pattern is similar in all, show-
ing minute crenation, which is most noticeable
in variant D but less so in variants C, A, and B,
respectively (see Fig. 3: A,, B,, C,, and D,). A
and D have similar epidermal cell size, type, and
arrangement as well as palisade cell size, form,

1987] OYEWOLE-— URGINEA INDICA 133
A p EET L L L RENT za "
4 7 8 9 10 t! 12 14.
*13 139
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UNAUA .
1 » Zaria I!
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š: ot

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3 4 5 é 7 8 9 (0 m 12 13 i4
L L 2 2 A. A L

2 [B], 3C] & A.D.

O denotes areas of major sampling of 1[A],

routes during trips.

FIGURE 1.

and arrangement. They differ in leaf margin, being
short and acute in A, the adaxial and abaxial
epidermal layers being separated to the margin
by the palisade, which aborts on a marginal epi-
dermal cell, while leaf margin in D is projected
with both epidermal layers coming together out-
side the palisade and terminating with a marginal
epidermal cell. They also differ in the distribu-
tion of phloem tissue in the vein, being bicol-
lateral in A but only collateral in D. In B and C,
the epidermal cells are short and isodiametrical,
the palisade cells are also short and less tightly

Map of Nigeria showing areas of major sampling of the large form of U. indica.

arranged, and the veins have large xylem vessels
with conspicuous bicollateral phloem tissue. The
leaf margin shows progressive elongation, being
long and more acute in B than in A. In B, both
epidermal layers do not close completely beyond
the palisade before terminating in a marginal epi-
dermal cell; in C, leaf margin is projected with
rounded tip as in D and the epidermal layers
barely close up beyond the palisade before ter-
minating in a single marginal cell. Finally, the
arrangement of the large metaxylem vessels is
similar in A, B and D but differs in C.

Undehisced anther color

Pin
Dirty white

in
Dirty white

in
Dirty cream

134 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74
TABLE 1. Data on morphological features.
Taxa
Characters A B C D
Bulb shape Small, edium, Medium, ovoid Large, spherical
spherical spherical
Shoot color reen Pink Light green Pink
Inflorescence color Pink Pink Yellowish cream Pink
Leaf form Short, Short, re- Long, coiled Long, straight
coiled flexed
Perianth color Pinkish Pink with Pinkish brown Pink with yellowish
brown yellowish with green keel keel
ith green keel
green keel
Ovary shape Pyramidal Globose Pyramidal Globose
Ovary color reen Green Green Light green
Filament color Pink Yellowish Pinkish Yellowish
Style color ink i i Pink

Creamy white

Shoot height (cm) 45-60; 52.9 70-90; 76.9 100-140; 123.8 100-140; 119.9
Inflorescence height (cm) 15-20; 17.6 24—30; 27.0 50-60; 54.7 50-70; 59.5
Number of flowers 15-20; 17 15-20; 17 20-30; 24 0-30; 25
Pedicel length (mm) 22-23; 22.6 35-40; 37.1 30—40; 33.9 *(30-) 50-70; 60.3
Tepal length (mm) 12-14; 12.8 12-13; 12.6 15-16; 15.5 1-13; 12.1
Tepal width (mm) 44.5; 4.2 4-6; 5.2 3.5—4.5; 4.0 4.5; 4.5
Ovary length (mm) 4.5—5.5; 4.9 5-6; 5.3 4.5-5.0; 4.9 5-6; 5.5
Style + stigma (m 6.5-7.5; 7.0 5-6; 5.3 *(6.5—)7.0; 7.0 5-6; 5.5
Filament length (mm) 8.5-9.5; 9.0 5; 5.0 *(8.5—)9.0; 9.0 5-6; 5.6
Anther length (mm) 2; 2.0 2.5-3.5; 3.0 2.5: 2.5 3; 3.

Leaf length (cm) 20-25; 22.7 25-35; 31.2 50-80; 67.3 50-70; 61.1
Leaf width (cm) 0.8-1.3; 1.0 0.8-1.3; 1.1 1.4-2.6; 2.1 1.0-2.4; 1.6
Leaf index (l/w) 17-31; 23.6 24-43; 29.8 22-52; 32.6 26-56; 39.5

* [ndicates infrequent deviating measurements.

All four groups have a somatic chromosome
number of 2n — 20. They have similar but not
identical karyomorphology. An analysis of the
karyotypes is the subject matter of a separate
report (Oyewole, 1986). Artificial crossing be-
tween the four groups failed to produce any hy-
brid fruits.

DISCUSSION

Urginea indica has been described as a vari-
able species (Hepper, 1968; Morton, 1961). Mor-
phological variation was maintained even under
uniform cultivation, suggesting that the variation

d

i
with ecological preference rather than being
ubiquitous in all populations, as would be ex-
pected if the variations were due to polygenic
effects (Dobzhansky, 1951; Huxley, 1942; Math-

er, 1943). In U. indica, two distinct forms, B and
D, were found together in the same population
area without intermediates, indicating that the
differences between them are not environmen-
tally induced, while each group has a distinct
karyotype (Oyewole, 1986). Hence variation in
this case is not just a case of polymorphism.
The correlation between the external morpho-
logical variations and the leaf epidermal and me-
sophyll features strongly supports the idea that
this taxon is not just a single species. These an-
atomical features are genetically controlled and,
under the same environmental conditions, still
maintain their differences. The importance of
such anatomical features in species delimitation
has been amply emphasized by Carlquist (1959)
and Metcalfe (1963) and exhaustively demon-
strated in many other works (for example, Prat,
1932; Church, 1949; Serensen, 1953; Borrill,
1959, 1961; Oyewole, 1971; Adeyemi, 1981).

OYEWOLE— URGINEA INDICA

FIGU ay e Vegetative morphology of the four groups (A-D) of the large form of U. indica. Horizontal
^s ar represents 4 m.—3. Leaf surface patterns: 1 — Leaf surface, Koi quede cell structure and the palisade layer;
leaf margin; and 3— Leaf vein structure. Diagonal bar represents 25 um

136

Speciation, in the words of Dobzhansky (1951),
is “that stage of evolutionary process at which
the once actually or potentially interbreeding ar-
ray of forms becomes segregated in two or more
separate arrays which are physiologically inca-
pable of interbreeding.” Recent views on specia-
tion emphasize the relationship of the organism
and the environment as the controlling factor
(Hutchinson, 1959; Lewis, 1969). Thus adaptive
radiations often occur when a species enters an
unoccupied habitat with diverse open niches or
when a population acquires a new complex of
adaptive characters that enables it to exploit
available environment more efficiently (Steb-
bins, S as recorded for the Axonopus com-

es Gledhill, 1966). Hence it is clear
that U. indica, in which there are four morpho-
logically distinct, genetically isolated forms even
within the so-called large form, is not simply one
phenotypically plastic genotype. It is significant
that these forms exist side-by-side in nature or
at least within the same geographical location
and climatic condition while maintaining their
identity both reproductively and morphologi-
cally. Obviously their karyotypes resemble one
another. However, they are biological entities. It
is untenable to regard the hitherto U. indica as
a single species (Lewis, 1969); rather it must be
recognized as a species complex. The evolution-
ary history of U. indica may possibly be similar
to that of A/buca nigritana and the U. altissima
complex in the same family (Gledhill & Oye-
wole, 1972; Oyewole, 1975, respectively).

LITERATURE CITED

ADEYEMI, F. A. 1981. Biosystematic Studies of Some
Nigerian Taxa of Anthericum Lin. and Chloro-

phytum Ker-Gawl (Liliaceae). P D. Thesis. Uni-

fais of Ibadan,

RILL, M. 1959. A biosystematic study of some

Glyceria species in Britain. I: Taxonomy. Wat-

sonia 3: 291-298.

961. Epidermal characteristics in the dip-

loid subspecies of D. glomerata. J. Linn. Soc. Bot.

8

56: 453-458.
CARLQUIST, S. 1959. Anatomy and systematic posi-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

s of Centaurodendron and Yunquea (Compos-
tae). Britonnia 10: 78-93.

CHURCH, G. L. A D. study of Gly-
ceria and Puccinellia. Amer. J. Bot. 36: 155-165.

DoBzHANSKY, T. 1951. Genetics and the Origin of
Species, 3rd edition. Columbia Univ. Press, New
Yor

GLEDHILL, D. 6. Cytotaxonomic revision of the
A W) Beauv. complex. Bol.

WOLE. "1972. The taxonomy of
Albuca in West Africa. Bol. Soc. Brot. 46: 149-
170.

HrPPER, F. N. 1968. Notes on West African mono-
cotyledons. Kew Bull. 21: 493-498.

HUTCHINSON, G. E. 1959. Homage to Santa Rosalia
or why are there so many kinds of animals? Amer.
Naturalist 93: 145-159.

HuxLEY, J. 1942. Evolution: The Modern Synthesis.
Harper and Brothers, p York.

Lewis, H. 1969. Speciatio y and evolution.
Taxon 18: 21-25.

MATHER, K

. 1943. Polygenic inheritance and natural
selection. Biol. Rev. 18: 32-64.
METCALFE, C. R. Comparative anatomy as a
modern botanical discipline. Advances Bot. Res.
8.

OYEWOLE, S. O. I Biosystematic Studies in the
Genus A/buca L. with Particular Reference to Those
Species Occurring in Nigeria. Ph.D. Thesis. Uni-
un of Ibadan, Nigeria.

axonomic treatment of the Urginea
altissima (L. : Baker complex in Nigeria. Bol. Soc.
rot. 49: -17

poe studies of the genus
Africa. IV. Population dif-

ype
indica (Roxb.) Kunth. Ann. Missouri Bot. Garden

7-14
PRAT, H. 1932. piderme des Graminees: étude
anatomique et EE P Ann. Sci. Nat. Bot.
14: 117-324.
e T. 1953. revision of the Greenland
es of Puccinellia. Parl. Medd. Gronl. 136(3):
46.

Nn G. L., JR. 1971. Processes of Organic Evo-
lution. Prentice Hall, Englewood Cliffs, New Jer-

sey.
THISELTON-Dy ER, W. T. (editor). 1898. Pp. 424—426
in Flora of Mrs Africa, Volume 8. L. Reeve
& Co., London

CYTOTAXONOMIC STUDIES IN THE GENUS URGINEA
STEIN IN WEST AFRICA. IV. POPULATION
DIFFERENTIATION AND KARYOTYPE VARIATION IN
URGINEA INDICA (ROXB.) KUNTH!

S. O. OYEWOLE?

ABSTRACT

Qualitative and quantitative studies of eS - over 250 individual plants of the variable

Sedes Urginea indica (Roxb.) Kunth were carried o

he +

The plants were sampled from 23 collection
ot tips were used for mitotic preparations. Ten

otypes were ` recognized, four of which represented the first phenotype while the remaining Six

ggre essive in its exploitation of various

the demands of each ecological niche and a device to isolate the individual gene pools.

Morphological variability is an undisputed at-
tribute of a species composed of sexually repro-
ducing individuals in a large panmictic popula-
tion. However, the assumption that continuous
populations were spatially fluid, panmictic, and
genetically homogeneous has been assailed by
the results of extensive works on both plants and
animals (Epling & Dobzhansky, 1942; Selander
et al., 1969; Bradshaw, 1972; Jones, 1973; Schaal,
1975). This assumption has now been largely
replaced by - Anita that, particularly i in plants,

many exten l
semi- d ne This, according to Linhart
et al. (1981), may be due to the effect of diver-
sifying selection in heterogeneous environments
and/or highly restricted gene flow as a result of
spatial isolation. There is abundant evidence in
support of each of these two phenomena in the
process of speciation. Evidence of restricted gene
flow has led to the assumption that, within con-
tinuous populations, there exist small clusters of
genetically related individuals (Bradshaw, 1972;
Levin & Kerster, 1974). In spite of this, the ex-
istence of distinct correlated discontinuities in
the phenotypic characteristics of a continuous

the existence of distinct segmentation in the ge-
netic structure of the population (cf. Oyewole,
1971). Such segmentation can be maintained only
by a number of factors, chief among which is an
intrinsic isolation mechanism.

The present work analyzes the results of stud-
ies of the genetic structure of morphological vari-
ants of the variable species or species complex,
Urginea indica (Roxb.) Kunth. The morpholog-
ical differentiation among the large form is the
subject of the third in a series of studies of the
genus (Oyewole, 1986).

MATERIALS AND METHODS

Populations were sampled in the wild, and
plants were cultivated in experimental sites
(Oyewole, 1986). Collection sites are illustrated
in Figure 1. The distribution of the species, be-
tween latitudes 7?N and 10°N, spans the decid-
uous woodland and savanna of the Southern
Guinea Savanna vegetation zone. Over 250 plants
were collected from 23 sites in 17 sampling areas.
Plants from distinct populations were grown to-
gether under the same experimental conditions;
seven distinct morphological groups were rec-
ognized. Four of the forms (A, B, C and D) rep-
resent the large form, while E, F and G belong
to the dwarf form.

Root tips were harvested between 8 and 9 A.M.,
pretreated for one hour in sat. aq. 1,4-dichlo-
robenzene, fixed, and treated for mitotic squash
preparations following the popio eth-
ods (Darlington & LaCour, 1969). Chromosome
counts were taken at random Malas every prep-
aration. Chromosome measurements were taken
from not less than 100 cells, at full mitotic meta-

! I gratefully acknowledge the technical assistance I received from the Department of Botany, University oo
ss des Kenya, especially the patience with which Scolastica W. Karari, departmental secretary, typed th

manu

2 esa m of Biological Sciences, University of Ilorin, Nigeria.

ANN. MISSOURI Bor. GARD. 74: 137-143. 1987.

138 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74
L ah. p a L. L L i À — ra
4 7 8 E) IO ft 12 (4.
[13 151
4
-12
[| 4
EU
(o
- [0
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aOa
tO 10; © N
1 s * 63 g
: 1
7
A
FG
(^ ^i, e A2, AA), A AL, °
e í
Zs t8, 4G XD, TE, PF oG) s.
3 4 Š G 7 8 9 (o II (2 13 (4.
FicunE |. Map of Nigeria showing areas of major sampling sites.
phase, in each morphological group. Homolo- kenir

gous chromosomes were easily identified in each
set of measurements from comparative chro-
mosome lengths and the relative lengths of chro-
mosome arms. Measurements were recorded in
order of magnitude for the haploid set. Data were
pooled for each group, from which average chro-
mosome length, the relative chromosome-arm
length, and the position of the centromere were
determined.

ower buds of appropriate age were collected
between 7 and 10 A.M., immediately incised and
fixed, and the anthers squashed and stained. As
many inflorescences as were available in each
group were sampled, and meiotic stages from
pachynema to telophase II were examined in not
less than 100 pollen mother cells (PMCs) in those
groups that flowered (not all the groups have
flowered in cultivation).

Figure 2 contains somatic metaphase comple-
ments of the various groups. All the groups have
2n = 20. Karyotype data is summarized in Table
1. Each morphological group is represented by a
different karyotype. The total length of chro-
matin material, at metaphase, of each karyotype
differs from the others. The karyomorphology is
similar, although intrinsic differences abound
(Fig. 3). Chromosome classification, using the
chromosome index (ratio of long arm to short
arm), is according to Levan et al. (1964). The
chromosomes in each complement were classi-
fied as long (6.0 um and above), medium (4.0-
5.9 um) and short (below 4.0 um). Details of the
meiotic study will be presented in a subsequent
part of this series.

Group A. This group is represented by four

OYEWOLE— URGINEA INDICA KARYOTYPE VARIATION 139

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in one sampling site about 80 km to the east of
A

E

A, karyotype consists of chromosomes whose
average lengths vary between 3.67 um and 11.0
um, with an average total chromatin length of
119.34 um per complement. The complement
consists of three long, six medium, and one short
pairs of chromosomes, all with terminal and sub-
terminal centromeres. Endomitosis frequently
Occurs in the root cells. Meiosis is normal, and
10 bivalents are regularly formed.

Even though each complement of A,, A, and
A, could be resolved into pairs of similar chro-
mosomes, members of such pairs are by no means
identical. None of the individuals of these three
karyotypes has flowered since they were brought
into cultivation; hence these hypothetical pair-
ings could not be verified in actual meiotic pair-
ing.

A, consists of chromosomes whose average
lengths range from 3.31 um to 10.56 um, with
an average chromatin length of 116.36 um per
complement. The complement consists of six
long, 10 medium, and four short chromosomes,
all with subterminal-terminal centromeres. The
longest two of the chromosomes have a centric
region as wide as the short chromosome arm
length.

A, chromosomes range in average length from
2.5 um to 7.5 um, with an average chromatin
length of 85.26 um per somatic complement.
There are five long chromosomes in the com-
plement, four of which resolve into two pairs
while the fifth is associable with a shorter chro-
mosome. This long chromosome has a conspic-
uous secondary constriction (arrowed in Fig. 2,
A4). The whole complement consists of five long,
four medium, and 11 short chromosomes, all of
which have their centromeres in the subtermin-

al-terminal region, except the smallest two pairs
which have submedian centromeres.

A, chromosomes vary in average length from
2.5 um to 7.5 um. They have an average chro-
matin length of 84.76 um per somatic comple-
ment. The complement consists of four long, six
medium, and 10 short chromosomes, all with
subterminal-terminal centromeres except a short
pair with median centromere.

Group B. This is represented by only one
karyotype. Chromosomes vary in average length
from 3.0 um to 9.5 um, with an average chro-
matin length of 106.36 um per somatic comple-
ment. The complement consists of three long,
three medium, and four short pairs. All the chro-

OYEWOLE— URGINEA INDICA KARYOTYPE VARIATION

Ailit anto Ay
È Jinna A2
GUA by than sees uns A3
EITAYTTITTTTIN A4

[ Diu Hillia B
Whun C

inus Mini D

T Ilii lilii ua E
MMM F

HTTP G

URE 3. Idiograms ofthe various karyotypes rep-
Eu by the groups. Horizontal bar represents 30

mosomes have subterminal-terminal centro-
meres. The longest pair has a wide centric region.
Meiosis is normal except for the infrequent early
separation of members of a small pair

Group C. Chromosome length varies from
an average of 4.88 um to 13.1 um, with an av-
erage chromatin length of 148.44 um per somatic
complement. There are five long and five me-
dium pairs of chromosomes in the complement.
All the chromosomes have subterminal-termi-
nal centromeres. The longest pair has a centric
region as wide as its short arm. Multivalents are
frequently formed during meiosis.

Group D. Populations of this group are sym-
patric with those of Group B. The karyotype con-
sists of chromosomes with subterminal-terminal
centromeres. The average chromosome length
varies from 3.5 um to 9.82 um, with an average
chromatin length of 108.48 um per somatic com-
plement. There are three long, three medium,
and four short pairs of chromosomes. The two

142

longest pairs have a wide centric region. Meiosis
is regular.

roup E. The karyotype of this group con-
sists of chromosomes with subterminal-terminal
centromeres. Chromosomes vary in length from
4.25 um to 13.5 um and have an average chro-
matin length of 142.52 um per somatic comple-
ment. There are four long and six medium pairs
of chromosomes. The first and third long pairs
have a wide centric region. Meiosis is regular.

Group F. The karyotype consists of chro-
mosomes that vary in length from 4.1 um to
12.05 um, with an average chromatin length of
135.97 um per somatic complement. The com-
plement is made up of four long and six medium
pairs, all with subterminal-terminal centro-
meres. The fifth pair has a secondary constriction
on the long arm. Meiosis is regular.

Group G. The karyotype consists of chro-
mosomes whose average lengths vary between
4.14 um and 12.33 um, with an average chro-
matin length of 137.28 um per somatic comple-
ment. The complement consists of four long and
six medium pairs, all with subterminal-terminal
centromeres. Meiosis is regular.

DISCUSSION

The similarity in the morphology ofthe karyo-
types is obvious. The differences in the karyo-
morphology of the different populations seem
minute, but they are basic and do underlie the
differences in the external morphology of each

retained. It is, however, evident that changes have
occurred (or are occurring) in this taxon that ma
be correlated with the morphological differentia-
tion of the populations. The recognition of dif-
ferent karyotypes that correspond to different
; the presence
of more than one karyotype in an otherwise mor-
phologically uniform unit raises interesting ques-
tions about evolutionary phenomena. This mor-
phologically uniform unit, Group A, is interesting.
$ ete see Kobe Ku fate,

panied by external morphological differentia-
tion, but it is partly correlated by habitat pref-
the A,, A, and A, complements clearly

plements represent natural hybrid swarms that

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

introgressively identify with A,, which is likely
to be one of the putative parents. The different
karyotypes are correlated with differences in ex-
ternal morphology and ecological preferences.
Those forms that inhabit different ecological
niches may have differentiated in response to
differences in ecological demands, while those
which inhabit same or similar ecologic niches
must be fundamentally different genetically in
order to retain their individual identities mor-
phologically. In both cases, karyotype differen-
tiation seems to have resulted in reproductive
isolation by which the different forms are main-
tained in nature

he differences in the amount of chromatin
material may have resulted from (or led to)
karyotype differentiation. Difference in the chro-
matin material is correlated with both morpho-
logical differentiation and ecological preferences
(note A, and A,, B and D are in the same niches;
A, and A,, and E, F, and G are in similar niches;
and C and E are in different niches).

The morphological variability of this species
has long been recognized, but no prior attempts
have been made to distinguish the different forms
beyond the arbitrary categorization of “large”
and "dwarf." This is probably due to the fact
that no field collection ever contains both floral
and vegetative features together, as well as to the
wide east-west distribution of the species. The
aggressive exploitation of different niches by dif-
ferent biotypes has resulted in the differentiation
of the morphological forms within the species'
broad areas of distribution. The isolation of spe-
cific biotypes, forming small clusters of individ-
uals, in such ecological niches probably led to
the accumulation of favored genes and/or mod-
est chromosomal changes and eventually to the
specific karyotypes that are associated with spe-
cific morphological forms as well as with specific
ecologic niches (Wright, 1940; Bush et al., 1977;
Bengtsson, 1980). There is no doubt that mor-
phological differentiation in these populations is
more obvious than differentiation in chromo-
some morphology, suggesting that chromosome
repatterning may have been mild and might have
involved only small segments in gene/gene block
rearrangement. Hence this species seems to com-
prise a stable polymorphism in which the differ-
ent forms have attained reproductive isolation
and genetic stability, and hence each form has
retained its morphological identity. This case is
therefore different from that of Agrostis tenuis

1987]

(Bradshaw, 1959) or Elymus rechingeri (Heneen
& Runemark, 1962) but seems to be similar to
that of the diploid neospecies of Clarkia (Lewis,
1973)

Flower formation and fruit development are a
common feature with most of the morphological
forms, especially in nature. Preliminary studies
of meiotic behavior have shown regularity of pol-
len formation in most of them under cultivation.
However, this apparent sexual reproduction is
coupled with vigorous vegetative propagation by
axillary bulb formation in varying degrees in all
forms. It is therefore necessary to have a closer
look at the reproductive biology of the entire
species in order to ascertain the extent of actual
sexual reproduction and the mechanism of pol-
lination in each form. Hybridization experi-
ments between the different forms, which will
hopefully shed light on their genetic divergence,
are in progress. Thus far, preliminary results of
such experiments show successful n cross-
ing between only two forms, A, an

LITERATURE CITED

BENGTSSON, B. O. 1980. Rates of karyotype evolution
in placental saci Hereditas 92: 37-47.
1959. Population differentiation
cal differ-

HAW, A.
in Agrostis pns Sibth. I. Morphologi
entiation. New Phytol. 58: 208-227.

1972. Some evolutionary consequences of
being a f Evol. Biol. 5: 25-47.
BusH, G. L., S. M. Case, A. C. WILSON & J. L. PATTON.
1977. Rapid speciation and chromosomal evo-
lution in mammals. Proc. Nat. Acad. Sci. 74: 3942-

BRADS

OYEWOLE— URGINEA INDICA KARYOTYPE VARIATION

143

ar e C. D. & L. F. LACougR. 1969. The Han-
ing of Chromosomes, 6th edition. George Allen
Lon

.& T. DOBZHANSKY. 1942. Genetics of nat-

ural populations. VI. Microgeographical races in
Lynanthus parryae. Genetics 27: 317-331.

HENEEN, W. K. & H. RUNEMARK. 1962. Chromo-
some polymorphism and morphological diversity

n Elymus pie fa u Sedis 48: 545-564.
dones J. S. Ecological genetics and natural
selection in molluscs. Science 182: 546-551.

LEVAN, A., K. FREDGAR & A. A. SANDBERG. 1964.
Nomenclature for centromeric position on chro-
mosomes. Hereditas 52: 201-220.

Levin, D. A. AND H. W. KERSTER. 1974. Gene flow
in seed plants. Evol. Biol. 7: 179-220.

Lewis, H. 1973. The origin of diploid neospecies in
Clarkia. Amer. Naturalist 107: 161— i

LINHART, Y. B., J. B. MITTEN, K. B. STURGEON & M.
L. Davis. 1981. Genetic ara in space and
time " a population of ponderosa pine. Heredity
46: 407—426.

OYEWOLE, E O. Biosystematic Studies in the
Genus A/buca L. with Particular Reference to Those
Species Occurring in Ix . Ph.D. Thesis. Uni-
versity of Ibadan, Nig

86. Cytotaxonomie studies in the genus

Ur, rein ea Stein in West Africa. III. The case of Ur-

on pe Ra fi Kunth i in n Nigeria. Ann. Mis-
-136.

ran mi E e p s structure and local dif-

ferentiation Ç: Liatris cylindracea. Amer. Natu-
ralist 109: 511-528.
SELANDER, R. K : Y. YANG & W. G. Hunt. 1969.

olymorphism in esterases and hemoglobins in
wild populations of the house mouse. Studies in
Genetics 5: 271—328.
WRIGHT, S. 1940. Breeding structure of populations
in relation to speciation. Amer. Naturalist 74: 232-
248.

NEW TAXA OF OENOTHERA L. SECT. OENOTHERA
(ONAGRACEAE)!

WERNER DIETRICH? AND WARREN L. WAGNER?

ABSTRACT

everal new ionis are described in advance of a complete monograph of Oenothera sect. Oenothera
imannia. u

the plains to eastern Colorado and Tex

unz to O. laciniata subsp. p

outcrossing species, O.

are differentiated from the oo ae fees O. pubescens, for a total of three species

assigned to subsect. Nutantigi

Subsection Raimannia is n divided ge two series: series Candela, established for the species

with densely flowered spikes bearing two

ers opening per day, inflorescences without

ore flow
lateral branches, acute to rounded petals, and straight floral tubes (O. rhombipetala, O. heterophylla,

O. clelandii, O. bifrons, and O. curtissii); and

to ema rgin
exicana, O. laciniata

described for relictual populations i in southeastern

series
inflorescences usually interrupted by lateral branches, usually only o
nate petals, and floral tubes curved upward
, O. drummondii, and O. humifusa). The new iiec Oenothera flare is

aimannia is restricted to pla D with loose
ne flower ope ng pe
prior to anthesis (O. grandis, O. fa lfurr.

n bivalents

o me differentiated from its presumed S O. grandis and pem
the morphologically similar and dein atric O. laciniata. The

ew combinat mondii su

thalassaphila is made for the populations in Baja California oie set as O. G ee

Over 20 years of cultivation of nearly 150
strains for experimental hybridizations, cytolog-
ical examination, study of breeding systems, and
extensive study of herbarium materials, has led
to revision of the species referred to subg. Rai-
mannia (Rose ex Britton & A. Brown) Munz by
Munz s 1965). Extensive analysis of cross-

as resulted in realignment ofa group

RS 75 species into sect. Oenothera, which in-
cludes the species referred by Munz to his subg.
Oenothera and subg. Raimannia (Stubbe & Ra-
ven, 1979). In turn, this large group of basically
intercrossable species was further subdivided into
ve smaller crossing groups that form highly fer-
tile hybrids and usually have compatible plas-
tids. The first of these groups comprises the South
American species, which Dietrich (1977) placed
into subsect. Munzia W. Dietrich (45 species).
The second, subsect. Oenothera (13 spp.), was
outlined by Raven et al. (1979). The third and
p is subsect. Emersonia
n & W. L. Wagner

(four spp.), revised in 1985 by Dietrich et al. The
fourth and fifth groups, subsect. Raimannia and
the new subsect. Nutantigemma described in this

paper, will be the subject of an upcoming detailed
revision. Prior to publication of the revision, the
present paper makes the new subsection and
species available for regional floras and concur-
rent work on Oenothera, including DNA restric-
tion mapping and studies of flavonoids, pollen,
and seed anatomy.

SUBSECTION NUTANTIGEMMA

Populations occurring in montane sites from
the western United States south to South Amer-
ica, referred by Munz (1935, 1965) to a variety

laciniata. This widespread entity is a permanent
translocation heterozygote treated by us as O.
pubescens Willd. ex Spreng.

Two new Mexican species, closely related to
Oenothera pubescens, one from the cape region
of Baja California and one in the southern Sierra
Madre Occidental, were detected during the study
of herbarium specimens and cultivation of nu-
merous strains in the experimental gardens
(Stubbe & Raven, 1979). They are described here
as O. breedlovei (bivalent-forming) and O. tam-

! This work has been supported by a series of grants from the National Science Foundation to Peter H. Raven.
? Botanisches Institut der apis META Universitatsstr. 1, D-4000 Düsseldorf 1, West Germany.

? Bernice P. Bishop Museum, P.O. B

ANN. MissouRi Bor. GARD. 74: 144-150. 1987.

19000-A, Honolulu, Hawaii 96817, U.S.A.

1987]

rae (presumably bivalent-forming). Oenothera
tamrae is known only from the type collection
and has not been grown in the experimental gar-
den. All three species are placed into the new
subsect. Nutantigemma.

Oenothera L. sect. Oenothera subsect. Nutanti-
gemma W. Dietrich & W. L. Wagner, sub-
sect. nov. TYPE: O. pubescens Willd. ex
Spreng.

Oenothera sect. ribs ia sensu Ser. ex DC., Prodr. 4:
46. 1828, pro part

a co sensu Wooton & pese Contr. U.S. Natl.

erb. 16: 150. 1913, M

br sensu Rose, 2 ‘Natl . Herb. 8: 330.

1905, pro parte; sensu T Sprague $ Riley, Bull. Misc.
Inform. 200. 1921,

a d fa mania an u (Rose ex Britton &

A. wn) Munz, Amer. J. Bot. 22: 645. 1935,

pro r.
nios sect. Anogra sensu Tidestrom, Fl. Ariz. &
x. 273. 1941, pro part
Oenothera Sue Raimannia sect. Raimannia sensu
ose ex Britton & A. Brown) Munz, N. Amer
I. 5: 105. 1965, pro
Oenothera sect. Oenothera subsect. Raimannia sensu
(Rose ex Britton & A. Brown) W. Dietrich, Ann.
Missouri Bot. Gard. 64: 612. 1977 [1978], pro
parte.

Her haps annuae vel probaliter biennes, erectae, ro-

sulatae, simplices vel caules principales ramosi ramis

We areas rosula adscendentibus. Folia rosulae anguste

eb'anceolata, profunde partita vel n integra, acuta;

el elliptica, a nguste

oblanceolata vel oblanceolata vel lanceolata vel an-
acut

folia strigillosa vel villosa, raro glanduloso- pubescen-

mplices vel in

tiaca post anthesin. Styli 276.5 cm longi, stigmate sub
th : Py e 1 + 1 th M4 ed x. —

Capsulae cylindricae, 1.8-4.6 cm longae. Semina am-
bito elliptica vel rotundata, brunnea vel atrobrunnea,
saepe atromaculata, 0. .9— : x mm longa, 0. 6- 1 mm cras-
sa. Planta | el struc-
turaliter UE LIEN heterozygotica complexa.
Numerus gameticus chromosomaticus, n = 7

The principal reason for according these three
species (O. pubescens-group of Stubbe & Raven,
1979) subsectional status is the sterility of hy-

DIETRICH & WAGNER—NEW TAXA OF OENOTHERA

145

brids in crosses between subsections Raimannia
and Nutantigemma. They range from western
Texas west to Arizona and southeastern Cali-
fornia, south to Mexico and Guatemala, and one
species extends to the Andes of Colombia, Ec-
uador, and Peru south to the province of Junin.
The nodding flower buds on the species of sub-
sect. Nutantigemma clearly differentiate them
from those of subsect. Munzia and subsect. Rai-
mannia. This feature is shared with the white-
flowered sect. Kleinia Munz, sect. Anogra (Spach)
Endl., sect. Ravenia W. L. Wagner, occasionally
with Oenothera caespitosa Nutt. subsp. nava-
Jjoensis W. L. Wagner, Stockhouse & Klein [sect.
Pachylophus (Spach) Endl.], and with the yellow-
flowered sect. Eremia W. L. Wagner. In Oeno-
thera fruticosa L. [sect. Kneiffia (Spach) Endl.],
which has yellow flowers, the shoot apices, but
not the flower buds, are bent downward; a similar
condition occurs in O. speciosa Nutt. [sect. Xy-
lopleurum (Spach) Endl.]. Nodding buds appear
to represent, at least for the most part, plesio-
morphy rather than convergence.

Another important argument for considering
this group a subsection is that these three species
are distributed completely allopatrically from
subsect. Raimannia. The species of series Nut-
antigemma grow exclusively in montane habi-
tats, from 3,900 m down to approximately 1,500
m, while the species of subsect. Raimannia grow
only at lower elevations. Oenothera breedlovei is
self-compatible and bivalent-forming; O. pubes-
cens is a permanent translocation heterozygote
forming a ring of 14 chromosomes in meiosis I
and with 40-70% pollen fertility; O. tamrae is
probably an outcrossing bivalent-former, based
on 90-1009 fertile pollen in the type collection.

KEY TO THE SPECIES OF SUBSECT.
NUTANTIGEMMA

la. Stigma elevated above the anthers at anthe-
sis; pollen ca. 90-100% fertile; Laguna Mts.,
ide California, Mexico, and Nayarit, Mex-

= Lower leaves deeply parted almost to the

midrib; mature buds up to 5 mm in di-

ameter: Laguna Mts., Baja California,
Mexico O. breedlovei

2b. Lower leaves not parted to the midrib;
mature buds rad mm in diameter; Na-

yarit, Mexic O. tamrae
1b. Stigma surroun hace I" the EE see ca.
40-70% fertile; Arizona, Texas, co (ex-
E s California), [i ti Colombe

o Per O. pubescens

146

Oenothera breedlovei W. Dietrich & W. L. Wag-

ner, sp. nov. TYPE: Mexico. Baja California:

glong Interior val-

jer: [La Laguna], south of Pico La Aguja,

Sierra La Laguna, 6,300-6,700 ft., 22 Oct.

1977, D. E. Breedlove 43362 (holotype, MO-
2695034).

Herbae annuae vel biennes, erectae, rosulatae, 2-3
dm altae. Folia profunde partita vel remote vadoseque
dentata. Gemmae vetustiores nutantes. Tubi florales
2.4—4 cm longi. Sepala 1.2-2.2 cm longa, saepe rubro-
egli villosa et glanduloso-pubescentia; apices se-
paloru mm longi. Petala 1.6-3.5 cm longa,
flava. Cou: 1.8-4.6 cm longae, E s Samina
1.2-1.6 mm longa, 0.6—0.7 mm crassa, saepe Mm
Numerus gameticus chromosomaticus, n = 7; planta
chromosomatice structuraliter homozygotica, npa
compatibilis.

Erect annual or probably biennial herbs, form-
ing rosettes; stems 2-3(-5) dm long, simple or
with a branched main stem and arcuating lateral
branches arising from the rosette, usually flushed
with red, densely strigillose, sometimes also scat-
tered villous. Rosette leaves narrowly oblanceo-
late, 4-12 cm long, 0.5-2 cm wide, pinnately
o remotely and shallowly dentate, the
apex acute, gradually narrowed to the petiole;
cauline leaves very narrowly elliptic to lanceo-
late, 2-5 cm long, 0.5-2 cm wide, usually pin-
nately parted, the apex acute, the base narrowly
cuneate, short-petiolate to sessile; bracts lanceo-
late to narrowly ovate, 1.5-3 cm long, 0.5-1.5
cm wide, deeply parted to remotely shallowly
dentate, the apex acute, the base broadly cuneate
to narrowly cuneate; leaves and bracts densely
strigillose and sparsely villous. Inflorescence usu-
ally with lateral branches, the young buds erect,
the older ones nodding, becoming erect just be-
fore opening. Flowers usually | per spike opening
near sunset each day. Floral tube flushed with
red, 2.4—4 cm long, ca. 1 mm diam., sparsely to
densely strigillose, villous, and glandular puber-
ulent. Mature buds oblong, 3-5 mm diam. at the
base. Sepals yellowish, often flushed with red,
also red maculate and striped at the margins,
scattered to densely villous and scattered to
densely glandular puberulent, 1.2-2.2 cm long,
the free tips 0.5-1 mm long, erect and appressed
in bud, strigillose to villous. Petals yellow, broad-
ly obovate to very broadly obovate, retuse, 1.6—
3.5 cm long, 1.6-3.7 cm wide. Filaments 1.1-2
cm long; anthers 4-12 mm long; pollen ca. 90-
100% fertile. Ovary 1-2.5 cm long, ca. 1.5 mm
diam., densely strigillose; style 4—6.5 cm long,
the visible part 1.7-2.5 cm long; stigma elevated

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

above the anthers at anthesis, the lobes 3-6 mm
long. Capsule cylindrical, 1.8—4.6 cm long, 3-3.5
mm diam., densely strigillose. Seeds ellipsoid to
broadly ellipsoid, brown to dark brown, often
with darker flecks, 1.2-1.6 mm long, 0.6-0.7 mm
diam., the surface pitted. — FD but
modally outcrossing. Gametic chromosome
number, n = 7 (7,, at meiotic metaphase I).

This new species, known only from Laguna
Mts., southern Baja California, Mexico, is named
in honor of Dennis E. Breedlove (California
Academy of Sciences), who has added greatly to
our knowledge of Mexico through his extensive
and excellent collections and to whom we are
indebted for collecting material of this and many
other Mexican Oenothera species for cultivation
at the Botanical Institute of the University of
Diisseldorf and at the Missouri Botanical Gar-

Oenothera tamrae W. Dietrich & W. L. Wagner,
sp. nov. TYPE: Mexico. Nayarit: Sierra Madre,
near Santa Teresa, territory of Tepic, 8 Aug.
1897, J. N. Rose 2133 (holotype, US-301038;
isotypes, NY, UC).

Herbae annuae vel biennes, erectae, rosulatae, 2-4
dm altae. Folia partita seu remote obtuseque dentata
vel quasi integra. Gemmae vetustiores nutantes. Tubi
florales 3.5-4.2 cm longi, villosi et glandoloso-pubes-
centes. Sepala 1.8-2.5 cm longa, rubro complana et
rubro-fasciata ad margines; apices sepalorum 0.
longi. Petala 2-3.5 cm longa, flava. Capsulae 4—4.5 cm
longae, strigillosae et villosae. Semina 1-1.1 mm longa,

7 mm crassa

Erect annual or biennial herbs, probably form-
ing rosettes; stems 2-4 dm long, simple or with
obliquely ascending lateral branches arising from
the rosette, densely strigillose and sparsely to
densely villous. Cauline leaves narrowly elliptic
or narrowly lanceolate to lanceolate, 4-8 cm long,
0.8-1.8 cm wide, pinnately parted or remotely
and bluntly dentate to subentire, the apex acute,
the base narrowly
rowly lanceolate to lanceolate,
0.7-1.5 cm wide, remotely and bluntly dentate,
the apex acute, the base narrowly cuneate, sessile;
leaves and bracts strigillose. Inflorescence simple
or with lateral branches, nodding. Usually 1 flow-
er per spike opening probably near sunset each
day. Floral tube flushed with red, 3.5-4.2 cm
long, 1.5-2 mm diam., sparsely villous and
sparsely glandular puberulent. Mature buds cy-
lindrical to narrowly ovoid, 5-7 mm diam. at
the base, nodding before anthesis. Sepals yellow-

T
>
v
°

1987]

ish, often flushed with red and striped red at the
margins, the pubescence as on floral tube, 1.8-
2.5 cm long, the sepal tips ca. 0.5 mm long, erect
in bud, strigillose. Petals yellow, very broadly
obovate, retuse, 2-3.5 cm long, 3-4 cm wide.
Filaments 1.5-1.7 cm long; anthers 6-9 mm long;
pollen ca. 90-10096 fertile. Ovary 1.8-2.6 cm
ong, ca. 2 mm diam., densely strigillose and
densely villous, the apex also glandular puber-
ulent. Style 5.3-6.4 cm long, the visible part 1.8—
2.2 cm long; stigma elevated above the anthers
at anthesis, the lobes 5-8 mm long. Capsule cy-
lindrical, 4—4.5 cm long, 3-4 mm diam., the pu-
bescence as on ovary but less dense. Seeds broad-
ly ellipsoid, brown with dark red flecks, 1-1.1
mm long, ca. 0.7 mm diam., pitted. Chromo-
some number unknown.

This rare new species, known only from the
type locality, near Santa Teresa in the Sierra
Madra, Nayarit, Mexico, is named in honor of
Tamra Engelhorn Raven, botanist and wife of
Peter H. Raven. The description is based entirely
on the type collection made by J. N. Rose i
1897.

SUBSECTION RAIMANNIA

The yellow-flowered species assigned by Munz
(1935, 1965) to his subg. Raimannia from the
central and eastern United States are now con-
sidered to comprise subsect. Raimannia. Stubbe
and Raven (1979), when considering the realign-
ment of sect. Oenothera, included O. grandis
(Britton) Smyth, O. laciniata Hill, O. drummon-
dii Hook., O. humifusa Nutt., O. heterophylla
Spach, and O. rhombipetala Nutt. ex Torrey &
A. Gray in subsect. Raimannia. Work toward
the overall revision of the subsection has led to
the recognition of two series within it.

Oenothera sect. Oenothera subsect. Raimannia

ose ex Britton & A. Brown) W. Dietrich

series Candela W. Dietrich & W. L. Wagner,

series nov. TYPE: O. rhombipetala Nutt. ex
Torrey & A. Gray.

d sect. III uha. Nouv. Ann. Mus. Hist. Nat.
347. 1835, pro p
Oenothea 8E ecco ias sensu S. d Proc. Amer.
. Arts 8: 574. 1873, pro

PT Ren sensu r Bull. Teer Bot. ‘Club 23: 172,
1896, pro part

Raimannia sensu Rose, Contr. U.S. Natl. Herb. 8: 330.
1905, pro parte.

Oenothera subg. Raimannia sensu Munz, Amer. J. Bot.

DIETRICH & WAGNER—NEW TAXA OF OENOTHERA

147

22: 645. 1935, pro parte; sensu iig & Riley,
Bull. Misc. Inform. 200. 1921, prop

Oenothera subg. Raimannia sect. RA sensu
Munz, N. Amer. FI. II. 5: 105. 1965, pro parte.

erbae annuae vel biennes, probaliter breviter pe-
ae apu e rosula Kaa e ua,

ramis later
iem

g
ae elliptica vel late elliptica, 1.1-1.9 mm longa,
0 mm crassa, plerumque atromaculata.

Series Candela comprises a clearly defined
group of five closely related species that occur in
sandy soil in open places such as fields, prairies,
roadsides, and open woods, from southern South
Dakota, Minnesota, and Michigan south to Tex-
as and southeastern New Mexico and east to
Louisiana, Alabama, northern Florida, and

outhern Georgia. They all have the presumably
derived characters of relatively densely flowered
spikes on which two or more flowers open every
evening, unlike the species of series Raimannia,
which nearly always produce only one flower per
branch each day, a plesiomorphic characteristic.
The spikes of series Candela never have lateral
shoots, as 1s often the case in series Raimannia.
The fully grown buds of series Candela are nar-
rowly oblong, whereas those of series Raimannia
are lanceoloid in the large-flowered structural
homozygote species and oblong to ellipsoid in
the small-flowered heterozygote species. In series

C t.
also clearly differentiate series Candela from se-
ries Raimannia; those of series Candela are acute
to rounded at the apex, in contrast with those of
series Raimannia which are truncate to emar-
ginate. Further, the capsules of series Candela
are relatively short and thicker toward the base,
while those of series Raimannia are on average
longer and nearly cylindrical. All species of series
Candela appear to be biennials. In contrast to
this specialized habit, the species of series Rai-
mannia have evolved an annual habit except O.
drummondii and O. humifusa, which inhabit sand
dunes and have retained the generalized peren-
nial habit.

Three of the species, Oenothera heterophylla,
O. bifrons D. Don, and O. f-
incompatible bivalent-formers, and O.
W. Dietrich, Raven & W. L. Wagner and O.

148

curtissii (Rose) Small are complex structural het-
erozygote species, presumably derived from an
ancestor similar to O. rhombipetala.

Oenothera sect. Oenothera subsect. Raimannia
(Rose ex Britton & A. Brown) W. Dietrich
series Raimannia (Rose ex Britton & A.
Brown) W. Dietrich & W. L. Wagner, comb.
& stat. nov. Based on Raimannia Rose,
Contr. U.S. Natl. Herb. 8: 330. 1905, ex
Britton & A. Brown, Ill. Fl. N. U.S., 2nd
edition, 2: 596. 1913. Oenothera subg. Rai-
mannia (Rose ex Britton & A. Brown) Munz,
Amer. J. Bot. 22: 645. 1935. Oenothera subg.
Raimannia sect. Raimannia (Rose ex Brit-
ton & A. Brown) Munz, N. Amer. FI. II. 5:
105. 1965. Oenothera sect. Oenothera sub-

Gard. 64: 612. 1977 [1978]. TYPE: O. lacin-
iata Hill.

Onagra sensu Moench, Methodus 1: 675. 1974, pro
Oenothera sect. Onagra sensu Ser. ex DC., Prodr. 4:

; , pro parte.
Oenothera sect. pig cba sensu Fischer & Meyer, In-
dex Sem. rt. Petrop. 2: 44. 1835, pro parte.
are bons I Spach, Hist. Nat. Vég. 4: 353. 1835,

dan BE. ae III & IV sensu Spach, Nouv. Ann. Mus.
Hist. Nat. 4: 347. 1835, pro parte.
Oenothera § P E sensu S. Watson, Proc. Amer.
. Arts 8: 574. 1873, pro parte.
Per en sensu Raimann, Nat. Pflanzenfam. III. 7:
14. 1893, pro parte; sensu Small, Bull. Torrey
Bot. Club 23: 172. 1896, pro
Oenothera sect. Oenothera sensu Tidestrom, Fl. Ariz.
& Mex. 272. 1941, pro parte.

Erect to procumbent, annual or perennial herbs

yo
ceolate to oblanceolate, narrowly elliptic, elliptic,
narrowly obovate or narrowly oblong, parted to
subentire, the lobes + dentate, the apex acute,
the base narrowly cuneate to acute to almost ses-
sile; all leaves strigillose or strigillose and villous,
sometimes also glandular puberulent. Inflores-
cence lax, often with lateral branches, usually
only 1 flower per spike opening near sunset each
day. Floral tube 1.5-5 cm long, yellowish, often

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

flushed with red, rarely also red-flecked, strigil-
lose or glandular puberulent, or villous and glan-
dular puberulent, usually curved upward. Sepals
greenish to yellowish, often flushed with red or

red- -stripe att e margins d-flecked,

the pubescence usually as on floral tube, the sepal
tips 0.3-5 mm long, erect and appressed or
spreading in bud, sometimes separated. Petals
very broadly obovate, truncate to emarginate at
apex, 0.5-4.5 cm long, 0.5-5.5 cm wide, yellow,
sometimes pale yellow, fading red to orange after
anthesis. Style 2-7.5 cm long; stigma elevated
above the anthers at anthesis or surrounded by
the anthers and pollen shed directly onto it. Cap-
sule cylindrical, 2-5.5 cm long, 2-5 mm diam.
Seeds ellipsoid to subglobose, brown, sometimes
with darker flecks (O. drummondii, O. humi-
fusa), 0.8-2 mm long, 0.3-0.9 mm diam. Self-
incompatible (1 sp.) or self-compatible and mod-
ally outcrossing (1 sp.) to modally autogamous
(2 spp.), or permanent structural heterozygotes
(2 spp.). Gametic chromosome number, n = 7
(7,,, ©14 or intermediate chromosome configu-
rations at meiotic metaphase I

Series Raimannia of subsect. Raimannia is
comprised of six species occurring in open, sandy,
and disturbed places, sometimes on dunes, from
North Dakota south to Texas and east to the
Atlantic Coast, in Mexico along the Gulf Coast,
and disjunct in southern Baja California. The
species exhibit considerable variation; among
them, however, only Oenothera drummondii can
be subdivided into two geographically separated
subspecies. Typical of the species of series Rai-
mannia are loose inflorescences, which often have
lateral branches, and upward-curving flower buds.
Comparisons with series Candela were made in
the discussion of that series.

The distribution of series Raimannia is essen-
tially the same as that of series Candela but ex-
tends farther east, to the Atlantic Coast, and ex-
tends south to the state of Campeche, Mexico,
along the coast of the Gulf of Mexico. Oenothera
drummondii Hook. subsp. thalassaphila (Bran-
degee), comb. nov. is disjunct, occurring along
the Pacific Eid at the southern tip of Baja Cal-
ifornia, Mex

x. puits (Britton) Smyth, O. edie
riae (described as new below),
Spach, and O. drummondii form bivalents
whereas O. laciniata Hill and O. humifusa Nutt.
are permanent structural heterozygotes. In this
section only O. grandis is self-incompatible; all

1987]

other species, both Divalent- formers and com-
plex structural het are self-compatible

and largely autogamous.

Oenothera falfurriae W. Dietrich & W. L. Wag-
ner, sp. nov. TYPE: Grown from seeds and
cultivated in the Botanical Garden of Düs-
seldorf, Germany, 2 July 1981, cult. no. 81-
115 from seeds collected in U.S.A. Texas:
Brooks Co., 13.3 mi. S of junction of High-
ways 281 and 285 in Falfurrias, 10 May 1978,
K. Allred & R. Shaw 2021 (holotype, MO-
3332203; isotypes, DUSS, M, MO)

Herbae annuae, erectae vel parum decum-
bentes, rosulae foliis paucis, 1-4 dm altae. Folia
ita

.5—4 cm longi,
villosi et glanduloso-pubescentes. Sepala 1—2.2
cm longa, viridi-flava, immaculata vel rubro-
maculata; apices sepalorum minuti, 0.5-2 mm
longi. Petala 1.3-2.5 cm longa, flava vel pallide
flava. Capsulae 2-4.5 cm longae, 2-2.5 mm cras-
sae, strigillosae, villosae et glanduloso-pubes-
entes. Semina 0.8-1.4 mm longa, 0.3-0.

crassa. Numerus amas chromosomaticus.
n = 7: planta chromosomatice structuraliter ho-
mozygotica, autocompatibilis.

Erect to decumbent annual herbs, forming a
rosette with only a few leaves; stems 1-4 dm
long, usually simple, densely to sparsely strigil-
lose, villous and sometimes also glandular pu-
berulent. Rosette leaves oblanceolate, 5-12 cm

bracts elliptic, narrowly ovate to lanceolate, 2—
4.5 cm long, 0.5-2.5 cm wide, dentate or sub-
entire to pinnately lobed, narrowed to the base,
subsessile; all leaves densely to sparsely villous
and glandular puberulent, especially on the mid-
rib of the lower surface and along the margin,
usually also sparsely to moderately strigillose.
Inflorescence lax, simple or with lateral branches,
usually only 1 flower per spike opening near sun-
set each day, erect at anthesis. Floral tube 2.5-
4 cm long, densely to sparsely villous and glan-
dular puberulent. Mature buds lanceoloid to nar-
rowly ovoid or oblong-ovoid, 0.4—0.6 cm diam.

DIETRICH & WAGNER — NEW TAXA OF OENOTHERA

149

at the base. Sepals green to greenish yellow,
sometimes with red spots, the pubescence as on
the floral tube, 1—2.2 cm long, the sepal tips 0.5-
2 mm long, erect in bud, strigillose and villous.
Petals yellow, broadly obovate, 1.3-2.5 cm long,
1.4-2.7 cm wide, the apex truncate to slightly
retuse. Filaments 10-17 mm long; anthers 4—5
mm long; pollen ca. nen fertile. Ovary 1-
1.7 cm long, ca. 1.5 mm diam., densely villous,
d E and glandular Sera aim style 3.5-5

m long, the visible part 1.2-3 cm long; stigma
deis elevated above the anthers at anthesis,
the lobes 3-7 mm long. Capsule cylindrical, 2-
4.5 cm long, 2-2.5 mm diam., the surface pitted.
Seeds brown, ellipsqid, 0. an l. 4 mm long, 0.3-
0.6 mm diam. Self ible, modally autog-
amous. Gametic chromosome mimber: n=7(7y
at meiotic metaphase I).

Oenothera falfurriae, named after Falfurrias,
Brooks County, Texas, where the type was col-
lected, is endemic to open sandy sites in south-
eastern Texas. Its range is nearly the same as
those of O. bifrons and O. mexicana. When Die-
trich first detected the species, the specimens were
treated as hybrids between O. grandis and O.
laciniata since they were somewhat intermediate
between these species. Seed samples collected by
K. Allred and R. Shaw made it possible to cul-
tivate this species at the Botanical Institute in
Dusseldorf, and it soon became obvious that the
plants were by no means hybrids, but instead
represented an undescribed bivalent-forming
species. All plants examined formed 7,, in mei-
OSIS, Bs no individuals grown from seed resem-
bled either O. laciniata or O. grandis. The in-
i ca collection numbers of Allred and Shaw
represent population samples of several plants
from which seeds were taken and sowed sepa-
rately: 2016, 2020, and 2021 contained O. fal-

furriae and O. laciniata; 2018 contained O. fal-
furriae and O. mexicana. Seeds taken from plants

of O. falfurriae produced only O. falfurriae and
those of O. laciniata produced only O. laciniata.

Oenothera falfurriae differs from O. grandis in
its self-compatibility and smaller petals, which
are intermediate in size between those of O. /a-
ciniata and those of O. grandis. Stigmas in the
closed mature buds are only slightly raised above
the anthers, suggesting that self-pollination is
common in O. falfurriae. Also, the shape of the
buds is more or less oblong, in contrast with the
lanceoloid buds of O. grandis, and the sepals in
O. falfurriae are very delicate and pressed to-

150

gether in bud, whereas in O. grandis they are
often spreading, longer, and thicker.

tan 3

fal is narrowly
presumably relictual. It appears to maintain itself
distinct from the other species of series Rai-
mannia with which it grows sympatrically — O.
grandis, O. laciniata, and O. mexicana — by pos-
sessing a unique plastome. Artificial crosses made
by Dr. Behn at the Botanical Institute in Düs-
seldorf showed that crosses between O. drum-
mondii or O. humifusa and O. falfurriae as the
staminate parent produced pale seedlings that
failed to grow beyond the cotyledon stage. Sim-
ilarly, the seeds of crosses between O. grandis
and O. falfurriae did not germinate at all (Behn,
pers. comm.). Also, since crosses between O.
drummondii, O. humifusa, or O. grandis and O.
laciniata produce completely green and viable
offspring, we can assume that similar crossing
barriers exist between O. falfurriae and O. laci-
niata, based on the pattern of such relationship
in Oenothera sect. Oenothera generally.

Oenothera drummondii Hook. subsp. thalassa-
phila irap be Dietrich & W. L. Wag-
ner, comb. Based on Oenothera tha-
lassaphila dde Univ. Calif. Publ. Bot.
10:185. 1922. Oenothera drummondii Hook.
var. thalassaphila (Brandegee) Munz, Amer.
J. Bot. 22: 651. 1935. rvPE: Mexico. Baja
California Sur: San José del Cabo, 12 Mar.
1892, T. S. Brandegee 216 (lectotype, UC-
107674; see Munz, Amer. J. Bot. 22: 651.
1935).

The separation of Oenothera drummondii
subsp. thalassaphila, which is restricted to dunes
of coastal southern Baja California del Sur, Mex-
ico, from subsp. drummondii depends on a com-
bination of characters since no single morpho-
logical feature separates them clearly. Oenothera
drummondii subsp. thalassaphila always grows

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

for several years, as is demonstrated by the con-
sistent presence of nonflowering shoots and large
taproots on the older plants. By contrast, O.
drummondii subsp. drummondii is basically an
annual, seldom overwintering for a second sea-
son; it usually has only a few nonflowering shoots
or none, and the development of its taproot is
considerably weaker than in subsp. thalassaphi-
la. In general, the habit of subsp. drummondii is
more upright than that of subsp. thallassaphila,
which has prostrate to ascending stems. In ad-
dition, the calyx of subsp. thalassaphila often has
red spots and lacks glandular hairs, whereas the
calyx of subsp. drummondii only rarely has red-
dish spots and is often glandular puberulent. The
sizes of the capsules and seeds are also modally
distinct: in subsp. thalassaphila the capsules are
2-4 cm long and 2.5-5 mm in diameter, and the
seeds are 1.5-2 mm long and 0.7-0.9 mm in
diameter; in subsp. drummondii the capsules are
2.5-5.5 cm long and 2-3 mm in diameter, and
the seeds are 1.1-1.7 mm long and 0.5-0.8 mm
in diameter.

LITERATURE CITED

a W. 7 [1978]. The South American
ecies of Oe es sect. Oenothera (Raimannia,
| fein. Onagraceae). Ann. Missouri Bot. Gard.

64: 425-626.

RAVEN & W. L. WAGNER. 1985. Re-
vision of Oenothera sect. Oenothera subsect.
uci eur SEP nied Syst. Bot. 10: 29-48.
Muwz, P. A.

. 1979 [1980].
An outline of the systematics of Oenothera sub-
sect. Euoenothera (Onagraceae). Syst. Bot. 4: 242-
252

STUBBE, W. & P. H. Raven. 1979. A genetic contri-
bution to the taxonomy of Oenothera sect. Oe-
nothera (including subsections Euoenothera,
Emersonia, Raimannia and Munzia). Pl. Syst.
Evol. 133: 39-59

A NEW COMBINATION AND NEW SUBSPECIES IN
OENOTHERA ELATA KUNTH (ONAGRACEAE)!

WERNER DIETRICH? AND WARREN L. WAGNER?

ABSTRACT

The new combination of Oenothera elata Kunth subsp. hookeri (Torrey & A. Gray) W. Dietrich &
ri

County,

ed fro razos
Texas. It appears to be a rare relictual entity most closely related to O. ati subsp. monte ys which
occurs disjunctly some 680 km to the west of subsp. texen

These names are made avallable in anticipa-
tion of their use in regional floras and concurrent
studies of flavonoids, cytology, and pollen mor-

phology in advance of a detailed revision of Oe-
nothera subsect. Oenothera (Dietrich & Wagner,
in prep.). Detailed presentation of data and dis-
cussions will be given in the revision.

Munz (1949, 1965) divided the large-flowered,
bivalent-forming, outcrossing populations of sect.
Oenothera subsect. Oenothera from the western
United States south to Panama into two species,
O. hookeri Torrey & A. Gray and O. elata Kunth.
During the past ten years we have reevaluated
the variation pattern of these plants by a detailed
study of a large number of herbarium specimens,
fieldwork, and study of plants cultivated in the
experimental gardens at Diisseldorf. These stud-
ies have shown that the two entities are extremel

act

diameter, and modally in several other features.
It was therefore suggested by Raven et al. (1979),
in an outline of the systematics of Oenothera sect.
Oenothera subsect. Oenothera (formerly Eu-
oenothera), that they should be considered con-
specific

Oenothera elata can be subdivided into four
subspecies: subsp. e/ata with a scattered distri-
bution from Guanajuato, Mexico, to Costa Rica
and Panama in Central America; subsp. hirsu-
tissima (A. Gray ex S. Watson) W. Dietrich, which
Occurs in the western United States from Wash-

ner, 1983); subsp. texensis subsp. nov., known
only from one collection in Brazos County, Tex-

as; and subsp. hookeri (Torrey & A. Gray) comb.
et stat. nov. occurring in moist coastal and slight-
ly inland sandy and bluff sites in California from
Marin County south to San Diego County.

The four subspecies of Oenothera elata can be
distinguished with the following key.

KEY TO THE SUBSPECIES OF
OENOTHERA ELATA

la. Stem, leaves, and ovary (capsule) exclusively

)
with red; ps s tips of
the capsule distinct ee
2b. Stem usually green; the = e of th

capsule indistinct
z Ma ture buds (excl. oo ae nar-
rowly lanceolate in outline 5-5 cm

"Pa ve tips 2-3 mm long; petals
m long; capsule 5-6.5 c
ur bcn undulate; leaves mem-
branous; plant in d asa to
. texensis

- ceolate in outline, a cm

more than 10 dm tall ............. bsp. elata
Ib. Stem and ovary (capsule) predominantly with
erect pubescence (short and long villous), stem

without glandular hairs . . subsp. Airsutissima
4b. Stem in the region of the inflorescence
with glandular hairs
5a. Sepals green or - flushed with red,
without or with indistinct pustulate
hairs, sparsely to scattered villous;

' This work has been supported by a series of grants from the National Science Foundation to Peter H. Raven.
ae d Institut der Universitat Düsseldorf, Universitatsstr. 1, D-4000 Düsseldorf 1, West Germany.
ernice P. Bishop Museum, P.O. Box 19000-A, Honolulu, Hawaii 96817, U.S.A.

ANN. Missouni Bor. GARD. 74: 151-152. 1987.

152

plant in cultivation id than
dm p. hirsutissima
; Sepals always flushed us red, w ith

wn
c

vation not more than 8 dm tall .....
subsp. hookeri

The plants described here as O. elata subsp.
texensis are known only from a single herbarium
specimen and from material cultivated at Düs-
seldorf. In 1981 Dietrich collected a seed of an
unusual Oenothera from a TRT herbarium spec-
imen. From the large flowers it appeared to rep-
resent O. grandiflora L'Hér., but when plants
were grown in Düsseldorf they clearly repre-
sented O. elata. Further, based on its strigillose
pubescence it appeared to be subsp. Airsutissima;
however, other characters clearly separated it
from both strigillose pubescent subspecies of O.
elata, subspp. elata and hirsutissima. The stems
of the Brazos County plants are always green, the
capsules are 5—6.5 cm long, the buds are narrowly
lanceolate, and the leaves are membranous. These
plants grow tall in cultivation, to 2 m or more.

Oenothera elata Kunth subsp. texensis W. Die-
trich & W. L. Wagner, subsp. nov. TYPE:
Grown from seeds taken from herbarium
specimen TRT-205991 and cultivated at the
Botanical Garden of the University of Düs-
seldorf, 12 Sept. 1984, cult. no. Stubbe 84-
204; original source, U.S.A. Texas: Brazos
Co.,ca. 17 km NW of Navasota River bridge
on Hwy. 6 in vicinity of Peach Creek cutoff,
25 Oct. 1978, P. M. Catling & K. L. Intosh
(holotype, MO-3332204; isotypes, DUSS,
M)

Herbae biennes, erectae, in culturam usque ad 20
si. Folia undulata

m
flava, 4.5-5.5 cm longa. Stylus longus, stigmate sub
anthesi supra unde edipi) Capsulae 5-6.5 cm lon-
gae, strigillosae. Numerus gameticus chromesomati-
cus, n = 7; 1 uis ires zem homozygotica (7
bivalenta in metaphasium primum meiosis), autocom-
patibilis

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Munz (1949, 1965) recognized two subspecies
of his Oenothera hookeri from moist coastal or
slightly inland sites in California: subsp. hookeri
and subsp. montereyensis. He referred plants with
a bushy habit, blunt buds, free sepal tips 1-2.5
mm long, and sepals usually 2-2.5 cm long to

subsp. montereyensis, whereas plants with a less
bu hed habit, attenuate buds, free sepal tips

S

referred to subsp. hookeri.
shown that these characters represent intra- and
inter-populational variation and, further, ap-
pear to vary independently. The bushier habit of
some coastal plants appears to be an adaptation
to wind and salt spray. All of these populations
are here treated as members of one coastal sub-
species of Oenothera elata distinguished from
the other three subspecies of O. elata primarily

y its densely glandular puberulent and long-
villous buds.

Oenothera elata Kunth subsp. pp dol
& A. Gray) W. Dietrich & W. Wagner,
comb. et stat. nov.

Oenothera hookeri Torrey & A. Gray, Fl. N. Am
49 0. TvPE: California [without stir sa n
cality] ouis s.n. (holotype,

Oenothera hookeri Torr. & A. Gra subsp. monterey-
ensis Munz, Aliso 2: 14. 1949. TYPE: United States.
California: Monterey Co., 0.2 mi. S of mouth of
Alder Creek, 6 Nov. 1934, C. B. Wolf6223 (RSA-
12778, holotype (not seen); isotypes, GH, NY,

OM).

LITERATURE CITED

Munz, P. A. 1949. The Oenothera hookeri group.
Aliso 2: 1-47.
————. 1965. ney Fl. N. Amer. II. 5: 1-278

RAVEN, P. H., W. D dole ipiis Tees
An outline of ib atics of Oenothera sub-
sect. peepee ae Syst. Bot. 4: 242-

WAGNER, W.L. New species and combinations
in the genus Oenothera heey ia Ann. Mis-
souri Bot. Gard. 70: —196.

A REVISION OF MEZILAURUS (LAURACEAE)!

HENK VAN DER WERFF?

ABSTRACT

The neotropical genus Mezilaurus (Lauraceae), which consists of 18 species and is best represented

in the drainage area of the A
from Licaria and reinstated on its
caatingae van der

mazon, is revised. Clinostemon, a genus of two species recently separated
own, is included here in Mezilaurus. Eight sp
Werff, M. duckei van der Werff, M. glaucophylla van der Werff, M. micrantha van

ecies, Mezilaurus

der Werff, M. opaca Kubitzki & van der Werff, M. due ped van 5 e M. pyriflora van der

Werff, and M. quadrilocellata van der Werff are described as new. One

me, M. thoroflora van

der Werff, and a new combination, M. mahuba Tree van ton Werff a are pere a

Mezilaurus comprises 18 species, occurring
from Costa Rica to southern Brazil. The majority
of the species are found in the drainage area of
the Amazon River and adjacent Guayana. The
type species, M. navalium, is restricted. to the

to insufficient material) are shrubs from the cer-
rado vegetation. Fourteen species are Amazo-
nian and have been reported from a variety of
habitats. Ten Amazonian species, ranging from
shrubs to tall trees, occur in terra firme vegeta-
tion, frequently in xeromorphic forests on white
sand. One species, M. mahuba, is restricted to
flooded forest. Three species are known to occur
in secondary vegetation (M. thoroflora, M. syn-
andra and M. lindaviana), although it is not clear
whether these are typical of secondary vegetation
or were left standing when the primary forest was

vesting. Lauraceae, widely used for timber, are
therefore frequently found as isolated trees in
pastures or similarly disturbed habitats. No hab-
itat information is available for the Colombian
species, which is only known from Chigorodó in
northern Antioquia. The Costa Rican species oc-
curs in wet lowland forest near the Pacific Coast.

Most species of Mezilaurus are collected in-
frequently, and I have seen more than ten col-
lect ctions for only two species (M. itauba and M.
many Mezilaurus species are

ave

doubt explains the paucity of collections. The

genus is greatly undercollected and much more
material is needed for a better taxonomic un-
derstanding.

The main use of Mezilaurus is for timber. The
species are locally well known and their hard
wood is much used for boat building and con-
struction. Mez (1889) mentioned that the fruits
of M. itauba are edible. On the label of Fróes
12152 (Mezilaurus pyriflora) it is stated that the
wood causes injuries to the skin, presumably a
kind of dermatitis.

The present revision was undertaken because
it became clear that the genus C/inostemon should
be merged with Mezilaurus. This led to a closer
look at recent collections, during which several
undescribed species were found. I now recognize
18 species in Mezilaurus, which more than dou-
bles the number of species recognized by Kos-
termans (1938

MATERIALS

This study is based both on older collections,
already cited by Kostermans (1938), and recent
collections personally selected during visits to

major American and European herbaria or re-
ceived on loan with other unidentified Laura-
ceae. In my experience the genus was unrecog-
nized in many herbaria. Unfortunately, I have
not yet had the opportunity to visit the leading
Brazilian herbaria. I fully expect that such visits
will yield additional taxa, not only from the Am-

in this group of small-flowered Lauraceae.

! I thank the curators of AAU, BM, BR, C, F, G, HUH, K, L, NY, U, and US for the loan of specimens. Dr.
A. Gentry suggested improvements in the text, and Dr. J. Dwyer corrected the Latin descriptions. John Myers

made the illustrations.

? Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

ANN. MissouRi Bor. GARD. 74: 153-182. 1987.

TAXONOMIC HISTORY

The first species of Mezilaurus was published
by Allemáo (1848) as Silvia navalium. Meissner
(1864) recognized that Silvia Allemào was a later
homonym of Silvia Vellozo and proposed the
new name Silvaea. Unfortunately, Silvaea
Meissner is a later homonym of Si/vaea Phillipi.
O. Kuntze (1891) proposed another new name,
Mezia, to replace Silvia Allemào, but the name
Mezia O. Kuntze was predated by Mezia
Schwacke, a genus of Malpighiaceae. Finally,
Taubert (1892) proposed the name Mezilaurus,
which Mez (1892) accepted. He included seven
species in the genus. Pax (1897), overlooking the
publication of Mezilaurus, proposed Neosilvia as

new name for Silvia Allemao. Neosilvia is
therefore a superfluous name. The name Mezi-
laurus was not universally accepted at first. Even
Mez (1904, 1920, 1924) used the name Silvia
again. Ducke (1930, 1935) published several new
species under the generic name Silvia. Koster-
mans's revision (1938) definitively established
the use of Mezilaurus Taubert

During the nineteenth century, several species
now placed in Mezilaurus were described in oth-
er genera, mostly in genera now included in Li-
caria. Meissner (1864) described three species in
Acrodiclidium and one in Oreodaphne (a syn-
onym of Ocotea). Bentham in Hooker (1878)
transferred two of these to Misanteca (a synonym
of Licaria) and Bentham and Hooker (1880)
placed the other two (including the type species
of Mezilaurus) in Endiandra, an Old World ge-
nus. Mez (1889) recognized six species in Me-
zilaurus (as Silvia), the type species, the four
species described by Meissner (1864), and a sixth
species now included in Licaria. Kostermans
(1938) accepted eight species of Mezilaurus, four
of Mez's species and four described since 1889
by Mez and Ducke. Allen (1964) described two
new species of Mezilaurus from Venezuela.

Kuhlmann and Sampaio (1928) published the
monotypic genus Clinostemon based on Acro-
diclidium mahuba Samp. Kostermans (1938) did
not accept this genus and placed it in Licaria.
Later Allen (1948) described a new species closely
related to Licaria mahuba and noted the resem-
blance to Mezilaurus. Recent investigations (Ku-
bitzki et al., 1979) have shown that these two
species do not belong in Licaria and are more
closely related to or congeneric with Mezilaurus.
In this paper both are included in Mezilaurus.

The eight species recognized in Kostermans's

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

monograph are all maintained in this publica-
tion. The increase in the number of Mezilaurus
species is partly due to the inclusion of Clino-
stemon in Mezilaurus and partly due to recent
collections that represent undescribed species.

GENERIC RELATIONSHIPS

One of the main taxonomic difficulties in Lau-
raceae is that many of the genera are poorly de-
fined. This is reflected in the various infrafamilial
classification schemes and the frequency with
which species are transferred between various
genera. During the last 30 years generic relation-
ships within the Lauraceae have been discussed
in three papers (Kostermans, 1957; Hutchinson,
1964; and Richter, 1981)

Kostermans (1957) attached much importance
to the fruit (with or without cupule) and less
importance to the number of anther cells; he
placed Mezilaurus in the subtribe Beilschmie-
diinae of the tribe Perseeae, close to Endiandra,
and noted, as did Bentham and Hooker (1880),
a similarity to Endiandra. He also stated that
Endiandra and Mezilaurus differ in anther shape
and the positions of their leaves. Additional dif-
ferences are the very fine reticulation of the leaves
and the large, spreading tepals in many species
of Endiandra (comparable to tepals in Nectan-
dra), which are very unlike the small, erect, scale-
like tepals in Mezilaurus. These differences great-
ly outweigh the similarities between Endiandra
and Mezilaurus (number of fertile stamens,
number of anther cells, fruit more or less without
cupule) and Endiandra is probably not a close
relative of Mezilaurus

Hutchinson (1964) considered the number of
anther cells more important than the develop-
ment of the cupule. He placed Mezilaurus next
to Misanteca, but the distribution data given un-
der Misanteca strongly suggest that Hutchinson,
as did Bentham and Hooker (1880), included in
his Misanteca two Brazilian species here includ-
ed in Mezilaurus.

As can be seen from the taxonomic history of
Mezilaurus, its species frequently have been
placed in Licaria (or its synonyms Acrodiclidium
and Misanteca). These two genera have in com-
mon that they are the only neotropical Lauraceae
with three fertile, two-celled stamens. The gen-
eral flower shape of species belonging to these
two genera can be quite similar and a generic
separation based solely on flowers can be ve
difficult. Fortunately, other characters readily

1987]

identify the genera: Mezilaurus has the leaves
always clustered at the tips of the branches, Li-
caria never; in Mezilaurus the cupule consists of
a small, platelike disk, in Licaria it grows into a
rather large, double-rimmed cup, and the inflo-
rescence of Mezilaurus is a double raceme (Figs.
3, 8C), an inflorescence type never found in Li-
caria. Kubitzki et al. (1979) partly enumerated
these differences and discussed the placement of
Licaria mahuba (Samp.) Kostermans and L. ma-
guireana Allen. They concluded that these species
did not belong in Licaria and resurrected the
generic name inostemon to accommodate
them. Clinostemon was considered a close rela-
tive of Mezilaurus, the only difference being the
presence of staminodia in Clinostemon and their
absence in Mezilaurus. The two Clinostemon
species also share edi obovate leaves with an
abruptly rounded bas

The taxonomic DR of absence/pres-
ence of staminodia in defining genera of Laura-
ceae is open to discussion. In some genera stam-
inodia are consistently present (Persea, Phoebe);
in others they may be present or absent (Aiouea,
Aniba, Licaria, Ocotea). This suggests that a ge-
neric separation based only on presence/absence
of staminodia is weak, especially because the
staminodia are small, ca. 0.5 mm, and not easy
to find. The discovery oftwo undescribed species
in Colombia and Costa Rica with the leaf shape
and size of a Mezilaurus, but with staminodia
like Clinostemon, is another reason to place Cli-
nostemon in synonymy under Mezilaurus.

Richter (1981) amply discussed the wood and
bark anatomy of the Lauraceae. He found that
within the Lauraceae, Mezilaurus occupied an
isolated position and was easily recognized both
on wood and bark characters. He also found that
the wood (bark was not available) of the two
Clinostemon species was either undistinguisha-

iz
ae erah earlier in iie ed et ES (1979). The
great similarity in wood ana plus the iso-
lated position of Mezilaurus/ Posi aodio in the
Lauraceae is the second reason to merge these
two genera. Richter (1981) also found that Li-
caria and Endiandra are, as far as wood anatomy
is concerned, not closely related to Mezilaurus.
The genus most closely related to Mezilaurus by
wood characters is Anaueria, a monotypic genus
incompletely known from a few collections in
Brazil and Peru. Kostermans (1952) and Hutch-

WERFF—REVISION OF MEZILAURUS

155

inson (1964) both placed Anaueria in Beilsch-
miedia; Kostermans mentioned, without giving
details, that he did so as a result of studying
additional herbarium material from Rio de Ja-
neiro. Neither Anaueria nor Beilschmiedia is
likely to be confused with Mezilaurus, since they
have flowers with six or nine fertile stamens and
never have clustered leaves.

Two species described in this paper merit ad-
ditional comments. Mezilaurus ines ie
and M. glaucophylla have an unusual di
tion, being only known from northern eade
and Costa Rica. All other Mezilaurus species oc-
cur in the Amazonian forests or other parts of
Brazil. Secondly, M. quadrilocellata and M.
glaucophylla have staminodia (as do the species
formerly placed in Clinostemon) and the leaf
shape of Mezilaurus species (quite unlike the
species formerly included in Clinostemon). Thus,
they link Mezilaurus with Clinostemon. More-
over, they are the only species in the genus with
four anther cells on each stamen. Given the im-
portance frequently attached to the number of
anther cells and the number of fertile stamens,
the presence of three four-celled anthers (not
found in any other New World Lauraceae) could
be sufficient for the recognition of a new genus.
However several other genera include species with
two-celled and four-celled anthers, and because
other characters (leaves clustered at branch tip,
shape of the inflorescence) point toward Mezi-
laurus, I include these species in Mezilaurus.

As a result of the transfer of Clinostemon and
the inclusion of Mezilaurus quadrilocellata and
M. glaucophylla, my concept of Mezilaurus is
wider than has been used by previous authors. I
regard as diagnostic characters the leaf position
(clustered at the tips of branches), the small,
platelike cupule (but fruits from most species are
not yet known), the type of inflorescence (a dou-
ble raceme), and the presence of three fertile sta-
mens. Richter (1981) discussed diagnostic wood
s bark characters.

n, Mezilaurus shows in floral char-
acters a strong resemblance to Licaria. However,

differ in characters of leaf position, inflorescence
type, and number of fertile anthers. A close re-
lationship between Mezilaurus and Endiandra is
very unlikely. Currently available information
indicates that Mezilaurus, including Clinoste-

156

mon, is endemic to the Neotropics, and that it
occupies an isolated position in the family

MORPHOLOGY AND TAXONOMIC CHARACTERS

Mezilaurus species range from small trees or
shrubs (the cerrado species) to tall forest trees
much valued for their timber. The twigs are gen-
erally thick, show conspicuous leaf scars, and are

The leaves in all species are pin-
nately veined. The lateral veins frequently arch
upward and become connected with the more
distal lateral vein. The texture of the leaves is
variable; most species have chartaceous leaves,
but a few have coriaceous leaves in which sec-
ondary and tertiary venation is poorly visible.
Conspicuous gland dots in the leaves occur rare-
ly. The leaves generally turn dark upon drying.

Characteristic for the genus is the fact that the
leaves are always clustered at the tips of the
branches. Young shoots grow initially rapidly
without beanie leaves; after this elongating
period, leaves are formed at the tip of the young
branch. Such branches may have several clusters
of leaves, representing different growing seasons.
I will call this growth pattern long shoot-short
shoot. Under unfavorable conditions (several
Mezilaurus species are reported from white sand
forests or caatinga forests) the difference between
the long shoots and short shoots becomes less
pronounced and the growth pattern may seem a
succession of short shoots, with only one cluster
of leaves at the tip of the branches. However, I
think the difference between long shoot-short
shoot or short shoot growth pattern is quanti-
tative, not qualitative.

Species with clustered leaves of the long shoot-
short shoot pattern occur regularly, but not fre-
quently, in several other neotropical genera of
Lauraceae (Aniba, Endlicheria, Nectandra, Oco-
tea, Phoebe, and Pleurothyrium). However, only
in Mezilaurus is this clustered leaf pattern char-
acterisitic or dominant. The non-Mezilaurus
species with clustered leaves are rarely confused
with Mezilaurus; even in vegetative state they
are readily separated by conspicuous pubescence
or leaf color differences. Only one species, Ocotea
rubra, very closely resembles Mezi/aurus in ster-
ile state. However, its flowers with nine four-
celled stamens and fruit with a large cupule make
identification easy

early all species of Mezilaurus have elliptic
to obovate leaves. The base of the leaves, how-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

ever, offers some useful characters. Four species,
ezilaurus mahuba, M. pyriflora, M. thoroflora,
and M. duckei, have large leaves (to 60 cm long)
which become gradually narrowed toward the
base; at the base the leaves are abruptly nar-
m: becoming rounded or even cordate. Three
pecies, M. subcordata, M. quadrilocellata, and
M glaucophylla, have an obtuse or rounded leaf
base with a distinct petiole, 2-6 cm long. In these
three species the leaf shape is slightly obovate or
ovate. In all other species (with the exception of
M. crassiramea), the leaf base is gradually atten-
uate or decurrent on the petiole. Most of these
species have a petiole. Three species, however,
M. caatingae, M. crassiramea and M. decurrens,
have sessile leaves or nearly so. In some collec-
tions of M. crassiramea the leaf base is gradually
narrowed, in others it is rather abruptly nar-
rowed. In this species the petioles, if present, are
less than 1 cm long
Pubescence. There is not much variation in
pubescence within Mezilaurus. Two species, M.
crassiramea and M. lindaviana, have erect pu-
bescence on the lower surface of the leaves; this
can be quite sparse in M. /indaviana, however.
The other species have varying amounts of ap-
pressed pubescence on leaves, stems, terminal
buds, and inflorescences. These varying amounts
of pubescence have little diagnostic value.
Inflorescence. The inflorescence of Mezilau-
rus consists of acompound raceme (dibothryum,
see Weberling, 1981, 1985). This inflorescence
type is present without modifications in M. cras-
siramea, M. lindaviana, M. mahuba, M. pyri-
flora, M. thoroflora, and M. duckei. Short tertiary
axes are sometimes present in M. glaucophylla
and M. quadrilocellata. In the other species the
inflorescences are smaller and the flowers are not
evenly distributed along the lateral branchlets of

clustered flowers are a derive
worth noting that the species with a well-devel-
oped dibothryum have larger (often much larger)
inflorescences and have usually smaller flowers
than the species with clustered flowers. A di-
bothryum is a rare inflorescence type among oth-
er neotropical Lauraceae (if it ever occurs outside
of Mezilaurus).

The inflorescence type of one Mezilaurus
species, M. decurrens, is not known due to the
fragmentary nature of the single available spec-
imen.

1987]

Flowers. In the following discussion I con-
sider only mature flowers. Because it is difficult
to tell whether a flower is mature or not in Me-
zilaurus, I define a mature flower as one with
opened anther cells. In buds or young flowers
diagnostic characters are often difficult to see.

The flowers of Mezilaurus are unusually dif-
ficult to dissect. In addition to their small size,
the floral tube contains much mucilage, which
has usually hardened during drying. Softening
the flowers requires boiling for at least one hour.
After softening, the mucilage becomes viscid and
sticky and the dissected floral parts frequently
stick tightly to the floral tube. Therefore, I have
rarely relied on characters of the floral parts to
separate taxa, especially because other characters
are available.

Dimensions given for flowers and their parts
should be accepted with some reservation. In
order to dissect the flowers, it is necessary to boil
them. During the boiling the flowers swell; the
degree of size increase depends on the duration
of boiling. I have recorded flowers that measured

3 mm dry as swelling to 1.7 mm after boiling.
Similar increases in size were found for the sta-
mens as well. Only rarely have I used flower sizes
in the key and, in these cases, flower sizes are
taken from dry flowers. i i are acu
small and equal. The exceptions are M.
cophylla and M. pedem in S the
outer three tepals are smaller than the inner three.

With the exception of three species (Mezilau-
rus caatingae, M. palcazuensis, and M. mahuba),
all species have clearly pedicellate flowers. Stam-
inodia (which are not easy to find due to their
small size) are present in six species (M. duckei,
M. glaucophylla, M. mahuba, M. pyriflora, M.
quadrilocellata, and M. thoroflora), and only M.
mahuba has glands at the base of the fertile sta-
mens.

The most interesting variation of floral struc-
tures 1s found in the shapes and positions of an-
thers. In most neotropical Lauraceae the anthers
have the shape of erect stalked plates with the
anther cells on the inner or outer surface (for
instance, in Persea, Phoebe, Nectandra, and Oco-
tea). In several of the genera with two-celled an-
thers the plate shape of the anthers is less pro-
nounced or lost and the anther cells are situated
at or near the tip of the anthers (Aniba, Licaria).
In several species of Licaria the stamens are
shaped like columns with the anther cells situ-
ated near the apex and here the difference be-

WERFF—REVISION OF MEZILAURUS

157

tween filament and anther has disappeared. In a
few species of Mezilaurus (M. duckei, M. glau
cophylla, M. pyriflora, and M. edv san en a
similar arrangement is found. Here the stamens
remain included in the erect tepals and the anther
cells are situated at the tips of the stamens. In
M. glaucophylla and M. quadrilocellata, the only
species with four anther cells on each stamen,
the tips of the stamens are flattened and form a
small platform on which the anther cells are sit-
uated. Thus, by looking from the outside in the
flowers, one sees the 12 anther cells as small
pores. In these four species the anther cells are
situated iim and also open apically; that is,
the flaps open upwar

Mezilaurus ot is the only species with
the anther cells situated introrse or introrse-lat-
eral; in all remaining species the anther cells are
extrorse, situated on the outer face of the stamen.
Here the anther cells are more or less exserted,
tend to be relatively large, and, most interesting,
the stamens develop a dorsal ridge on which the
anther cells are situated. The ridge is usually about
as long as the anther cells and appears as an
outward-facing hump near the tip of the stamen.
The anther cells open towards the crest of the
ridge, where the flaps are situated back-to-back.
Such anther cells have been described as opening
“laterally” in the literature, but in order to avoid
confusion with laterally situated anther cells
(which occur, for instance, in Pleurothyrium), I
will call this type of opening ‘“‘back-to-back.”

n decurrens the dorsal ridges are not
strongly dexeloned and the anther cells are hard-
ly exserted. In the remaining species the anther
cells are clearly exserted.

e most extreme development is found in M.
ahuba, where the dorsal ridges with the anther

fro

greatly exserted anther cells does not correspond
with variation in other characters. In fact, the
extremes (M. pyriflora with immersed anther
cells, M. mahuba with greatly exserted anther
cells) are very similar in other characters such as
leaf shape, leaf size, and type of inflorescence.

In a few species (M. crassiramea, M. linda-
viana, M. palcazuensis, M. sprucei, M. subcor-
data, and M. synandra) the filaments of the sta-
mens are fused in a ring or a short tube. This is
a useful character, but because it is hard to rec-
ognize, I have not used it in the key. It is most
easily seen on young fruits. When the filaments
are free, they are visible at the base of the fruit;

158 ANNALS OF THE MISSOURI BOTANICAL GARDEN

when the filaments are united, they are visible
as a small cap on top of the fruit.

REPRODUCTIVE BIOLOGY

In a study of the reproductive biology of some
neotropical Lauraceae, Kurz (1983) and Kubitz-
ki and Kurz (1984) reported that a Clinostemon
species here described as Mezilaurus duckei
showed synchronized dichogamy with a pro-
nounced protogyny.

In this system, two flowering morphs are found.
In the A morph, flowers open in the morning
and expose the receptive stigmas. During this
phase, no pollen is released. Around midday, the
stigma wilts and is no longer receptive. The male
phase, when the anthers shed the pollen, takes
place in the afternoon. An individual flower thus
lasts only one day. In the B morph, flowers open
in the afternoon, when the stigmas are receptive.
Pollen is released during the morning of the fol-
lowing day.

During the fieldwork, Kurz was able to observe
only two flowering Mezilaurus trees and these
turned out to be both A morphs. However, he
studied A and B morphs of other Lauraceae
species and found that for fertilization, cross-
pollination between A and B morphs was nec-
essary. Selfing of A or B morphs did not result
in seed set. It is likely that these findings apply
to Mezilaurus as well.

The Mezilaurus flowers observed by Kurz were
visited by four species of small (2-3 mm) Trigona
bees (Meliponinae). Mezilaurus flowers are not

[Vor. 74

known to produce nectar and it is likely that
pollen is the only reward for their pollinators.

TAXONOMIC TREATMENT

Mezilaurus Taubert, Bot. Centralbl. 50: 21. 1892.
TYPE: M. navalium (Allemao) Taubert.

Silvia Allemào, Dissertatio, Rio de Janeiro. 1848, non
Conc. Silvaea Meissner, DC. Prodr. 15: 84.

wa
Natürlichen Pflanzenfamilien, Nachtrag zu Teil
II-IV. 1897, nom. super
Clinostemon Kuhlm . & Samp., B s. Nac. Rio de
Janeiro 4(2): 57. 1928. TYPE: ys ep a w (Samp.)
uhlm. & Samp.

Shrubs to tall trees, mostly South American,
but with one species in Costa Rica. Leaves al-
ternate, usually congested at the apex of the twigs,
entire. Petioles often swollen at base. Inflores-
cences axillary, sometimes seemingly terminal,
few- to many-flowered, forming a compound ra-
ceme (dibothryum); flowers clustered at the tips
of the inflorescence branchlets in several species.
Bracts and bracteoles deciduous. Tepals 6, equal
or nearly so, small, scalelike, usually erect. Fertile
stamens 3, representing the third whorl, 2-celled
(in two species 4-celled). Staminodia present or
absent. Staminal glands present in one species.
Filaments free or connate; anther cells usually
extrorse and exserted, situated on a dorsal ridge.
Ovary ellipsoid to ovoid, included in the flower
tube. Fruit ellipsoid, situated on a small, plate-
like cupule.

KEY TO MEZILAURUS

la. Stamens 4-celled; N. Colombia and Costa Rica

2

ter tiary

FF

quadrilocellata

N. Colombia .......
2b. Inflorescence gray ron idi glabrous or with few basal hairs; tertiary ventu on upper

leaf surface raised; Cost

M. P

=

Stamens 2-celled; S. America 'E " the Andes
3a

Leaves d narrowed toward base, usually abruptly rounded there, generally large, ex-

nes 25c

+} 4

uetlal

Amazon

1 To1 3a E POLI
J

from fl

M. mahuba
5

4b. esum pedicellate; anthers not hooklike exserted; not occurring in flooded forest
Sa. Leaves elliptic or slightly obovate, rounded at tip; anthers not exserted at anthesis

M. duc s

Sb. Leaves strongly obovate, acute or acuminate at tip; anthers exserted or not ........
6a. Ant

hers exserted at anthesis; pedicels 4-8 mm long

Anthers included at anthesis; pedicels 1.5-2 mm lon

3b. Leaves decurrent or obtuse at base, generally ue rarely exceeding 20 cm
Leaves

M. oda
SM M. pyriflora
7

7a pubescent below; twigs visibly pubescent
a owers pubescent; shrub or id tree in cerrado vegetation U U M. crassiramea
Flo abrous; tree in rain fore M. lindaviana
7b. Leaves glabrous below; twigs not visibly pubesce 9
f tip acute (in M. micrantha apex is iid but present) 10
9b. Leaf tip rounded 12

1987]

10a. Flowers sessile; Per

WERFF—REVISION OF MEZILAURUS

159

10b. Flowers pedicellate; "Brazil or Peru .
lla. Leaves coriaceous; pedi
11b. Leaves — wa: pedicels (5)10-15 n long

abrous

12a. Flow

M. palcazuensis
11

M. micrantha
M. sprucei
13

cels ca. 2 mm

Flowers sessile; caatinga forests along Rio Negro ... j caatingae

13b. Flowers en rain forests of S. Brazi

12b. Flowers pubesce

ll u. ge

14a. Leaf base yee or rounded; anther cells (lateral) uen N

14b. Leafb

11 š *4 th +

subcordata
1

a.

16b. Leav

l

wn
o

l

M
S

Mezilaurus caatingae van der Werff, sp. nov.
TYPE: Brazil. Amazonas: Rio Negro, Sao Fe-
lipe, caatinga on sandy soil, tree, 15 m, 27
Sept. 1952, Fróes 28761 (holotype, MO).
Figures 1, A & B; 2

Arbor, 15 m. Ramuli teretes, glabri. Folia coriacea,
$ ne ‘ad apices. ramulorum, glabra, margine invo-
0-15 x
Poe super laevia nitidaque, subtus opaca costa
elevata nervis lateralibus et venatione immersa. Petioli
. 1 cm lon

axillaris

pubes scens (basis inflorescentia tantum adest). Flores
labri vel basi leviter minute adpresse pubescentes,
ronis, sessiles, conferti ad apices ramulorum inflo-
rescentiae sie m longi. Tepala 6, aequalia, erecta,
pa 0 m). Stamina 3, per anthesim exserta, |
, extrorsis lateraliter des
ntibu us. Filamenta libra, pubescentia, antheris ae-
e re sm Tubus per glaber. Ovarium gla-
brum, globosum, 0.5 m ylus 0.6 mm longus, per
anthesim a Fructus Med

Tree, 15 m. Twigs terete, glabrous. Leaves
clustered at the tips of biais glabrous, cori-
aceous, obovate, 10-15 x 4—5 cm, the tip round-
ed, the base gradually debi to the short pet-
iole, the margin inrolled, the upper surface
smooth, slightly lustrous, the lower surface with
a raised midvein; lateral veins (8-12) and reti-
culation + immersed; petioles ca. 1 cm, the lam-
ina decurrent as two narrow ridges. Inflorescence
axillary, minutely appressed pubescent, panicu-
lately branched, broken on the specimen seen.
Flowers glabrous or with some minute pubes-
cence at the base, sessile, clustered at the tips of
the inflorescence branchlets, 2.2 mm long, ob-

15a. Tepals erect; : flower tube not ‘constricted at apex flowers
cup-shaped ..

. Flowers 1.5 x 1.5 mm; upper leaf surface PE
M.

ALULUISUC

Leaves sessile or nearly so; anthers scarcely exsert-

m M. decurrens
1 cm long; anthers
M. itauba

. Tepals incurved; Ere tube constricted at apex; flowers
depressed globos
17a. Flowers 2 x iege

m; upper leaf surface shiny and

with raised reticulation ynandra

the reticulation not raise opaca

conic. Tepals 6, very small, 0.2 mm long, equal,
erect. Stamens 3, exserted at anthesis, | mm long,
the anthers 2-celled, extrorse, opening back-to-
back; filaments free, as wide as anthers, pubes-
cent. Floral tube glabrous. Ovary globose, gla-
brous, 0.5 mm. Style slender, 0.6 mm long,
exserted at anthesis. Fruit unknown.

In leaf shape Mezilaurus caatingae resembles
M. itauba and M. decurrens. The sessile flowers

separate it immediately from these species, how-
ever. Vegetatively, the recurved leaf margin is
diagnostic. This character also occurs in M. mi-
crantha, but this species has a bluntly acute leaf
apex, never rounded as in M. caatingae.

Mezilaurus crassiramea (Meissner) Taubert ex
Mez, Arbeiten Kónigl. Bot. Gart. Breslau 1:
112. 1892. Oreodaphne crassiramea Meis-
sner, DC. Prodr. 15(1): 117. 1864. Silvia
crassiramea (Meissner) Mez, Jahrb. Kónigl.
Bot. Gart. Berlin 5: 106. 1889. Mezia cras-

siramea (Meissner) Kuntze, Revis. G

2: 574 TYPE: Brazil. Goias: Sects

d’Ourada, Pohl 1463 (W, K, G-DC, U—this

the only specimen seen). Figures 1, C & D; 2.

Small trees, to 6 m tall. Twigs terete, thick, the
older ones with a thick and conspicuous cork
layer, the young tips with a dense, light brown
tomentum. Leaves clustered at the tips of the
twigs, almost sessile (petioles to 3 mm long), ovate
or slightly obovate, the tip rounded, the base

160 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

Figure 1. A-B. Mezilaurus Ap in — A. Leaf. — B. Flower. C-D. M. crassiramea. —C. Leaf. — D. Flower.
E-F. M. decurrens. — E. Leaf. — F. Flower. G-H. M. itauba. — G. Leaf. — H. Flow

rounded or gradually narrowed and abruptly
rounded, with margins frequently recurved, 10 x
5 cm, the upper surface pubescent, but becoming
glabrous with age, the lower surface rather dense-

ly pubescent with pale brown hairs; lateral veins
10-15 pairs, leaving the midvein under almost
90*: veins and the final reticulation immersed on
upper surface, raised on lower surface. Inflores-

1987]

WERFF—REVISION OF MEZILAURUS

O 100 200 300 400 500 600 miles

80 70 60

50 40

FIGURE 2. Distribution of Mezilaurus caatingae (), M. crassiramea (8), M. decurrens (A), and M. duckei

cences in the axils of small bracts, seemingly ter-
minal, 7-12 cm long, about as long as leaves,
pedicellate, puberulous or with very short pu-
bescence, the fl ged spicatel
lateral branchlets; branchlets and flowers sub-
tended by puberulous, ovate bracts, these ca. 0.1
mm long, the flowers brown puberulous, ca. 2
mm long; pedicel ca. 2 mm. Tepals 6, equal, ca.
0.8 mm long, erect, ovate. Stamens 3, exserted;
anthers 2-celled, the cells small, on dorsal ridge
of anthers, opening back-to-back; filaments fused
into a tube. Ovary ovoid, ca. 1 mm, the style

along the

gradually narrowed, ca. 1 mm, exserted beyond
anthers. Staminal glands and staminodia lacking.
Young fruit globose, 6 mm diam., seated on
swollen pedicel, the tepals persisting.

Vernacular name. Cumbuquinha (fide Rat-
ter).

Additional specimens examined. BRAZIL. GOIÁS:
Serra Dourada, Anderson 10003 (F, NY, MO, US).
MATO GROSSO: 7 km SW of Xavantina, Ratter et al.
805 (MO); ca. 270 km N of Xavantina, Ratter 1293
(MO).

162

Mezilaurus crassiramea is a well-defined
species known from a few collections in cerrado
vegetation. Diagnostic characters are the thick
corky twigs, the pubescent leaves, and puberu-
lous flowers. It can, as many cerrado species,
withstand fire; the Ratter collections come from
trees with charred or fire-blackened trunks.

Mezilaurus decurrens (Ducke) Kosterm., Med-

ed. Bot. Mus. Herb. Rijks Univ. Utrecht 25:
40. 1936. Silvia decurrens Ducke, Trop.
i 35

Yale No. 20999 (lectotype, RB, not seen;
isolectotype F, fragm. U). Figures 1, E & F;
2

Large tree. Twigs minutely puberulous toward
apex, the terminal bud with yellowish, appressed
pubescence. Leaves clustered at the tips of
branches, glabrous on both surfaces, elliptic or
narrowly elliptic, 15-25 x 5-7.5 cm, the tip
rounded, the base gradually decurrent onto the
petiole, this 1-2 cm long; laminae coriaceous,
opaque, the reticulation not raised, rather lax.
Lateral veins not strongly developed, 7-12 pairs,
immersed above, slightly raised below. Midrib
thick, dark, raised above, more conspicuously so
on lower surface. Inflorescences axillary, subter-
minal, pyramidal, 10-18 cm long (fide Koster-
mans), appressed pilose. Flowers subglobose or
obconical, ca. 2-2.5 mm long, 2 mm wide, ap-
pressed pubescent. Tepals 6, equal, erect, trian-
gular, wider than long. Fertile stamens 3, 2-celled,
1-1.5 mm long, slightly exserted, pubescent; fil-
aments connate, wider than anthers; anther cells
extrorse, opening back-to-back. Ovary subglo-
bose, densely pubescent (except base), ca. | mm
long, the style slender and briefly exserted. Fruit
unknown.

Mezilaurus decurrens is rare and known to me
with certainty only from the type collection. At
first glance it appears quite similar to M. itauba,
but differs in several subtle characters. The leaves
of Mezilaurus decurrens do not have the raised
reticulation and minute gland dots of M. itauba,
and their leaf bases taper more gradually into the
tiga Better differences are found in the flow-
ers: in M. decurrens the anthers are scarcely
ed and the filaments are connate, whereas
in M. itauba the anthers are greatly exserted and
the filaments free.

A few collections that I place in Mezilaurus

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

itauba are close to M. decurrens in leaf outline
(Ducke 681, Foldats 3613), but I attach more
diagnostic value to the raised reticulation and
the greatly exserted anthers

Mezilaurus duckei van der Werff, sp. nov. TYPE:
Brazil. Amazonas: Reserva Florestal Ducke,
Aleusio 98 (holotype, US). Figures 2, 3.

Arbor, 20 m alta. Ramuli crassi, cicatribus conspi-

a :
uper obscura, subtus pude Peti li crassi, ca.

e pyram
persistentibus et puberulis. Flon ores spicatim secus ra-
mulos inflorescentiae dispositi, frequenter reflexi, tur-
binati, ca. ] mm
vel pubescentia laxa ad basim. Tepal
— ca. 0.3 mm longa. Stamina fertilia 3, ca.

m longa, filamentis pubescentibus, liberis, latioribus
eae antheris glabris 2-locellatisque. Staminodia 6,
parva, ca. 0.2 mm longa. Ovarium conicum, glabrum
ca. 0.4 mm longum, stylo ca. 0.7 mm longo. Fructus
ignoti.

Tree, 20 m. Twigs thick, 1 cm diam. 5 cm
below the tip, with conspicuous leaf scars, the
tip very finely brown tomentellous. Leaves clus-
tered at the tips of branches, slightly hien
25-30 x 8-10 cm, the tip rounded, gradua
narrowed toward the base, the base abruptly nar-
rowed; upper surface glabrous, opaque, venation
immersed, the tertiary venation scarcely visible;
lower surface minutely puberulous when young,

secondary and tertiary venation raised, the mid-
rib strongly raised and thick; petioles 0.5 cm thick,

cences axillary, ca.
branched, the branchlets 3-4 cm long, the u
ones slightly shorter than the lower ones, mi-
nutely brown puberulous. Bracts and bractlets
pilose, persisting at anthesis, the bracts 1.5 mm
long, bractlets ca. 0.7 mm long. Flowers pedi-
cellate, pedicels ca. 2 mm long, glabrous. Flowers
glabrous, cup-shaped, | mm long, tepals 6, small,
0.3 mm long, erect. Fertile stamens 3, ca. 0.5
mm long, the filaments free, pubescent, wider
than the glabrous, 2-celled anthers. Staminodia

, ca. 0.2 mm long. Ovary conical, glabrous, ca.
0.4 mm long, the style ca. 0.7 mm long. Fruit
unknown.

1987]

WERFF—REVISION OF MEZILAURUS

FIGURE 3. Mezilaurus duckei. — A. Habit. — B. Flower.

163

164

Common name. Itauba abacate.

Paratype. BRAZIL. AMAZONAS: Reserva Florestal
Ducke, tree nr. 116, Rodrigues 8203 (NY).

This species is named after Adolpho Ducke,
an outstanding botanist and collector who made
numerous excellent collections of Lauraceae in
Amazonian Brazil. It is fitting that this new species
is only known from the forest reserve dedicated
to Ducke

Menzilaurus glaucophylla van der Werff, sp. nov.
TYPE: Costa Rica. Prov. San José: Zapatón
de Puriscal, tree, 9 m, Zamora & Poveda

1014 (holotype, MO; isotypes, F, CR). Fig-

ure 4

r, 20 m. Ramuli teretes, cicatribus insignibus

viter €—— vel triangulares, cinereo-pubescentes,
. Folia ad apices ramulorum conferta,
obovata, 2. subtus glauca, apicibus basibusque

illares, pyramida atae, cinereo-strigosae. Pedicelli ad 1.5
mm longi, glabri vel basim aliquot pilis. Flores glabri,
il ees in sicco ca. 1 mm longi et lati. Tepala
6, per anthesin erecta vel paullo i incurvata; inaequalia,
3 exteriora interioribus breviora, late deltoidea. Sta-
mina 3, 4-locellata, filamentis latitudine antheris ae-
quantibus, ca. 1 mm longa, ca. 0.7 mm lata, tepalis
exterioribus opposita. Staminodia 3, ca. 0.5 mm longa,
li ind a tepalis interioribus opposita. Tu-
bus flor vadosus, intus glaber. Ovarium glabrum,
silipscideum. sensim in stylo attenuatum, ovarium
stylusque ca. | mm longus. Fructus ignotus.

Tree, to 20 m tall. Twigs terete, with conspic-
uous leaf scars, minutely pubescent, the terminal
bud white appressed pubescent; bark on older
twigs soft and flaking. Leaves clustered at the
tips of the branches, alternate, obovate, charta-
ceous, glaucous and laxly and minutely puber-
ulous below, the tip rounded or acute, the base
obtuse or acute, 15-25 x 8-12 cm, the midrib
and lateral veins immersed, but the tertiary ve-
nation slightly raised above, glabrous except for
the puberu midrib; midrib and lateral vein

rescences axillary, pyramidate, to 15 cm long,
gray strigose. Pedicels ca. 1.5 mm long, glabrous

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

or with few gray, appressed hairs at the base,
subtended by 2 small, deltoid, strigose bracts, ca.
0.2 mm long. Flowers glabrous, more or less cup-
shaped, ca. | mm long and wide when dry. Tepals
6, at anthesis more or less erect, the outer three
smaller than the inner three, broadly deltoid. Sta-
mens 3, all 4-celled, the filaments as wide as the
anthers; glabrous or with a few hairs at the base,
ca. 1 mm long, 0.7 mm wide, situated opposite
the outer tepals; tips of the stamens curved in-
ward; anther cells positioned on the upper part.
Inner tepals pushed apart at anthesis and expos-
ing anther cells in their sinuses. Staminodia 3,
alternating with stamens, lanceolate, strigose, ca.
0.5 mm long. Ovary glabrous, ellipsoid, gradu-
ally narrowed into style, ovary and style ca. 1
mm long. Floral tube shallow, glabrous inside.
Fruit unknown.

types. CosrA RICA. PUNTARENAS: Osa Penin-
E in E t W. of Rincón, Hammel et al. 15214 (MO;
duplicates to be distributed).

There is no doubt that Mezilaurus glauco-
phylla and M. quadrilocellata are closely related.
They differ from the other Mezilaurus species in
the following characters: distribution (the only
species north of the Andes); presence of four-
celled anthers; leaves more or less glaucous be-
low; unequal tepals; and the inflorescence not
strictly a dibothryum, but sometimes with short
tertiary axes. These two species could be regard-
ed as forming a new genus, based on their four-
celled stamens and unequal tepals, but it should
be mentioned that several other Mezilaurus
species have small tepals that one cannot very
well judge to be equal or not. If additional dif-
ferences separating M. glaucophylla and M.
quadrilocellata from the other Mezilaurus species

are found (in cupule shape, for instance), it might
be better to treat them as a separate genus, but
for the time being, I prefer to include them in
Mezilaur

In addition to the THeteneps mentioned in
Table 1, the few a
that Mezilaurus elaucophylla has larger, "innt
leaves and larger inflorescences than M. quad-
rilocellata.

suggest

Mezilaurus itauba (Meissner) Taubert ex Mez,
Arbeiten Kónigl. Bot. Gart. Breslau 1: 112.
1892. Acrodiclidium itauba Meissner, DC.
Prodr. 15(1): 86. 1864. Endiandra itauba
(Meissner) Benth. & Hook., Gen. Pl. 3: 154.
1880. Silvia itauba (Meissner) Pax, Natür-
lichen Pflanzenfamilien 3(2) 123. 1889.

1987] WERFF—REVISION OF MEZILAURUS 165

FicunE 4. Mezilaurus glaucophylla. — A. Habit. — B. Flower and bud seen from above.

Mezia itauba (Meissner) Kuntze, Rev. Gen. Acrodiclidium itauba Meissner var. amarella Meissner,
à DC. Prodr. 15(1): 86. 1864. Type: Brazil. Para:

Š " : A. PE:
Pl. 2: 574. 1891. TYPE: Brazil. Para: Santa- Santarem, Spruce 646 Qectotype not chosen, Dij;
rem, Spruce 643 (lectotype, K, fide Koster- i
mans, BM, C, fragm. F). Figures 1, G& H; 5. Oreodaphne hookeriana Meissner, DC. Prodr. 15(1):

166 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74
TABLE 1. Comparison of Mezilaurus glaucophylla with M. quadrilocellata.
M. glaucophylla M. quadrilocellata
Inflorescence gray strigose rufous tomentellous
Pedicels few basal hairs, otherwise glabrous rufous tomentellous

Tertiary venation on upper leaf surface
Terminal bud

aise
densely gray pubescent

immerse
brown tomentellous

131. 1864. Type: Brazil. Para: Santarem, Spruce
669 (not seen.
ezilaurus anacardioides deri Taubert ex Mez,
Arb. Bot. Garten Breslau 1: qas Acrodi-
clidium anacardioides Meissner, m r. 15(1):
a kyi Misanteca anacardioides eee
. & Hook., Gen. Pl. 3(2): 155. 1880. Silvia
d (Meissner) Mez, Jahrb. Bot. Gart.
Berlin 5: 108. 1889. Mezia anacardioides (Meis-
sner) Kuntze, Revis. Gen. Pl. 2: 574. 1891. TYPE:
Venezuela. Amazonas: San Carlos de Río Negro,
Spruce 2961 (lectotype not chosen, BM, G).
Silvia polyantha Mez, Bull. Herb. Boiss. 2 sér. V: 233.
. TYPE: Brazil. Amazonas: Moura, Rio Ne-
gro, Ule 6055 (holotype, B, not seen, isotype, G).
Silvia rondonii Mez et Hoehne, Bot. Archiv VI: 230.
4. TYPE: Brazil. Mato Grosso: near Tres Bu-
ritys, Kuhlmann 1976 (K, not seen).

Large trees to 35 m tall, rarely shrubs. Twigs
terete, glabrous or nearly so, the terminal bud
appressed pubescent, the bark rather thin and

aking. Leaves clustered at the tips of branches,
firmly chartaceous or coriaceous, glabrous at ma-
turity, elliptic or slightly obovate, ca. 15 x 5 cm,
the base gradually narrowed into petioles, the tip
rounded, the reticulation on both surfaces slight-
ly raised, the midrib and lateral veins (7-12 pairs)
immersed above, raised on lower surface, the
lateral veins arching upward and fading near the
margin. Lower leaf surface densely and minutely
gland dotted; petioles glabrous, with swollen bas-
es, 1-2.5 cm long. Inflorescences axillary, sub-
terminal, narrowly pyramidate, 5-10 cm long,
laxly appressed pubescent. Flowers subumbel-
lately arranged at tips of lateral branchlets, laxly
and minutely appressed pubescent. Pedicels mi-
nutely appressed pubescent, 2-4 mm long. Bracts
and bractlets deciduous. Flowers hemispherical,
1.5-2 mm long, the 6 equal tepals erect (rarely
spreading), wider than long, the anthers exserted.
Fertile stamens 3, 2-celled, ovate-elliptical, ca.

the cells large, opening back-to-back. Ovary el-
lipsoid, pubescent, the style exserted. Flower tube
pubescent. Staminal glands and staminodia lack-
ing. Fruit an ellipsoid berry, ca. 2 x 1 cm, sub-
tended by a small, platelike cupule.

Selected additional specimens examined. SURINAM.
Boschreservaat, sectio O, tree No. 760, Boschwezen
3088 (NY). BRAziL. PARÁ: Rio Tapajos, Villa Braga,
Ducke RB 17537 (G, US). PARÁ: Serra dos Carajas, M.
G. Silva 2909 (MO); Rio Jari, Monte Dourado, N. T.
Silva 1041 (NY). PERU. MADRE DE DIOS: Tahuamanü,
Diaz 17 53-96 (MO). Borivia: San Francisco, 50 km
from Pto. Rica Pando, E. Menesk 626 (MO)

Mezilaurus itauba is the most frequently col-
lected and widest ranging species of the genus.
In addition to the countries listed, it has been
reported from French Guiana, based on a Mé-
linon collection I have not seen. Although there

s some variation in degree of pubescence and
leaf shape throughout its range, M. itauba is ad-
equately characterized by free staminal fila-
ments, pubescent flowers, and glabrous leaves
rounded at the tip and gradually narrowed ba-
sally. The anthers, with large cells, are also long
exserted for their size. The numerous gland dots
on the lower leaf surface are best seen on dena
young leaves; on mature, more coriaceous ve
they are often scarcely visible.

Alencar 55 (MO) is included in M. itauba with
hesitation; it differs somewhat in leaf shape, but
in the absence of floral differences I regard it as
M. itauba. However, when more collections are
sci it might turn out to be a new species.

ood of Mezilaurus itauba is hard and
much pie for construction. Mez (1889) reported
that the berries are edi

Mezilaurus lindaviana Schwacke et Mez, Arb.
B arten Breslau 1: 112. 1892. TYPE: Bra-
zil. Amazonas: Rio Branco, Schwacke 7080
(lectotype, chosen by Kostermans, B, not
seen). Figures 6, A & B; 7

Misanteca duckei Samp., Commissao Linhas Telegr.
Estrat. Matto Grosso Amazonas, Publ. 56 (An-
f 1917. Silvia duckei

M wurdackiana C. K. Allen, Mem. New York

rden 10: 56. 1963. TYPE: Venezuela. Bo-

a Hato La Vergaretia, Wurdack & Guppy 91
(holotype, NY, isotype, US).

1987]

WERFF—REVISION OF MEZILAURUS 167

O 200 400 600 800 1000km
L ee eee
—— ===

O 100 200 300 400 500 600 miles

30 70 60

FIGURE 5. Distribution of Mezilaurus itauba.

Tree, to 25 m. Twigs thick, round, witha rather
thick corky layer, the tips with brown velutinous
pubescence. Leaves alternate, clustered at the tips
of the branches, chartaceous or coriaceous, ob-
ovate, 8-17 x 4-8 cm, the tip rounded or very
shortly acute, the base cuneate or abruptly
rounded; young leaves hirsute, becoming gla-
brous above (except on midrib and primary veins)
at maturity; venation impressed above, the mid-
rib, secondary and tertiary venation raised be-
low; secondary veins about 10 pairs; petioles
short, thick, brown-tomentose, 5-8 mm long. In-

50 40

florescences axillary, mostly subterminal on
branches, pyramidal, tomentellous, about as long
as the leaves; branchlets patent, 1-3 cm long, the
flowers spicately arranged and often somewhat
recurved. Flowers white or yellow-green, fra-
grant, glabrous, 1.2 mm long. Tepals 6, equal,
triangular, ca. 0.2 mm long, 0.4 mm wide. Fertile
stamens 3, 1 mm long; filaments (ca. 0.5 mm
connate, narrower than anthers, pubescent; an-
thers shortly exserted, 2-celled, 0.5 mm long, the
cells situated on a dorsal ridge, opening back-to-
ack. Ovary ellipsoid, 0.8 mm long, with slender

168

FIGURE 6.
F. M. micrantha. —E. Leaf.—

1 E i +h +h bre |

mens t basal
glands. Staminodia lacking. Fruit ellipsoid, 2 x
1.2 cm, subtended by a small platelike cupule
(ca. 3 mm diam.).

QC] "TT | Pe LPT

ANA:
Kanuku Mountains, For. Dept. Brit. Vies 5804 (NY),
same inpet Smith 3208 (F. MO, US). BRAZIL.

NAS: Rio ids nco, Boa Vista, Ducke 1336 (F, NY,
US) A ing of Macapa, Fróes & Black
27453 (NY). es Pinsha de Santarem, Fróes 30976

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

3cm

A-B. “yayawa ee —A. Leaf.—B. Flower. C-D. M. mahuba. —C. Leaf. — D. Flower. E-

(NY, US); Rio Jari, Monte Dourado, E. Oliveira 4749
(NY), same location, N. T. Silva 996 (NY, US).

Mezilaurus lindaviana is somewhat variable
in leaf shape but is clearly characterized by the
combination of pubescent leaves and glabrous
flowers. Mezilaurus crassiramea, a similar species
with pubescent leaves, has pubescent flowers and
is known only as a shrub or a small tree in cerrado
vegetation.

1987]

WERFF—REVISION OF MEZILAURUS

169

200 400 600 800 1000km

eS es eee eee eee |

O 100 200 300 400 500 600 miles

80 70

40

FIGURE 7. Distribution of Mezilaurus lindaviana (6), M. mahuba (A), and M. micrantha (gp.

Silva 2403 (NY, 2 sheets, MO) is included in
M. lindaviana as an aberrant collection; it differs
from other collections of that species in having
sparse appressed pubescence. Other characters
(leaf shape, size, flowers) point to M. /indaviana
and I feel that a single collection with unusual
pubescence need not be given taxonomic status.

Kostermans (1938) cited as type of M. linda-
viana Schwacke 7080 — Glaziou 19798 and gave
Serra d'Antonio Pereira in Minas Gerais as the
type locality. Schwacke and Mez (Mez, 1892)
cited only Schwacke 7080 as type collection and

gave as type locality “in campis ad Rio Branco."
It is likely that Glaziou distributed duplicates of
the Schwacke collection under his own name with
incorrect locality data, as he did with other col-
lections (Wurdack, 1970). Therefore, I ignore the
reference of M. lindaviana as occurring in Minas
Gerais, as cited by Glaziou (1905-1913) and
Kostermans (1938).

Mezilaurus mahuba (Samp.) van der Werff, comb.
nov. Basionym: Acrodiclidium mahuba
Samp., Commissao Linhas Telegr. Estrat.

170

Matto Grosso Amazonas, Publ. 56 (Annexo

5, Bot. Parte X): 14. 1917. Clinostemon ma-

huba (Samp.) Kuhlm. & Samp., Bol. Mus.

Nac. Rio de Jan. 4(2): 57. 1928. Licaria ma-
T

(Samp.) Lundell, Wrightia 4:
TYPE: Brazil. Para: Gurupa, Varzea do Rio
Amazonas, Ducke MG 16538 = RB 17582
(isotype, U). Figures 6, C & D; 7.

Large trees. Twigs thick, glabrescent, with dense
brown tomentum when young. Leaves large, 20-
40 x 12-15 cm, obovate, clustered at the tips of
branches, glabrous above with the exception of
the puberulous midrib, softly pubescent below,
the apex rounded, the base cuneate; venation
immersed on upper surface; midrib, lateral veins
(15-23 pairs) and tertiary venation raised below;
petioles thick, 5 mm diam., 2-3 cm long, densely
and minutely tomentose. Inflorescences subter-
minal, densely puberulous, large (20-35 cm long),

minute, incurved. Fertile stamens 3, all 2-celled;
filaments densely strigose, with 2 glands attached
a little above the base; anthers glabrous, strongly
curved outside the flower tube. Staminodes 9,
ca. 0.5 mm long, lanceolate, strigose. Ovary gla-
brous, globose, ca. 0.5 mm long, the style ca. 1
mm long, curved at the tip. Fruit ellipsoid, 3.5
cm long, seated on a small, glabrous disk.

. — examined. BRAZIL. AMAPA:
o Juruxi-Mazagao, B. V. Rabelo 2715 (MO). PARA:
is Ducke RB 1 7583 (U, US), Ducke 1234 (NY,
MO, US); Trapiche Hypolito, Krukoff 5870 (BR, NY,
MO); Ilha de Pará, Mori et al. 16510 (MO), Belem,
Murça Pires 1488 (NY).

AL 7 L 2 4°

of its peculiar anthers. When naq it is espi
similar to M. thoroflora. The latter, however, has
shorter petioles and a fine and closely appressed
pubescence on young twigs and leaves, whereas
in M. mahuba the petioles are longer and the
pubescence consists of spreading hairs. Mezilau-
rus mahuba is a species known only from sea-
sonally inundated forest in the states of Pará and
Amapa

pa.

The few available collections of Mezilaurus
mahuba clearly show two phases in the floral
development, probably corresponding with the
male and female phase as described by Kubitzki
and Kurz (1984) for several other species of Lau-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

raceae. The three sheets of Krukoff 5870 all have
flowers with the anthers recurved and tightly
pressed against the flower, almost hiding the an-
ther cells (female phase), while the three sheets
of Ducke 1234 have only flowers with the anthers
spreading and free of the flower, fully exposing
the anther cells (male phase). The inflorescences
of these specimens are large, and it is interesting
that the flowers on an inflorescence all appeared
to be in the same phase of development, quite
unlike what I have seen in species of Ocotea and
Nectandra with large inflorescences.

Mezilaurus micrantha van der Werff, sp. nov.
TYPE: Brazil. Amazonas: Manaos, Reserva
Florestal Ducke, Rodrigues & Coelho 7555
(holotype, NY). Figures 6, E & F; 7.

Arbor, 20 m alta. Ramuli teretes, glabri vel prope
mm

atra. Venat

, leviter e d esposa

triangularia, er
0.6 mm longa, per anthe
liberis, den ibus. Antherae 2- locellatae, lo-
cellis extrorsis, tae ventrali staminis omnino pubes-
centi. Ovarium glabrum, sensim in stylo attenuatum,
ovarium stylusque | mm longus. Fructus ignoti.

Tree, 20 m tall. Twigs terete, glabrous or, near
the apex, with some appressed pubescence, the
terminal bud sericeous; the bark rather thick.
Leaves clustered at the tips of the branches, co-
riaceous, drying blackish, glabrous at maturity,

ut when young with some appressed pubes-
cence, elliptic, ca. 10-15 x 3.5-5 cm (exclusive
of petiole), the base gradually narrowed atten-
uately into the petiole, the tip blunt but not
rounded, the margins revolute; veins and retic-
ulation not or scarcely raised on upper surface;
midvein and main lateral veins (5-8 pairs) raised
on lower surface, but reticulation not obvious.
Petioles to 2 cm long, glabrous at maturity. In-
florescences axillary, subterminal, to 5 cm long,
pyramidate, with some appressed pubescence.
Flowers arranged subumbellately at the ends of
the lateral branches, appressed pubescent. Ped-

1987]

icels minutely appressed pubescent, ca. 2 mm
long at anthesis. Bracts and bractlets deciduous.

equal, triangular, erect. Stamens 3, 0.6 mm long,
exserted 0.3 mm at anthesis, the filaments free,
densely put ; anthers 2-celled, glabrous, the
cells extrorse, opening back-to-back; ventral side
of the anther entirely pubescent. Floral tube pu-
bescent. St staminodia lacking
Ovary glabrous, gradually narrowed into style,
the ovary and style ca. 1 mm long, the style
exserted at anthesis. Fruit unknown.

Paratypes. marge AMAZONAS: Manaos, Reserva
Floresta Ducke, W. Rodrigues 3190 (NY); AM-1, Km
, W. Rodrigues 7066 (NY).

Mezilaurus micrantha is rather similar to M.
itauba; it differs in having smaller flowers, blunt
but not rounded leaf tips, lack of gland dots on
the leaves, slightly revolute leaf margins, and
nearly black dried leaves. These characters are
not strong individually, but taken together they
allow identification of flowering as well as sterile
collections. The flowers of this species are among
the smallest I have seen in the genus, hence its
specific epithet.

Mezilaurus navalium (Allemáo) Taubert ex Mez,
Arbeiten Kónigl. Bot. Gart. Breslau 1: 112.
1892. Silvia navalium Allemáo, Dissertatio,
Rio de Janeiro. 1848. Silvaea navalium (Al-
lemao) Meissner, DC. Prodr. 15(1): 84. 1864.
Endiandra navalium (Allemao) Benth. &
Hook., Gen. Pl. 3: 154. 1880. Mezia nava-
lium (Allemáo) Kuntze, Revis. Gen. Pl. 2:
574. 1891. TYPE: Brazil, Rio de Janeiro, A/-
lemáo, s.n. (holotype, R, not seen). Figures

B, 9

Tall trees, to 25 m. Branches terete, glabrous,
the tips with appressed, short hairs, the terminal
bud densely gray strigose. Leaves clustered at
branch tips, narrowly elliptic, 10-12 x 3-3.5 cm,
somewhat coriaceous, glabrous on both surfaces
or with few appressed hairs along the midrib, the
tip rounded, the base sharply acute, lateral veins
not strongly developed, 10-15 pairs, the upper
surface dull, smooth, lower surface with slightly
elevated reticulation; petioles ca. 1 cm long. In-
florescences axillary, glabrous or with few scat-
tered hairs, 3-5 cm long, the flowers clustered at
the ends of the lateral branches. Flowers gla-
brous, ca. 2 mm long. Pedicels 2-3 mm long,
glabrous. Braots deciduous. Tepals 6, equal, erect,

WERFF—REVISION OF MEZILAURUS

171

scalelike, ca. 0.3 mm long. Stamens 3, ca. 1.5
mm long; filaments free, strigose; anthers exsert-
ed, glabrous, the 2 large cells positioned on a
dorsal ridge, opening back-to-back, slightly di-
vergent, exposing the exserted stigma. Ovary el-
lipsoid, glabrous, ca. 2 mm long, including the
exserted stigma. Staminal glands and staminodia
lacking. Immature fruits subtended by the small
tepals, occasionally the stamens visible at the
base of the young fruit.

Additional specimens examined. BRAZIL. RIO DE
JANEIRO: Theresopolis, Glaziou 11470 (C, G, U), Gla-
ziou 11473 (C); Petropolis, Glaziou 12124 (C, G), Gla-
ziou 12125 (C, G, MO, U, US).

Mezilaurus navalium is the only Mezilaurus
species known from the Atlantic rain forests in
southern Brazil. The wood is hard and much
used for naval construction. Diagnostic charac-
ters, in addition to its distribution, are the gla-
brous flowers and leaves with rounded or blunt
apices.

Mezilaurus opaca Kubitzki & van der Werff, sp.
nov. TYPE: Peru. Depto. Loreto: Prov. Re-
quena, Distr. Jenaro Herrera, trocha al Rio
Yaveri, cerca al Arboretum de Jenaro Her-
rera, 15 m tree in low forest, flowers greenish
yellow, 20 Aug. 1976, Revilla 1226 (holo-
type, MO; isotype, HBG). Figures 8, C & D;
12

0 m. Ramuli glabri, sed apicibus foliiferis
e tertia immersa;

aequalia, parva, ca. 0. x mm, minute va

2-loculatis. Ovarium glabrum, ovoideum, sensim in

stylo attenuatum. Fructus ignoti.

Tree, 15 m. Twigs glabrous, rather thick (5-7
mm diam. immediately below the leaves), with
conspicuous leaf scars, the bark gray. Tips o
branches with dense, brown, sericeous pubes-
cence. Leaves alternate, clustered at tips of
branches, ca. 20 x 9 cm, young ones with some
appressed pubescence, glabrous at maturity,
chartaceous, elliptic, the tip rounded or bluntly

172

Ficure 8. A-B. Mezilaurus navalium. — A. Leaf. — B.
M. palcazuensis. — E. Leaf. — F. Flower.

acute, the base gradually narrowed onto the pet-
iole, green, opaque above, the midvein elevated,
the secondary veins slightly elevated, the tertiary
venation more or less immersed, not easily vis-
ible; lower surface also opaque but with the ve-
nation more elevated; petioles 3—4 cm long, with
narrow wings of the decurrent laminae. Inflores-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Flower. C-D. M. opaca. —C. Habit. — D. Flower. E-F.

cences compound racemes in the axils of decid-
uous bracts, slightly longer than the petioles, when
young rather densely appressed pubescent, at an-
thesis less so. Bracts and bracteoles deciduous.
Flowers clustered at the tips of inflorescence
branchlets, with some appressed pubescence, de-
pressed globose, constricted at the apex, ca. 1.5 x

1987]

WERFF—REVISION OF MEZILAURUS

173

—

200 400 600 800 1000km

r—r——— sh
O 100 200 300 400 500 600 miles

80 70 60

FIGURE 9.
(4), and M. navalium (©)

1.5 mm (including the exserted stamens). Tepals
6, equal, pointing inwards, wider than long, ca.
0.4 mm wide, ca. 0.2 mm long, with some ap-
pressed pubescence. Floral tube short, ca. 0.5
mm long, with a pubescent ring in the upper part.
Stamens 3, 2-celled, ca. 1 mm long, the anthers
glabrous, exserted at anthesis, the cells on a dor-
sal ridge, the valves opening back-to-back; fila-

into the style; style exserted beyond stamens;
stigma a small plate. Fruit unknown.

50 40

Distribution of Mezilaurus palcazuensis (@), M. pyriflora (8), M. quadrilocellata (), M. sprucei
O).

Mezilaurus opaca is rather similar to M. syn-
andra, but differs from that species by its smaller
flowers and the leaf characters mentioned in the
key. Béguin et al. (1985) reported M. synandra
from the Jenaro Herrera Arboretum. It is pos-
sible that this specimen represents M. opaca, but
I have not seen it.

Mezilaurus palcazuensis van der Werff, sp. nov.
TYPE: Peru. Cerro de Pasco: Selva Central,
Palcazu Valley, elev. 300-600 m, 7 Dec.

174

1984, Hartshorn, Quijano & Mateo 2691
(holotype, MO). Figures 8, E & F; 9.

Arbor, 25 m. Ramuli teretes, glabri vel prope apicem
adpresse pubescentes, cicaticibus foliarum conspicuis
funds is. Gemma terminalis sericea. Folia chartacea,

ad adulta glabra, juvenalia

mid sse x iru vel anguste obovata, ap-

ice acuta, basi at l ed cm. Costa ele-

: i aud conspi-

M forescentia ore
niculata, 4cm lon

cua. Peti oli ca. | cm

F
n ils

sessilibus vel e sessilibus. Flores minuti aree
s, obconi .l.5m n gi. Tepala

tamina 3, mm
longa, per anthesin exserta. Antherae 2- locellatae, gla-
brae, extrorsae
parvam pubescentem in superficie. ventrale. Ovarium
globosum, glabrum. Fructus ignoti

Tree, 25 m. Twigs terete, glabrous or with some
appressed pubescence near tip, with conspicuous
leaf scars. Terminal bud sericeous. Leaves char-
taceous, clustered at the tips of branches, with
some appressed pubescence when immature, gla-
brous when mature, obovate or narrowly ob-
ovate, the tip acute, the base gradually narrowed
into the petiole, 10-15 x 3-4 cm; costa raised
on both surfaces; secondary veins and reticula-
tion immersed, not obvious; petioles ca. 1 cm
long. Inflorescence axillary, 4 cm long, panicu-
late, minutely appressed pubescent, the flowers
clustered at the tips ofthe lateral branches, sessile
or nearly so. Flowers minutely appressed pubes-
cent, sessile, ca. 1.5 mm long. Tepals 6, erect,
0 ong. Fertile stamens 3, 2-celled, exsert-
ed at anthesis; anthers extrorse, situated on a
dorsal ridge, opening back-to-back; filaments
united, glabrous. Staminal glands and stamino-
dia lacking. Fruit unknown.

Mezilaurus palcazuensis is only known from
the holotype, consisting of a small twig with few
leaves and one inflorescence. It is therefore quite
likely that the description does not embrace the
morphological variation of this species. Note-
worthy features are the acute leaf tips and the
sessile flowers, both unusual characters in the
genus. The leaves are also thinner than in other
species of Mezilaurus, but with only one speci-
men available, it is not certain whether this is a
distinguishing character.

Similar sessile flowers occur also in M. caa-
tingae, known from caatinga forest along the Rio
Negro, which differs in having larger flowers and
coriaceous, rounded leaves with inrolled mar-
gins.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Mezilaurus pyriflora van der Werff, sp. nov. TYPE:
Brazil. Amazonas: Sao Paulo de Olivenga,
basin of creek Belem, 26 Oct.-11 Dec. 1936,
Krukoff 8711 (holotype, NY; isotype, MO,
GH). Figures 10, A& B; 9

5 m alta. Ramuli crassi, ad 1 cm diametro,

et subtus eleva per im-

5
°
e
=
2
p
e.
"S
e

lateralibus. Inflorescentia axillaris, subterminalis, an-
guste pyramidalis, ramulis basalibus perlongioribus
quam terminalibus, adpresse pubescens. Flores parvi

mm x 0.9 mm), glabri, pyriformes. Pedicelli pu-
bescentes, 1.5-2 mm longi. Stamina fertilia 3, inclusa,
0.8 mm i

celli pa

Ovarium depresse globosum, glabrum, 1.2 m
ongum. Ped a pA Glandulae filamentorum
nullae. Fructus igno

Tree, to 25 m tall. Twigs thick (diam. 1 cm 4-
5 cm below tip), terete, with conspicuous leaf
scars, the tip with gray, minute and appressed
pubescence, becoming glabrous with age. Ter-
minal bud densely gray pubescent. Leaves clus-
tered at tips of branches, firmly chartaceous, ob-
ovate, the basal half of the lamina widening very
gradually, the apical half rather abruptly wid-
ened, large (40-60 x 14-18 cm at maturity), the
tips acute, gradually narrowed towards the base,
but the base abruptly narrowed, rounded or al-
most subcordate, mostly glabrous, with some ap-

venation scarcely visible above, raised below;
petioles ca. 1 cm long, 5 mm thick, densely ap-
pressed pubescent, the lamina decurrent as two
narrow ridges. Inflorescences axillary, subter-
minal, pyramidal, the basal branchlets much
longer than the terminal branches (the longest
ca. 10 cm long, decreasing to ca. 1 cm), the main
axis and branchlets with appressed pubescence;
bracts and bracteoles pubescent. Flowers ar-
ranged spicately along branchlets, glabrous, often
reflexed, pear-shaped, ca. 0.9 mm long, 0.9 mm
wide. Pedicels pubescent, especially near the base,
1.5-2 mm long at anthesis. Fertile stamens 3, ca.

1987] WERFF— REVISION OF MEZILAURUS 175

FicuRE 10. A-B. Mezilaurus pyriflora. — A. Leaf. — B. Flower. C-D. M. quadrilocellata. — C. Leaf. — D. Flower.
E-F. M. sprucei. —E. Leaf. — F.

0.8 mm long, included; filaments free, pubescent; Paratype. BRAZIL. AMAZONAS: Fróes 12152 (NY).
anther cells minute, terminal, opening towards

the tip. Ovary isis E glabrous, ca. Mezilaurus pyriflora is known only from two
1.2 mm wide, ca. ong. Staminodia 6, collections from the vicinity of São Paulo de Oli-

representing the outer 6 um mens, ini ca. venga. Leaf shape, the short, pubescent pedicels,
m long. Staminal glands lacking. Fruit un- the included, short anthers and the pear-shaped
known. flowers (with the apical valves of the anthers

176

mimicking the dried crown of a pear) separate
this species from M. mahuba and M. thoroflora.
The Fróes collection in NY bears the annotation
“wood causes injury to the skin.”

The two collections of Mezilaurus pyriflora had
been annotated as Euphorbiaceae and Ochna-
ceae; because Krukoff's collections were widely
distributed, it is possible that duplicates of M.

ified in additional herbaria

Lud Gd

Mezilaurus quadrilocellata van der Werff, sp. nov.
TYPE: Colombia. Antioquia: Chigorodó.
Tree, 20 m. Flowers white. 100-200 m, M.
Garcia Barriga 17626 (holotype, GH; iso-
types, AAU, US). Figures 10, C & D; 9

Arbor, 20 m. Ramuli crassi, 4-5 mm diametro, te-
retes, glabri apicibus foliiferis tomentellis. Folia con-
ferta ad apices ramulorum, glabra, elliptica vel leviter
obovata, chartacea, basi cuneata, apice rotundata, ca.

5x 8cm, petiolis 2-4 cm longis, tomentellis, margine
cartilaginea, leviter incrassata. Venatio super immersa,
subtus tomentella costa manifeste elevata, nervis la-
teralibus (6—8) elevatis, nervis basalibus marginem at-
tingentibus. Inflorescentiae axillares, foliis breviores,
anguste pyramidatae, 6—9 cm longae, tomentellae. Flo-
res parvi, m longi, pedicellis 2-3 mm longis,
tomentellis. Tepala 6. 3 interioria 3 exterioribus duplo

o ,c nga, interiora o m
nga, omni abra. Stami longa,
lamentis pubescentibus, sine glandu antheris

bescentia. Ovarium ellipsoideum, glabru
gum, sensim in stylum attenuatum. Fructus ignoti.

Tree, 20 m tall. Twigs thick, 4-5 mm diam.
immediately below the leaves, terete, glabrous
except for the rufous tomentellous leaf-bearing

apex. Leaves clustered at tips of branches, gla-
brous, elliptic or slightly obovate, ca. 15 x 8 cm,
the base cuneate, the apex rounded, the margins
y thickened, the venation

immersed above, the midrib prominently raised
and tomentellous on lower surface, the lateral
veins (6—8) less prominently raised, the basal ones
reaching the margin, the upper ones arcuate and
not reaching the margin; tertiary venation slight-
ly raised, petioles 2-4 cm long, rufous tomen-
tellous. Inflorescences axillary, shorter than the
leaves, narrowly pyramidate, 6—9 cm long, to-
mentellous. Flowers small, ca. 1.5 mm long; ped-
icels 2-3 mm long, tomentellous; flower tube and
tepals glabrous outside. Tepals 6, the outer ones
half as long as the inner ones, erect, with tips
curved inward; outer tepals ca. 0.6 mm long,
ovate, inner ones ca. 1.2 mm long, ovate, gla-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

brous. Ovary ellipsoid, gradually narrowed into
the style, 1 mm long, glabrous. Fertile anthers 3,
ca. 1 mm long; filaments rather densely pubes-
cent; anthers 4-celled, glabrous, the tips bent in-
ward, forming a flat shield exposed at anthesis;
anther cells situated on this shield. Fertile an-
thers alternating with 3 small (ca. 0.5 mm), slen-
der, densely pubescent staminodia. Fruit un-
known

Mezilaurus quadrilocellata is known only from
the type collection in northern Colombia, not far
from the Panamanian border and the Caribbean.
Further discussion is given under M. glauco-
phylla.

Mezilaurus sprucei (Meissner) Taubert ex Mez,
Arbeiten Kónigl. Bot. Gart. Breslau 1: 112.
1892. Acrodiclidium sprucei Meissner, DC.
Prodr. 15(1): 86. 1864. Silvia sprucei (Meiss-
ner) Mez, Jahrb. Kónigl. Bot. Gart. Berlin
5: 119. 1889. Mezia sprucei (Meissner)
Kuntze, Revis. Gen. Pl. 2: 574. 1891. TYPE:
Brazil. Amazonas: San Gabriel de Cacho-
eira, Rio Negro, May 1852, Spruce 2323
(lectotype, chosen by Kostermans, K; iso-
types BM, C, NY, U). Figures 10, E & F; 9.

Mezilaurus maguireana C. K. Allen, Mem. New York
Bot. Garden 10: 58. 1963. TYPE: Venezuela. Ama-
zonas: Río Guainía, Maroa, Maguire et al. 41698
(holotype, NY; isotype, GH).

Small tree, to 10 m tall. Twigs terete, glabrous,

raceous, glabrous, elliptic, 12 x 5 ) cm,
the base acute, the apex acute or acuminate; lat-
eral veins 10-15 pairs, slightly elevated or im-
mersed on upper face, raised on lower surface,
the reticulation slightly raised on both surfaces;
petioles to 3 cm long, flat, bordered by a narrow
ridge, the very base of the petiole round and

branchlets, ca. 2 mm long, the pedicels to 1.5 cm
long. Tepals 6, equal, erect, ca. 0.5 mm lon

Stamens 3, ca. 1.2 mm long, exserted; anthers
free and somewhat divergent, 2-celled, the cells
large, opening back-to-back; filaments pubes-
cent, connate. Ovary ellipsoid, ca. 0.8 mm long,
abruptly narrowed into the slender, ca. 1 mm

1987]

long, style. Flower tube pubescent within. No
staminal glands or staminodia. Fruits not seen.

po Bate examined. VENEZUELA: 30 km

f Puert achucho, Guanchez 255 (TFAV).
AMAZONAS: ee Carlos de Río Negro, Clark & Ma-
guirino 7784 (MO), 8091 (MO); Cerro Neblina base
camp, Gentry & Stein 46836 (MO); BRAZIL. AMAZONAS:
Río Negro above Camanaus, Prance et al. 16042 (NY).
PERU. LORETO: Iquitos, near Picuruyacu, Revilla 106
(G, MO); Requena, Jenaro Herrera, Vasquez & Jara-
millo 984 (MO).

Mezilaurus sprucei can be recognized easily by
its glabrous, acute or acuminate leaves, lax in-
florescences, and especially by the flowers with
long pedicels. The Revilla collection from Peru
has short pedicels (5 mm long) but agrees in other
characters with Mezilaurus sprucei.

I tentatively place Mezilaurus maguireana in
synonymy under M. sprucei. The type does not
agree completely with the typical M. sprucei; the
inflorescences are stiffer, the flowers have pedi-
cels only 5 mm long, the reticulation is less raised
on the upper leaf surface, and the leaf apices are
less acuminate. These differences are only of de-
gree and the few collecti of M. sprucei at hand
probably do not show the i range of variation
in the species. In what I consider important char-
acters (acute leaves, long petioles, glabrous, sub-
umbellately arranged flowers, and connate fila-
ments), M. maguireana agrees with M. sprucei.

I found that in old flowers the anthers become
divergent, although the filaments remain con-
nate.

ee dodge (Ducke) Kosterm., Med-

i . Rijks Univ. Utrecht 25:

40. 1936. Silvia ao Ducke, Arch.

Jard. Bot. Rio de Janeiro 5: 115. 1930. TYPE:

Brazil. Pará: dry upland forest of Jumanda

River, Ducke RB 19974 (holotype, RB; not
seen; isotype, U). Figures 11, 12.

Medium-sized tree to 20 m. Twigs terete, with
conspicuous leaf scars, the terminal bud densely
and minutely appressed pubescent, this disap-
pearing rapidly as twig matures. Leaves clustered
near apices of branches, rarely few leaves per-
sisting on older twigs, the leaves glabrous, cori-
aceous, slightly ovate or elliptic, ca. 15 x 7 cm,

ioles 3-5 cm, rarely only 1 cm long.
Inflorescences axillary near tips of twigs, pyram-
idal, rather laxly flowered, appressed tomentel-

WERFF—REVISION OF MEZILAURUS

177

lous, 5-12 cm long; branchlets slender, distant,
to 1.5 cm long, the flowers clustered near their
tips. Bracts and bracteoles deciduous. Pedicels
slender, tomentellous, 1-1.5 mm long. Flowers
globose, puberulous, ca. 1.5 mm long. Tepals 6,
equal, erect, ca. 0.6 mm long. Stamens 3; ca. 1
mm long; filaments ca. 0.7 mm, pubescent, con-
nate; the 2-celled anthers exserted like small horns
from the flower tube, glabrous; anther cells large,
lateral-introrse, opening toward the tip. Ovary
vin ca. ] mm long; style exserted, the stig-
nute. Staminal glands and staminodia
T Fruit (fide Kostermans, 1938) ellipsoid,
.5-3 cm long, 1.5 cm diam., subtended by a
small, platelike cupule (4-5 mm diam.) with sub-
persistent tepals.

Additional specimens examined. PERU. MADRE DE
DIOS: Tambopata, Gentry et al. 46116, 45952 (MO).

Mezilaurus subcordata is collected rarely and
known to me only from an isotype and two recent
collections in Peru. Very possibly it is not a true
disjunct—it may also occur in the intervening
area, since a tree with green flowers 1.5 mm long
can easily be overlooked. The Peruvian collec-
tions come from a tree plot in which every tree
was sampled, regardless of whether it was fertile
or not.

The description is based on the U isotype. The
Peruvian specimens differ in having glabrous
flowers and slightly thinner leaves. I find these
differences too weak for recognition of a new
taxon, but additional collections may show the
Peruvian plants to be distinct.

Mezilaurus subcordata can be recognized
readily by its long petioles and rounded leaf bas-
es; it is also the only species in the genus with
lateral-introrse anther cells.

Mezilaurus synandra (Mez) Kosterm., Meded.

Manáos, dry upland forest near Pensados,
Ule 8835 (lectotype, B, fide Kostermans, >
seen; isolectotype, L). Figures 13, A& B

Tree, to 15 m tall. Twigs terete, glabrous, often
with conspicuous leaf scars, the terminal buds
yellowish strigose-sericeous. Leaves clustered at
tips of branches, glabrous, elliptic or slightly ob-
ovate, 8-20 x 4-10 cm, the base cuneate or acute,
the tip rounded, lateral veins 10-15 pairs, + im-
mersed above, slightly raised below, the tertiary

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FIGURE 11.

venation reticulate and slightly raised on both
surfaces, the midvein raised below, triangular in
diameter; petioles conspicuous, 3-5 cm long, gla-
brous, flat or with 2 narrow ridges on upper side.
Inflorescences subterminal, much shorter than
leaves, ca. 3 cm long, the flowers in small (4—5
flowered) clusters at the ends of few lateral
branchlets, minutely strigose. Bracts and bract-
lets early deciduous, not seen. Flowers sparsely
and minutely strigose, depressed globose, as wide
as long or wider than long, the flower tube nar-
rowed toward the tip (ca. 2 mm wide and 1.7
mm long on unpressed flowers). Tepals 6, equal,

erect, scalelike, ca. 0.4 mm long, ca. 0.8 mm
wide. Stamens 3, ca. 1.9 mm long, exserted, the

Mezilaurus subcordata. — A. Habit. — B. Flower.

anthers glabrous, 2-celled, the cells opening lat-
eral-apically or apically; filaments connate, pu-
bescent. Ovary pupescent ea. ES mm dong; ae
ca. 0.3 mm long. St

lacking. Fruit ovoid, ca. A cm long, 1 cm wide,
subtended by a small, platelike cupule.

Additional specimens examined. BRAZIL. AMAZONAS:
anáos, Igarape da Cachoeira, Baixa do Tarumã, Cha-
gas s.n. = MG 21.108 (NY); Manáos, Parque 10 de
Novembre, Coelho s.n. = INPA 3934 (NY); Manáos,
Pensador, Ducke 233 (F, NY), Ducke 233, second col-
pe (US); Manáos, Ducke RB 23964 (G, US); Man-
s, Pensador, Ducke RB 25092 (US); Manáos, Cach-
oeira, Alta do Tarumã, Rodrigues & Lima 30 Y).
Reported by Béguin et al. (1985) from the Arborétum
Jenaro Herrera, Loreto, Peru

1987] WERFF—REVISION OF MEZILAURUS 179

o 200 400 600 800 1000km
€ ere

| r———mcCàs1xa”
| O 100 200 300 400 500 600 miles

80 70 60 50 40

FIGURE 12. Distribution of Mezilaurus opaca (), M. subcordata (W), M. synandra (B), and M. thoroflora
(e).

Mezilaurus synandra is only known from dry, Mezilaurus thoroflora van der Werff, nom. nov.

low forest on terra firme, near Manaus and one Basionym: Licaria maguireana Allen, Bull.
collection in Peru. Several collections indicate Torrey Bot. Club 75: 315. 1948. Misanteca
that it occurs in secondary vegetation. When maguireana (Allen) Lundell, Wrightia 4: 100.
flowering, M. synandra can be recognized easily 1969. Clinostemon maguireanum (Allen)
by its short inflorescences and broad, depressed- Kurz, J. Arnold Arbor. 60: 520. 1979. TvPE:
globose flowers, which otherwise occur only in Guyana: Mazaruni Station, Forestry Dept.
M. opaca. Fruiting or sterile collections are rath- Brit. Guiana 2956 (F220) (holotype, NY;
er similar to M. itauba, which has, however, a isotype, K). Figures 13, C & D; 12.

larger inflorescence, generally smaller leaves, and

the upper leaf surface dull, not shiny, with less Tree, ca. 25 m tall. Twigs thick, terete, the tip

prominently raised reticulation. densely and minutely appressed pubescent, be-

180 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

3cm

FIGURE 13.

A— B. Mezilaurus synandra. — A. Leaf. — B. Flower. C-D. M. thoroflora. — C. Leaf. —D. Flower.

coming glabrous with age. Terminal bud yellow-
ish pubescent. Leaves clustered at the tips of
branches, coriaceous, obovate, gradually nar-
rowed toward base, the base abruptly rounded,

the apex acute or shortly acuminate, 30-60 x
12-18 cm, pinnately veined, 15-25 pairs of lat-
eral veins, these immersed above, but elevated
below. Midrib thick, ca. 7 mm wide, raised on

1987]

both surfaces. Tertiary venation not visible above,
but raised below. Lamina glabrous above, except
along midrib, sparsely appressed pubescent be-
low, especially near the base and along midrib.
Inflorescences in axils of aborted leaves, subter-
minal, appressed gray-pubescent when young,
with few hairs at anthesis, 30-50 cm long, lateral
branchlets 2-3 cm long over the entire length of
cylindrical inflorescence, these with scattered
hairs, bracts and bractlets, strigose, persisting at
anthesis. Flowers clustered toward the tips of
Üranchlets, glabrous, ellipsoid, ca. 1.5 mm long,

wide, the anthers ca. 0.5 mm exserted,
pedicels 4-8 mm long. Tepals 6, equal, erect,
scalelike. Fertile stamens 3, ca. 1 mm long, the
filaments free, pubescent, ca. 0.5 mm long, the
2-celled anthers glabrous, exserted, the large cells

bescent, the outer six slightly sagittate, the inner
three lanceolate. Infructescence ca. 60 cm long,
the fruit ellipsoid, ca. 1.7 x 1 cm, subtended by
a thin platelike cupule, ca. 0.6 cm diam.

Paratype. Guyana, Mazaruni Station, Forest Dept.
Brit. Guiana 2704 (K).

Mezilaurus thoroflora is known only from a
few collections in Guyana. Diagnostic for this
species are the long-pedicelled flowers, the large
exserted anthers, and the uniform, short, lateral
branchlets of the inflorescence. Vegetatively, M.

roflora

The illustration in Kubitzki et al. (1 979) of M.
thoroflora (as Clinostemon maguireanum) is not
representative of material I have seen. The two
collections from the Forest Department of Brit-
ish Guiana do not have such pronounced cordate
leaf bases and do not have the lower branchlets
of the inflorescences much longer than the upper

ones, as shown in Kubitzki et al. (1979).
Kostermans (1938) cited two collections
(Monteiro Costa 323, Kaufmann 605, both in F)
under M. lindaviana. In my opinion these spec-
imens belong to M. thoroflora or are very close
to it. They have considerably smaller leaves than
the type of M. thoroflora, but leaf shape is quite
similar. Monteiro Costa 323, a flowering speci-
men, shows exserted anthers and staminodia, and
this clearly indicates M. thoroflora. Pedicels are
shorter and leaves smaller than in M. thoroflora,

WERFF—REVISION OF MEZILAURUS

181

but it is likely that the few available collections
do not show the full range of vegetative variation.

Because the combination Mezilaurus magui-
reana already exists, it was necessary to create a
new epithet for Licaria maguireana. The
epithet thoroflora is derived from the Crest

"thoros," semen, and “‘flos,”’ flower, in reference
to rembia between the small, long-pedi-
celled flowers and spermatozoa.

IMPERFECTLY KNOWN SPECIES
Mezilaurus sp. A.

A collection made by J. da Silva Costa (RB
1807 96) in the State Mato Grosso, Brazil, prob-
ably represents an undescribed species. The lo-
cality data suggest it was collected in cerrado
vegetation as a 9 m tree. The most distinguishing
characters are found in the leaves, which are
densely gland-dotted on the upper surface, char-
taceous, and have a few appressed hairs on the
lower surface. The young flower buds are gla-
brous. This is clearly not the other cerrado species,
M. crassiramea, which has pubescent leaves, and
the gland-dotted upper leaf surface has not been
found in other Mezilaurus species. I prefer to
wait with a formal description until flowering
material is available.

LITERATURE CITED

ALLEMAO, F. 1848. Dissertatio, Rio de Janeiro.
ALLEN, C. K. 1948. Lauraceae. In B. Maguire et al.
Plant explorations in Guiana in 1944, chiefly to
the Tafelberg and the Kaieteur plateau — III. Bull.
7

9 a e. In B. Maguire et al. The
of the Guayana Highland, V. Mem. New
uid) 44-1

. D. R. SPICHIGER & J. MiBGEe. 1985. Las

3: 146-164. London.

Ducke, A. 1930. Plantes nouvelles ou peu connues
de la Région Amazonienne IV. Arch. Jard. Bot.
Rio de Janeiro 5: 101-188.

35. Notes on the itaüba trees: the Ama-
zonian species of the genus Silvia Allem. Trop.
Woods 42: 18-21.

1905-1913. Plantae Brasiliae centralis

. The Genera of Flowering
Plants. Clarendon Press, Oxfor

KOosTERMANS, A. J. G. H. 1938. LO TR of the Lau-
raceae III. Recueil Trav. Bot. Néerl. 35: 56-129.

182

————. 1952. A historical survey of the Lauraceae
III. ; Sci. Res. (Jakarta) 1: 141-159.

Lauraceae. Reinwardtia 4: 192-256.

a K. & H. Kurz Synchronized di-
chogamy and dioecy in neotropical Lauraceae. Pl.
Syst. Evol. 147: 253-266.

. RICHTER. 1979. Reinstate-
ment of Clinostemon (Lauraceae). J. Arnold Ar-
bor. 60: 515-522.

KUHLMANN, J. G. & A. DE SAMPAIO. 1928. Clino-
stemon, Kuhlm. et A. Samp., n. gen. de Lauraceas
da Amazonia. Bol. Mus. Nac. Rio de Janeiro 4:

KUNTZE, O. 1891. Revisio Generum Plantarum II.
Zi

Kurz, H. 1983. 5 ue Gat-
tungen neotropischer Laur und Revision der
Gattung Licaria patei, "Thesis, Laives
of Hambur;

MEISSNER, C. F. 1864. Lauraceae. /n A. de Candolle,
Prodromus Systematis Naturalis Regni Vegeta-
bilis 15: 1-260.

MEZ, C. 1889. Lauraceae Americanae. Jahrb. Kónigl.

Bot. Gart. Berlin 5: 1-556.

March

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

. 1892. Spicilegium Laureanum. Arbeiten
Kónigl. Bot. Gart. Breslau 1: 71-166.
. 1904. Additamenta monographica 1904. Bull.
Herb. Boissier. sér 2. 5: 233-244.
9 Additamenta monographica 1919. Re-
pert. Spec. Nov. Regni Veg. 16: 305-309.
1924. Additamenta monographica 1924. Bot.
Arch. 6: 230-231.
Pax, F. 1897. Die Natürlichen Pflanzenfamilien.
Nachtrag und Register zu Teil II-IV.
Anatomie des sekundáren Xy-
lems und der Rinde der Lauraceae. Sonderbande
des Naturw. Ver. Hamburg, 5, Verlag Paul Parey,

Hamburg.
TAUBERT, P. 1892. [Review of: Revisio Generum
Plantarum by O. Kuntze.] Bot. Centralbl. 50: 17-
24.

WEBERLING, F. 1981. Morphologie der Blüten und

der EC sies Verlag Eugen Ulmer, Stuttgart.

. 1985. r In foreszenzmorphologie der Lau-
as

rroneous X dahi in Glaziou
collections of Vid Bean iniu] Taxon 19: 911-
913.

Volume 73, No. 4, pp. 653-831 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN, was published on 30
1987.

|. Portraits of Botanists ——

The Missouri Botanical Guia has begun issuing a series of patos“ “Portraits
Salas are reproduced from the Garden’s Archives, and brief information a

CONTENTS

Systematic Embryology of the Anisophylleaceae Hiroshi Tobe & Peter H. Raven _... 1

nosae: Papilionoideae) David A. Neill

Australian Acacia Peter Bernhardt ...

Flower Longevity and Protandry i in Two cae of Gentiana (Gentianaceae)

C. J. Webb & Jan Littleton ..

Notes on the Breeding XAR of Sacoila senweatia (Aublet) Garay (Orchidaceae)

Paul M. Catling ....

Flower and Fruit Sui in DES Spe Belarus Sinbad:
- Javier Herrera

w... U. T... U...

Reproductive Systems and SOME SEN of Oxalis pes-caprae L. G Their en

on the Genesis of a Noxious Weed Robert Ornduff

Trapliners in the Trees: Hummingbird Pollination of Erythrina Sect. Erythrina (Legumi- £

A Comparison of the Diversity, Density, and F. oraging Behavior of Bees and Wasps on ` iy
51 j
€
79.
85

pel : Flora of the Venezuelan Guayana—ll Julian A. Steyermark
SE A New Spee of f Jatropha ere) from Nicaragua Grady

gx.

ges B. Faden & D. R. Hunt

—

»psis with XI eer ' (Commelinaceae)

$ Webster i d 17 š 2

5e Chromosome Numbers B Madagascar Plants — Elisabeth R abakonandrianina & Gerald s

p

Gr. 1:

; Cytotaxonomic. Studies i in the Cems: Urginea Stein i in West Africa. IL Karyotype Evolution

in Urginea altisima (L. ) Baker FO Oyewole

' Cytotaxonomic Studies i in the Genus Gris Stein in West Africa. HL The Casa of Urginea di

| indica. (Roxb.) Kunth i in Nigeria. SN Io Oyewole

; Cytotaxonomic Studies i in the Genus Urginea Stein in West (Doe IV. P.

entiation -and EAS Virin + in S an > uite une K

no MA

126.

To:
opulation Differ- n.

š Š 2 seene.

Taia Warren

Volume 74

Number 2

Volume 74, Number 2
Summer 1987

Annals of the
Missouri Botanical Garden

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Volume 74
Number 2
1987

Annals
of the
Missouri
Botanical

Garden

WZ

VASCULAR EPIPHYTISM: TAXONOMIC PARTICIPATION
AND ADAPTIVE DIVERSITY!

DaviD H. BENZING?

ABSTRACT

Vascular epiphytes carci few qualities beyond occurrence in tree crowns that identify them as a

those of terrestrials native to comparably arid or infertile sites. Moisture supply, m

arily because their phylogenetic origins and life styles in forest canopies

feature, determines where a particular type of epiphyte will survive. Epiphytes constitute about 10%

orchids for epiphytism are identified. R

families in tree crowns are less clear, but some possibilities are offered.

relations and, Bm mineral nutrition may explain why epiphytism has rarely, if ever, preceded

branch parasiti

EPIPHYTISM: DEFINITION AND BREADTH

Epiphytes are plants that spend much or all of
their lives attached to other plants. Qualifying
forms range from microbes to angiosperms; both
aquatic and terrestrial vegetation provide me-
chanical support. Interaction with phorophytes
can range from incidental to physiologically in-
timate; primarily rain-fed “atmospheric” bro-
meliads and orchids (Figs. 3, 4) are anchored by
a few roots (sometimes only one), whereas con-
tact is almost complete for the largely endophytic

dwarf mistletoes. (The designation “epiphyte” is
here reserved for free-living vascular species; it
does not include hemiparasites.) This report has
two purposes: to describe taxonomic participa-
tion in epiphytism and to explore the extraor-
dinary proliferation in arboreal habitats by some
families and higher taxa. First, epiphytes are ex-
amined in terms of basic characteristics and im-
portant biological distinctions.

Plants considered to represent a specific eco-
logical category usually share key qualities that

! The first four papers of this issue were presented as part of the 32nd Annual perd Symposium, The

Biology of Epiphytes, held at the Missouri Botanical Garden 18-19 October 1985, an nd s
11392, Gerrit Davidse, Principal Investigator

grant BSR 831

—— in part by
nto the Missouri

Botanical Garden.
? Oberlin College, Oberlin, Ohio 44074, U.S.A.

ANN. Missouni Bor. GARD. 74: 183-204. 1987.

ounda

184

set them apart from other vegetation. Occurrence
on the same general type of substratum and, more
importantly, utilization of comparable resource
bases by similar mechanisms require consider-
able, often conspicuous, convergence. All botan-
ical carnivores possess traps designed to attract
and process fauna for food; all lianas feature
slender habit and novel vascular anatomy; all
vernal ephemerals from temperate deciduous
forests generate simplified shoots bearing helio-
philic, short-lived foliage. In contrast, anchorage
in tree crowns— sometimes even inability to sur-
vive terrestrially—exhibited by so many of the
approximately 25,000 epiphytic species appears
to have little unifying basis. No growth form,
seed type, identity of pollen vector, water/carbon
balance regimen, source of nutrient ions, nor re-
source procurement mode is shared by all me-
chanically-dependent species. Moreover, char-
acteristics of epiphytic and soil-based flora
overlap broadly, as do important aspects of their
habitats.

Growing conditions for epiphytes, as indicated
by rooting media, microclimate, and stability and
dispersion of substrata, are diverse and similar
to those for much terrestrial vegetation. Like soils,
which range from equable to dry and infertile,
canopies of everwet to microphyllous to cactus
forests 1 Dose me bi severe stress on resident
flora. Problem d dispersal in tree
crowns are probably similar to those experienced
by cliff dwellers and plants native to other pre-
cipitous sites. A naked bark surface must com-
plicate water and mineral balance just as do thin
soils and rock faces. At the other extreme, how-

ever, the everwet forest's rotting trunks or
branches (Fig. 2), debris-filled knotholes, and ant
nests (Fig. 7) probably offer to certain humi-
philous epiphytes resources equal to those avail-
able at humid, fertile, terrestrial sites. In effect,
heterogeneity and similarity to other habitats
complicate definition of the epiphytic biotope
and thwart attempts to understand why tree
crowns offer the only recourse to such a variety
of vascular plants. From another vantage point,
canopy-dependent species seem to be no more
versatile than soil-rooted plants; most will sur-
vive only under narrowly proscribed circum-
stances (e.g., rigid confinement to twigs rather
than larger axes, humus as opposed to “uncon-
ditioned”’ bark, dark instead of better-illuminat-
ed sites)

Many taxonomies have been formulated to
classify mechanical dependence. Criteria most

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

often used are extent of arboreal presence (fa-
cultative vs. obligate forms), nature of depen-
dency on supporting vegetation (mistletoes vs.
epiphytes), exposure requirement (heliophiles vs.
sciophytes), habit (e.g., tank forms, myrmeco-
phytes), and type of substratum (e.g., humi-
phobes, ant-nest forms). A further criterion rec-
ognizes two types in a special group that normally
taps soil part of the time—the hemiepiphytes.
Primary hemiepiphytes, a group which includes
stranglers, begin life on bark and later produce
soil roots; secondary hemiepiphytes (Fig. 11)
germinate on the ground and then become can-
opy-dependent as older roots and basal stems of
vining shoots decay. Features more directly re-
flecting peculiarities of vegetative function as-
sociated with epiphytic life (e.g., presence of ab-
sorptive trichomes, carbon fixation pathway)
have not generally been employed to distinguish
epiphytic vegetation despite their utility for ex-
plaining type of microsite and how existence in
that kind of space is sustained.

ne criterion above all others may prove
meaningful to those concerned with the me-
chanics of epiphytism: temporal access to mois-
ture. While adequate mineral nutrition and ex-
posure are no less critical than are suitable water
relations to long-term survival of an epiphyte,
avoiding drought injury is the more immediate
challenge. Constant adjustment to rapidly chang-
ing moisture supply must be accomplished via
appropriate stomatal and photosynthetic re-
sponses. There are just two types in this new
taxonomy: continuously-supplied (CS) and pulse-
supplied (PS) forms. Moisture supplies are steady
for CS species tapping deep, absorbent media
(Figs. 2, 5) or catchments created by the plant
itself (Figs. 8, 9). The most self-contained mem-
bers of the canopy-dependent flora (PS epi-
phytes) draw water from relatively nonabsorbent
substrata or other sources subject to quick drying
(Figs. 3, 4).

Epiphyte life history has been influenced by
many selective forces, including substratum dis-
tribution and stability. Patchiness in the epi-
phytic biotope ranges from gross patterns due to
hyperdispersion of tropical-forest trees to finer-
grained discontinuity in the scattered arrays of
suitable bark within individual hosting crowns.
Mortality continues to be high during and fol-
lowing seedling establishment. Disturbance is le-
thal when supporting bark fragments exfoliate,
inhabited twigs fall, infested trees collapse, an
(less common but broader in extent) natural dis-

1987]

BENZING— VASCULAR EPIPHYTISM

185

ed on.

ing in a rotting bra

zomes pearing traps of sik ales i ucl a tank brom ges NE id
ot as, Venezuela.—

artially dissected shoot of the bromeliad Brocchinia tatei on Cerro Neblina, Venezuela,

mic.— 2. Young specimen of
ing electron micrograph of the
bi

of Campylocentrum pachyrrhizum and two small bromeliads growing on a small branch in south Florida.

asters ravage whole communities. Effective patch
life (patch duration relative to the interval need-
ed for a resident plant to reproduce) must be
especially short for a multitude ofepiphytes whose
fecundity is limited by drought and poor nutrient
sources. It is this group that possesses an unusual

combination of stress and r-selected characters
(Benzing, 1978). In contrast, CS species, with
their relatively regular resource supplies, should
exhibit less emphasis on shortening the life cycle
and channeling scarce commodities into repro-
ductive rather than vegetative tissue.

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

1987]

ADAPTIVE VARIETY IN CANOPY DEPENDENTS

Some adjustments to conditions in tree crowns
are predictable and obvious— viz. capacities for
aerial dispersal and holdfast — but others are more
cryptic and varied. Depending on taxon and hab-
itat, epiphytes exhibit various mechanisms for
carbon fixation and procurement of moisture and
nutrients. Essential elements are drawn from re-
markably diverse sources, including several not

ormally available to plants. A survey in some
Indian forests (Sengupta et al., 1981) revealed
greater nitrogenase activity in the phyllosphere
of epiphytes than of terrestrials, including host-
ing trees. Impoundments built of roots or shoots
provide access to nutrients via intercepted can-
opy fluids and litter (Figs. 8, 9) and the necessary
detritivores and saprophytes these catchments
attract (Benzing, 1986a). Carnivory is rare in the
epiphyte synusia (Fig. 1), while trophic myr-
mecophily (Fig. 12) is common and may, in fact,
be entirely restricted to canopy habitats (Thomp-
son, 1981; Givnish et al., 1984). Where substrata
are deeply penetrable and moist, absorptive or-

special permeabilities to match the character of
the media (organic and acidic) they so often ex-

oit.

Water/carbon balance is also assured by con-
siderable mechanistic variety. At one extreme,
some canopy- -dependent plesdanhyes: alterne

croclimate dictates: Quite the opposite, im-

poundments provide continuous drought relief
for hundreds of species of bromeliads (Fig. 8),
fewer orchids, some other ts (Fig. 9), and
ferns. Turnover of the entire leaf surface cued by
impending seasonal drought is exemplified by
members of several orchid genera (Fig. 2). Cras-
sulacean acid metabolism (CAM) may be better
represented in the forest canopy than in any other
habitat type; stable carbon and hydrogen isotope
data as well as more direct measurements of fix-
ation pathway indicate that every known vari-
ation on this mechanism exists in epiphyte flora.
Occasional subjects—the shootless orchids, for

BENZING— VASCULAR EPIPHYTISM

187

instance—exhibit novel machinery for gas ex-

ange (Cockburn et al., 1985). Unusual osmotic
qualities and related stomatal sensitivities ap-
pear to rank among the most unique of the func-
tional peculiarities of epiphytism (Benzing,
1986a). Survival is often predicated on extensive
storage capacity, economical water use, and abil-
ity to rebound rapidly from drought-imposed
stress. Water balance is aided by special absorp-
tive tissues that prolong contact with transitory
fluids via mini-impoundment (e.g., the velamen
of orchid roots; Fig. 10; Benzing & Pridgeon,
1983). Like other stress-tolerant plants, PS epi-
phytes grow slowly, a characteristic which mod-
erates resource requirements but heightens vul-
nerability to habitat patchiness, disturbance, and
other phenomena that oblige heightened fecun-

ty.

Habits of epiphytes are often specialized in
ways other than those associated with impound-
ment and myrmecophily. Abbreviation and
merger of functions is common in the especially
xerophytic forms, particularly the PS epiphytes;
the most spectacular reductions occur in Bro-
meliaceae and Orchidaceae (Benzing & Ott,

and paralleled by complementary shoot m
fication. Just about every intermediate condition
between profuse and very sparse rooting exists
throughout the complex of more than 500 species.
Corresponding shoot changes center on a pro-
gressively diminished impoundment capacity and
an indumentum elaborated to the point where,
among atmospheric forms, absorbent trichomes
densely cover all foliar and most stem surfaces
(Fig. 3). At this stage of specialization, no other
organs are needed for uptake function. Orchi-
daceae (subtribe Sarcanthinae) exhibit a com-
parable progression except that here a telescoped,
leafless shoot now supports production of green
roots and the periodic axillary scape (Fig.

The loss of axial differentiation seen exclu-
sively in PS epiphytes may not be coincidental.
Species that grow attached to impenetrable me-
dia or hang free in the atmosphere occupy hab-
itats as uniform as those colonized by prevas-

—

FIGURES 5-8.
Neblina, Venezuela.— 6.

the base of a tree in undisturbed lowland rain forest in Amazonas, Vene uela.
in ant-nest carton in lowland Venezuelan rain forest.
orida

—5. A mixed colony of angiospermous epiphytes rooted in deep
dense growth of bryophytes supporting ipd tolerant hymenophyllaceous ferns at
7. Seed

humus on a palm on Cerro

lings of epiphytes rooted
—8. Tillandsia ey M a tank bromeliad, in south

188 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

,
` -

_ CORTEX

1987]

cular plants or modern aquatics. In such an
environment, PS epiphytes need not maintain
the division of labor between shoot and root that
would be required for life on soil and, in fact,
may sacrifice fitness by retaining a body plan
inherited from terrestrial ancestors. Material and

of

energy econor
vegetative Exo in organ system while
other parts are lost may promote fitness where
stress and high mortality place a premium on
fecundity. Complex functional tradeoffs obliged
by extreme a reduction in Bromeliaceae and

J
1

(Benzing & ps 1981; Benzing et al., 1983
Somewhat less spectacular abbreviations leading
to single-leaf ramets parallel increased exposure
and stress tolerance in additional, predominantly
epiphytic, orchid lines (e.g., Dendrobium, Epi-
dendrum ).

Although extensive studies have been made of
orchid pollination, seed germination, and seed-
ling nutrition, and some work has been done on
mistletoes (e.g., Bernhardt, 1983), bromeliads
(Smith & Downs, 1974), and a few other epi-
phytes, life history characteristics of canopy-de-
pendent vegetation as a whole remain obscure.
What data are available indicate that nonuni-
formity is the rule once again. An exception is
the usual pattern of extended iteroparity; only a
few Tillandsia and Vriesea species flower just
once. Pollinators include all the common vectors
except wind. Because air turbulence is adequate
to disperse most epiphyte seeds despite their
greater mass compared with pollen, absence of
wind pollination must relate to other factors, most
likely to the expense of required reproductive
material or hyperdispersion of populations. Syn-
dromes featuring specialized animals or unusual
foraging activities are more common here than
in other tropical synusiae. Breeding systems vary
from those obliging strict outcrossing via traplin-
ers to autogamy. Some minor patterns may be
emerging, however. Orchids seem to be predom-
inantly allogamous, while many ant-nest inhab-
itants and Marcgraviaceae are selfers (Gentry &
Dodson, 1987). The widely-held view that, on
average, epiphyte populations are more frag-

BENZING — VASCULAR EPIPHYTISM

189

mented than those of terrestrials and hence sub-
ject to special genetic structuring and propensity
for cladogenesis needs to be verified. If they are
not, neither a uniform nor unique reproductive
pattern should be expected.

Seed is dispersed by various agencies. Among
families, the vectors are most often animals (via
fleshy fruits), but when species are counted,

g wind-borne species
(more than 80% of the total), seeds bearing elab-
orate appendages are uncommon. Attempts to
relate seed form and mass to mobility in aerial
habitats can be complicated because disparate
agents sometimes provide service for the same
species. Dischidia seeds bear a plumose coma but
also an oily eliasome to attract ants (short-range
vectors; Docters van Leeuwen, 1954). Nest-in-
habiting Codonanthe has a fleshy berry contain-
ing myrmecochorous seeds. Other subtleties of
epiphytic reproduction are just as easily over-
looked — one flat side on otherwise fusiform Hyd-
nophytum seeds reduces vulnerability to dis-
lodgement by stemflow (pers. obs.). Seed size
differential between terrestrial and epiphytic rel-
atives is not consistent. Madison (1977) reported
more massive seeds in soil-based compared with
canopy-based Araceae and Cactaceae, but epi-
phytic gesneriads definitely do not produce light-
er seeds than do numerous terrestrial relatives
(H. Wiehler, pers. comm.). Surprisingly, micro-
sperms of terrestrial orchids are more buoyant
than those of test epiphytes (Stoutamire, 1974).
Rockwood's survey (1985) of 365 species in eight
families from diverse communities in Costa Rica,
Panama, and Peru yielded additional, unexpect-
ed results. Average seed mass of 59 epiphytes
(orchids excluded) was lighter than that of tree
seeds but heavier than that of terrestrial herbs
and shrubs.

Table 1 tabulates angiosperm families con-
taining more than 50 epiphytic species and lists
characteristics that contribute to their success in
tree crowns. Figure 13 summarizes adaptive di-
versity within free-living, canopy-dependent,
vascular flora relative to local moisture supply.
Species with character states assigned to the far

—
FIGURES 9-12.

section through the root of E pidendrum ] rise illustrating the major tissues, x 250.—

—9. A “trash basket" Anthurium in lowland rain forest in Amazonas, Venezuela.— 10. Cro

11. Hemiepiphytic aroid

in rain forest at Río Palenque, Ecuador.
within the swollen hypocotyl.

o

[Vor. 74

ANNALS OF THE MISSOURI BOTANICAL GARDEN

190

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right of the grid (indicated by lines) are usually
PS types. Those on the extreme left experience
continuous supply punctuated only by the un-
common, normally brief, drought. Epiphytes
bearing character states assigned predominantly
to the left or right of Figure 13 can occur together.
Pulse-supplied and CS forms inhabit humid sites,
although only PS epiphytes occupy tree crowns
in drier locations. Twig and free-hanging “mist”
types in rain forests, for instance, receive mois-
ture and nutrients exclusively via periodic flow,
so their supply is intermittent. A nearby tank or
humus epiphyte experiences considerably less
discontinuity between rain events. Species situ-
ated on highly exposed sites in humid forests or
on most anchorages in arid woodlands exhibit
slow growth, long-lived photosynthetic organs,
and foliar or root surfaces modified to prolong
contact with passing fluids (i.e., thick velamina
or absorbing trichomes). Vegetative bodies are
often much abbreviated, another means of im-
proving economy. Woodiness seems to be sus-
tainable only where moisture is abundant.
Features associated with regular resource sup-
ply are diverse. Hemiepiphytes are continuously
supplied, at least while rooted in soil; habits may
be woody or herbaceous. Nutrient i drawn
from various types of humus by roots and ab-
sorptive leaves of various CS forms. Carnivory
is possible for the tank former. Ant-nest gardens
are scarcer in the drier forests, but trophic myr-
mecophily is widely available. Too little is known
about mycorrhizae and nitrogen nutrition in the

tosynthetic pathway, first supplemented with
CAM-cycling and then ever greater reliance on
nocturnal fixation, seems to play a diminishing
role from left to right. Precise ordering of CAM
variations in Figure 13 and the proposed signif-
icance of CAM-cycling as a mechanism that
*poises" the plant for drought (Ting, 1985) should
be considered provisional at this time. Poikilo-
hydry is feasible only on relatively humid sites
for reasons described below. Deciduousness (e.g.,
Catasetum: Fig. 2) occupies an intermediate po-
sition, occurring where drought is predictable
(seasonal) and not too severe. High solute po-
tential has been documented among CS and PS
epiphytes (Harris, 1918; Spanner, 1939; Sinclair,
1983) although its effect on stomatal sensitivity
has not been examined extensively. Vegetative
features seem to have no bearing on type of pol-

BENZING— VASCULAR EPIPHYTISM

191

linator used. Dispersal syndromes correlate only
weakly with habitat humidity or resource pro-
curement mechanisms. Anemochory is found
everywhere, but a pulse-supplied nature in-
creases the probability that an angiosperm will
be wind-dispersed. Propagules that pass safely
through a bird's gut are generally larger (more
expensive) than the smaller anemochores, a re-
lationship that may help explain why wind-dis-
persed Tillandsia and Orchidaceae dominate the
most stressful epiphytic habitats. Without small
seeds, low productivity would probably result in
low generative capacity for the PS epiphyte. The
few monocarpic bromeliads are relatively well-
provisioned impounders rather than stress-tol-
erant atmospherics; the latter’s resource pro-
curement capability is probably inadequate to
support semelparity (Benzing & Davidson, 1979).
In summary, epiphytes engage in a broadly
similar rather than a narrowly proscribed way of
life. Many other ecological groups are better de-
fined because members subjected to less varied
environmental constraints conform to more co-
herent adaptive syndromes. Inconsistency in the
epiphyte synusia is a function of habitat breadth
(numerous possibilities for adaptive specializa-
tion)and th ftree
crown occupants. More than one daina com-
bination can be associated with specific types of
rooting media and climates, and no single key
feature or suite of features is evident. Vegetative
henomena that seem to be proportionately bet-
ter represented i in canopy- than in soil-dependent

patchiness and disturbance. Modes of pollen and
seed conveyance are also various among epi-
phytic plants, with few, if any, characters unique
to these taxa. Indeed, most features of canopy-
dependent plants occur among terrestrial flora as
well, although not necessarily in the same com-
binations.

SYSTEMATIC OCCURRENCE AND THE
MULTIPLE ORIGINS OF EPIPHYTISM

Approximately 10% ofall vascular plant species
are at least occasionally epiphytic, but distribu-
tion among higher taxa is uneven (Gentry &
Dodson, 1987). Broad involvement is particu-

[VoL. 74

ANNALS OF THE MISSOURI BOTANICAL GARDEN

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BENZING— VASCULAR EPIPHYTISM 193

1987]

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194

larly apparent at the higher taxonomic levels.
Fully 44% of all vascular plant orders and 16%
of families (sensu Cronquist, 1968) contain
one or more tree-crown populations. Arboreal
monocots far outnumber dicots, even exceeding
Filiopsida (44 vs. 2796) for proportional occur-
rence in forest canopies. Of the two large lycopod
genera, Lycopodium is about one-third epiphytic
(143 spp.) while Se/aginella —a considerably
larger, heterosporous group— contains only five
such species (Beitel, 1979). A majority of Psi-
lotophyta regularly inhabit tree crowns. Gym-
nosperms, on the other hand, are largely confined
to soil, in part no doubt because of their heavy
seeds and costly wind pollination. Orchidaceae
have been more successful than any other lineage
in canopy habitats. Two out of three epiphyte
species are orchids; about 7096 of the family is
canopy-dependent. Two other large monocot
groupswithar ra-
ceae, especially Anthurium, Tiladandron. nd
Rhaphidophora, and Bromeliaceae, more than
half of which anchor on bark. Canopy-dwelling
dicots are disproportionally represented by Cac-
taceae, Ericaceae, Gesneriaceae, Melastomata-

rsatility within
participating clades is not consistent. Compo-
nent taxa may be diverse or uniform in the mech-
anisms that foster habitation in tree crowns (Ta-
ble 1).

In all, about 80 vascular families contain at
least one canopy-dependent member, but there
are conspicuous, occasionally puzzling, omis-
sions. Some very large, ecologically diverse groups
with numerous tropical members are little rep-
resented in, or even absent from, canopy floras.
Leguminosae may be constrained by fruit and
seed types that offer little potential for aerial dis-
persal. Poaceae (and all Cyperales) may be al-
most entirely terrestrial owing to consistent wind
pollination and the graminoid habit, which does
not lend itself to evergreen drought endurance.
Plausible reasons for the paucity of epiphytes

this family. These CAERpHODP e
n the
more advanced taxa. The same is true of ferns,
but again, there are exceptions. Two members
of the primitive order Ophioglossales root in hu-
mus impounded by persistent palm leaf bases

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

and Platycerium fronds. If Psilotum and Tme-
sipteris of the bigeneric Psilotophyta are indeed
sole remnants of the Silurian/Devonian psilo-
phyte complex, epiphytism is possible for the
most primitive vascular relics.

Epiphytism has probably evolved indepen-
dently from terrestrial stock in every participat-
ing family of seed plants and most families of
ferns. It has arisen more than once in most of
those spermatophyte taxa containing canopy-de-
pendent species on different continents (Gent
& Dodson, 1987), indicating lI Spit
for arboreal life is rather fundamental in s

major clades. Most diverse of ic Wi bros
types, phylogenetically speaking, are the humi-
philes whose drought liability obliges access to
moist organic debris such as that covering older
bark exposures in everwet forests. Virtually every
family containing epiphytes includes at least one
such species; most contain no other type. Where
more exacting conditions (e.g., exposed bark and
twig surfaces) prevail, few families are repre-
sented, although these are sometimes speciose
(Benzing, 1978). Neotropical ant nests provide
specialized substrata for a small myrmecocho-
rous flora which often offers extrafloral nectar.
Perhaps additional traits as yet unrecognized that
might affect capacity to root in cartons are mostly
limited to Araceae, Bromeliaceae (specifically
subfamily Bromelioideae), Cactaceae, Gesneri-
aceae, Marcgraviaceae, Orchidaceae, and Piper-
aceae. Some Asclepiadaceae, Melastomataceae,
and Rubiaceae are part of similar but less well-
defined symbioses in Australasia (Janzen, 1974).
Those trophic myrmecophytes that exhibit un-
equivocal modifications for ant occupancy all be-
long to Asclepiadaceae, Bromeliaceae, Melas-
tomataceae, Orchidaceae, Polypodiaceae, and
Rubiaceae (Huxley, 1980). Rubiaceae seem to be
most specialized for the relationship (Fig. 12).
Stranglers—about 300 in all—come from just
eleven families of dicots, most notably Moraceae
(Ficus). Secondary hemiepiphytes have vining
habits, and indeed the majority belong to groups
with a scandent tendency (e.g., Araceae, Cyclan-
thaceae, Marcgraviaceae). Bromeliaceae largely
account for the tank formers (Fig. 8). Trash-bas-
ket habits (Fig. 9) are present in a modest number
of families; the numerous PS epiphytes are al-
most exclusive to Bromeliaceae (mostly Til-
landsia) and Dreidaceue (many subtribes).

Both parallelis
much sada in the rise of epiphytism. Par-

o,

1987]

allelism is illustrated by CAM, which features a
y mechanism (incorporation of CO, via 8-car-
boxylation) fundamental to regulation of ionic
and osmotic balance through metabolism of ma-
late and perhaps other organic ions (via a bio-
chemical pH-stat). Heightened enzyme activity,
appropriate malic acid mobility and storage ca-
acity, phase-shifted stomatal behavior, and oth-
er associated characters enabling high water use
efficiency have emerged in close to 30 families
and more than once among some of those con-
taining epiphytes— Bromeliaceae and Orchida-
ceae, for instance. Inherently slow relative growth
rates, great longevity of whole plants and their
individual parts, along with underlying oligotro-
phic physiology in canopy- and soil-based vege-
tation alike, probably also reflect common po-
tential realized under similar selective impetus.
Somewhat more unique features based less on
homology than on novel design opportunity are,
among others, devices to effect impoundment
and/or accommodate ant occupancy. On bal-
ance, gresource use appear
to be based on relatively few. widely available,

tures effecting resource procureme mo
numerous and varied origins, s. likely based on
convergence.

PREDISPOSITION AND ADAPTIVE
CANALIZATION

Compared with vertebrates, vascular plants

hibit varied habits, tolerances, and life histories.
Quite commonly, natives of disparate habitats
even belong to the same genus. Difference in evo-
lutionary pattern partly reflects the higher plant's
greater physiological plasticity aided by contin-
uous turnover of both vegetative and reproduc-
tive organs. Selection may occur among ra-
owe even tissues within individual
rgans— during the life of a single clone (Walbot
& Cullis, 1985). Evolutionary tempo is rapid
enough to effect speciation in as little as 104—105
years—or perhaps a few decades! —in Orchida-
ceae (Gentry & Dodson, 1987). Reputedly, only
a small number of genes need to be involved to
alter structure, function, and ecological tolerance
(Gottlieb, 1984; but see Coyne & Lande, 1985).

BENZING— VASCULAR EPIPHYTISM

195

Some angiosperm lineages are quite conserva-
tive, however, as if for them, too, entry into cer-
tain adaptive zones restricts access to others.

Cases of evolutionary canalization abound in
Magnoliophyta. Somatic conservatism is fre-
quent within genera and an invariant adaptive
mode can typify entire, albeit small, specialized
families (e.g., Sarraceniaceae, Lemnaceae). More
impressive evidence for constraints on direction
but not speciation in plant evolution is provided
by greater redundancy in other families. Exam-
ples include halophytism in Chenopodiaceae,
ruderalism in Brassicaceae, drought-enduring
xerophytism in Cactaceae and, of course, epi-
phytism in Orchidaceae. No single life style char-
acterizes the entire clade in any of these exam-
ples, but particular themes turn up too often to
deny underlying family-wide dispositions. Suc-
cess in a particular environment is predicated on
a novel set of features that imparts ability to
counter major external constraints. Once the es-
sential components were perfected and integrat-
ed, whether by serendipity or some optimizing
mechanism, each ancestral stock became better
equipped to enter one, and less so another, adap-
tive zone. Since accessible space was widely
available in each case, considerable cladogenesis
ollowed, sometimes through agencies unrelated
to vegetative performance, hence native substra-
tum. Orchidaceae achieved unequalled specia-
tion in humid, neotropical forest canopies be-
cause of behavior peculiar to male euglossine
bees.

Adaptive syndromes are comprised of struc-
tural, functional, and phenological components
which can be identified through consistent oc-
currence in participating species. Ruderals suc-

tats. Components of this well- ma re ah ii
are: | ical and
root tissue; vigorous S nud iran d
small, long-lived, often light-sensitive, propa-
gules; and self-compatibility. Here, such traits as
low shoot/root ratios, woodiness, and extensive
succulence are never found because they are in-
consistent with ephemeral life histories. Al-
though the habits of vascular halophytes range
from large trees (mangroves) to succulent terres-
trial and submerged herbs, certain physiological

phenomena are common to all because potentia
for desiccation is always one of the predominant

196

problems in saline media. Osmotic compensa-
tion and compartmentation of specific solutes
provide the only means for maintaining water
relations in hyperionic habitats. In every case,
potentially toxic ions, taken in from the envi-
ronment to lower tissue water potential, must be
sequestered in vacuoles at concentrations above
those encountered in glycophytes, and they must
be balanced across the tonoplast by osmotica of
plant origin. Salt balance can be fine-tuned fur-
ther by taxon-specific features. Chenopod halo-

hytism is aided in salty surroundings by mul-
ticelled vesicular hairs which excrete excess Na*
an ^ succulence provides for ion dilution.
The presence of organic solutes, including pro-
line, in other salt-tolerant taxa instead of the be-
taines found in Chenopodiaceae reveals that

ments is not complete. Likewise, the
abundant malic acid synthesized by all CAM
plants at night for subsequent daytime genera-
tion of an internal CO, supply is usually decar-
boxylated through only one of the three existing
reactions: mediation by NAD- or NADP-depen-
dent malic enzyme or PEP carboxykinase.

On balance, floristic variety generally reflects
the equability and heterogeneity of a particular
adaptive zone, all other variables being equal.
Fertile, moist soils and abundant light have
. wide convergence—ruderals are a good

ample. Weeds in crop fields belong to many
nde reflecting easy access to a collection of
required character states. Tropical forest cano-
pies have been similarly colonized but only to a
certain degree. Whereas many lineages have
evolved traits that allow growth on the organic
arboreal “soils” of everwet forests, few have in-
vaded more demanding zones, as demonstrated
by narrower taxonomic participation in PS epi-
phytism (Benzing, 1978).

Inherited —in effect, phyletic — constraints de-
termined which ancestors of modern families
could generate epiphytic derivatives. Inherent
barriers are complex and little understood, but
their existence in plants is becoming recognizable
(e.g; Kochmer & Handel, 1986; Hodgson &
MacKey, 1986). Potential to express key com-
ponents of an adaptive syndrome was not suffi-
cient to assure its establishment. Universal oc-
currence of CAM-like function, for instance, did
not guarantee adoption of CAM by all xero-
phytes nor by all aquatic macrophytes in the soft-
water lakes where nocturnal fixation is favored

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

by limited carbon supply (Keeley, 1981; Rich-
ardson et al., 1984). Moreover, each character of
a syndrome must be free of pleiotropic and link-
age relationships with maladaptive or internall
incompatible traits. Biomechanical compatibil-
ity is essential; for instance, plants with vigorous
cambia may rarely employ CAM, probably be-
cause costly woodiness is not sustainable without
greater capacity for carbon gain.

ittle is known at this point about inertia in
the genetic, epigenetic, and biomechanical inter-
play among phenomena responsible for salt bal-
ance in halophytes, resource uptake and pro-
cessing in short-lived eutrophs, or water economy
in CAM plants. But occurrence among related
genomes does demonstrate that some characters
affecting vegetative performance and thereby
suitability in particular habitats — e.g., sites of ni-
trate reduction (shoot as opposed to root), ability
to adjust osmotically under various degrees of
stress, and presence of particular ion pumps and

More narrowly restricted
throughout Magnoliophyta, often to a single
family or order, are features such as capacity to
form a particular type of mycorrhiza, nodulate
with Rhizobium, synthesize cardiac glycosides or
betaines rather than anthocyanin pigments, and
utilize sorbitol rather than sucrose for phloem
transport.

Access to vital characters during plant evolu-
tion has been affected by related impacts on pho-
ton, water, and nitrogen use efficiency (Raven,
1985a). Constraints on evolutionary opportunity
have been strongest where drought, shade, or
infertile substrata accentuate premiums on water,
energy, and nutrient economy respectively. Type
of nitrogen source, for instance, imposes different
demands, depending on where (in which organ)
processing takes place, how much water and en-
ergy is needed per unit of product, and the en-
vironmental context (is light or moisture scarce
or abundant?). Calculation of comparative costs

creted by NO, assimilators. Although the am-
monium-to-protein pathway is least expensive
in terms of energy consumption, overriding fac-
tors may still dictate another choice even where
NH, is in greatest environmental supply. Soil
is the usual sink for H: generated by NH,* use.

1987]

Indeed, owing to the immobility of protons in
phloem, terrestrial plants process most of their
acquired NH, in roots, a potential limitation
with special relevance for epiphytism. If this is
the universal rule, then what compensation, if
any, accompanied root system reduction in ad-
vanced Bromeliaceae? Evidence indicates that
NH, is the predominant form of nitrogen in at
least some tropical-forest canopies (e.g., Curtis,
1946). Conceivably, the absence of similar mor-
phological diminution in nonbromeliad lineages
is in part related to their less flexible nitrogen
metabolism, although the role bromeliad foliar
trichomes have played in obviating absorption
by roots cannot be ignored in such comparisons.
Perhaps slow-growing plants like the atmospher-
ic bromeliads metabolize N at such low rates that
complications are avoided. Either the internal
biochemical pH-stat is adequate for disposal or
excess protons are dumped from trichomes when
shoots are wette

Similarly, land plants have evolved several
mechanisms to effect osmotic balance. Here, cost
escalates with deployment of inorganic (e.g., Na*,

Cl), then organic nonnitrogenous, and finally
nitrogenous solutes, especially on infertile me-
dia. Again, nitrogen is an important currency but
perhaps no less so than the energy and water
spent for its acquisition and processing. In a
somewhat different vein, fungal biomass has to
be supported by the mycorrhizal plant; but re-
turns in phosphorus, and sometimes water and
other nutritive ions, justify the investment in all
but the most fertile habitats. Comprehensive cost
accounting, through knowledge of functional in-
compatibilities and the hereditary and epigenetic
phenomena controlling access to key traits, is
necessary to interpret patterns of radiation. It is
at these levels of plant performance that many
of the tradeoffs, economies, and accommoda-
tions mo the evolutionarily-stable strat-
egy oc

HISTORICAL BASIS FOR CANOPY
DEPENDENCE

It is currently impossible to explain fully why
one particular lineage developed canopy depen-
ence while another did not. But partial answers
are available in some cases; several of the more
notable ones are discussed below. Four questions
provide focus: Why are proportionally more ferns
than seed plants epiphytic? Why do so many

BENZING— VASCULAR EPIPHYTISM

197

monocots, particularly orchids, dwell in cano-
pies? Why have several families of dicots with

bvious advantage by basic habit or water
balance profile hun a ie so widely there? Fi-
nally, why is branch parasitism relatively un-
common?

Ferns. Homosporous pteridophytes are suc-
cessful in tree crowns, where they usually occupy
lower strata, because of small diaspores. A sec-
ond, less obvious, factor is a capacity to tolerate
substantial drying and deep shade. Poikilohydry
is pronounced in exceptional taxa (Polypodium
polypodioides), but many other filicaleans exhibit
desiccation tolerance superior to that possessed
by most seed plants. A fern's pattern of drought
resistance, unlike that of most CAM plants, is
particularly compatible with occurrence deep in
the forest. For instance, ultrathin fronds of Hy-
menophyllaceae (Fig. 6) probably photosaturate
at very low fractions of full insolation and they
can survive considerable desiccation. Greater ex-
posure might be tolerable, and upper as well as
lower strata heavily colonized, but for the trade-
off associated with poikilohydry. Resurrection is
adequate for countering the occasional brief
drought every umid forest ascent now and
then, but frequent d another matter.
A regulated water ebum e on thick (ex-
pensive, opaque) epidermal barriers and greater
diffusive control is critical on markedly arid sites
because photosynthesis is more likely than is res-
piration to be curtailed by severe water deficits.
d moisture supply be too intermittent and

=
i O

9 !
Raven (1985b) cited rates of physiological pro-
cesses, including photosynthesis and transpira-
tion at the low end of the ranges reported for
tracheophytes, as a reason why ferns are so well
equipped to inhabit shady, drought-prone loca-
tions.

Nevertheless, a modest invasion of drier lo-
cations has been possible for higher ferns. One
enabling mechanism here is drought avoidance
via seasonally deciduous foliage (e.g., PAlebo-

ium aureum). Occurrence in some stressful
Australasian sites is possible for evergreen Pyr-

Drymoglossum piloselloides, Pyrrosia longifolia,
and P. angustata (Wong & Hew, 1976; Sinclair,
1983; Hew, 1984) along with drying character-

198

istics more reminiscent of a resurrection plant
than a typical desiccation-resistant xerophyte. A
thorough examination of ferns with regard to
microclimates, substrata, and water and carbon
relations in both gametophyte and sporophyte
stages will be necessary to place discussion of the
evolution of pteridophytic epiphytism on a firm-
er foundation.

Liliopsida as a whole. Orchids account in large
measure for the immense numbers of epiphytic
species, but monocots would be disproportion-
ately common in tree crown habitats even with-
out them. Bromeliaceae and Araceae rank sec-
ond and third. The ratio of monocot to dicot
species in tree crowns is 5: 1, but itis 1 : 4 overall.
Although Araceae, Bromeliaceae, and Orchida-
ceae are the most successful families in forest
canopy habitats, there is no common adaptive
theme. Two photosynthetic pathways in many
variations, tank and trash-basket impound-
ments, myrmecophytism, foliar trichomes, vel-

phytic monocots. A peculiar body plan, shared
to some extent with the higher ferns but less so
with dicots, may have offered special class-wide
opportunity.

Monocots in particular, but dicots as weil, are
often sectoralized in the sense that individual
shoots operate as collections of relatively inde-
pendent, serially aligned, physiological units
(IPUs; sensu Watson & Casper, 1984). Tracer
studies have shown that meristems may receive
fixed carbon mainly from nearby leaves, some-
times only those attached at the subtending nodes.
Longitudinal segmentation seems to be more
characteristic of dicots, and may also help ex-
plain why several different habits and associated
ecological tendencies have been emphasized in
one or the other ofthe two classes. Partitionment
into vertical compartments is evidenced by
movement of labeled photosynthate among
leaves and associated buds along a single or-
thostichy. Xylem supply is similarly restricted.
Secondary thickening may eventually obliterate
conductive barriers to lateral flux imposed by
the dicot primary body, peice where eu-

ure

as previously noted,

r set of limitations,
especially in arid habitats. Monocots with their
more reticulate '**atactostele," best known today

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

in palms (a group portraying the **Raphis Prin-
ciple"; Tomlinson ) appear to possess
unique capacity for functional integration on one
hand and, on the other, habits that permit ex-
tensive vegetative renewal in mimimal space.
Perhaps especially fortuitous is the unusual abil-
ity to coordinate remote sources and sinks, and
a related capacity to localize effects of damage
and maintain vascular supply to organs that a
more rigid system might be forced to abandon
or underutilize. This kind of flexibility was il-
lustrated by itl
mature ramets, returned to sink status by shading
or defoliation (e.g., Callaghan, 1984; Welker et
al. icd. remained alive rather than self-pruned,
as commonly occurs in forest trees. Historically,
peers of this sort may have fostered ex-
traordinary architectural plasticity n special
implications for novel evolution. Recall that sev-
eral large epiphytic monocot lineages have
undergone major vegetative reorganization un-
equaled in dicot counterparts.

The exceptionally well-developed horizontal
segmentation just described was undoubtedly
important in the evolution of stress-tolerant epi-
phytic monocots, perhaps ferns as well. The ad-
vantages of a rhizomatous sympodial habit and
potential physiological autonomy of the single
phyton (a morphological unit composed of a leaf
and associated adventitious root(s), bud(s), and
subtending stem segment) were most accentuat-
ed during orchid phylogeny. Sequential produc-
tion of reduced shoots (in effect, shoots com-
posed largely of single expanded phytons in
extreme cases) is a recurrent theme in this family.
Adult shoots of Dendrobium ultissimum consist
of nothing more than strings of stubby, closely
placed, leafless pseudobulbs. The broader suit-
ability of architecture based on serial renewal via
determinate shoots generated from closely-placed
meristems is illustrated by occurrence beyond
the monocots. Somewhat less condensed ver-
sions of the same general arrangement exist
mong mechanically-dependen , some
Gesneriaceae) and ‘Lycopodium. Differentiation
of roots into feeder and holdfast types (Fig. 9),
a useful division of labor for the vine or epiphyte,
seems to be more common in monocots than
elsewhere.

onorchid monocots. Bromeliaceae, with far
fewer species and almost exclusively neotropical
distribution, nevertheless rival Orchidaceae for
variety of epiphytic life styles. Tank habits have

dao5oo5vo vv hen

eo

1987]

evolved independently in two subfamilies, and
in all three if Brocchinia is correctly assigned to
Pitcairnioideae (Benzing et al., 1985). A rosulate
shoot was required for each transference of ab-
sorptive role from root to shoot. Ancestry was
apparently mesic in both Tillandsioideae and
nk shoots are associated with

ating soil substitutes in leaf bases, are funda-
b CAM plants. Specialization for PS
iphytism, in effect for greater stress tolerance,
Y: proceeded farthest in Tillandsioideae by way
of the derived atmospheric forms (Benzing et al.,
1985). Here, absorbing trichomes (Fig. 3) are per-
fected to the highest degree whilst the vegetative
apparatus is reduced to simplest form.

Epiphytic Bromeliaceae, more than most, were
clearly adapted to endure rigorous conditions by
the presence of a suitable epidermal appendage
and habit in precursors. Here, epiphytism is based
on a modified shoot (Fig. 8) with the foliar tri-
chome as its keystone feature (Table 1). Absorp-
tive function might be possible in glabrous leaf
bases when long-term contact with moist, nutri-
tive tank fluids is maintained, but the rapid up-
take required to sustain a rootless, nonimpound-
ing PS bromeliad would be impossible without
an extraordinary foliar indumentum. Myrme-
cophytism, and perhaps a single case of carnivory
(Catopsis berteroniana; Givnish et al., 1984), are
also associated with specialized trichomes and
inflated leaf bases (Benzing, 1970). Hypotheses
concerning how the bromeliad foliar epidermis
may have acquired its current function and im-
portance are described elsewhere (Pittendrigh,
1948: Benzing et al., 1985). (Briefly: contrary to
Pittendrigh’s proposition that absorptive func-
tion would only emerge under drought selection,
Benzing et al. posited a mesic, infertile, ancestral
habitat where the foliar epidermis evolved pri-
marily to promote acquisition of nutrient ions
from impounded humus or perhaps animal prey.)
Bromeliad seeds are disseminated by birds (Bro-
melioideae) or wind (Pitcairnioideae and Til-
landsioideae). Pollination syndromes are diverse
and apparently not associated with either tank
or atmospheric habit.

Aroid, by comparison with bromelioid or or-
chidoid, epiphytism is neither as advanced nor
as versatile. There are no reports of CAM here,
and overlapping foliage that might mitigate
drought lacks the water-tight quality possessed

BENZING— VASCULAR EPIPHYTISM

199

by inflated bromeliad leaf bases. Trash-basket
catchments (Fig. 9) sometimes trap falling litter
but little moisture. Pots fail to produce velam
ving the most drought;
tolerant Orchidaceae, nor is there any indication
that these organs can match foliage in photosyn-
thetic vigor. Seasonally deciduous leaves on green
or tuberous stems occur in Philodendron and Re-
musatia respectively, but these are minor themes
with few participating species. Arboreal exis-
tence in Araceae is based predominantly on two
mechanisms, both humus-based: impoundment,
seen in short-stemmed Anthurium and some
Philodendron (Fig. 9); and secondary hemiepi-
phytism (Fig. 11), a more widespread phenom-
enon most often M. xin in Philodendron.
nd t d vining hab-
its incorporating progressive dieoff of proximal
stem regions, appear to be the central vegetative
features responsible for aroid expansion in can-
opy habitats (Table 1). Both sympodial (e.g.,
Philodendron) and monopodial (Pothos) habits
are found in the hemiepiphytes. Ant nests are
utilized by some Anthurium and veo ei
Water and nutritional relations exhibit no ob-
vious unusual modHE onis for arboreal life but
neither have b losely. Baccate fruits
are an integral part of the syndrome, but they
occur throughout the family without habitat re-
striction. Pollinators range from beetles to eu-
glossines. Specialized pollen vectors may have
encouraged enlargement of Anthurium and pos-
sibly other genera. Cyclanthaceae, the only other
nonorchid monocot family with a sizable epi-
phyte contingent (about 6696 of Asplundia),
mostly penetrate the forest canopy as rooted
climbers and secondary hemiepiphytes. True
epiphytism occurs in Sphaeradenia and Stele-
stylis (G. Wilder, pers. comm.). Stems and inter-
nodes are shorter than those of related hemiepi-
phytes.

Orchid monocots. Orchidaceae owe their nu-
merical superiority among epiphytes to an ex-
ceptionally propitious set of vegetative and re-
productive features and to extraordinary
cladogenesis promoted by specialized pollina-
tion syndromes (Benzing & Atwood, 1984). Veg-
etative mechanisms vary tremendously accord-
ing to the taxon's native substratum and
ecoclimatic conditions. But there are several im-
portant attributes common to all canopy-depen-
dent family members that, in some form, pre-
disposed early stock for arboreal life. For example,

200

epiphytic Orchidaceae possess specialized roots
varying in photosynthetic performance and water
relations, depending on structure and metabo-
lism; uptake is enhanced in all cases by a non-
living velamen which imbibes precipitation and
contained solutes for subsequent sorption through
transfer cells in an underlying exodermis (Benz-
ing et al., 1983; Benzing & Pridgeon, 1983; Fig.
10). This same mantle effectively retards desic-
cation injury from both short-term and extreme
drought (Benzing, 1986a). Velamina simply em-
bolize air in order to break the hydraulic contin-
uum that, if left intact, would allow matric forces
in adjacent drying substrata to dehydrate living
epiphyte tissue. A green cortex supplements shoot
photosynthesis and is the major site of carbon
gain for the plant in exceptional cases (Fig. 4).
Hyperovulate gynoecia and aggregated pollen
characterize most of the family. Microspermy—
up to millions of tiny, lightly provisioned seeds
per capsule—requires fungal intervention for
germination. High-fidelity, long range, but often
inefficient pollinators promote ethological iso-
lation leading to plant speciation (Benzing & At-
wood, 1984).

Pre-epiphytic orchid stock probably possessed
structurally modified, locally suberized epider-
mis/hypodermis layers as do most extant terres-
trial family members and some other monocots
(e.g., Zea, Allium, Amaryllis). Similar root spe-
ae is sl eee m dicots. Microspermy
and associ were prob-
ably also CREE ina terrestrial context, as sug-
gested by the current habitats in which all other
such Hetesoiznphic: plant hoops occur (e-g..

Production of numerous tiny diaspores, subsis-
tence on transitory resource supplies, and main-
tenance of high water and nutrient use efficien-
cies set the stage for migration to forest canopies,
including many uninhabitable by less stress-tol-
erant epiphytes. Evolution of pheromonelike
fragrances and specialized floral morphology
tightened relationships with specific hymenop-
terans and dipterans, and assured extensive pro-
liferation of several clades that happened to be
canopy- -dependent. Large es ba related
ort-range
vectors with r no known propensity for exclusive
foraging suggest that substratum-specific factors

have also been important. As in a number of

other diverse communities of sessile organisms,
coexistence of densely packed epiphyte popula-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

tions may be favored on substrata subject to in-
termediate disturbance (Connell, 1978; Benzing,
1981, 1986b). Radiation in other speciose, can-
opy-dependent taxa visited by opportunistic or
sedentary pollinators (e.g., Anthurium, Peper-
omia) may be due to similar phenomena.
Dicots. Magnoliopsida are, on the whole,
poorly disposed to epiphytism — just 2% of species

ry
is often predicated on a single theme and no
other family incorporates the diverse resource

hibited by epiphytic Bromeliaceae and Orchi-
daceae. Except in Marcgraviaceae, dicot terres-
trials always outnumber confamilial epiphytes.
In tree crowns, Peperomia ranks first in size
among successful dicot genera and even families,
a statistic fostered by pantropical distribution,
the presence of CAM variations (Sipes & Ting,
1985), and high-volume production of small ad-
hesive fruits. Habits range from shrubby to mi-
nute and creeping. Moraceae also owe much of
their major epiphytic presence to a single genus
(Ficus) with a similar broad range; here the stran-
gler habit provides the vegetative basis for suc-
cess. Rampant speciation within a relatively nar-
row adaptive profile has again been encouraged
by circumtropic range and host-specific polli-
nators—in this case, the fig wasps. According to
Ramirez (1977), the moraceous strangling habit
evolved *'as a response to lack of light at the
forest level." Necessary attributes for success in-
cluded presence of viscid hyaline coats on seeds
that would germinate only on moist humus, long
aerial roots, water-use-efficient seedlings, and
dispersal by winged vertebrates. Marcgraviaceae
and Clusiaceae are additional single-strategy
families, emphasizing secondary and primary
hemiepiphytism respectively. Stranglers also oc-
cur in Schefflera (Araliaceae), Posoqueria (Ru-
biaceae), Metrosideros (M eae) and else-
where, but they are atypical among confamilial
canopy-based taxa

Most epiphytic Asclepiadaceae belong to
closely related, succulent, vining Dischidia and

oya. Flasklike leaves of D. rafflesiana and sev-
eral other species enclose nests; ants provide dis-
persal service for many more. Forms with less
specialized foliage regularly root in or grow against
ant debris, providing clues as to how ant-fed rel-
atives evolved. Dome-shaped leaves of D. col-
lyris grow tightly pressed against bark, providing

1987]

shelter for Iridomyrmex colonies (Huxley, 1980).
Photosynthesis involves CAM and/or CAM-cy-
cling (Kluge & Ting, 1978). Cactaceae became
equipped for the canopy-dependent synusia
through drought selection in terrestrial habitats.
Fleshy, small-seeded fruits and climbing habit
would eventually favor life in canopies. Move-
ment into tree crowns appears to have involved
some reversals. Originally aphyllous, stems of
the most advanced epiphytic forms (which hap-
pen to be natives of humid forests—e.g., Zygo-
cactus, Ripsalis) have lost their armature and
become much flattened, or narrowed if still terete
(e.g., Hatiora), presumably to improve perfor-
n shade. Family-wide CAM is probably

mands requires uptake by long-lived roots from
more or less continuous supplies in tree fissures
or soil (the secondary hemiepiphytes). Despite
the extreme drought tolerance of many terrestrial
relatives (e.g; Maxillaria, Ferrocactus), epi-
phytic Cactaceae seemingly never colonize the
most demanding bark and twig exposures.

Less obvious is the basis for high epiphyte suc-
cess in Ericaceae, Gesneriaceae, Melastomata-
ceae, and nonmyrmecophytic Rubiaceae. Most
canopy-dependent members in all four families
grow exclusively on humus mats in humid for-
ests. Woody habits and sclerophyllous foliage,
sometimes complemented by storage tubers,
characterize Ericaceae and Melastomataceae.
About one-half of the rubiaceous epiphytes
(Hydnophytum, Myrmecodia) supplement min-
eral nutrition and store moisture via ant-inhab-
ited, swollen hypocotyls (Fig. 12). Basically
herbaceous Gesneriaceae feature broader growth-
form variety, and several genera contain CAM
plants. Like some Peperomia, these gesneriads
exhibit trilayered mesophyll that may signal un-
usual photosynthesis. Substrata are more diverse
in this family, ranging from ant-carton to less
specialized humus. Codonanthe and related gen-
era, along with some hemiepiphytic cacti, are
probably the best drought-insulated of the dicot
epiphytes. Baccate fruits provide seed mobility
in most cases, although Rhododendron, a few
gesneriad genera, and large proportions of Me-
lastomataceae and Rubiaceae ripen wind-borne
seeds. Ant-associated species are myrmecocho-
rous.

Representation in canopy habitats varies
among these families, ranging from 4% to 35%

BENZING— VASCULAR EPIPHYTISM

201

of all genera in Rubiaceae and Ericaceae respec-
tively (Madison, 1977). The ericad statistic 1s all
the more ing the family’s size
ane numeraus temperate taxa. The other three

and neal ly OI exclusively moist-
tropical, hence have had greater access to arbo-
real habitats. Breadth and depth of specialization
for canopy dependence is further indicated by
comparing the total of exclusively epiphytic gen-
era containing two or more species with the num-
ber which include soil-based species as well. Ges-
neriaceae are most canopy-adapted by this
measure with 13 genera meeting each criterion,
while Ericaceae is least so with only four terres-
trial-free genera out of 22 containing epiphytes;
the largest of the four contains only eight species.
Vaccinium is especially noteworthy for its wide
range throughout Old and New World boreal to

Melastomataceae and Rubiaceae are 8:12 and

The influence of geography in determining
which families would contribute the most species
to the forest canopy flora (Gentry & Dodson,
1987) is apparent in Magnoliopsida. Epiphytic
Gesneriaceae are centered in the neotropics where
tribe Episcieae and particularly genera like Dry-
monia, Columnea, Dalbergaria, and Trichantha
have radiated extensively in tree crowns. Spe-
ciation here, as in epiphytic Liliopsida (Araceae,
Bromeliaceae, Orchidaceae), appears to involve
specialized American pollinators (humming-
birds and euglossines). Paleotropical counter-
parts are far fewer but still number more than
100 species. Canopy-dependent ericads are also
disproportionally neotropical and ornithophi-
lous. All but a small fraction of the peperomias
are American, although little is known about the
group’s reproductive biology. Because 73% of
Costa Rica’s large fern flora roots in tree crowns
(Wagner & Gomez, 1983), it appears that neo-
tropical epiphytism is not all pollinator-related.
Wide availability of moist montane forest in the
northern Andes and Central America, which is
reputedly amenable to fine niche partitionment
(Gentry & Dodson, 1987), has also been impor-
tant for expansion of American forms. Melasto-
mataceae and Rubiaceae number among the
few heavily epiphytic p families that
failed to generate more n Old World
epiphytes. Scrutiny of le aasan systems and
pertinent vegetative characters might help ex-

202

plain why mechanically-dependent New World
portions of these two families have failed to
undergo as much cladogenesis as have many co-
occurring epiphytic pteridophytic and angio-
spermous groups.

Chemical peculiarities of substrata or unusual
mycorrhizae may be responsible for uneven epi-
phyte development among higher monocot and
dicot taxa. Ericaceae, Melastomataceae, and Or-
chidaceae exhibit family-wide affinity for acidic,
often humid and organic, infertile soils. Little is

probably often scarce. Substrata in everwet forest
where most epiphytes live tend to be sodden, at
least moderately acid, and certainly organic. Use
of NH,* rather than oxidized nitrogen by plants
native to such substrata may have been a pre-
disposing character for epiphytism. Ericads are

notably deficient in nitrogen reductase, a sign of

long dependence on reduced nitrogen. Ericaceae
were perhaps especially well positioned for can-
opy invasion through formation of mycorrhizae
of a type seen in some extant terrestrials that
mobilize nitrogen and phosphorus from sterile,
organic soil (Stribley & Read, 1975; St. John et
al., 1985). Terrestrial Orchidaceae are also
strongly mycorrhizal, but the advantages, if any,
that fungi provide canopy-dependent adults re-
main little studied and controversial (Hadley &
Williamson, 1972; Sanford, 1974; Benzing &
Friedman, 1981). Broad surveys of epiphyte roots
and nitrogen-cycle microbes in canopy substrata
could prove rewarding. A general work-up of tree-
crown media as nutrient sources is much needed.

Mistletoes. Restricted taxonomic participa-
tion in branch parasitism is made all the more
intriguing by the widespread abundance of mis-
tletoes, some of which range farther north than
any vascular epiphyte. Virtually the entire com-
plement—about 1,300 species— belongs to San-
talales. Why many thousands of plants from so
many other higher taxa root nowhere but on bark
yet never invade host vasculature is puzzling.
Occurrence in Santalales of terrestrials with
haustoria, including primitive Loranthaceae, but
no epiphytes suggests that branch parasitism arose
from root parasitism. Perhaps more direct aerial
transitions were precluded by biomechanical
constraints. Mistletoes transpire profusely even
during drought in order to acquire sufficient ni-
trogen from host xylem (Ehleringer et al., 1985).
This is the antithesis of an epiphyte’s usual con-
servative water use pattern. Failure by so many

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

epiphytic lineages to achieve parasite status sim-
ply because invasive organs are difficult to ac-
quire seems unlikely in light of the diverse soil-
based species that tap any nearby roots. More
plausible is the theory that continuous function
during a historical crossover between the hig

solute potential/low maximum turgor/sensitive
stomata pattern of the epiphyte and the three to
five times more concentrated osmotica (Harris
& Lawrence, 1916; Harris, 1918) and correlated
foliar conductance patterns of the parasite would
require buffering, which is less available in can-
opy than in terrestrial habitats. Most of the root

function in the conventional manner, drawing
upon soil moisture and thus reducing the liability
to water loss of transitional forms.

SUMMARY

Epiphytes do not constitute a narrowly defined
group of plants on either taxonomic or functional
grounds, nor is their habitat usually unique in
its important physical characteristics. Growing
conditions in both humid and arid forest cano-
pies overlap with those at ground level, as do
adaptive mechanisms in canopy- and soil-de-
pendent flora. Substrata exhibiting the most
unique and powerful constraints on plant success
occur in the driest exposures. Circumvention of
dependence on soil by epiphytes is varied. Modes
of moisture and nutrient acquisition are the most
unusual, especially among PS forms. Ways in
which resources are conserved and deployed are
often variations on, and perhaps sometimes
identical to, patterns present in terrestrial rela-
tives. No single key feature—like CAM, im-
poundment habit, or animal dispersal—nor
combination of key characters underlies epiphyt-
ism. Solutions are many, a condition favoring
broad taxonomic composition of mechanically-
dependent synusiae. In contrast, failure of a large
tropical group to evolve epiphytic members may
relate to the presence of a few well-entrenched,
maladaptive features. Evolutionary inertia prob-
ably also reflects little-understood genetic con-
straints and functional incompatibilities that
block emergence of enabling syndromes even
though some of their mechanistic components
are already well established.

Cladogenesis and adaptive radiation for life on
varied substrata have been more closely allied
in some epiphytic taxa than in others. Most fam-
ilies, even some containing hundreds of epi-

1987]

phytic species, have entered forest canopies
through a single vegetative theme. Propitious re-
lationships with pollinators, intermediate dis-
turbance, and narrowly circumscribed substra-

Orchidaceae are exceptional for their diverse
vegetative mechanisms. These two families do
well under equable conditions, but they are es-
pecially dominant in stressful environments be-
cause they can maintain enough generative pow-
er to compensate for mortality imposed by habitat
patchiness and disturbance.

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DIVERSITY AND BIOGEOGRAPHY OF NEOTROPICAL
VASCULAR EPIPHYTES!

ALWYN H. GENTRY AND C. H. DopsoN?

In his classic work Schimper (1888), empha-
sizing the taxonomic diversity of epiphytes, list-
ed 33 families and 232 genera of epiphytes. Until
very recently, subsequent authors have generally
accepted Schimper’s figures (Richards, 1957; Jo-
hansson, 1974). However, epiphytism (here used
in a broad sense to include hemiepiphytes, see
Kress, 1986) was recently reported to exist in 65
different vascular plant families (56 families ex-
luding ferns), 38 of these with epiphytes in the
Neotropics (Madison, 1977). Our own data (Ap-
pendix 1) and additional records compiled by
Kress (1986) now record 83 vascular plant fam-
ilies with epiphytic species.* At least 876 genera
include at least one epiphytic species and there
are perhaps 29,000 epiphytic species, ca. 10% of
all vascular plants (Table 1). Thus at first sight
epiphytism seems a very widespread and suc-
cessful life-style, which very many unrelated taxa

ave evolved.

However, a closer examination suggests that
even though there are both many species and
higher taxa of epiphytes, few of the higher taxa
account for most of the species. Burger (1985),
for example, emphasized that relatively few lin-
eages have been able to enter the epiphytic niche,
presumably because of the complex suite of ad-
aptations needed. Thus even though it is true
that the evolution of an epiphytic habit has been
a relatively common feature of vascular plant
evolution, it is equally true that very few of the
taxa that have evolved an epiphytic habit have
radiated successfully to produce other epiphytic
species (Table 2). In most of the epiphyte-con-
taining families, epiphytism is a rather insignif-
icant anomaly. Indeed, eliminating a mere 85
such “oddball” species from the roster of the
world’s epiphytes removes 31 families from the

epiphytic ranks. Only 32 seed plant families have
as many as five or more epiphytic species, 26 of
these with epiphytes in the Neotropics. It is on
the 42 families (Table 3) that contain =
in the Neotropics that this paper will

Even though this analysis of epiphyte posset
and distribution is largely focused on the Neo-
tropics, a few comparisons with the Paleotropics
are instructive. There are actually slightly more
families with epiphytes in the Paleotropics (43)
than in the Neotropics (42), with all of the pa-
leotropical epiphytic families having epiphytic
dipped in Australasia but only 15 in Af-

and Madagascar. If only the 32 seed plant
families with five or more epiphytic species are
considered, there are also roughly equal repre-
sentations of epiphyte-containing families in the
Neotropics (26) and Australasia (25) but only
about half as many in Africa (14).

At the species level the story is very different.
There are many more epiphytes in the Neotrop-
ics, at least half again as many as in Australasia
and six times as many as in Africa. Althoug
similar numbers of genera and families evolved
epiphytism in the different regions, subsequent
speciation as epiphytes was dramatically greater
in the Neotropics. A major objective of this pa-
per, then, will be to try to explain why there is
so much epiphyte diversity in the Neotropics.

EPIPHYTE FAMILIES

One approach to an overview of neotropical
epiphyte diversity is a taxonomic one. Table 3
summarizes the neotropical epiphytic seed plant
flora by family. Of the 42 families represented
by at least one habitually epiphytic species in the
Neotropics, the Orchidaceae are by far the most
important with ten times as many epiphytic neo-

! We thank the National Science Foundation (INT- ps 6840; BSR-834- ei for — of our Ecuadorian

fieldwork. Drs. H. Bedell, T. Croat, B. Hammel, J. Lut

binson provided unpublished data on disteibuHon ‘of epiphytism in bru

r, L. B. Smith, M. Dillon, and
taxonomic specialties. We

n, S. Renner, P. Tay

H. R
especially thank B. Hammel for making available oe data on the habit composition of the La Selva
; B. Burger

r, T. Croat, B. Hammel, M. Madis

cata Botanical Garden, P.O. Box 299, St. L

n, S. er
and E. Zardini for much technical ipe in its prepar
ouis, Missou ri 63166,

and E. Zardini for reviewing the manuscript;

U.S.A.
3 Th set used in this paper was s derived a Su that of Kress (1986) and is mostly based

on Madison’ s figures as modified in Appendix

ANN. Missouri Bot. GARD. 74: 205-233. 1987.

206

BLE |. Taxonomic distribution of vascular epi-
s (modified from Madison, 1977 and Kress, 1986).

Fami- Gen-
lies era
with with Species
i- f Epi-
phytes phytes phytes
Pteridophytes 13 92 2,593
Gymnosperms 2 2 4
Monocots
Without orchids 80 2,657
Orchids (fide Madi-
son) 500 20,000
Orchids (compiled
ressler,
1981) 460 15,000
Orchids (fide Kress,
6 440 13,951
Total (fide Kress,
1986) 17 520 16,608
Total (our estimate) 17 540 22,657'
Dicots 51 262 4,251
Total 83 896 29,505

! Includes Madison’s orchid figure (see text).

tropical species as runners-up Araceae and Bro-
meliaceae. All three of these most speciose neo-
tropical epiphyte families are monocots. One
other monocot family, Cyclanthaceae, also has
a significant number of epiphytes. Commelina-
ceae, Rapateaceae, and Philesiaceae, although
with few species, have an exclusively epiphytic
genus (respectively, Cochliostemon, Epidryos,
Luzuriaga, and (in our experience) Philesia). As
summarized by Madison (1977), the rest of the
epiphytic monocot flora of the Neotropics con-
sists of single species of Burmannia and Yucca
and a few Central American species of Smila-
cina.

Since orchids are so overwhelmingly the most
diverse group of epiphytes (about 7096 of their
species are epiphytic), estimates of orchid diver-
sity are critical to an evaluation of epiphyte di-
versity. Unfortunately, orchids are amazingly
poorly known taxonomically (compare the ca.
12 orchid taxonomists with the ca. 200 system-
atists specializing i in the similar-sized Compos-

ler, 1981) or 17,000 (Airy Shaw, 1973) to 30,000
(Madison, 1977) or 35,000 (“some authors" fide
Dressler, 1981). In the best available review
Dressler (1981) counted almost 20,000 species

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

and suggested 20,000-25,000 as the best esti-
mate of orchid species numbers. Since 7096 of
the total number of orchid species should ap-
proximate the number of species of epiphytic
orchid, there should be between ca. 12,000 (from
Airy Shaw's estimate) and ca. 20,000 (from Mad-
ison's estimate). Unfortunately, the 8,000 species
"slop" between these two estimates is as great as
the total number of epiphytes in all other families
combined! Madison (1977) used the higher figure
for his calculations, whereas Dressler (1981) gen-
erally opted for more conservative estimates of
orchid species numbers. We have mostly taken
the higher values since we know of many cases
where Dressler's figures for species numbers are
significant underestimates but none where he has
overestimated. For example, Dressler suggested
that there are 830 species in subtribe Epiden-
drineae, whereas in Ecuador alone there are 500
species in the single genus Epidendrum and we
think 1,200 species would be a better subtribal
estimate.

Moreover, new orchid species are being dis-
covered at an astonishing rate, especially in the
northern Andean region, again suggesting that
Dressler's estimates of species numbers will have
to be adjusted dramatically upward. For exam-
ple, about 2,315 orchid species are now known
from Ecuador, ca. 700 of these described only in
the last 15 years. Nevertheless, more than 1,500
additional Ecuadorian orchid morphospecies"

ave not been identified with any
Even if the 300 unaccounted for names should
all prove applicable to the unidentified speci-
mens at hand, it is inevitable that most of the
unidentified taxa will prove undescribed and the
list of orchids for Ecuador alone will increase to
well over 3,000 species.

The neotropical epiphytic dicot flora is more
diverse in families but much less diverse in species
than the monocots. Twenty-nine dicot families
have at least one habitually epiphytic species in
the Neotropics. The largest of these are Pipera-
ceae (ca. 500 spp.), Gesneriaceae (483 spp.), Me-
lastomataceae (227 spp.), Ericaceae (ca. 300 spp.),
Cactaceae (133 spp.), Guttiferae (ca. 90 spp.),
and Marcgraviaceae (87 spp.). In addition there
are perhaps 110 neotropical species of Moraceae
stranglers in the genera Ficus and Coussapoa.
The only other dicot families with more than 20
epiphytic species in the Neotropics are Aralia-
ceae, Bignoniaceae, Compositae, Rubiaceae, and
Solanaceae (Table

In addition, Bombacaceae, though with few

1987] GENTRY & DODSON-— NEOTROPICAL VASCULAR EPIPHYTES 207
TABLE 2. Largest epiphyte families (in part from Madison, 1977).
No. Genera
wit No. Epiphytic Percent
Family Epiphytes Species Total No. Species Epiphytes

Orchidaceae 460 20,000 30,000 67

(—13,951') (—19,128') (73)

Bromeliaceae! 26 1,144 ,500 46
raceae 15 1,100? 2,5004? 42
cia 53 1,023 1,100 93
Piperaceae 2 710 3,000 24
Melastomataceae 33 ca. 647? 4,770? 14
Gesneriaceae 28 598* 3,000* 20
Moraceae (incl. stranglers) 3 521 1,400 37
ricace 28 4785 4,000 23
l TN 2 400 600 67
Aspleniaceae! 1 400 675 59
Dryopteridaceae! 10 292 1,920 15
Rubiaceae 21 217 6,000 4
Lycopodiaceae! l 200 400 50
Davalliaceae! 8 139 150 10
Asclepiadaceae 6 135 2,000 T
Cactaceae 25 133 2,000 [Í
Cyclanthaceae 7 125° 205 61
Vittaria 9 112 11 100
Guttiferae 6 92 1,000 9
Marcgraviaceae 7 897 117 76
Araliaceae 5 73 700 10

* Wiehler, 198
5 Luteyn, pers. comm.
* Hammel, pers. comm.

? Bedell, pers. comm.

species, has an epiphytic genus (Spirotheca). The
other 16 epiphytic neotropical dicot families are
represented by only occasional epiphytic species
of predominantly terrestrial genera.

In addition to these angiosperm families, there
are two gymnosperm families with single epi-
phytic neotropical species in generally terrestrial
genera and at least 838 epiphytic fern species
belonging to 32 different genera, a common epi-
phytic Psilotum, and some epiphytic species of
Lycopodium.

REPRODUCTIVE BIOLOGY

A number of salient characteristics that may
be critical to success as epiphytes are shared by
many different neotropical epiphytic taxa. Mad-
ison (1977) nicely summarized many of the fea-
tures of epiphyte reproductive biology and this
discussion is largely based on his. From the view-
point of dispersal biology, there are three main

types of epiphyte propagule. The great majority
of epiphyte genera and species have tiny dust-
like wind-dispersed sporochores, often with
highly sculptured epidermis, to aid in air flota-
tion. Such seeds, representing an extreme in r-se-
lection and a high risk gamble on chance es-
tablishment, are found in the two most successful
epiphyte groups, orchids and ferns, as well as in
such taxa as Begonia (although at least one Af-

perhap
capsular melastomes, although seeds of capsular
melastomes may be | mm long and are not strict-
ly comparable (Renner, pers. comm.). In closed-
canopy tropical forests such seeds are virtually
unique to epiphytes. While tiny sporochore seeds
are found in some tropical weedy herbs, they are
unknown among tropical forest lianas (except
mostly hemiepiphytic Adelobotrys) and trees (ex-
cluding tree ferns), although the pterochore seeds
of genera like Chimarrhis may not be any larger.

208

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

TABLE 3. Epiphytic neotropical seed plant families (in part from Madison, 1977).

Number of
Neotropical
Epiphytic Neotropical Genera
Family Species with Epiphytes Distribution of Epiphytes
Gnetaceae l Gnetum also epiphytic in Malaysia
Zamiaceae l mia Costa iie pM only in
Neotro
Agavaceae l Yucca Mexico; ann in Neotropics
Araceae 1,034 Anthurium, Caladiopsis, Mon- also epiphytic in Africa and
stera, Rhodospatha, Philo- Asia
dendron, Stenospermation,
yngonium
Bromeliaceae 1,144 (fide 18 genera entirely or predomi- only in Neotropics

Burmanniaceae

Commelinaceae
Cyclanthaceae

Dioscoreaceae
Liliaceae

Orchidaceae
Philesiaceae
Rapateaceae
Apocynaceae
Alzateaceae

Araliaceae

Asclepiadaceae
Begoniaceae

Bignoniaceae
Bombacaceae

Burseraceae
Cactaceae

Campanulaceae
Compositae

Crassulaceae
Ericaceae

Gentianaceae
Gesneriaceae

Kress, 1986)

125

4

11,000 (fide
Madison)
3

nantly epiphytic; 5 others
with some epiphytes
Burmannia

Cochliostemon, Campelia
Asplundia, Dicranopygium,
Evodianthus, Ludovia
Sp P Stelestylis,

cocarpus

Smilacina

ca. 80 genera entirely or pre-
dominantly epiphytic
Luzuriaga, Philesia

Epidryos, Stegolepis
Mandevilla

Alzatea
Schefflera, Oreopanax

Cynanchum
Begonia

Schlegelia, Gibsoniothamnus
Spirotheca

Bursera

25 genera entirely or predomi-
nantly epiphytic

Burmeistera

es r
Neomirandea, Pseudogynox-
ys, Sen necio (Pentacalia), Sin-
clairia, Tuberostylis

Echeverria

18 genera entirely or predomi-
nantly epiphytic; several
with epiphytic species
oyria

12 genera entirely or predomi-
nantly epiphytic; 4 other

also epiphytic in New
Gui

uinea
only epiphytic in Neotropics
only in Neotropics

Ecuador

also epiphytic in —
Pacific, and Madagasca

also epiphytic in Africa me
Australasia

also epiphytic in New Zea-
land

only in oo

Costa R

Costa Rica-Colombia

also i in Africa and
Austr

mostly snobi in Malaysia

also epiphytic in Africa and

sia

only in Neotropics

only in Neotropics; mostly
Andean

Costa Rica

Rhipsalis also epiphytic in
Africa and Ceylon

neotropical; mostly Andean

n New Zealand and Mad-
agascar

also few epiphytic in Hima-
layas and Madagascar

also epiphytic in Australasia

South America
also epiphytic in Africa and
Australasia

GENTRY & DODSON —NEOTROPICAL VASCULAR EPIPHYTES 209

1987]
TABLE 3. Continued.
Number
Neotropical
Epiphytic Neotropical Genera
Family Species with Epiphytes Distribution of Epiphytes
genera with some epiphytic
species
Griseliniaceae 3 Griselinia Chile and Brazil; also epi-
phytic in New Zealand
Guttiferae ca. 90 Clusia, Clusiella, Havetiopsis, only epiphytic in Neotropics
edem matopus, Quapoya,
enggeria
Lentibulariaceae 12 Utricularia also 2 epiphytic in Africa
and Australasia
Marcgraviaceae 89 all 7 genera entirely or pre- only in Neotropics
dominantly Pone or
miepiphyt
Melastomataceae 227 7 genera sores or largely epi- also epiphytic in Africa and
hytic; 5 others with some Asia
epiphytic cles
Moraceae 111 Coussapoa, Ficus subg. Uro- Ficus stranglers also in Afri-
stigma, and 1 Pourouma ca and Australasi
Myrsinaceae ca. 12 Cybianthus, Grammadenia, also epiphytic in Africa and
Myrsine (Rapanea ia
Onagraceae 3 Fuchsia only epiphytic in Neotropics
Piperaceae ca. 500 Peperomia, Piper also epiphytic in Africa and
Asia
Rubiaceae ca. 57 Balmea, Coprosma, Cosmi- also epiphytic in Australasia
buena, Hillia, Malanea, Ma-
nettia, Psychotria, fm
Eon nium, Schrader.
apotaceae 1 melia Costa Rica
Saxifragaceae ca. 3 PM Phyllonoma only in Neotropics
olanaceae ca. 30 Juanulloa, Lycianthes, Markea also epiphytic in Malaysia
(+ segregates), Solanum (Solanum, Lycianthes)
Urticaceae ca. 15 Pilea

also epiphytic in Indo-Ma-
laysia

The second most prevalent dispersal mode
among epiphytes is via birds. Most bird-dis-
persed epiphytes have indehiscent berry fruits
but a few, including Drymonia and Clusia, have
dehiscent capsules with arillate seeds. In either
case the seeds tend to be smaller and more nu-
merous than in related nonepiphytic taxa (Mad-
ison, 1977). In some families there is a marked
change in pigs mode accompanying the shift
to epiphytis Bignoniaceae all epiphytic
species Su two probably ogi dispersed excep-
tions) are bird-disperse only one nonepi-
phytic species (Synapsis Mentum: is (see Gentry,
1983). In Melastomataceae 8596 of the epiphytic
species have berry ird as compared with 60%
of the nonepiphytic species (Renner, 1986).
The third major diaspore dispersal syndrome

in epiphytes is wind-dispersal via winged or
plumed seeds (pterochory and pogonochory, re-
spectively). Interestingly, plumed seeds as com-
with winged seeds greatly predominate
among epiphytes, whereas the opposite holds true
for trees and lianas, at least in mature forest
species. Some of the important epiphyte taxa with
pogonochore diaspores are Bromeliaceae
subfamily Tillandsioideae, Asclepiadaceae, Ges-
neriaceae, and Rubiaceae. In Rubiaceae some
epiphytic genera have small winged seeds while
others have true pogonochores; the difference be-
tween these dispersal modes tends to break down
in such groups, with some species having such
narrow reduced n: that these effectively ap-
proximate large hai
Finally there are a m epiphyte taxa with such

210

miscellaneous dispersal syndromes as bat-dis-
ersal (some strangler figs), exozoochory via
sticky diaspores (some Peperomia), and the not
readily classifiable “sloppy corn-on-the-cob"
ingestion of some cyclanth fruits by Callicebus
and other primates (Terborgh, pers. comm.).

In general, epiphyte seeds are smaller and more
numerous than those of nonepiphytic relatives.
For example, Renner (1986 and pers. comm.)
noted that in Melastomataceae mostly epiphytic
Blakea and Topobea have ca. 1,000 seeds per
fruit compared with a few deren seeds per fruit
in typical nonepiphytic genera such as Miconia
and Clidemia. Madison (1977) estimated that
seeds of epiphytic Anthurium are typically ca. 2
mm long as compared with 4-8 mm long in ter-
restrial Anthurium species. There are also excep-
tions to this pattern. For example, Rockwood
(1985) pointed out that in Gesneriaceae epi-
phytic species actually have significantly larger
seeds than do shrubs and herbs. According to
Rockwood's (1985) analysis, epiphyte seed size
tends to be bimodal; those groups with dust-
seeds or other wind-dispersed seeds have the
smallest seeds of any habit type while taxa not
dispersed by wind have seeds averaging larger
than those of herbs, vines, and shrubs, similar
to lianas, and ga smaller than trees. Neverthe-
less, since the great iid of epiphytes are
wind- dipened. pon iphyte habit class as a
whole is generally characterized by the smallest
seeds of any habit class. A dispersal strategy em-
phasizing many small seeds and chance estab-
lishment is typical of the r-selection syndrome
often found in weedy species. Epiphytes would
seem to be most unusual in being r-selected com-
ponents of mature forest ecosystems.

POLLINATION

Madison (1977) emphasized animal-pollina-
tion as a characteristic trait shared by all angio-
sperm epiphytes. While true, this is hardly re-
markable in a tropical context since, with virtually
no exceptions (Myriocarpon (pers. obs.), Trophis
(Bawa et al., 1985), and just possibly Sorocea
and a few Chamaedorea species (fide Bawa et
al, 1985), all lowland tropical forest angio-
sperms are animal-pollinated. Nevertheless, epi-
phytes as a whole surely have a more pronounced
trend toward highly specific and specialized pol-
lination systems than do nonepiphytes, if for
no other reason than that so many epiphytes are
orchids. In addition to the well known orchid

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

pollination specializations (e.g., Dodson, 1967;
van der Pijl & Dodson, 1966; Williams, this
ymposium), many aroids have similar Euglos-
sine-attracting scent systems. Five of the largest
neotropical orchid genera have specific bee pol-
linators attracted by specific scents and Anthur-
ium is the largest nonorchid genus in the
Neotropics. While Pleurothallis, the largest
neotropical epiphyte genus, does not participate
in the presumably speciation-promoting Euglos-
sine-pollination syndrome, it is pollinated by the
large and diverse fly genus Bradesia (CD, pers.
obs.) and similar coevolutionary patterns may
be involved; among other small-flowered and in-
conspicuous but highly diverse orchid genera,
Stelis and Lepanthes are probably pollinated by
Drosophila and similar flies, and Telipogon is
pollinated by pseudocopulation with tachinid
flies, another very large and diversified insect
taxon

an

Unique to the Neotropics, hummingbird-pol-
lination is also much more prevalent among epi-
phytes (and terrestrial herbs) than in trees or free-
climbing lianas. Epiphytic taxa among which
hummingbird- pollination is prevalent include
Bromeliaceae, Gesneriaceae (espe-

um aviaceae (Norantea,
sensu lato), Rubiaceae (Ravnia, Manettia), and
Cactaceae (e.g., Schlumbergera). The correlation
between hummingbird-pollination and epiphy-
tism is well shown by Bignoniaceae. Of the two
epiphytic genera of Bignoniaceae, one (Gibson-
iothamnus) is entirely hummingbird-pollinated
and the other (Schlegelia) also has several hum-
mingbird-pollinated species; hummingbird-pol-
lination is rare elsewhere in the family. Other
specialized pollination systems shown by epi-

l, pe
rat-pollination in B/akea chlorantha (Lumer:
1980). Perhaps more striking than the diversity
of highly specialized pollination systems among
neotropical epiphytes is their lack of the small,
inconspicuous, generalist-pollinated flowers that
characterize the great majority of trees in the wet
forests where epiphytes are prevalent. The only
epiphytic taxa characterized by such flowers are
Araliaceae, Moraceae, Piperaceae, Myrsinaceae,
and Urticac
epiphytic m
cialist pollinators (Stiles, 1981), are taken as an

1987]

example, this pattern may be clearly seen at the
community level: in lowland tropical forests
hummingbird-pollination is almost exclusively
confined to herbs and epiphytes.

Another relevant aspect of epiphyte pollina-
tion biology is that such phenomena as self-com-
patibility and autogamy are apparently much
more prevalent than typical in tropical lowland
taxa. For example seven of seven species of B/a-
kea, Topobaea, and Adelobotrys tested at Mon-
LI Costa Rica were self-compatible (Lu-
mer, 1980; Renner, 1986) vs. 34 of 43 tested
oa melastome species in the Manaus area
(Renner, 1984). In Marcgraviaceae this is carried
to an extreme with all species tested being au-
togamously pollinated in bud despite the elab-
orate floral adaptations (Bedell, pers. comm.).

In summary, epiphyte reproductive biology
appears to be a unique mix of r-selection and
specialization. Unlike other components of ma-
ture forest communities, epiphytes share many
reputedly r-selected traits with weedy herbs, es-
pecially in their dispersal ecology. Yet at the same
time most epiphytes have highly specialized pol-
lination systems, strong niche specificity, and
many other traits more characteristic of k-se-
lected mature forest species.

DISTRIBUTIONAL PATTERNS

To this point, we may conclude that, although

taxa have been more than marginally successful
at speciation and adaptive radiation as epi-
phytes. However, the few taxa that have suc-
cessfully radiated as epiphytes have done so very
prolifically. Even though remarkably similar
numbers of plant families have achieved epi-
phytism in the Neotropics and Paleotropics, the
process of epiphyte speciation would seem to
have been much accelerated in the former, to
judge from the very many more neotropical
species of epiphytes.

w uld now like to examine some trends
In epiphyte distribution that may help to under-
stand not only neotropical epiphyte biogeogra-
phy but also some of the continental differences
in epiphyte occurrence.

MOISTURE

One of the most striking distributional pat-
terns shown by epiphytes is a tremendous de-
crease in both numbers of species and individ-

GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES

211

uals in drier habitats. Although this pattern seems
obvious, it is by no means well-documented. In-
deed Walter (1985: 57) claimed that epiphytes,
contrary to such other habit groups as lianas, are
found in dry as well as wet tropical forests. At
the other extreme, Schimper (1903) suggested
that in areas with marked dry seasons epiphytes
are either completely wanting or rare and that
presence of epiphytes outside the rain forest is
always a sign that the dry season is not long or
is accompanied by copious dew. Data for 1,000
m? samples of western Ecuadorian dry forest
(Capeira, 804 mm per year) and moist forest
(Jauneche, 1,855 mm) quantify the extent of this
difference (Gentry & Dodson, 1987; Table 4, Fig.
1). If our results are indicative, most plants in a
wet forest are epiphytes. At Rio Palenque such
a sample included 4,517 epiphytic plants rep-
resenting 63% of all individuals sampled. At
Capeira a mere ten epiphytic plants were includ-
ed in a similar sample of dry forest, representing
0.2% of the sampled individuals. The moist for-
est Jauneche site was intermediate with 116 epi-
phytes constituting 4% of the individual plants
sampled. The difference in epiphyte density be-
tween wet and dry forest is almost 500 fold.

Moreover the decrease of epiphyte density in
dry forest contrasts greatly with the situation for
other habit groups. The number of herbs more
than doubles from our moist and wet forest sam-
ples to our dry one. Contrary to Walter’s asser-
tion (1985: 57), lianas double from wet to dry
forest; they are much commoner yet in our moist
forest sample, the latter presumably atypical since
Jauneche happens to be the most liana-rich site
in the Neotropics (of 45 similar samples). Shrub
density also increases somewhat from wet to dry
forest; unlike lianas, shrubs are only about half
as abundant in the intermediate moist forest as
in dry and wet forest. In contrast, the number of
individual trees = 10 cm DBH, and thus the ap-
parent density of the forest, changed little (52,
64, and 69 trees 210 cm DBH in wet, moist,
and dry forest, respectively).

Epiphytes are also important contributors to
the species richness of neotropical wet forests.
Indeed there are 35 epiphyte species in our 0.1
ha. wet forest sample, accounting for over a third
of the sampled species. This compares with only
13 epiphytes (8% of the species) in the compa-
rable moist forest sample and three (2% of the
species) in the dry forest one (Fig. 2)

The importance of the epiphytic contribution
to species diversity is equally apparent when en-

212 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

TABLE 4. Number of species and individuals of different habit types in 1,000 m? samples of three forests in
western Ecuador. Río Palenque is wet forest, Jauneche is moist forest, Capeira is dry forest (from Gentry &

Dodson, 1987).

Río Palenque Jauneche Capeira
No. No. No. No. No. No.
Habit Group Spp. Ind. % Spp. % Ind. % Sp. 96 Ind. 9
Herbs (incl. palmettos) 50 14 1,220 17 18 11 944 34 50 29 2,854 53
Shrubs 39 11 531 7 16 10 279 10 13 8 742 14
Epiphytes (incl. hemi-
epiphytes) 127 35 4,517 63 13 8 116 4 3 2 10 52
Climbers (incl. lianas;
excl. hemiepiphytes) 36 10 117 2 58 34 484 17 58 34 895 16
Lianas 22.5 cm (excl.
herbaceous + hemi-
epiphyt.) 12 3 28 43 25 124 4 19 11 58 l
Total tree spp. (incl. Ju-
ven.) 114 3l 653' 9 64 38 960 34 48 28 927 17
Trees <2.5 cm DBH (=
saplings + seedlings) 87 24 559! 8 48 28 672 24 38 22 750 14
Trees <10 cm, >2.5 cm 86 24 217 3 35 21 245 9 32 18 108 2
Trees 210 cm DBH 32 9 52 1 30 18 64 2 29 17 69 l
Total herbs (epiphytes +
herbs—woody cpi-
162 5,525 31 1,060 53 2,864
Shrub layer (shrubs +
saplings) 126 1,090 65 948 51 1,492
Underlayers (< ca. 3 m)
(herbs + shrubs +
saplings) 176 2,310 83 1,892 101 4,346
Total 365 7,210! 169 2,783 173 5,428

! Excluding dense patch of 123 Quararibea asterolepis seedlings.

tire florulas are compared (Table 5, Fig. 3). AI-
most one fourth of all the Río Palenque plant
species are epiphytes (Dodson & Gentry, 1978).
Similarly, in another lowland wet forest at La
sea Costa Rica, dh of the species are epi-
phytes (Hammel, pers. com ven in moist
ed sites like Barro Cian ‘land: Panama
(Croat, 1978) and Jauneche, Ecuador (Dodson
et al., 1985), epiphytes constitute 12-1696 of the
total flora. Only in dry forests are epiphytes rel-
atively insignificant, accounting for 2-4% of the
species of Capeira (Dodson & Gentry, 1987) and
Santa Rosa National Park, Costa Rica (Janzen
& Liesner, 1980).

We conclude, contrary to Walter (1985), that
epiphytes d tically in drier areas
than does any other habit group, but contrary to
Schimper’s (1903: 198) emphasis, a few vascular
epiphytes are characteristically present in even
the driest neotropical forests (e.g., Capeira with
804 mm of annual precipitation).

Familial makeup of the epiphytic flora also

changes with precipitation (Table 6). Many more
families have epiphytic representatives in wet
forests than in drier ones (Fig. 4), and many epi-
phytic taxa are confined entirely to wetter forests.
In the Neotropics the same families tend to be
represented by epiphytes under similar climatic
conditions. In the driest forests, the only epi-
phytes are orchids and bromeliads, perhaps the
two most specialized epiphytic families. Ferns,
peperomias, and Cactaceae join orchids and bro-
meliads in slightly moister conditions. The next
epiphytic families to appear with increasing hu-
midity are aroids, Moraceae (stranglers), and
Gesneriaceae, joining representatives of all the
dry forest families, each of whose number of epi-
phytic species is maintained or increased. The
two local florula sites with over 2,500 mm of
precipitation, Barro Colorado Island (Croat,
1978) and Rio Palenque (Dodson &
1978), me remarkably similar epiphytic floras.

he same seven epiphytic families are most
species rich at both sites, in roughly the same

1987] GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 213

100 9
80 4
EEB epiphytes
4 WET L* trees
63 4
DRY L herbs
60 1
53 * shrubs
°
2) 504 o? climbers
= °
ç °
EJ
=
>
Es] MOIST
£ 40 1
34 34
4
4 e
e
ra
16 pr
20 4 17 4
€
>
4 4
Pa <
1 »
4 4
9 4 |",
P. e
Pa P.
Capeira Jauneche Río Palenque

FIGURE 1. Percent of individual plants belonging to different habit groups in 1,000 m? samples of three
western Ecuadorian forests.

50 :
ENEB epiphytes
DRY MOIST WET
»
Bo trees
4
40 - n
r3 35 CJ herbs
4 4
3 io * shrubs
a »
°
M Ta & 31 Ó .
o 29 ° G ~ ° š climbers
° 304 28 o? « < °
a o » P
° < <
[7] e o » »
» o 4 25 4
= < ° > >
2 > o 4 4
4 o » o P”)
> o 4 [e] 4
4 ° P ° P.
4 96 > $9 »
204 » o < ° <
< ° » o »
» ° < ° <
4 ° » o 2
» ° < o
4 ° » ° >
> o 4 ° <
4 ° P ° P
P o 4 [e] 4
4 ° > ° >
> ° < ° pi
10 4 Ç o? e ao <
4 o » ° "i
» o 8 4 o
4 8 ° > ° Pa
> o 4 o
4 ° > o 2
o * 1°, »* Ta »
2 1" * bar >. 96 >
i * lo < ° 4
Capeira Jauneche Río Palenque

FIGURE 2. Percent of species belonging to different habit groups in 1,000 m? samples of three western
Ecuadorian forests.

214

MOIST

saprophytes

shrubs parasites &

treelets

&

of species
herbs,

%

epiphytes climbers trees

SANTA
ROSA

CAPEIRA JAUNECHE B.C.. PALENQUE SELVA

FiGURE 3. Percent of species in local florulas be-
Peper to different habit groups.

order, just as they are in Hammel’s unpublished
La Selva, Costa Rica, species list. Remarkably,
there are 81 epiphytic species of orchids at Río:
Palenque and 82 at Barro Colorado Island (BCI),
35 epiphytic aroids at Rio Palenque vs. 30 at
BCI, 18 species of epiphytic Bromeliaceae, and
14 species of epiphytic or strangler Moraceae at
both sites. Only epiphytic ferns and fern allies
are noticeably better represented at BCI than at
Rio Palenque and only epiphytic Gesneriaceae
and Cy the revers
sites have three epiphytic cacti and one epiphytic
melastome plus one or two epiphytic Araliaceae,
Solanaceae, and begonias. Only Rubiaceae and
Saxifragaceae have epiphytic members at BCI

ut not Rio Palenque; only Urticaceae, Pole-
moniaceae, Bignoniaceae, and Ericaceae have

TABLE 5.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

epiphytic representatives in the Rio Palenque
flora but not the Barro Colorado one.

Again, the same epiphyte families predomi-
nate at other wet sites in western Ecuador (Table
7). The exact same seven most species rich fam-
ilies at BCI, La Selva, and Rio Palenque are the
most species rich at Centinela, again in roughly
the same order. Even at Tenafuerste (alt. 1,000
m) the five families richest in epiphytes are ex-
actly the same ones that are most species rich at
the other Ecuadorian wet forest sites.

At the extreme wet end of the precipitation
gradient there are other, as yet unquantified,
changes in the epiphytic flora. In the wetter part
of Chocó (precipitation > 8,000 mm) berry-
fruited epiphytes like Melastomataceae, Ara-
ceae, Marcgraviaceae, and Ericaceae, along with
arillate-seeded Guttiferae, appear to predomi-
nate to a much greater degree than at Rio Pa-
lenque, while ferns (Sota, 1972) and dust-seeded
orchids are more t poony fepresenieo. Possibly

to establish themselves in the face of such su-
aasan wa rainfa

here m a aodceable habit change in the
pou. d of the wettest sites. Most of the
predominant Chocó epiphytic families are ac-
tually hemiepiphytic, many of them woody as
well. In the same sites with abundant hemiepi-
phytes, free-climbing lianas become noticeably
less prevalent. While the average of climbers
> 2.5 cm in diameter for 0.1 ha. samples at two
pluvial forest sites in the Colombian Chocó was
exactly the same (68) as that for a series of 20
similar samples from neotropical lowland moist
and wet forests, half the sampled pluvial forest
climbers were hemiepiphytic vs. an average of

Habit compositions of complete local florulas. Capeira, Ecuador and Santa Rosa National Park,

Costa Rica, are dry forest; Jauneche, Ecuador and Barro Colorado Island, Panama, are moist forest; Río Palenque,

Ecuador, is wet forest.

anta Barro
Capeira Rosa Jauneche Colorado Palenque
Habit Category No. 96 No. 96 No. % No. % No. %
Trees => 10 cm DBH 69 15 141 21 112 18 290 22 165 16
Small trees + large shrubs 28 6 64 10 60 10 151 11 99 9
Herbs 4 subshrubs 242 52 317 48 224 37 389 30 376 36
Epiphytes (including Pis dE 9 2 24 4 72 12 216 16 238 23
Parasites and saprophyt 4 l 6 l 4 l 12 l 6 l
Lianas 46 10 52 8 81 13 149 11 87 8
Small vines 66 14 63 9 55 9 109 8 84 8
Total species 464 667 608 1,316 1,055

1987]

GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES

215

TABLE 6. Familial composition of epiphyte floras in lowland forests with different precipitations.

Santa
La Selva, arro Jauneche, Rosa
Costa pios bonia Ecuador* Costa Capeira, Makokou,
Rica! or? 1,855 Rica? cuador* Gabon’
Family 4,000 mm 2,980 mm Py 750 m mm mm 1,550 mm 804 mm 1,755 mm
Orchidaceae 109 81 82 33 8 5 21
76 35 30 10 — — 10
Ferns and allies 59 28 43 5 7 — 26
Piperaceae 12 19 10 4 l l —
Bromeliaceae 29 18 18 6 3 2
eae 13 14 14 9 2 — 7+
Gesneriac 16 12 4 2 —
Cyclanthaceae ll 8 1 — — —
Marcgraviaceae 8 5 2 — — — —
Guttiferae 11 4 2 — — — —
Cactaceae 6 3 3 3 3 l l
ace 2 3 — — — — —
Araliaceae 2 2 l — — — l
Bignoniacea 3 2 — — — — —
Melastomataceae 2 l l — — — —
Polemoniaceae — l — — — — —
Solanacea 1 l 2 — — — —
Urticaceae l l — — — — —
Begoniaceae 2 l l — — — —
laceae 4 — 1 — — — —
Saxifragaceae — — l — — — —
mmelinaceae l — — — — —
Total epiphytes 368 238 216 72 24 9 66+
Percent of flora 24 23 16 12 4 2 6+
No. families with epiphytes 20 18 17 8 6 4 6

Ee Hammel, pers. comm.
pos 4 Gentry, 1978.
978.

ds
7 Hladik & ain in prep.; Florence & Hladik, 1980 and cited references.

2.71 l limbers sampled at the moist
dub wet sits (Gentry, 1986). in a sense, hemi-
epiphytic climbers seem somehow to replace free-
climbing lianas in the wettest lowland forests and
also in middle elevation cloud forests.

ALTITUDINAL

The epiphytic flora also changes in both di-
versity and composition on an altitudinal gra-
dient. The general tendency is for epiphytes to
be better represented in intermediate elevation
cloud forests. In the Andes the peak in epiphyte
diversity appears to be between 1,000 m and
2,000 m, but it lies somewhat lower in Costa
Rica and Panama. Few data are available, but a
comparison of incomplete data sets for several

Ecuadorian sites (Table 7) documents this trend
for the lower part of the gradient. The sites for
which relevant data are available are Centinela
(600 m) and Tenafuerste (1,000 m), both on the
western slopes of the Central Ecuadorian Andes,
and Mera (1,000 m) on the eastern slope. Of the
well-documented sites, Centinela has the most
species of epiphytes, 337, or 3596 of the flora.
This compares with 238 epiphyte species ac-
counting for 2396 of the flora at nearby Rí

lenque (alt. 200 m). The data for the two 1 000
m sites are less complete, with many species re-
maining to be discovered. Tenafuerste, with much
less cloud forest effect than Centinela or Mera,
has the poorest epiphyte representation, only 31%
of the flora. The extreme is pluvial Mera where
few collections have been made as yet. Three

NUMBER OF EPIPHYTE FAMILIES AND SPECIES VS. PRECIPITATION

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

N
" bs
o >
o a
o a
à 400; 120 o
- £
= @ number of families E
a * number of species š u
a P o a
o o =
300 115 E
= g 3
o ] 5
D [ " 5
n £
E ra 5
E : ° š
2oo 8 = fo 5
7 ° c z
is] 2
' S `
. - n
m °
° °
[6] —
à 2 a
100} o e 45
, E:
m
E
1000 2000 3000 4000
Annual precipitation (mm)
FIGURE 4. Increase in numbers of epiphyte species and families in local florulas as a function of precipitation.

hundred twenty-two orchid species are already
known and we expect that there may be as many
as 200 additional epiphytes as well.

In contrast, Peru’s Huascaran National Park,
3,500-5,000 m in altitude, has only seven epi-
phytic species, constituting a mere 196 of the
park's flora (Smith, pers. comm.).

The differences between Mera and Tenafuerste
point out that altitudinal and moisture effects on
epiphytes are complexly interrelated. Similarly
Gilmartin (1973) showed that the 17 species of
Bromeliaceae that occur on both sides of the Ec-
uadorian Andes occur at lower altitudes on the
moister eastern slopes.

Density of epiphytes, although also greatest at
intermediate altitudes, does not closely parallel
diversity; although unquantified, we feel that epi-
phyte density in the Andes tends to be greatest
around 2,000-2,500 m, again occurring at some-
what lower elevations in Costa Rica and Pana-
ma. Due to the high densities, epiphytes are often
most conspicuous at these relatively high alti-
tudes, even though relatively few species may be
present. In middle elevation cloud forests, epi-
phytes may make up as much as 30% of the foliar
biomass and 4596 of the foliar mineral capital of
a forest (Nadkarni, 1984).

Contrary to what happens along the moisture

gradient, familial composition of the epiphyte
flora changes very little along an altitudinal gra-
dient, or at least on that part of it for which we
have data. The same families are important in
roughly the same order. The seven families with
the most epiphyte species at Centinela (600 m)
are the same seven that have the most species at
Río Palenque (200 m). The five families with the
most species at Tenafuerste (1,000 m) are the
same five that have the most species at Río Pa-
lenque and Centinela. The most noteworthy dif-
ference between these sites is the absence of Mo-
raceae stranglers at Tenafuerste; Gesneriaceae are
also conspicuously less diverse at the 1,000 m

more epiphyte diversity at intermediate altitudes
(four spp. at Centinela and two at Tenafuerste
but only one at Río Palenque; see also Renner,
1986). A few families (e.g., Bignoniaceae) dis-
appear from the middle elevation epiphyte flora
and most other families have decreasing num-
bers of epiphytes at higher elevations.

Our only high altitude data set is for Huas-
carán National Park, Peru (Smith, pers. comm.),
where the seven epiphyte species, all restricted
to the lower part of the park between 3,500 and

1987]

4,000 m, belong to four families. The four fam-
ilies with epiphytes— Piperaceae, Bromeliaceae,
Orchidaceae, and ferns—are all in the top five
epiphytic families in the wet Ecuadorian sites.
Of the usually prevalent epiphyte families, only
Araceae is lacking. Perhaps more interesting, the
epiphyte families at Huascarán are exactly the
same ones that are represented at Santa Rosa,
Costa Rica, except that Cactaceae is missing. Ap-
parently at environmental extremes, either alti-
tudinal or VOULU only these same fam-
ilies that are otherw most successful as
epiphytes are able to survive.

The very interesting but controversial sugges-
tion has been made that in the tropics diversity
is generally greatest at middle elevations along
an altitudinal gradient. This has been shown for
ed Bay herps (Scott, 1976), insects (Janzen,

; Janzen et al., 1976), and suggested for
T Greater equability is a likely controlling
factor for this putative ‘“‘mid-elevation bulge” in
species diversity. However, data for 0.1 ha. sam-
ples of plants 22.5 cm DBH suggest that plant
species diversity decreases more or less uniform-
ly from the most diverse lowland wet forest sites
to the least diverse high altitude ones (Gentry,
1982a, 1987c). If a mid-altitude bulge in plant
species richness really does occur, it must be due
largely to epiphytes. Unfortunately our data sets
from middle and upper elevation forests are too
incomplete to be definitive. Indeed one of us
(CD) thinks that because of the increase in epi-
phytes there are more plant species at middle
elevations than in lowland tropical forest while
one of us (AG) thinks that the decrease in species
numbers of such other habit groups as lianas and
trees with altitude outweighs the increased num-
ber of epiphytes. In either case the role of epi-
phytes in the plant community is presumably
greatest in middle elevation forests.

SOIL FERTILITY

To our knowledge no attempt to relate epi-
phyte diversity to soil fertility has been made
previously. Indeed one might suppose that since
epiphytes are intrinsically “insulated” from di-
rect dependence on soil nutrients they would be
relatively unaffected by changes in soil fertility.
For example, Janzen (1974a) discussed the sym-
L'a: 14° LC hat t » | i. A

nce

ants in low-diversity poor soil “kerangas” hab-
itats in Borneo, with the implication that epi-
phytes are unukialiy well represented in such

GENTRY & DODSON — NEOTROPICAL VASCULAR EPIPHYTES

217

TABLE 7. Familial composition of epiphyte floras
in wet forests at different altitudes in Ecuador

Rio Pa- Centi- e
lenque nela fuerste

Family 200 m 600 1,000
Orchidaceae 81 133 68
Araceae 35 52 26
Ferns 28 38 28
Piperaceae 19 19 11
Bromeliaceae 18 23 18
M eae 14 10 —
Ges ace 12 16 8
Cyclanthaceae 8 5 3
Marcgraviaceae 3 2
Guttiferae 4 9 3
Cactaceae 3 2 l
Ericaceae 3 9 9
cto 2 4 —
Bignon 2 2 —
Melastomataceae 1 4 2
Polemoniaceae 1 l —
Solanacea 1 2 1
Urticaceae l 1 l
Acanthaceae — 1 =
Rubiace — l —
Total 238 337 181
Percent of flora 23 35 31

habitats. Whitmore (1984) also emphasized the
frequency of epiphytes in these forests.

It is increasingly well-documented that major
changes in the diversity and floristic composition
of other components of tropical plant commu-
nities are associated with changes in soil fertility
(e.g., Ashton, 1976, 1977, 1978; Huston, 1979,
1980; Gentry, 1987b; Gentry & Emmons, 1987).
For example, there are generally fewer tree, liana,
and terrestrial herb species in neotropical forests
on poorer soils (Gentry, 1981; Gentry
mons, 1987). We have few data with which to
relate epiphyte community composition to soil
fertility. One of us (AG) has compiled species
lists for a series of sites on different substrates in
the Iquitos, Peru, area which share a similar rain-
fall and climatic regime. Of these, the site with
the poorest soil (Mishana, on almost pure white
sand) has the fewest epiphytes (31 epiphyte
species plus a few “‘indets.”’ in a relatively inten-
sively inventoried area vs. 38 identified and many
unidentified at less intensively studied better-soil
Yanamono), suggesting that epiphyte diversity
varies with soil fertility as does the diversity of
other habit groups. However, to date the sam-

218

pling of epiphytes at these sites has been much
less intensive than at our Ecuadorian local florula
sites and is probably too haphazard and incom-
plete to make these data very meaningful.

Data are also available for understory com-
position and levels of flowering and fruiting for
a broad array of neotropical (and paleotropical)
sites (Gentry & Emmons, 1987). If the data for
epiphytes are extracted from that data set, a very
strong reduction in numbers of fertile species of
understory epiphytes on poorer soils is apparent,
paralleling the overall trend of decreased num-
bers of flowering and fruiting understory Species

loss of soil fertility: as soil fertility decreases,
terrestrial herbs, epiphytes, understory shrubs,
and lianas disappear from the understory in that
sequence, leaving virtually only tree saplings and
seedlings in the most severely stressed forests
(Gentry & Emmon 7).

Anecdotal ae also indicates that epi-
phytes are much less diverse and abundant on
poor soils. We have observed many fewer epi-
phytes in poor soil parts of Central Amazonian
Brazil, southern Venezuela, and elsewhere in the
Guiana shield area than in parts of the Neotrop-
ics with richer soils. Large-scale biogeographical
analysis also indicates that epiphytic taxa are
poorly represented in these areas compared with
richer soil areas nearer the Andes and in Central
America (Gentry, 1982b). In a somewhat differ-
ent context, Janzen (1977) has suggested that the
paucity of epiphytes in dipterocarp forests results
from the generally nutrient-poor Southeast Asian
soils. Among the evidence for this hypothesis
cited by Janzen (1977) is the observation that
trees cultivated in Malesia along roads, where
dust stirred up by passing vehicles increases the
nutrients — to oie soe eh in an LEE
unusually poor-s large
epiphyte loads compared with the native eee

On balance it seems clear that the epiphytic
plant community is very sensitive to soil fertility,
with fewer epiphytes and fewer epiphytic species
in forests on poorer soils. Indeed epiphyte di-
versity may be even more sensitive to change in
soil fertility than is tree or liana diversity, a sug-
gestion that would accord with the idea (Gentry
& Emmons, 1987) that plants (presumably in-
cluding epiphytes despite their lack of direct con-
tact with the soil) that are barely able to eke out
a marginal existence should be more susceptible

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

to the effects of relatively slight decreases in en-
vironmental favorability.

LATITUDE

It is well known that the presence of vascular
epiphytes is a characteristic of tropical forests as
compared with temperate ones. We know of only
four vascular epiphytes that occur north of Flor-
ida in the temperate United States— Tillandsia
usneoides, T. recurvata (only in southernmost
Arizona), Epidendrum conopseum, and Polypo-
dium polypodioides. Even Tillandsia usneoides,
the northernmost vascular epiphyte, does not
reach the Mason-Dixon line. The only continen-
tal United States epiphyte species that is not
widespread in the tropics is Tillandsia simulata
Sm., endemic to Central Florida but sometimes
lümped with 7. bartramii Ell. (The Mexican range
of Epidendrum conopseum is also somewhat lim-
ited.) Even in very wet areas that would be full
of epiphytes in the tropics, only lower plants have
adopted the epiphytic habit. In intermediate sub-
tropical areas a gradient of increasing epiphytism
at lower altitudes is evident. The decrease in vas-
cular epiphytes with increasing latitude can be
clearly seen in Florida where oe South
Florida (latitude 25°N) has 46+ (= 2.8%) epi-
phytic Pup (Long & PEA 1971), Central
Florida 1 (= 1.9%) (Wunderlin, 1982), and
Florida E d State Park in northern Florida
(30*50'N latitude) has only two (= 0.4%), Til-
landsia usneoides and Polypodium polypodioides
(Mitchell, 1963).

Curiously, the decrease in vascular epiphytes
with increasing latitude is not symmetrical on
both sides of the equator. A number of epiphytic
species and even a few endemic genera of epi-
phytes occur in south temperate forests. Endemic
temperate South American epiphytic genera in-
clude the monotypic fern Synammia, the mono-
typic cactus Pfeiffera, three monotypic Gesne-
riaceae genera (Asteranthera, Sarmienta, and
facultatively epiphytic Mitraria), and the Lili-
aceae (or Philesiaceae) Luzuriaga and Philesia.
In the Australasian region, New Zealand is es-
pecially noteworthy for its autochthonous epi-
phytes including genera like the monotypic fern
Anarthropteris, the liliaceous Collospermum (also
reaching Fiji and Samoa), the only epiphytic
species of families like Cunoniaceae (with two
different genera having epiphytic members), and
genera like Microlaena (Gramineae) and Metro-
sideros (Myrtaceae). There is even a largely epi-
phytic south temperate family shared by New

1987]

Zealand and Southern Argentina—Chile—Grise-
liniaceae (sometimes included in Cornaceae). In
the north the only noteworthy temperate epi-
phytes are in the Himalayas where aberrant epi-
phytic species of otherwise terrestrial genera like
Ilex, Tripogon, Euonymous, Sedum, and Tha-
lictrum occur. This latitudinal asymmetry was
already noted by Schimper (1903), who pointed
out that north temperate epiphytes are merely
range extensions of widespread tropical species,
whereas many unusual and distinctive epiphytic
taxa occur in the South Temperate region.

At the community level the same trend is ap-
parent. For example, Parque Nacional El Rey in
Argentina, at 24°45’S latitude, has a species list
(L. Malmierca, pers. comm.) of well over 500
vascular plants including 47 species of epiphytes:
20 ferns, four orchids, three species of Rhipsalis
and Peperomia, and no fewer than 17 bromeliads
including 14 tillandsias. In contrast, Florida Cav-
erns ih rig at 30?50'N latitude has only two
epiphy its similar-sized flora of 485 native
species (Michel 1963). Even in rather dry south
temperate vegetations vascular epiphytes can be
extremely prevalent, a situation apparently with-
out parallel in the North Temperate region. For
example, in the Valley of Lerma near Salta, Ar-
gentina (1,200 m, ca. 700 mm ppt.), there are at
least 14 angiosperm epiphytes including at least
ten Tillandsia species in a flora of over 750 species
(Novara, 1984).

Farther south in the Valdivian region of Chile
vascular epiphytes, mostly belonging to endemic
genera, are conspicuous elements of local floras,
ranging from six species (396 of the native flora)
at relatively dry Parque Nacional Tolhuaca
(38?15'S) (Ramírez, 1978) to 17 species (17% of
the native flora) at very wet Fundo de San Martín
(39°30'S) (Cárdenas, 1976; Riveros & Ramirez,
1978); even at 41°S there are 15 vascular epi-
phyte species in Puyehue National Park (Munoz,
1980). New Zealand forests have even more epi-
phytes than the Chilean ones; even well south of
40°S, about 30 vascular epiphytes are typically
included on local species lists (Dawson, 1980).
However, at comparable latitudes in North
America there are no vascular epiphytes

perate forests is unclear but presumably relates
to the relatively mesic, more or less oceanic cli-
mates that prevail in the Southern Hemisphere.
There are more epiphytes at 25°S (47 epiphytes
constituting ca. 8.5% of the flora of Parque El

GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES

219

Rey, L. Malmierca, pers. comm.) as compared
with 25°N (46 epiphytes ~~ ca. 2.8% of
the South Florida flora, L & Lakela, 1971).
At least in South America, fi prevalence of many
species of the genus Tillandsia in southern forests
(e.g., 14 at Parque El Rey), perhaps due purely
to biohistorical reasons, is another important
factor. A similar pattern occurs with lianas, which
are better represented in New Zealand tha
north temperate forests (Dawson, 1980).

CONTINENTAL TRENDS

There are several conspicuous differences be-
tween the epiphytic floras of different continents.
Obviously, predominantly extratropical conti-
nents u TN yaaa p However,

thin the trop-
ics between the Neotropics, tropical Africa, and
tropical Australasia. The African epiphytic flora
has been widely noted to be very impoverished
compared with the other two regions, presum-
ably reflecting a loss of mesic-adapted species
during the dry periods associated with the Pleis-
tocene glacial advances at higher latitudes (Rich-
ards, 1973; Madison, 1977). According to Mad-
ison there are only ca. 2,400 epiphytic species in
Africa, less than a sixth as many as in the Neo-
tropics and a quarter as many as in tropical Aus-
tralasia. Even though several families and genera
with epiphytes in Madagascar were omitted from
Madison’s (1977) epiphyte summary (see Ap-
pendix), their inclusion does not appreciably in-
crease the number of African epiphyte species.

Curiously, the depauperate nature of the Af-
rican epiphytic flora is not obvious at the com-
munity level. For example, the 59 epiphyte
species at Makokou, Gabon, constitute 5% of the
total Makokou flora (Table 8; compiled from
Hladik & Gentry, in prep.; Florence & Hladik,
1980, and included references). While 5% of a
moist forest flora might seem fewer epiphytes
than would be expected in the Neotropics, Ma-
kokou is quite dry, with only 1,785 mm ofannual
rainfall, and its 66 (6%) epiphytic species (in-
cluding stranglers) are quite in line with the 7
(12%) epiphytic species at Jauneche and 24 (4%)
at Santa Rosa (Table 9). Johansson (1974) stud-
ied a relatively moist region in the Nimba moun-
tains of northern Liberia and reported 153 vas-
cular epiphyte species (excluding six filmy ferns
and 23 “facultative” epiphytes) in his study area,
up to 44 species in a single 750 m plot, and up
to 22 species on a single tree. In even wetter areas

220 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74
TABLE 8. Habit distributions of Makokou, Gabon plant species.
Gymno-
Habit Ferns sperms Monocots Dicots Total
Epiphytes 26 0 31 2 59
Parasites + saprophytes 0 0 3 6 9
Climbers l 2 8 248 259
Trees 0 0 5 390 395
Herbs, shrubs, and treelets 42 0 125 251 418
Total species 69 2 172 897 1,140

like southwestern Cameroon epiphytes are al-
most as prevalent as ls many similar areas of the
Neotropics (AG, p obs.).

That tropical Heide: is also floristically
impoverished with respect to the Neotropics has
only recently been realized (Raven, 1976; Gen-
try, 1982b). Epiphytes account for much of the
overall difference between the two regions with
half again as many epiphytes in the Neotropics
as in Australasia (15,500 vs. 10,200) according
to Madison's (1977) figures. Moreover, at least
in those lowland Asian forests we have visited,
there also seem to be many fewer epiphytic in-
dividuals than in comparable neotropical forests.
Richards's (1936) remark is typical: “One of the
most striking features of the Sarawak rain forest,
especially when compared with that of tropical
South America, is the poverty of the epiphytic
vegetation both in species and individuals."
Madison suggested that the fewer tropical Asian

phytes might stem simply from lack of the
extensive loud forest habitats of the Neotropics,

a
1977) emphasized that m
tropical Asian forests tend to have relatively nu-
trient-poor soils, and, if so, our suggestion of a
positive correlation between epiphytes and soil
fertility might help explain the relatively low epi-

phyte diversity and biomass in tropical Asia.
Another important continental level difference
in epiphytes is taxonomic. Orchids and ferns are
the predominant vascular epiphytes nearly
everywhere, but the other elements of the epi-
phytic flora are often very different on different
continents (Table 10). Johansson (1974) gener-
alized that the African epiphytic flora is made
up almost entirely of pteridophytes and orchids,
whereas these groups are joined by bromeliads
and Cactaceae as important epiphytic taxa in

in Australasia and 15 with epiphytes only in the

TABLE 9. Representation of different habits in local florulas.

Barr
Santa Jau- Colo- Río Pa-
Capeira osa eche rad lenque Selva! Makokou

Habit No. % No. % No. 96 No. % No. % No. % No. 9
Epiphyte (incl.

stranglers) 8 2 19 3 72 12 216 16 238 23 368 25 66+ 6+
Parasites +

saprophytes 4 l 6 l 4 l 12 1 6 | 8 1l 9 l
Cli s 112 24 115 18 136 22 258 20 171 16 182 12 259 23
Trees >10 cm

DBH 69 15 142 21 112 19 290 22 165 16 310 21 389 34
Terrestrial

herbs,

shrubs, tree-

lets 270 S8 381 S8 280 47 540 41 475 45 622 42 418 37
Total species 463 667 604 1,316 1,055 1,490 1,140

' Data from B. Hammel (pers. comm.).

1987]

GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES

TABLE 10. Taxonomic distribution of epiphytic taxa in some lowland florulas.

Ferns Monocots Dicots Total
No. No. No No. No. No.
Site Spp. 96 Fam. Spp. % Fam Spp. 96 Spp. 96

Neotropics

Capeira, Ecu — — 2 7 88 2 2 22 9 2

Santa Rosa, C.R 7 29 2 11 46 3 6 25 24 4

Jauneche, Ecu. 5 7 3 49 68 4 18 25 72 12

Barro Colorado, Pan. 43 20 4 131 61 12 42 19 216 16

Rio Palenque, Ecu. 28 12 4 142 60 14 69 29 238 23

La Selva, C.R 59 16 5 226 61 15 83 23 368 25
Africa

Makokou, Gabon 26 44 2 31 53 3 9+ 3 66+ 6+
Asia

Flora of Java 200 24 3 520 63 ll 109 13 829 —

Neotropics (in the case of Campanulaceae there
are also 11 Hawaiian species), Africa has only
one family uniquely epiphytic. That family, Cos-
taceae, is a dubious segregate of Zingiberaceae,
the latter with epiphytes in Asia. At least nine
seed plant families have epiphytic species in both
the Neotropics and Australasia (but not Africa),
but not a single one has epiphytes in both the
Neotropics and Africa-Madagascar but not Aus-
tralasia. There are 14 families with epiphytic
species in all three of the world's main tropical
regions. In total there are 33 (or 34 ir pepo
is nlv
continent as compared with 23 with pins doe
on more than one continent: clearly most seed
plant families have epiphytes on only a single
continent.

Many of the “epiphytic” families included in
the above analysis are actually terrestrial families
with one or two aberrant species adapted to epi-
phytism. There are only 32 seed plant families
with five or more epiphytic species. If the con-
tinental representations of these 32 families are
compared, seven (Bromeliaceae, Cyclanthaceae,
Rapateaceae, Bignoniaceae, Campanulaceae,
Marcgraviaceae, and Guttiferae) have epiphytes
only in the Neotropics; six (Myrtaceae, Nepen-
thaceae, Pittosporaceae, Loganiaceae, Balsami-
naceae, and Zingiberaceae, sensu stricto) have
epiphytes only in Australasia; 14 have epiphytes
in all three tropical regions; and five (Asclepia-
daceae, Ericaceae, Rubiaceae, Solanaceae, and
Urticaceae) wi ge a only in the Neotrop-
ics and Austr;

Put B way, LES are 42 neotropical seed

4 AY
J families

it eaninhvtec

plant families with epiphytes (Table 3) but epi-
phytism is minimal in 19 of these (one to four
epiphytic species in the Neotropics). Of the fam-
ilies with at least five epiphytic species in the
Neotropics, seven— Bromeliaceae, Cyclantha-
ceae, Rapateaceae, Campanulaceae (also in Ha-
waii), Bignoniaceae, Marcgraviaceae, Guttifer-
ae—are epiphytic exclusively in the Neotropics,
and another, Cactaceae, has only one epiphytic
species widespread in the Paleotropics. Fourteen
of the families with some neotropical epiphytic
species have epiphytes in all three tropical re-
gions— Araceae, bi ordi Liliaceae, Aralia-
ceae, Begoniaceae, Com sitae, Crassulaceae,
Gesneriaceae, Lent tomata-
ceae, Moraceae, ME ne Piparace eae (plus,
marginally, Cactaceae). In addition to exclusive-
ly south temperate Griseliniaceae and Philesi-
aceae, seven families—Gnetaceae, Burmanni-

aceae, Asclepiadaceae, Ericaceae, Rubiaceae,
Solanaceae, Urticaceae—have epiphytes only in
the Neotropics and Australasia.

The sharing of epiphytic taxa between Austral-
asia and the Neotropics but not with Africa is a
very different pattern from that normally found

giosperms, where close floristic relation-
ships between Africa and the Neotropics (reflect-
ing a shared early angiosperm Gondwanan flora,
Raven & Axelrod, 1974) or between Africa and
tropical Asia (reflecting the relatively direct mi-
gration route provided by today’s geography) are
the general rule.

Even within the same family paleotropical and
neotropical epiphytes are often not closely relat-
ed. For example, the paleotropical epiphytic

222

Gesneriaceae belong to subfamily Cyrtandroi-
deae while neotropical ones are mostly in the
endemic subfamily Gesnerioideae (Wiehler
1983). Madison (1977) noted that epiphytism
has arisen independently in at least three differ-
ent groups of aroids. Most epiphytic neotropical
orchids belong to subtribes Pleurothallidinae,
Maxillarinae, and Oncidinae, while most paleo-
tropical ones belong to Dendrobiinae and Bul-
bophyllinae. The few epiphytic Central Ameri-
can species of Cynanchum are quite unrelated to
the many paleotropical epiphytic species of As-
clepiadaceae. Most epiphytic neotropical Erica-
ceae belong to subfamily Vaccinioideae, half the
paleotropical ones to Rhododendroideae. Most
epiphytic neotropical Rubiaceae belong to tribe
Cinchoneae (subfamily Cinchonioideae), most
paleotropical ones to Psychotrieae (subfamily
deae).

`

Rubioideae

th h a ee

ven g piphytic sp
in all three continental regions, these may not be
closely related to each other. Altogether only 30
epiphyte-containing seed plant genera are found
in more than one of the three main tropical re-

gions, and only 14— Liparis, Malaxis, Vanilla,
Polystachya, Bulbophyllum, Schefflera, Begonia,
Rhipsalis, Vaccinium, Utricularia, Ficus, Myr-
sine, Peperomia, Senecio —are pantropical.
Twelve of the genera that occur as epiphytes on
more than one continent are large, diverse, ter-
restrial genera in which epiphytism has arisen
occasionally. For example, there are at least two
independent origins of epiphytism in Utricularia
(P. Taylor, fide Madison, 1977). Genera like
Gnetum, Myrsine, Burmannia, Schefflera, Be-
gonia, Senecio, Gaultheria, Vaccinium, Piper,
Psychotria, Solanum, and Pilea have indepen-
dently derived and mostly quite unrelated epi-
phytic species in | Old and New Worlds. There

with epi-
phytic: species in | Africa and Asia: predominantly
epiphytic Medinilla (Melastomataceae) and the
orchid genera Acampe, Oberonia,

l
(M visini naceae). We are oi with only Schefflera,
Rhipsalis, Ficus, Peperomia, Liparis, Malaxis,
Vanilla, Polystachya, aad Bulbophyllum as gen-
era in which epiphytism is widely prevalent on
all three continents. Indeed the only genera in
which epiphytism in both the New and Old
Worlds seems to represent a true synapomorphy
are three south temperate genera (Luzuriaga,
Griselinia, and Coprosma), Ficus, Rhipsalis,
probably Peperomia, and possibly Schefflera. The

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

clearest cases of a shared epiphytic ancestor are
Cactaceae where long distance dispersal of one
species of Rhipsalis is responsible for the pattern
seen today (Barthlott, 1983), and the peculiar
case of Ficus where the large successful pantrop-
ical subgenus (Urostigma) has specialized as
stranglers. The five pantropical epiphytic orchid
genera likely achieved their present distributions
via long-distance dispersal of their dustlike seeds.
Predominantly epiphytic Peperomia may have
been originally epiphytic; it is probably ancient
(cf. Burger, 1977) and in many aspects of its bi-
ology, in addition to its pantropical distribution,
it is anomalous among epiphytes.

Schefflera is predominantly epiphytic in the
Neotropics, predominantly terrestrial in the Pa-
leotropics; according to Madison’s (1977) esti-
mate only 65 species are epiphytic. Many species
grow both as epiphytes and terrestrials; epiphyt-
ism, though widespread, does not seem funda-
mentally intrinsic to Schefflera and may have
arisen independently in all three regions and
probably within a given region as we

erns, notorious for the ease of long-distance
dispersal of their dustlike diaspores, contrast with
the angiosperms in having most large epiphytic
genera preponderately pantropical. There are
even exclusively epiphytic pantropical fern gen-
era including Pleopeltis, Platycerium, Ctenopte-
ris, Xiphopteris, Psilotum, Vittaria, and (almost)
Polypodium, there is not a single exclusively epi-
phytic pantropical seed plant genus. Even in ferns
the majority of the epiphytic genera are restricted
to one geographical region. Only one epiphytic
fern genus is disjunct between the Neotropics and
Asia (Ophioglossum, a large genus with only two
epiphytic species separately arisen on the two
continents). Five epiphytic fern genera occur in
both tropical Asia and Africa but not the Neo-
tropics, three of them on Madagascar but not
continental Africa; again this is a pattern without
parallel in the seed plants.

We may conclude that, except for the ferns,
strangler figs, Peperomia, Rhipsalis, and a few
orchid genera, the epiphytic floras of the different
tropical regions are independently derived, even
in most cases where the same family or genus is
involved in different regions.

While certain taxa are preadapted to an epi-

largely on the peculiarities of that region’s evo-
lutionary milieu. Epiphytism has arisen very
many times in very many groups. However, cer-

1987]

tain regions have given rise to many epiphytes;
others have not. In this context, the partial ex-
planation offered by Madison (1977) for the
greater representation of epiphytes in the Neo-
tropics—that historical accident in the distri-
bution of families like Bromeliaceae, Cactaceae,

species—seems largely irrelevant. Indeed,
many families (and other supraspecific taxa) have
evolved epiphytism in the Paleotropics as in the
Neotropics; the difference is that in the Neo-
tropics evolutionary experiments with an epi-
phytic life-style have subsequently led to much
more profuse speciation. Madison (1977) also
thought that one element in explaining the con-
tinental difference of epiphyte diversity is a pau-
city of paleotropical nonorchid monocot epi-
phytes. However, our analysis emphasizes that
it is not the evolution of epiphytism itself in such
taxa that is the critical factor, but rather that
there has been little subsequent radiation. Why
have epiphytic taxa of Zingiberaceae, Costaceae,
Pandanaceae, or Liliaceae not evolved into pa-
leotropical versions of Bromeliaceae or Cyclan-
thaceae? We will try to analyze why this should
be so in the next section.

EvoLuTION or EPIPHYTE SPECIES
DIVERSITY

Why are epiphytes so much better represented
in some habitats than in others within the Neo-
tropics? Part of the answer to that question can
be adduced from the diversity gradients dis-
cussed above. From the patterns documented

somewhat tenuously established trend toward
greater diversity at middle elevations, these are
exactly the trends shown by angiosperms in gen-
eral (Gentry, 1982a, 1982b, 1987c). Ina previous
paper, based on data extrapolated from a large
array of published monographs of neotropical
taxa, Gentry (1982b) concluded that plant fam-
ilies belonging to different habit groups have fun-
damentally different distributional patterns.
Families composed mostly of canopy trees or
lianas have their greatest diversity in Amazonia
whereas families made up mostly of epiphytes,
shrubs, or palmetto-type herbs are largely extra-
Amazonian and are especially concentrated along
the lower slopes of the northern Andes and to a

GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES

223

lesser extent in southern Central America. For
epiphytes this concentration of species diversity
could have been predicted from the trends out-
lined above. But why are epiphytes (and some
other plants) so much more diverse in these re-
gions?

One reason that epiphytes are especially di-
verse in wet aseasonal forests is that they are able
to achieve a much finer niche partitioning, and
thus a higher alpha diversity, there. Western Ec-
uador provides a good example of how this phe-
nomenon operates. In the evergreen Rio Pa-
lenque wet forest nearly all of the epiphytes have
a characteristic and usually very restricted hab-
itat, occurring only in the understory, the middle
tory, or the canopy. Altogether 41 species of
vascular epiphytes at Rio Palenque are under-
story specialists: 19 species of Araceae, one of
Begoniaceae, one of Bignoniaceae, five of Cy-
clanthaceae, eight of Piperaceae, one of Solana-
ceae, and six ferns. However, there is not a single
understory-specialist epiphyte species in the
highly seasonal semideciduous moist forest at
Jauneche, only a few tens of kilometers away.
The presence of 41 species of understory spe-
cialist epiphytes at Rio Palenque accounts for
much of the difference between its diverse epi-
phytic flora and the relatively depauperate one

Nn

constant environment at enque rs
within-community microhabitat specialization
by epiphytes. Thus classical ideas about the
greater spatial heterogeneity of everwet tropical
forests (e.g., Baker, 1970) are certainly applicable
to epiphyte diversity patterns both within the
tropics and on a latitudinal gradient.

To some extent niche fine-tuning in constant
environments also occurs in nonepiphytes. A
good example is provided by Gasteranthus at
Centinela, Ecuador (see Gentry, 1987b). Six
species occur together sympatrically. All are ter-
restrial herbs and five are strictly endemic. The

Another species, G. crispus, grows only in sandy
creek beds of the north part of the ridge and is
not strictly sympatric with the other four species.
Of the four strictly sympatric species, one, G.
atratus, has switched from hummingbird- to bee-
pollination and has yellow flowers completely
distinct from the orange flowers of the other

224

species. Two of the strictly sympatric species both
have large flowers and grow in the deep shade
along creek beds, but G. macrocalyx blooms in
the wet season and G. pubescens in the dry sea-
son. The final species, G. carinatus, is morpho-
logically differentiated by slightly smaller flowers
and ecologically by growing only along the ridge
top. In Gasteranthus, as in the Rio Palenque epi-
phytic flora, very fine niche partitioning in a rel-
atively constant climate seems an important key
to maintenance of high species diversity. To the
extent that such niche partitioning is related to
equable montane cloud forest conditions, it might
be expected to be relatively favored in the Neo-
tropics.

A second explanation for the great epiphyte
diversity in the Andean area focuses on £-di-
versity resulting from the greater microsite dif-
ferentiation typical of mountainous regions. A
test of this hypothesis might come from com-
paring epiphyte diversity in areas with high and
low microsite differentiation. The two “nudos”
where the eastern and western Andean cordil-
leras come together briefly at opposite ends of
Ecuador provide such a test. The southern Nudo
de Loja (or Sabanilla) marks the beginning of the
Huancabamba biogeographic discontinuity (cf.
Berry, 1982); the northern Nudo de Pasto, most-
ly across the border in Colombia, marks the point
where the three Colombian cordilleras diverge.

n
different orientations of adjacent slopes. Teli-
pogon (Orchidaceae) provides a good example of
how such microsite heterogeneity can multiply
epiphyte species diversity. There are 37 Telipo-
gon species in Ecuador and five more on the
Colombian side of the Nudo de Pasto. There are
14 species on the Nudo de Loja and 16 species
on the Nudo de Pasto, as compared with only
14 species in the entire intervening 600 km. More
instructive, nine ofthe Telipogon species on Nudo
de Loja and five on Nudo de Pasto are locally
endemic, each found in a single valley or a single
slope. In the much larger intervening area there
are only nine endemic species, each restricted to
a small area of a few hectares, in several cases
representing unusual and restricted microhabi-
tats similar to those around the Nudos. It seems
clear that in 7elipogon diversity is associated
with local speciation and microgeographic spe-
cialization. Again, epiphytes are not the only

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

group to show such patterns, but they do seem
especially prone to this kind of microgeographic
speciation. Probably the association of accen-
tuated microgeography with mountainous ter-
rain, which is much more prevalent in the Neo-
tropics, peu some of the intercontinental
diversity differen

third, not ucc mutually exclusive,
potential explanation for high epiphyte diversity
in the northern Andean and southern Central
American cloud forests is the “evolutionary ex-
plosion" hypothesis advanced by Gentry (1982b)
in an attempt to explain why there are so man
more neotropical than paleotropical plant species.
A relatively small number of genera of epiphytes,
understory shrubs, and tto-type h
speciated profusely in the northern Andean re-
gion, in each case giving rise to very many locally
endemic and often rather poorly differentiated
species. Gentry (1982b: 587) interpreted the high
local endemism (cf. Gentry, 1987b) and apparent

plastic genera as An m, Piper, aven-
dishia as reflecting Hed balance" founde
effect phenomena 977. Templeton,

1980) with major genetic reorganizations or ge-
netic transilience (Templeton, 1980) optimized
by small and localized populations and by the
need for constant in a habitat par-
titioned by mountains, local rainshadows, ver-
tically shifting cyclically coalescing vegetational
zones, and frequent landslides. If genetic founder
effects associated with recolonization of the open
areas resulting from landslides or unusually fre-
quent tree falls in these geologically and ecolog-
ically dynamic regions are major determinants
of speciation events, then much speciation in
Andean-centered taxa could well be essentially
sympatric and largely random. This model thus
differs from the **microgeographic speciation”
model in that speciation could take place at a
much finer “sympatric” scale, e.g., as coloniza-
tion of a specific landslide, with many of the
resultant species ecologically indistinguishable
from each other, rather than each adapted to a

mism imputed to be associated with tropical
mountains it would be expected to be most ap-
plicable in the Andean region, by far the most
extensive mountain system in the world's trop-
cs.

We have indirect but highly suggestive evi-
dence for speciation associated with genetic tran-

m

1987]

GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES

Stanhopea jenishiana Reichb. f.

FIGURE 5.

silience in founder populations in several genera
of epiphytes. Embrya rodigasiana (Cogn.) Dod-
son provides a good example of a founder event.
This species is distributed mostly in western Co-
lombia, between 1,000 m and 1,500 m, ranging
from west of Medellin to near Buenaventura.
There is also a disjunct population in a small
area of southern Ecuador near Pangui, at 1,500
m above Bomboiza and Gualaquiza; prior to re-
cent widespread habitat destruction, the species
was very common in the disjunct and geograph-
ically very limited Ecuadorian part of its range,
where it was surely introduced by a long-distance
dispersal founder event. Masdevallia chonta-
lensis is a similar example. Well known and oc-
curring in a well-known habitat, it ranges from
Guatemala to Panama with a small disjunct pop-
ulation near Piñas in southern Ecuador. There
are several similar cases of unsuccessful founder
events by dust-seeded tropical epiphytes that have
temporarily established disjunct populations in
Florida, been duly recorded as members of the
Florida flora, and then subsequently disap-
peared. An example is Leochilus labiatus which
grew for a while near Fackahatchee.

The Stanhopea jenishiana complex, well char-
acterized by a suite of distinctive morphological
characters, provides an example of the next step
in such a process. Stanhopea jenishiana Reichb.
f. (Fig. 5) ranges from Cali to Popayan in the
Cauca Valley of Colombia, with a disjunct pop-

ulation near Piñas in southern Ecuador. There
are two other small geographically isolated pop-
ulations of this complex in western Ecuador, each
of them specifically distinct but clearly derived
from S. jenishiana and each likely resulting from
a single long-distance dispersal event— Stanho-
pea frymirei Dodson is endemic to a range of
relatively moist hilltops near the coast in Manabi
and Guayas provinces and S. embreei Dodson
occurs only around 1,000 m in the Bolivar-Canar
border area (Fig. 6).
he herbaceous and shrubby taxa that show

such patterns have relatively short generation
times, providing conditions appropriate for rap-
id evolutionary diversification. Under either the
microgeographic or founder effect hypothesis,
epiphytes, as the major herbaceous component
of wet tropical forests, might be expected to show
unusually rapid “evolutionary explosion" type
speciation. Relatively specific pollination sys-
tems constitute a second factor wansi bl
Gentry (1982b) as potentially promoting un
ally rapid speciation in **Andean-centered" ie
with shifts in specific pollinators accompanie
by coevolutionary fine-tuning of precise plant-
pollinator systems apparently a common evo-
lutionary theme. Again epiphytes, often char-
acterized by high pollinator specificity, should be
prime candidates for rapid speciation.

The Stanhopea jenishiana complex (see above)
is a good example of how founder events and

N
N
ON

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FIGURE 6. Two localized derivatives of Stanhopea jenishiana.—A. S. embreei Dodson.—B. S. frymirei
odson

shifts in pollination syndromes can combine to
give rise to new species. The presumed ancestral
species, S. jenishiana, has orange flowers and is
pollinated by bees of the genus Euglossa while
both of the locally endemic derivative species in
central Ecuador have white (E. embreei) or straw-
colored (E. frymirei) flowers and have switched
to pollination by Eulaema bomboides. All three
species have methyl cinnamate as the major scent
attractant but each has a different set of scent-
modifying compounds. Eulaema bomboides, a
very effective pollinator that specializes on flow-
ers producing methyl cinnamate, is endemic to
central Ecuador. We interpret this situation as
reflecting three different instances of long-dis-
tance dispersal. In the southern Ecuador Pinas
population, outside the range of E. bomboides,
pollination by Euglossa was maintained and no

vantage of Eulaema bomboides as a pollinator
occurred with essentially the same selection op-
erating in these two different founder popula-
tions. Different evolutionary solutions reflected
by the two distinct derivative species resulted.
Scelochilus, another epiphytic genus of orchid,
provides an even more intriguing indication of

how rapid speciation might be in such taxa. Sce-
lochilus is a genus of 34 species found mostly
epiphytic in guava trees; sparsely represented in
natural vegetations, it was apparently ideally pre-
adapted for the special conditions provided by
guava plantations. Fifteen species of Scelochilus
occur in Ecuador. In 1957 one of us (CD) made
an intensive study of populations of Scelochilus
in an extensive guava grove near the edge of wet
forest at 1,000 m altitude at km. 94 of the old
Guayaquil-Cuenca road in Ecuador. Two un-
described species of Scelochilus were present,

ost
diues ni but their flowers, though some-

hat variable in each species, were very distinc-
tive with no overlap whatsoever between the two
species. Indeed the original study, intended to
focus on hybrid introgression, had to be aban-
doned because the two taxa proved to be so con-
sistently differentiated. These species were de-
scribed as SS. frymirei and S. embreei (Fig. 7).
Fifteen years later, in idis a return visit was
made to the ve. In the intervening
years most of the itin Wasa had been cut and
converted to pasture. As a result the habitat was

1987]

very different, with a much dryer aspect and the
remaining guava trees rather old and decrepit.
Scelochilus was much rarer but ca. 50 plants were
located in the remaining guavas. Incredibly, nei-
ther S. embreei nor S. frymirei was present in
1982, but rather two different new species were
found, later described as S. gentryi and S. ro-
mansii. As in 1957, both of these species, veg-
etatively indistinguishable, were clearly differ-
entiated from each other by floral characters (Fig.
7). Both of the species present in 1982 are closely
related to S. frymirei. We suspect that they may
represent in situ speciation events, at least in the
case of S. romansii (S. gentryi has also been found
in several other localities in western Ecuador). If
so, natural speciation in Scelochilus can occur in
as little as 15 years!

An obvious alternative interpretation is that
specific limits in Scelochilus are too finely drawn,
with S. gentryi and S. romansii representing part
of the intraspecific variability within polytypic
S. frymirei. However, this possibility seems ob-
viated by the fact that the two co-occurring Sce-
lochilus species of 1982 pass the test of sympatry
as biologically differentiated populations, even
though they are more similar to each other than
to their putative ancestor S. frymirei. We favor
the interpretation that the kind of genetic reor-
ganization that reflects speciation in Scelochilus,
a ma

nonorchid epiphytes, is so labile that it can be
effected in incredibly short times. If this inter-
pretation is correct, then it is no wonder that
many epiphytic taxa have undergone what ap-
pears to be truly explosive speciation in the Neo-
tropics. Obviously, it is also possible that S. gen-
tryi and S. romansii immigrated to the guava
grove in question sometime between 1957 and
1982 with S. frymirei and S. embryi coinciden-
tally becoming locally extinct during the same
time interval. But the mere fact that natural spe-
ciation in fifteen years seems an equally plausible
explanation for these observations is surely sig-
nificant.

Another line of reasoning also supports the
idea that certain neotropical epiphytes have
undergone explosive evolution. It is probably safe
to assume that, as a general rule and despite many
potential exceptions, genera with many species
are those which have, on the average, undergone
the most rapid speciation, Thus an examination
of the largest epip might indicate some
a Sr ponds, both geographic and nop,
th Table 11

GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES

227

lists the 47 largest genera of vascular epiphytes
worldwide (those with 90 or more epiphytic
species) and provides the data for such an anal-
ysis. Again orchids are preeminent. Half (22) of
the 47 largest epiphyte genera are orchids. While
orchids might appear to be exceptional in their
unusual genetic plasticity and highly specific pol-
lination systems, they must unavoidably be re-
garded as the most successful practitioners of the
art of being epiphytic. Biogeographical analysis

T t
number of large neotropical and See ses
orchid genera is the same, there is a dramatic
difference in the number of species that they con-
tain. The nine neotropical genera are far larger,
accounting for 5,240 species (average 582 spp.
per genus) vs. 2,626 a (average 263) for the
ten paleotropical on

erns account for pum of the largest wa d
genera. Not surprisingly, in view of their dia-
spore vagility, nearly all of the large IE.
fern genera are pantropical, showing little evi-
dence of unusually rapid differentiation or spe-
ciation in the Neotropics. Only one fern genus,
Pyrrhosia, is restricted to the Paleotropics, none
to the Neotropics.

The remaining 15 of the largest epiphytic gen-
era are split evenly between dicots and mono-
cots. The two largest dicot genera, Peperomia
and Ficus, are pantropical, although the former,
at least, is better represented in the Neotropics;
the other largest epiphytic dicot genera are two
neotropical gesneriads (Drymonia and Colum-
nea, sensu lato), two ericads (Rhododendron and
Cavendishia), one each epiphytic in Old and New
Worlds, and two melastome genera (Medinilla
and Blakea), restricted respectively to the Old
and New Worlds (ane extremely similar to each

other as wellast

Topobea). In ganar: here seems a reasonable
numerical balance between Old and New World
representation in the largest dicot epiphyte gen-
era, except in Gesneriaceae. The situation is very
different among the nonorchid monocot epi-
phytes. Six of the seven largest genera— Anthur-
ium, Tillandsia, Philodendron, Vriesia, Guz-
mania, an mea —are exclusivel
neotropical; only Rhaphidophora is paleotropi-
cal. Thus the very many species that have evolved
among certain monocot epiphyte genera seem a
peculiarity of the Neotropics.

Why have epiphytes been especially suscep-

228 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

SCELOCHILUS EMBREEI Dodson SCELOCHILUS FRYMIREI Dodson

]—

SCELOCHILUS GENTRYI Dodson SCELOCHILUS ROMANSII Dodson & Garay

1987]

tible to the rapid, even explosive, speciation that
seems to have characterized a large element of
the neotropical flora? At least in the case of or-
chids, unusually specific pollination systems have
clearly played a major role in making possible
very active speciation. Like the orchids, the non-
orchid epiphyte families that have speciated the
most profusely in the northern Andes and south-
ern Central America—Gesneriaceae, Bromeli-
aceae, Ericaceae, and Araceae—all share, to a
greater or lesser extent, relatively specialized pol-
linators and specific palliation systems as com-
pared with the “average” tree, shrub, or free-
climbing liana (e.g., prevalent hummingbird or
euglossine pollination). On the other hand the
epiphytic taxa that have speciated most profusely
are characterized, as a group, by more general-
ized, higher risk dispersal strategies than typical

optimal genetic transilience related to multiply
replicated founder events in a dynamic and kal-
eidoscopically changing habitat as Gentry (1982b)
suggested, then epiphytes, characterized by their
unique combination of r- and k-selected repro-
ductive traits, might be uniquely equipped to
react to this adaptive milieu. Like weeds, they
have diaspores intrinsically adapted for quick
colonization of newly available habitats. In epi-
phytes, like weeds, the primary need for such
adaptation is presumably a response to the

nd unstable nature of their normal

adapting epiphytes for rapid speciation in an
environment in which the need to recolonize
dynamically changing microhabits is unusually
frequent. However, unlike weeds whose di-
versification is generally constrained by overly
generalized pollination syndromes (frequently
even with loss of sexual recombination), the rich
array of specialized pollination systems that
characterizes the successful epiphytic taxa would
seem to preadapt them for rapid, pollinator-re-
lated evolutionary diversification. In this con-
text, it is not likely to be an accident that the
preeminently successful epiphytic family, Or-

GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES

229

chidaceae, is precisely that family that has both
the most specific pollination systems and the
tiniest, most r-selected diaspores of any angio-
sperm. Moreover, from the viewpoint of “‘acci-
dental” speciation via founder effect phenome-
na, the unusual genetic similarity among the
potential-colonizer orchid propagules that arrive
together at a specific site—due to orchid seeds’
unique sharing of fathers as well as mothers,
thanks to their pollinia—might be expected to
accentuate the potential for genetic drift in col-
ations. Interestingly, the closest
a o

Andean cloud forest epiphytes is found in
baceous and palmetto taxa that most closely ap-
proximate the unique epiphyte combination of
specific pollination systems and an r-selected
high-risk dispersal mode.

CONCLUSION

Although many unrelated kinds of plants have
evolved epiphytic habits, most of these represent
isolated species or genera in otherwise terrestrial
families. Only three nonfern families — Orchi-
daceae, Cyclanthaceae, Marcgraviaceae (and
possibly also Bromeliaceae) — are predominantly

Orchidaceae, Bromeliaceae, Polypodiaceae, and
Araceae—and 8996 in eight families. Very few
families have been successful at undergoing ex-
tensive radiation as epiphytes. Indeed over two-
thirds of all epiphyte species belong to the single
family Orchidaceae, and to a large degree un-
derstanding epiphytic diversity is synonymous
with understanding orchids.

Although notably few families have been very

in a local flora or 6396 of the individual plants
in a given sample area.

Epiphytes have speciated most profusely in the
Neotropics, especially in northwest South Amer-

7. Postulated rapid KT in Scelochilus. Top two species bee described as new from 1

FIGURE 7.
tibns! ed guava grove at
two species, both ne
Plantarum “Tropic carum.

of Guayaquil-Cuenca road. In 19
and both vend d to S. frymirei. yr reer ie dh Orchids of Ecuador, Icones
)

957
y had been replaced by bottom

230

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

TABLE 11. Major epiphyte genera (in part after Madison, 1977; Dressler, 1981; Kress, 1986).
Genus No. Epiphytic Spp. Total No. Spp. Distribution
Pleurothallis 1,520 1,600 Neotropical
Bulbophyllum 1,000 1,000 Pantropical
endrobiu 900 900 Australasian
Epidendrum 720 800+ Neotropical
Peperomia 700 1,000 Pantropical
Anthurium' 600 850 Neotropical
Lepanthes 600 600 Neotropical
elis 540 600 Neotropical
Ficus 500 800 Pantropical
Maxillaria 570 600 Neotropical
Eria 500 550 ustralasian
Oncidium 475 500 Neotropical
splenium 400 650 Pantropical
Tillandsia 400 450 Neotropical
Grammitis 400 400 Pantropical
Philodendron' 350 475 Neotropical
Masdevallia 400 400 Neotropical
Medinilla 300+ 400+ Paleotropical
Liparis 300 350 osmo an
Oberonia 300 300 Paleotropical
Odontoglossum 285 300 Neotropical
Columnea (sensu lato) 262 265 Neotropical
Elaphoglos. 250 500 Pantropical
Hymenophyllum 250 300 Pantropical
Angraecum 206 206 Trop. A
Lycopodiu 200 400 Cosmopolitan
Polystachya 200 210 Pantropical
Ctenopteris 200 200 Pantropical
riesia 200 260 Neotropical
Phreatia 190 190 Australasian
Trichomanes 150 300 Pantropical
Polypodiu 140 150 Pantropical
Encyclia 130 130 Neotropical
Octomeria 130 130 Neotropical
Taeniophyllum 120 120 Paleotropical,
esp. Afr.
uzmania 120 140 Neotropical
Dendrochilum 120 120 Australasia
Aechmea 120 150 Neotropical
Rhododendron 112 850 ostly Asian
Coelogyne 100 100 rop. Asia
monia 100 110 opical
Pyrrosia 100 100 Paleotropical
Rhaphidophora 100 100 Paleotropical
Appendicula 100 100 Australasia
Blakea? 98 100 Neotropical
Thrixspermum 90 100 ustralasia
Cavendishia? 90 100 Neotropical

! Croat, pers. comm.
? Renner, 1986 and pers. comm.
? Luteyn, pers. comm.

1987]

ica and southern Central America. The explosive
speciation of relatively few epiphyte genera in
this region has been responsible to a large extent
for the excess floristic diversity of the Neotropics
as compared with the Paleotropics.

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1987a. An overview of neotropical phyto-
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1987b. wii ed in tropical versus tem-

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. 1987c. Changesin “ss gangsa diversity
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EMM

P1987. Geographical variation

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974a. Epiphytic myrmecophytes in Sara-
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74b. Tropical Ape IS rivers, animals,
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FF, M. FARI

" TA NAS, S. REYES, N. RINCON,
A. SOLER, P. Socks WL VERA.

1976. Changes

234

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[VoL. 74

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1987] GENTRY & DODSON—NEOTROPICAL VASCULAR EPIPHYTES 233

APPENDIX I.

Changes in data base from epiphyte list in Madison (1977).

Additions

450)
,200 species, not 850)
Anthurium (600/850)
Philodendron (350/475)
Cyclanthaceae (125 species, not 31)
Dicranopygium (few/44)
Evodianthus (1/1)
Stelestylis (4/4)
Dj

coreacea
Dioscorea (1/600) (Ecuador, pers. obs.)
Liliaceae
Rhodocodon (1/8) (Madagascar, pers. obs.)
Philesiaceae
Philesia (1/1) (Chile, pers. obs.)
Luzuriaga (3/3) (Chile, pers. obs.)
Alzateaceae
Alzatea (1/2)
Anacardiaceae
iini af 10) (SE Asia, fide Kress)
Apocy
Mandevilla (1/114) (M. pittieri, Costa Rica)
Begon
Begonia K 1,000)
Bignoniacea
Gibs oniothamnus (11/11)
Schlegelia (18
ombacaceae
Spirotheca (4/4)
seraceae
MN (1/80) (B. standleyana, Costa Rica)
fen eal (ca. 30 epiphytic species, not 3)
bum !GSinclairia) (2/90)
Mikani
desde da (20/24)
Nelso po (1/1)
Pseudogynoxy. 1)
— (Pentacalia) b l p (epiphytic in Neo-
s, Madagasca
URN ylis (2/2)

Crassulaceae
Kalanchoé epiphytic in Madagascar, pers. obs.
Marcgraviaceae (89 epiphytic species, not 94)
Marcgraviastrum (10/15
Marcgravia (ca. 50—55/55)

Melastomataceae (567 epiphytic species, not 483)
The paleotropical genera Baekeria, Dalenia, Dic-
cochaeta, Neodissochaeta, Om phalop us, and
Plechiandra each has at least one hemiepiphytic
species fide Renner, 1986.

yricaceae
Myrica (1/35) (SE Asia, fide Kress, pers. comm.)
apotaceae
xri (Costa Rica, fide Kress, pers. comm.)

Saxifrag;

H ydrangea (2/80) (epiphytic in Neotropics)
Solana

fapa (also epiphytic in Neotropics)

Deletions

Apostasiaceae = Orchidaceae

pou = oe

Bromelia —Ana

Cycla <” pei Ci arian

Balsaminaceae — rubr eod Qm epiphytic in Neo-

ignoniaceae — Radermachera
Gutt viiei -Clusia ari occurs in Neotropics
Marcgraviaceae — Car
Araliaceae — eade a inr Schefflera

Note: Additional occasional or sporadic epiphytes i in-

clude s Kress (e.g
Cyperu Pseudoeverardia (= Everardia), Arenaria,
E ora, Maranta, Pourouma) are also excluded.

NITROGEN FIXATION BY EPIPHYLLS
IN A TROPICAL RAINFOREST!

BARBARA L. BENTLEY?

ABSTRACT

Bluegreen 5 (Cyanobacteria) growing on the leaf surfaces of understory plants in a tropical

rainforest can ospheric n

nitrogen. Rates of fixation are strongly influenced by the presence of

glucose and pied: nutrients leached from the host leaf, by light intensity as it relates to the pho-
tosynthetic rates of the algae, and by desiccation especially as it is influenced by the co-occurrence of

epiphyllous bryophytes. A signific

ant portion of this newly fixed nitrogen is transferred to the host

leaf and may account for 10-25% of the total nitrogen in a leaf.

Although the major source of nitrogen for plant
growth and reproduction is from decomposition
of organic matter, most ecosystems can gain sig-
nificant quantities from precipitation and bio-
logical fixation of atmospheric nitrogen. In fact,
Burns & Hardy (1975) estimated that biological
fixation alone may account for 4396 of the nitro-
gen transferred worldwide. Subba Rao (1977)
suggested that the rates in tropical areas may be
even higher and thus fixation could have a major
role in the tropics where available nitrogen can
be limited.

In this paper, I will discuss one aspect of ni-
trogen fixation in a tropical ecosystem — that of
fixation by epiphylls in a rainforest understory.
Although epiphyll fixation can account for up to
2596 of the nitrogen in a host leaf (Bentley &
Carpenter, 1980), rates of fixation are quite vari-
able in both time and space. Here I document
some of the environmental factors that influence
fixation rates.

What are epiphylls? Epiphylls are epiphytes
that are restricted to the surfaces of leaves. In
tropical rainforests, most visible epiphylls are
leafy liverworts of the family Lejeuneaceae but
may also include mosses, lichens and, on some
occasions, seedlings of other epiphytes such as
bromeliads and orchids. In this study, we focused
on the microorganisms, especially the bluegreen
algae (Cyanobacteria) growing in association with
the visible forms (Fig. 1).

The epiphyll community is best developed in
regions of high rainfall and low evaporation and
is most diverse in tropical rainforests (Richards,

! [ would first like to express my appreciation to Maureen Dunn who gave expert HIIS e in both
n Mei obert Bl

80-10676 from the National Science Foundat

1964). Because they grow on photosynthetic sur-
faces, epiphylls are often considered to be det-
rimental to the host plant by interfering with light
penetration to the leaf (Richards, 1964). Indeed,
some epiphylls may actually be semiparasitic.
For example, the rhizoids of Radula flaccida, an
epiphyllous liverwort, penetrate the cuticle and
absorb nutrients from their host leaf (Berrie &
Eze, 1975).

Other workers feel that the presence of a
phylls increases the time that leaves remain w
and thus contributes to the growth of omens
pathogenic bacteria and fungi (Gregory, 1971;
Stahl, 1891). Long- term wetting may alse: seduce
the rates of and subsequent mineral
uptake by the roots (Stahl, 1893; McLean, 1919).
As early as 1891, Jungner suggested that rain-
forest plants may have adaptations such as leaf
“drip tips" to increase the rates of drainage off
the leaf and reduce the rate of colonization by
epiphylls. Subsequent work has failed to either
deny or confirm the adaptive role of these char-
acters (Stahl, 1893; Shreve, 1914; Seybold, 1957;
and pers. obs.).

Nitrogen fixation by epiphylls. Some species

Biological Station in Costa Rica, we found that
fixation rates are extremely variable, both within
and among species (Fig. 2). Interestingly, in con-
trast to Ruinen's data where fixation was by free-
living bacteria (primarily Beijerinckia), fixation
in the La Selva understory was most commonly

Black,

Young for assistance in the field. = research was supported by grants DEB 78- 12032 and DEB

? State University of New York, Stony Brook. New York 11794, U.S.A.

ANN. Missouni Bor. GARD. 74: 234-241.

1987.

1987]

BENTLEY — NITROGEN FIXATION BY EPIPHYLLS

Photomicrograph of epiphylls. The strand of the nitrogen- rem bluegreen dni is across the lower
h.

ue of n photograp

associated with the presence of bluegreen algae
in the genera Scytonema, Stigonema, and Hap-
alosiphon. High fixation rates were invariably
associated with a dense cover of bryophytes, sug-
gesting that the bryophytes provide a good sub-
strate for the nitrogen-fixing microorganisms.

Most of our work on fixation was done using
the acetylene reduction method for determining
nitrogenase activity (Bentley & Carpenter, 1980;
Prestwich & Bentley, 1981; Burris, 1972). While
this is an extremely easy field assay for estimating
fixation, it is not a direct measure of nitrogen
fixation, and cannot be used to answer the most
critical question in our study: does the newly
fixed nitrogen get into the host leaf? By using '°N
as a tracer, we were able to document that indeed
this is the case (Bentley & Carpenter, 1984). There
we showed that nitrogen fixation by epiphylls
could account for up to 25% of the nitrogen in
a host leaf.

In other studies with bluegreen algae, Stewart
(1963) and Jones & Stewart (1969) found that
the extracellular a products are pri-
marily a eptides, but less complex
compounds, E ammonium nitrite and

nitrate, can be present. The pathway for move-
ment of the new nitrogen is not through the sto-
mata, as might be assumed, but rather to the
epidermal cells via threadlike ectodesmata pen-

(Franke,
phology of the ectodesmata, Franke felt that the
movement of soluble materials into and out of
the leaf is a normal process, iic de orrelated
with foliar absorption of “foreign” substances
such as fertilizers and dr

Environmental factors affecting fixation. The
rates of fixation and transfer that we measured
were made under ideal conditions for the activ-
ities of microorganisms. Although the high tem-

d almost constant moisture in a trop-
ical rainforest can permit high fixation rates and
concomitant release of nitrogenous products, we
have also observed extremely high variance both
among and within species at the La Selva Station
(Fig. 2). Thus, it becomes important to ask what
environmental factors influence fixation rates by
epiphylls. Basically, the answer lies in three fac-
tors: time, moisture, and nutrients.

e
o
T

ng N FIXED/ hr

X+S

|
| PP,

SPECIES OF HOST

RE 2. Nitrogen fixation rates by epiphylls on

t.
The host species, from highest to lowest fixation rates,
are: Ocotea atirrhensis, Proteum pittieri, Chamaedorea
sp., Ficus sp., Swartzea sp., Miconia sp., Piper sp. 1,

sp., Costus sp., Syngonium 1 SP., Asterogyne sp., and
Geonoma s
Time. Colonization by epiphylls is a time

phenomenon. Young leaves are epiphyll-free
simply because the epiphylls have not hann
established. Long-lived leaves have
greater probability of having a well- tablished
epiphyll community. To document the role of
leaf longevity on fixation, we measured fixation
on leaves which had been previously marked for
up to 24 months (Bentley, 1979). Since we knew
the average survivorship of leaves on a plant, we
could then test for the correlation between the
age of a leaf and the rate of fixation. As can be
seen in Figure 3, fixation rates are significantly
correlated with leaf longevity. At the same time,
we measured fixation by leaves with differing
levels of epiphyll biomass. Again, there is a sig-
nificant correlation (Fig. 4).

Moisture. Interestingly, in the experiments
described above, the rates of fixation were cor-
related with the biomass of the bryophytes, even

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

though bryophytes per se do not fix nitrogen. On
the other hand, bryophytes are correlated with
and can influence the levels of moisture on the
leaf surface (Richards, 1964). Since moisture can
affect fixation rates in other systems (Balandreau
et al., 1974; Horne, 1972; Fogg et al., 1973), we
set up an experiment to test the effects of des-
iccation on fixation by epiphylls. In this case,
individual pinnae of Welfia georgii were moist-
ened with water 12, 4, 2, or 0 hours before in-
cubation. As can be seen in Figure 5, desiccation
has a very dramatic effect on fixation, yet the
recovery is quite rapid and is almost that of the
continuously moist control within four hours.

Although moisture conditions in a tropical
rainforest are relatively constant compared with
most other ecosystems, moisture conditions in
the understory do vary, both seasonally and diur-
nally (Longman & Jenik, 1974; Ruinen, 1961).
For example, water may continue to drip down
through the forest canopy long after a rain has
ceased, but the leaves of the upper canopy will
dry within minutes. Seasonal variation can range
p an increased frequency of dry periods in a
day days or weeks without any appreciable
Rm (Schnell, 1971). Thus, under field con-
ditions, moisture can have significant effects. We
documented this in another experiment, shown
in Figure 6. In this case we found that fixation
rates on rainy days y higher than
on dry days.

Although it is tempting to suggest that the
bluegreens and the bryophytes have some close
symbiotic relationship, the role of bryophytes is
probably no more than to maintain appropriate
moisture conditions on the surface of the leaf.
Epiphyll-laden leaves dry more slowly, not only
because capillary action holds water but also be-
cause bryophytes can draw water directly from
the interior of the leaf (Berrie & Eze, 1975). In
addition, the epiphyll-laden portion ofa leaf sur-
face will become wetter faster, again because of
capillary action drawing water from surrounding
areas. High fixation rates by algae are correlated
with the Presente ad bryophytes simply because
the
bryophytes are appropriate for the growth and
activity of microorganisms.

2A

Energy sources. Nitrogen fixation consumes
energy — free-living microorganisms may require
200 g of glucose for every gram of nitrogen fixed
(Mulder, 1975). Unlike other microorganisms,
bluegreen can use carbohydrates from photosyn-

1987]

BENTLEY — NITROGEN FIXATION BY EPIPHYLLS

237

25- i
= 20r °
hom
O
uj
><
TN I5}
Z
4
IOF
Ix ° "
5r
3
1 1 l l L L l 1 À À J
O 20 40 60 80 100

% MARKED LEAVES REMAINING AFTER 24 MONTHS

E 3. Nitrogen fixation rates by epiphylls on n with different leaf longevities. Highest rates are on

IG
Pa unit in which all leaves live longer t

months. Low ates were on leaves of Piper (80%

est r:
last less than two years). The rank correlation of on with longevity is significant (r = 0.95, P < 0.01).

thesis as well as exogenous carbon to supply the
nitrogen fixation process. In addition, bluegreens
may be able to use intermediate products of pho-
tosynthesis to provide electrons for nitrogen fix-

ation (Mulder, 1975).
T T T T T T T T
e
80} =
°
E
o 60 . =]
bo.
a "ü 4
uj
x
u 40r " 5
z °
o pe ia
€
eo
° °
20+ 4 ° ° 7
L- e° e 4
ee
È Er á oY s * ç, * s f 1 1
0.02 0.04 0.06 O. 0.10
ORY WEIGHT OF EPIPHYLLS
Th 1 à 4,5 c 1 111 m

tion and biomass is significant at the P < 0.05 level
(r = 0.39).

In the field, nitrogen fixation by bluegreen al-
gae is usually light dependent (Henricksson &
Simu, 1971; Horne & Viner, 1971; Stewart, 1973;
and see Fig. 6) and has the same relationship to
light as photosynthesis—decreasing with de-
creasing light intensity. However, low rates of
fixation can occur in the dark (Forman, 1975;
Dugdale & Dugdale, 1962), especially if the or-
ganisms are grown on substrates containing glu-
cose, fructose, or sucrose (Fay, 1965). Thus, many
workers feel that high rates of fixation under nat-
ural conditions are a function of both photo-
mediated internal processes as well as various
forms of exogenous carbohydrates

Exogenous carbohydrates may come from a
wide variety of sources. The major source for
epiphylls is in leachate from the host leaf (Tukey
& Morgan, 1962; Tukey, 1971). Since the amount
of carbohydrate present in leachate is also a func-
tion of the photosynthetic rate of the host leaf
(Tukey et al., 1957), the amount of exogenous
carbohydrate available to the epiphylls will also
be correlated with light intensity in the field.

To document the effects of light on the epiphyll
system, we set up two series of experiments: the
first to establish that levels of light commonly
found in the rainforest understory could influ-

238

eor
£
N
O
W
x
u
z
e 10h

[ —1
0/12 12/0 10/2 8/4
hrs DRY/hrs WET
FIGURE 5. The effects of desiccation on nitrogen

fixation by epiphylls. Leaflets from the same pinna of

georgii were either kept constantly wet (0 hr.
dry; 12 hr. wet), or were subjected to varying lengths
of drying (12 hr. dry, 0 hr. wet; 10 hr. dry, 2 hr. wet;
or 8 hr. dry, 4 hr. wet)

ence fixation rates, and the second to differentiate
between direct effects of light and effects of ex-
ogenous glucose on fixation rates by the blue-
greens. In the first set of experiments, samples
were incubated at three levels of light set to mim-
ic those found in the forest: over epiphyll-laden
leaves (12 uEinsteins m °), in a light gap (68
uEinsteins m ’), and in the dark. As can be seen
in Figure 7, fixation was highest at the highest
light intensity and was still significant even at the
usual light intensity for most epiphylls

In the second set of experiments, we measured
nitrogen fixation by leaf samples that had been
dipped in a p 1% glucose solution, a concentra-
tion chosen imic concentrations we mea-
sured in leaf s (Tukey et al., 1957). Again,
we incubated the samples at two light intensities.
In this case, we were able to separate out the
effects of light from the effects of exogenous glu-
cose. Note in Figure 8 that fixation is highest in
those samples which were both dipped in the
glucose solution and incubated in the light. The
lowest rates were for the control samples dipped
in water and incubated in the dark. Note, how-
ever, that fixation by the samples dipped in the
sugar solution but incubated in the dark contin-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

T T cs. EE
425
°
°
420 ~
N
E
= ü
<
x i
2 5 >
x Ë
* 2
š 2
š 0 z
=
T
°
=]

T T Y ME
AM6 7 8 9 10 Il 12 | 2 3 4 5 6PM
TIME OF DAY
T T T T T T T T T T
E
b " dd
50 # 425
3
bi
`
40} 420 “E
= `
= ul
à a
x B 4 >
K 30 5r
z
$ Ë
c yr z
20r K 40
Š
x a
lor 5
e
AM6 7 8 9 10 I| 12 | 2 3 4 5 6PM

TIME OF DAY
FIGURE 6. The effects of light intensity on nitrogen

Natural light intensities were abnormally high because
of the lack of cloud cove

ued to fix nitrogen at a reasonable rate for at least
eight hours. These results document that fixation
by epiphylls is influenced by both light directly
and by the presence of exogenous glucose. And
perhaps as important, they document that fixa-
tion can continue at night if exogenous glucose
is present on the leaf surface.

Mineral nutrients. Growth of nitrogen-fixing
microorganisms may also be limited by avail-
ability of mineral nutrients—most commonly
“upi phosphorus, calcium, molybdenum,
and iron (Mulder, 1975; Fogg, 1973; Mague,

977. Deficiencies in any of these can have an

1987]

150r

IOOF

nMOLES CH, /lO cm* LEAF

L 1 L 1 1 L M sub: 1 L...
2 4 6 8 10 l2 I4 16 I8 20 22 24
TIME (hr)

FiGuRE 7. Experimental variation of light intensi-
ties. Fixation rates are given in terms of nM ethylene
to illustrate continuing fixation throughout the 24 hours
of the experiment. Ethylene concentrations of samples
incubated in the dark do not change after 12 hours.

indirect effect on nitrogen fixation because each
is involved in general metabolic activities. In
addition, Mo and Fe are components in the ni-
trogenase molecule. Deficiencies in either of these
reduces rates of enzyme synthesis and thus di-
rectly affect fixation (Burns & Hardy, 1975).

Exogenous combined nitrogen, most notably
NH,*, will also inhibit nitrogen fixation (Stew-
art, 1973; Burns & Hardy, 1975; Fogg, 1973).
Again, the effect is the result of reduced nitroge-
nase synthesis. Interestingly, the effects of com-
bined nitrogen may explain why fixation rates
are low in nongrowing cells: as growth slows
down, nitrogenous products accumulate in the
cell, which in turn impacts on enzyme synthesis.
Thus, deficiencies in any substance required for
growth may reduce nitrogenase activity.

As with exogenous carbohydrates, the main
source of mineral nutrients is through decom-
position of organic litter on the forest floor, al-
though some may be available in leachate, stem-
flow, or throughfall water (Long et al., 1956). The

BENTLEY — NITROGEN FIXATION BY EPIPHYLLS

pe Lr © T T T T T T T T T
25r =-
20r- 4

£
s
i-
o I5F 4
o
—
z
o
c
IOF -
5 E
L 1 1 1 L 1 1 L 1 L L L
2 4 6 8 10 I2 14 16 I8 20 22 24

IGURE 8. The influence of light and exogenous
glucose on nitrogen fixation by epiphylls. Samples were
incubated in the dark (solid symbols) or light (open

d water (circles). The
light treatments are significantly different after 4 hours
(P < 0.05) and all treatments are significantly different
after 8 hours (P < 0.01).

latter, in fact, might be quite important in rain-
forest systems: often a large proportion of de-
composition occurs before the dead materials
reach the forest floor (Duvigneaud & Denaeyer-
de Smet, 1970; UNESCO, 1978). Thus, stem-
flow and throughfall water can contain significant
quantities of nutrient ions derived from “‘pre-
fall" decomposition as well as that leached from
living tissue.

o document the effects of mineral nutrients
under field conditions, we performed two types
of experiments. The first was to simply measure
the nutrients present in rainfall, throughfall, and
water flowing off the surfaces of epiphyll-laden
leaves, and the second was to determine if nu-
trients in surface waters iier me mation rates.
We chose to measure pI
simply because these two ions have the most
direct effects on fixation. As can be seen in Figure
9, both phosphate and ammonium are present
in the environment. Interestingly, the water flow-
ing off epiphyll-laden leaves actually has less ni-
trogen than the rainfall collected in the open.
This suggests that the uptake mechanisms on the
leaf surface are extremely efficient. As expected,
the concentrations of these two nutrients were

T -300
300 L x
| f
-200+ 4 100
100 F + 50
O T P O T P
NH, PO,

FIGURE 9. Concentrations of ammonium (ug/ml,
left ris and phosphate (ug/liter, right axis) in open-
collected (O) rainwater, throughfall (T), and from epi-
phyll-laden leaflets of Welfia georgii pinnae (P). Note
the Ri iid high variance (error bars) in the through-
fall value

most variable in water collected as throughfall.
Since these nutrients can affect fixation rates (Fig.
10), the concentrations of nutrients in the water
washing over the leaf surface can be yet another
component adding to the variances of fixation
Observed in the field.

Summary and conclusion. During the course
of this study, we have documented that epi-
phyllous microorganisms can fix significant
quantities of atmospheric nitrogen. This nitrogen
can be absorbed by the host leaf and thereby
contributes to the nitrogen economy of the host.
However, fixation rates are extremely variable
and are strongly influenced by environmenta
factors including substrate (leaf) longevity, light
and concomitant desiccation, both organic and
inorganic nutrients, and by the co-occurrence of
bryophytes. Thus, while fixation by epiphylls can
account for up to 2596 of the nitrogen present in
a host leaf, the contribution of new nitrogen by
epiphylls to an ecosystem is probably fairly small.
Nevertheless, the very patchiness of the process

ANNALS OF THE MISSOURI BOTANICAL GARDEN

en 43: —121.

[Vor. 74
400r
D PO,
ó CONTROL
+ NO,
x NH
300|- N
N
E
o
—
u
z
u 200}
>
=x
=
uj
E x
c ga
100+ x
Zee
1 1 sb 1
6 E 18 24
INCUBATION TIME
FIGURE 10. Effects of nutrient ions on nitrogen fix-

ation by epiphylls. Leaf samples were dipped into so-
lutions containing 176 ug/liter phosphate as NaH,PO,
(D), 429 m n as NaNO, (+), 125 ug/liter
ammonium X), or deionized water (Ó).
Fixation E are d as nM ethylene to document
that fixation continued at a constant rate throughout
the 24-hour incubation.

can free an individual plant from competition
with its neighbors, which in turn could allow
changes in community interactions. In other
words, it is the variance in the system which has
made this study so interesting.

LITERATURE CITED

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CARE. 1980. Effects of er

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& 1984. Direct transfer of newly- fixed
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BERRIE, G. K. & J. M. O. EZE The line
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Burns, R. C. & R. W. F. HARDV. 1975. Nitrogen
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1987]

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Fay, P. 1965. Heterotrophy and nitrogen fixation in
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Foe G.I E. 1973. Nitrogen fixation. Pp. 560-582 ¿n

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Or

FORMAN, R. R. 1975. Canopy lichens with blue-green
algae, a nitrogen PU in a Colombian rain forest.
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FRANKE, W. 1970. vip MN and the cuticular
penetration of leaves. Pestic. Sci. 1: 164-167

GREGORY, P. H. 7]. The leaf as a spore trap. Pp.
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. Nitrogen fixation

. The ecology i nitrogen fixation
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A. B. Viner. 1971. Nitrogen fixation and
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Jones, K. & W. D. P. STEWART. 1969. Nitrogen turn-
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4 y ricus der Pflanzen an

as Klima in den Gegenden der Regnerischen. Ka-

merungebure E ds 47: 353-360.

V. Sweet & H. B. TUKEY. 1956. The
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radio- pp qa Science 123: 1039-1040

LONGMAN, K. A. & J. JENIK. Eni n d Forest
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MAGUE, T. H. 1977. Ecological aspects of dinitrogen
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F. Hardy & A. H. Gibson (editors), A Treatise on
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MULDER, E. G. 1975. ioc s ecology of free-
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ing Micro-organisms. Cambridge University Press,
London

PRESTWICH, G. D. & B. L. BENTLEY. 1981. Nitrogen
fixation by intact colonies of the termite Nasuti-
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RICHARDS, P. W. . The Tropical Rain Forest, an
Ecological I Cambridge University Press,

ndon

Lon

RUINEN, J. 1961. The phyllosphere. I. An ecologically
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1975. Nitrogen fixation in the phyllosphere.

Pp. 85-100 in W. D. P. Stewart (editor), Nitrogen

Fixation by Free-living Micro- "von sms. Cam-
ridge University Se Cambridge.

ies C. E. ndbook for Tropical Biology

n Costa Rica: Data, gd s, Illustrations, and De-
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Ueber bunte Laubblatter, Ein Beitrag
zur Pflanzenbiologie Ann. Jard. Bot. Buitenzorg
13:

A im D. P. 1963. Liberation of extracellular
nitrogen by two nitrogen-fixing blue-green algae.
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1973. Nitrogen fixation by photosynthetic

microorganisms. Ann. Rev. Microbiol. 27: 283-

316.

SuBBA Rao, N. S. 1977. w a es voci as a
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& A. H. Gibson (editors), | ole on Dinitrogen
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1971. Leaching x n io

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& J. V. MORGAN. 1 e occurrence of

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H. B. RUE 1957. Leach-

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(Natural
Resources Research XIV). UNESCO, Paris.

ADAPTIVE RADIATION OF SALAMANDERS IN
MIDDLE AMERICAN CLOUD FORESTS!

DAVID B. WAKE?

ABSTRACT

Tailed amphibians, or salamanders, occur in the tropics only in the New World, where they are
concentrated primarily in Middle America and northwestern South America. All are members of the
family Plethodontidae, e lungless salamanders. As recently as 1926 only 30 species of tropical
salamanders were known, and all were placed in a single genus. Today 11 genera are recognized. All
occur in Middle Pepe and over 140 species have been described. Many local tropical regions are
very rich in numbers of species, and as many as 21 species may be present along a single altitudinal
gradient. Community organization of species of salamanders in the tropics differs from that in tem-
perate regions in that species of tropical salamanders te

es, wl y given species restricted to a narrow elevational band. Within ational cie
are segregated by major ha in) type, then by microhabitat, body size, and finally trophic and behavioral
fi Cloud sa le eleva tions, from about 750 m to 2,500 m, are especially rich in

e proportion of species occurring at lower elevations increases com-
pared with Mexican and Guatemalan transects.

Bromeliads and moss mats in the mid-elevational wet and rain forests are ideal microhabitats for
these insectivorous, direct developing amphibians. Bromeliads offer abundant food resources, egg

America has featured bo th
convergent and parallel evolution. The mid-elevational cloud forests, le their ie iius assem-
blages and highly dissected topography, have been of great significance in speciation, morphological
and behavioral diversification, geographical ecology, and historical l... of tropical pletho-
dontid salamanders.

The cloud forests of Middle America are home
to a unique group of vertebrates— lungless,
climbing salamanders that belong to the family
Plethodontidae. By “cloud forest" I refer rather
loosely to those forest assemblages which form

pending on many local and regional factors ue
as temperature of the water of the nearest ocean,
topography, rainfall patterns, and wind ir
(Grubb & Whitmore, 1966; Myers, 1969).

general, cloud forests form between elevations of
about 800 and 2,700 m. Both upper and lower
boundaries shift with climatic changes, with the
lowest occurring along humid slopes in the low

rs
cu species. Although they do not breed

n water, these organisms nevertheless require
ici conditions for activity, and the cloud forest

' Research reported in this paper has i£ a collaborative effort involving many "n usaspa especially James

whom have given helpful criticisms of the manuscript.

Marvalee

. Pa
H. Wake carefully reviewed the manuscript n: David veia and Nancy Staub also provided useful comments.
d à : à

y
I gratefully acknowledge the cooperation of govern

Museum of Vertebrate Zoology.

nal agencies in Mexico, Guatemala, and Costa Rica in

obtaining permission to See pon in their nee and Douglas Robinson and Pedro Leon for their

help and cooperation in Cost

I dedicate this paper to ee senna of L. C. Stuart, who generously offered his unparalleled knowledge of the
Guatemalan herpetofauna to me, and whose published work remains as an inspiration to future students of

Guatemalan and other tropical environments

? Museum of Vertebrate Zoology ep Department of Zoology, University of California, Berkeley, California
985.

94720, U.S.A. Paper received 7 July

ANN. MISSOURI BOT. GARD. 74: 242-264. 1987.

1987]

environments are effectively buffered from des-
iccating conditions as well as from extremes of
temperature. In addition, these forests typically
support abundant epiphytes that are used exten-
sively by salamanders.

There are more than 140 species of pletho-
dontid salamanders in the New World tropics;
about 80% occur in Middle America (Wake &
Lynch, 1976; Frost, 1985). What makes them
unusual is their great diversity in the New World
tropics and their total absence from the Old World
tropics. Middle America has been the setting for
an extensive, unique adaptive radiation that has
remained very localized.

The success of these species can be measured
against that of salamanders generally. World-
wide, there are about 350 species of salamanders
divided among nine families (Frost, 1985). Eight
families are restricted to North Temperate re-
gions. Over 200 of the species are members of
the family Plethodontidae, the only group of sal-
amanders to radiate in tropical regions. All trop-
ical species of salamanders are members of the
supergenus Bolitoglossa (Wake, 1966), which
contains 11 genera and about 40% of the species
of salamanders in the world. The supergenus is
exclusively New World in distribution and does
not occur north of Mexico.

The plethodontids have a curious distribution,
with two primary areas of evolutionary diver-
sification: North America with concentration in
the Appalachian region, and Middle America
(Fig. 1). In eastern North America are found three
major groups of plethodontids, two of which have
life histories involving an aquatic larval stage.
The Middle American region contains members
of a fourth major group, the tribe Bolitoglossini,
members of which have a uniphasic life history
featuring direct development without a larval
stage (Wake, 1966). The other two supergenera
in the Bolitoglossini occur in California and Or-
egon (and possibly in Alaska and Mexico), and
on Sardinia, the Italian mainland, and a tiny
portion of southeastern France.

I have argued elsewhere that the absence of an
aquatic larval stage facilitated occupancy of the
relatively densely crowded, predator-rich tropics
(Wake, 1966; Hanken et al., 1980). A unique
feeding mechanism and an associated behavioral
repertoire (Lombard & Wake, 1977, 1986; Roth

Wake, 1985a, 1985b), which could evolve its
particular characteristics only in a group lacking
aquatic larvae, may have aided in the successful
radiation of the tropical species. Those animals

WAKE-— ADAPTIVE RADIATION OF SALAMANDERS

243

FIGURE 1. Latitudinal tof
diversity in nthe salamander say Plethodontidae. The
bers g

scribed species for which descriptions are being pre-
pared have been included.

use a highly specialized, extremely fast tongue
projection mechanism to capture moving prey
at a considerable distance, and thus they are able
to feed on a wide array of invertebrate animals.

We still know relatively little about the tropical
plethodontids. The authoritative work of Dunn
(1926) listed but 30 species and placed them all
in a single genus. Taylor (1944) recognized the
generic diversity of the group (seven genera), but
four additional genera were described as recently
as 1983 (Elias & Wake, 1983; Wake & Elias,
1983), and about one-half ofthe 140 species have
been described since 1950. Not surprisingly, most
published work has dealt with taxonomy and
systematics, although there has been some re-
search on life history (Vial, 1968; Houck, 1977a,
1977b) and geographical ecology (Schmidt,
1936a; Martin, 1958; Wake & Lynch, 1976; Wake
et al., 1987)

The present paper attempts to evaluate the role

mation that my research collaborators and I have
gathered over the past 15 years. In particular, I
examine the results of transect studies from Mex-
ico to Costa Rica and concentrate on an area that
appears to have been of critical importance for
salamanders, the mid-elevation cloud forests.

EcoLoGICAL GEOGRAPHY AND SYSTEMATICS

Salamanders in the tropics have diversified in
three major regions, each characterized by a rel-

Q

° Nl ë
ACTIVE SS
saa

IGURE 2. Generalized map illustrating ap of
major evolutionary diversification within t e family
Plethodontidae during Cenozoic times. od major
fault t systems are indicated The family is thought to
h the old and tectonically relatively

adaptiv ve radiation in what is today Middle America,

ith ail the three
core regions indicated: the southeastern margin of the
Mexican Plateau, Nuclear Central America, and Ta-
lamancan Cent ral America. Two ' supergenera with e:

in a reinvasion of temperate North
America through association with land movements and
tectonic activity in the extended San Andreas fault sys-
tem. ore detailed analysis of geological history
in relation to salamander distribution see Hendrickson
(1986).

atively ancient tectonic core, high topographic
diversity, and high tectonic activity along some

Central America (Wake & Lynch, 1976; Fig. 2).
Each region is characterized by species richness
and a high degree of endemism (Savage, 1982).
For example, Chiropterotriton, Lineatriton,
Thorius, and Parvimolge are endemic to region
1, and Pseudoeurycea occurs mainly in region 1
with only a few species in region 2. Dendrotriton,
Bradytriton, and Nyctanolis are endemic to re-
gion 2, and the great majority of the species of
the beta assemblage of the large genus Bolito-
glossa occur there. The few species of Bolito-
glossa beta which occur in regions 1 and 3 are
members of distinct subgroups (see Papenfuss et
al., 1983). The genus Oedipina is centered in re-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FIGURE 3. C kei RAS arboreus, a bromeliad-
dwelling salamander from

meliad specialist: a long prehensile tail, relatively long

legs, large hands and feet with widely spread digits, and

frontally directed eyes. The fine divisions on the scale
m.

gion 3, and only one species extends as far north
as Chiapas. The alpha assemblage of Bolitoglossa
is centered in region 3 and areas to the south,
but a distinct species group extends northwest-
ward to the other two regions. The genus Noto-
triton is something of a puzzle (see below). It has
species in all three regions.

Early zoogeographic biases combined with in-
adequate collecting led to the perception of trop-
ical salamanders as northern invaders that had

"trickled" down into the tropics. Dunn (1926)
placed all 30 species then recognized in a single
genus, and several of his species groups later
proved to us polyphyletic. For example, he united

p, an nd all small,
bromeliad dwellers into another, thereby ob-
scuring the extensive parallelism and conver-
gence that have occurred. Even after Taylor's
(1944) generally progressive division ofthe genus
Oedipus into several genera, the bias of recent

ku...

FiGURE 4. Dendrotriton Vibia a bromeliad-
dwelling ian from the p

Ovando, Chiapas, Mexico. This bro
shares many gross structural s imilarities with species
that occupy similar microhabitats (Figs. 3, 5). The scale
bar is 25 mm

1987] WAKE—
FIGURE 3. dieat . er a bromeliad-
dwellin N Santa Cruz, Za-

capa, Guatemala, in the Sierra den las Minas. Compare
this species with unrelated bromeliad ase in Fig-
ures 3 and 4. The scale bar is 25 m

penetration from the north persisted. The genus
Chiropterotriton as recognized prior to 1983 pro-
vides an example. Its species mainly are bro-
meliad specialists living in cloud forests, and they
are superficially similar in external morphology
(Figs. 3-5). Both Rabb (1960) and Wake (1966)
recognized that members of the genus differed
substantially in osteology, but they chose to in-
terpret this as increasing divergence and spe-
cialization toward the south. This especially in-
structive case is relevant to the main theme o
this paper and is developed further below
Species once assigned to Chiropterotriton are
typically small, slender, ibe tailed, acrobatic
forms, that are s of cloud for-
ests in Mexico, Nuclear Central America, and

T T ER T
105 DU 1400 95* 90
z N
\ PM
` P j
t 5M
: ' ER
4
i t vM 25°
` >
3 aE
Pa Chiropterotriton ==

FIGURE 6. Probable distribution of the genus Chi-
ropterotriton. Most species occur in cloud forests, and
The gen-

a number

eralized ranges were derived by grouping known lo-
calities into suger contiguous units, based on for-
est distribut Se tion gathered by David M.
Darda and i autho

ADAPTIVE RADIATION OF SALAMANDERS

245

Denarotriton

ed to cloud forests. All occupy small geographic ranges,
and ; ccurs in more than one of the iso-
Re indicated here (Elias, 1984).

Costa Rica (Figs. 3-5). Based on their study of
comparative osteology, Lynch & Wake (1975,
1978) recognized that the species below the Isth-
mus of Tehuantepec formed a cladistically dis-
tinct group, which they termed CAiropterotriton
beta. The latter group was found to include at

Nototriton

1
IGURE 8. Distribution of the genus Nototriton. This
genus may not be a monophyletic group. The species

morphological and ecological features. All of the species
inhabit cloud forests, but the

recently discovered and as yet undescribed species.

246

Cerro San Felipe

Pine- Oak
_Fir Forest

GEE 7 »c7/sova/s
ENS Punquidentis

SSSSS P smithi

T. pulmonaris

||

Y

E I maoga

P. cochranae

BSSSS P belli

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

r

t
- M
,XPine-Oak Fir

a
>
Š
E
a
[7]
Cerro Pelón q.
p
A
A
Z

S P smithi
SSS P juarezi

Gl D

m d
Forest

o
a
o

K

a

4

P sp.nov 3

Pine-Oak Forest

Cloud

2000 5

TW

E= Norotriton sp.nov

ENS P wer/er/

eed B rufescens

Forest

SEEN 7 -pE
L^ mm C sp nov

4

°
A
Š
| `
`
ef Tropical Deciduous Forest | a
* Oaxaca | a
oo] — x
l m
ý Evergreen
Seasonal
Forest
`
` Low Deciduous Forest
s
< SW ^ |00 Kms NE —

-northeastward from Ciuda

Distribution of plethodontid salamanders along the Northern Oaxacan transect, about 100 km in
ort d de O

extensive cloud forest which offers habitat to numerous species of salamanders. Updated version of diagram

presented by Wake et al. (1987).

least two distinct subgroups, an interpretation

were given generic status as D
drotriton and Nototriton by Wake & Elias (1983),

who noted that Nototriton might prove to be
paraphyletic. Some species of each of the three
genera resulting from the subdivision of Chirop-
terotriton include members that are very similar
in external morphology. Because the species are
very small (frequently «40 mm in head plus
body length), the critical differences in mor-
phology can be difficult to detect. The species of
the three genera typically have small ranges, and
the widely disjunct ranges offer little od
for symp hese salamanders found
mainly in iced cloud forests, and nl of

sively allopatric (Figs. 6—8). Chiropterotriton oc-
curs as 11 or more discrete geographic isolates
in eastern Mexico, north of the Isthmus of Te-
huantepec. Dendrotritonis found only in Nuclear
Central America, mainly on Pacific slopes, but
also in some internal regions of Caribbean drain-
age. Nototriton is the southernmost of this group,

and it occurs in regions of Caribbean drainage.
An undescribed Oaxacan species of Nototriton

detailed studies of vertical zonation along a tran-
sect (Fig. 9). Our knowledge of the groups prob-
ably remains far from complete, even at the alpha
taxonomic level, but they clearly offer fascinating
opportunities for the study of convergent evo-
lution.

Lynch & Wake (1978) showed that bromeliad-
inhabiting species currently placed in the genera

endrotriton an

drotriton in external morphology (evaluated by
methods of multivariate morphometrics) than to
semi-fossorial congeners that live in moss mats
on soil banks. These distantly related species have
converged so that they share body forms that are
especially well suited for life in bromeliads. In-
deed, if Nototriton is, as I believe, paraphyletic
it is possible that we actually have underesti-
mated the true amount of evolutionary conver-
gence.

1987]

The three genera discussed above are com-
monly encountered inhabitants of cloud forests
and epiphytes. Other tropical salamander genera
also contain cloud forest inhabitants, and many
use epiphytes as their main microhabitats. Some
of these, such as the Nuclear Central American

ered and their habits are very poorly
1984). The monotypic Mexican genera Parvi-
molge and Lineatriton (the latter an elongate fos-
sorial form that utilizes moss mats to some de-
gree; Fig. 11) are relatively rare within their
restricted ranges, which lie at the lower margins
of cloud forests. Some species of the Mexican
genus Thorius occasionally occur in bromeliads;
an undescribed species from the northern slopes
of the Sierra de Juarez in Oaxaca seems to occur
primarily in bromeliads (undescribed species E,
fig. 9 in Hanken, 1983). The remaining genera
(Pseudoeurycea, Bolitoglossa, and Oedipina) have
numerous species that are inhabitants of cloud
fore

P sn is widespread in Mexican and
southwestern Guatemalan cloud forests, but most
species are terrestrial and are not often found in
epiphytes. The only described species that are
bromeliad specialists are P. firscheini (Werler &
Smith, 1952; Shannon & Werler, 1955b) and P
nigromaculata of Veracruzian cloud forests (un-
publ. data, contra Taylor, 1941). An undescribed
species from our Monier) Oaxacan transect (Fig.

9) uses arboreal bitats, and an additional

undescribed species from our San Marcos tran-
sect (Wake & Lynch, 1976: 30) uses bromeliads
consistently.

Bolitoglossa, with 68 currently recognized
species, has by far the greatest geographic range
of the tropical salamander genera (from Vera-
cruz, Mexico, to Brazil, Bolivia, and Peru). Many
of the species in Nuclear Central America are
cloud forest specialists, and they frequently occur
in bromeliads. Over one-half of the B. engel-
hardti encountered during an intensive investi-
gation of an elevational transect on the lower
slopes of Volcan Tajumulco, western Guate-
mala, were found in bromeliads, and B. franklini
is also a frequent inhabitant of bromeliads (Wake
& Lynch, 1976). Most records for Bolitoglossa
in bromeliads refer to members of the beta as-
semblage (e.g., Stuart, 1943). Some of these occur
north of the Isthmus of Tehuantepec (the north-
ern limits of Nuclear Central America), including
B. hermosa (Papenfuss et al., 1983), from the
Pacific slopes of Guerrero, Mexico, and the wide-

WAKE—ADAPTIVE RADIATION OF SALAMANDERS

247

spread Gulf-Caribbean slope species B. rufescens
(Taylor & Smith, 1945), which ranges from San
Luis Potosi, Mexico, to Honduras. The only two
species of the beta assemblage that reach Costa
Rica (B. alvaradoi, B. arborescandens) have been
taken in bromeliads (Taylor, 1954; unpubl. data
Occurrence of members of the large alpha as-
semblage of Bolitoglossa in bromeliads is less
well documented. The distribution of this group

found mainly in the lowlands, and there are no
records of the species being found in bromeliads
in cloud forests. Two members of the group, B.
platydactyla and B. mexicana, have been re-
corded from bromeliads, mainly at elevations of
<500 m (Taylor & Smith, 1945). There are scat-
tered reports of Bolitoglossa alpha in cloud forest
bromeliads in Talamancan Central America and
regions to the south (e.g., B. borburata near Ran-
cho Grande, Venezuela, Trapido, 1942; B. lig-
nicolor, Dunn, 1937; B. subpalmata, Robinson,
1977; B. taylori, Wake et al., 1970). But in Costa
Rica, where the assemblage is well represented,
there are surprisingly few records of its occur-
rence in bromeliads (Robinson, 1977), although
we now know that some species are common in
such microhabitats (see below).

Several species of the alpha assemblage of Bo-
litoglossa are associated with arboreal microhab-
itats in cloud forests. The only known adult of
Bolitoglossa diminuta was collected with an egg
mass in a mat of liverworts (Robinson, 1976;
recent examination of the tiny holotype, which
lacks a sublingual fold, suggests that this species
should remain in Bolitoglossa, contra Wake &
Elias, 1983). Other species associated with moss
mats covering tree trunks and branches include
B. marmorea of Panama (Wake et al., 1973) and
an undescribed species sympatric with B. dimi-
nuta.

The final genus, Oedipina, is widespread in
cloud forest habitats in Costa Rica, the center of
its diversity (Brame, 1968). These salamanders
are elongate, mainly fossorial species that include
some relatively specialized arboreal climbers in
lowland forests (e.g., O. parvipes). The species
that occur at intermediate elevations in cloud
forests typically are found in moss mats covering

owned vegetation and soil banks.

Information in the above paragraphs makes

248

clear that there has been an extensive adaptive
radiation of salamanders in the New World trop-
ics, but the age of this radiation remains un-
known. Since the initial effort of Dunn (1926),
subsequent studies have for the most part sug-
gested progressively earlier dates for the entry of
salamanders into the region (Martin & Harrell,
1957, is a striking exception), and until recently
an Early or Middle Tertiary origin of the group
was accepted (Wake & Lynch, 1976). But bio-
chemical and immunological studies in the last
decade have shown that even within genera there
has been very great genic differentiation, which
implies relatively great age for the separation of
the lineages studied (Hanken, 1983; Hanken &
Wake, 1982; Larson, 1983, 1984; Lynch et al.,
1983; Maxson & Wake, 1981; Papenfuss et al.,
1983; Wake & Lynch, 1982). Progress has been
made in defining monophyletic groups, but I be-
lieve that we have not yet achieved a robust cla-
distic hypothesis for the group (Wake & Elias,
1983), mainly because of the extensive parallel-
ism and convergence that have obscured pat-
terns. Nevertheless, Hendrickson (1986) has at-
tempted to interpret the history of the group by
combining what is known about likely cladistic
patterns with knowledge of the geological history
of the region in a vicariance biogeography study.
He suggested that salamanders which gave rise
to the tropical radiation first separated from those
in the Appalachia area by rifting of an ancient
Maya terrane from Appalachia or by a post-Mid-
dle Jurassic to Mid-Cretaceous marine transgres-
sion. In general he argues for much older times
a slater than previous authors, based both
nts from earth history and from his

belief rois he has not studied these sala-
manders directly) that the extensive radiation of
the tropical salamanders must have taken a long
time. I cannot discuss this provocative study in
detail here, but it is important to understand that
available evidence suggests that salamanders and
habitats have coevolved in areas that became
present-day Middle America for a very long time.

SALAMANDERS AND EPIPHYTES

The epiphytic component of cloud forests of-
fers two major classes of microhabitats for sal-

angiosperms). These microhabitats, particularly
bromeliads, are used on occasion by other ver-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

tebrates: frogs (especially Hyla and Eleuthero-
dactylus), lizards (especially Abronia), and snakes
(e.g., Bothrops schlegeli). However, with the ex-
ception of a few species of frogs whose tadpoles
are clearly adapted for life in the water of tank
bromeliads, only salamanders rely on epiphytic
plants as their main microhabitats, and sala-
manders are far more common in bromeliads
than are any other vertebrates.

The density of salamanders in bromeliads is
difficult to document. As many as 34 Dendro-
triton xolocalcae have been found in a single bro-
meliad in Chiapas, Mexico (Taylor & Smith,
1945), but until recently we have had few quan-
titative data to indicate the frequency of occur-
rence of salamanders in bromeliads. Although
one of the first reports of salamanders living in
bromeliads was from Costa Rica (Picado, 1913),
the general impression has been that salaman-
ders are not common in bromeliads (Robinson,
1977). Bromeliads have been thought to be more
important for salamanders in Mexico, Guate-
mala, and Honduras. In an early account, Gadow
(1908) reported that a Mexican species of Pseu-
doeurycea leads a “‘partly arboreal life, their fa-
vorite hunting and hiding-places being in the
clusters of epiphytic plants, such as tillandsias,
orchids and the climbing phyllodendrons.”’
Schmidt (1936a, 1942) described bromeliad-
dwelling salamanders as relatively abundant in
Guatemala and Honduras, apparently more so
than in Costa Rica and Panama (Dunn, 1937).

In contrast to the above generalization, sala-
manders in Costa Rica use moss mats more com-
monly than do salamanders farther to the north
and west. The genera Nototriton and Oedipina
use moss mats extensively in Costa Rica, but
apparently rarely do so in northern Middle
America. To the north and west Oedipina is
mainly fossorial, and Nototriton is associated
mainly with bromeliads (an exception may be a
poorly known, undescribed species from Chiapas
that has been taken in a moss-covered bank).

There is a general morpaoiogy that Character
izes most bromeliad igs
3-5). They typically are small animals hei
« 50 mm body length) with long, prehensile tails,
long limbs with widely spread digits, and fron-
tally directed eyes. They are acrobatic climbers
and are very adept in a three dimensional en-
vironment. Some larger salamanders use bro-
meliads on occasion, but the true oe usu-
ally approximate the above descriptio

Occupants of moss mats are less PH

1987]

in morphology. In general they are slender and
have relatively short legs. The Appendix con-
tains a list of species which have been reported
to occur in bromeliads and moss mats. It also
contains a few species known by me to have such
habits, but which as yet are not reported in the
literature. This list does not differentiate between
species that specialize on these microhabitats and
those that are but casual occupants.

COMMUNITY ORGANIZATION

For nearly 15 years my colleagues (James F.
Lynch and Theodore J. Papenfuss) and I have
been engaged in a broadly based survey of geo-
graphical ecology and community organization

f the salamanders of Mesoamerica (Wake &
Lynch, 1976; Wake et al., 1987). We established
a series of line transects in Mexico, Guatemala,
and Costa Rica and conducted intensive sam-
pling over a multi-year period along each of them.

r the most intensively studied of these

the continental divide, down t
of Volcán Tajumulco, to the Pacific coastal plain.
Here we have documented the presence of a rich
salamander fauna comprised of 15 species, rang-
ing from near sea level to nearly 4,000 m. This
transect has provided an opportunity to study
the ecological organization of this group of
species, relative to each other and to various
physical and biotic factors. An earlier study (Wake
& Lynch, 1976) presented a general overview of
our results, and a recent report (Wake et al., 1987)
updates the main patterns of distribution. A
summary of our primary results and data rele-
vant to the use of epiphytes by salamanders is
presented here.

Schmidt (19362) collected seven species of sal-
amanders on the slopes of Volcán Tajumulco
and inferred the presence of two additional
species. He outlined their main patterns of ver-
tical distribution and presented comparisons with
the nine species then known from Veracruz,
Mexico. His basic conclusion was that zonation
was sharper in Guatemala than in Mexico.

e (Wake & Lynch, 1976; Wake et al.,

ysi
nized four elevational assemblages of salaman-

WAKE—ADAPTIVE RADIATION OF SALAMANDERS

249

ders, including a lower cloud forest (1,600-2,400
m) group of four and an upper cloud forest (2,400—
2,800 m) group of seven species. These ten species
(one is present in both elevational belts) are re-
stricted not only in elevational zonation, but also
in geographic distribution; none of the species
occurs beyond the limits of the southwestern
Guatemalan volcanoes and the adjacent Sierra
Madre of Chiapas, Mexico. The region between
1,600 m and 2,800 m is occupied by Evergreen
Cloud Forest (above about 1,900 m) and Mon-
tane Rain Forest, using the terminology of Breed-
love (1981), who also has characterized these
formations floristically

Within elevational zones we examined differ-
ential use of major habitat types. For example,
some species are more common in edge situa-
tions, and some favor small clearings and open
spaces, while others are found throughout the

log-dwelling (within and under the bark of logs
and stumps), arboreal (within a leaf-axil micro-
habitat, including bromeliads), and fossorial
(within subterranean passageways). We found no
moss mat specialists and therefore did not rec-
ognize this microhabitat category. Within mi-
crohabitats we paid special attention to differ-
ences in body size and in trophic wampis
(morphology of the Jaws and teeth). The primary
modes of ecological segregation of de t species
are indicated in Table 1. Here a one-sided matrix
of potential co-occurrence of the species records
our assessment of primary segregation ordered
according to decreasing spatial proximity of the
segregated species: elevation, habitat, microhab-
itat, size, and trophic specialization. Nearly three-
fourths of the potential sympatric associations
of the 15 species are precluded by differences in
elevational distribution, and habitat or micro-
habitat differences separate all but nine of the
remaining paired associations. Of the nine pairs
of species which show elevational, habitat, and
microhabitat snp cight differ importantly
by size
Smari in size and morphology, ekteni that ne
species has about halfas many substantially tp
er maxillary teeth and enlarged jaw muscles.
Six species on the transect commonly occur in
arboreal microhabitats (in order of frequency in
such microhabitats, from most frequent to least):
Bolitoglossa occidentalis, Dendrotriton bromelia-

Primary modes of ecological segregation among species that occur along the San Marcos transect. S
appearance from high to low elevations. In order of decreasing spatial proximity: E
specializations. From Wake & Lynch (1976).

TABLE 1.

pecies arranged in approximate order of

habitat; M

250

— size; T — trophic

microhabitat; S

elevation; H =

P. rex

1.

2. P. sp.

(E)M

3. P. brunnata

5. B. rostrata
6. B. resplendens

7. D. bromeliacia

ANNALS OF THE MISSOURI BOTANICAL GARDEN

(H) M

9. B. franklini

13. B. salvinii
14. O. ignea

15. B. flaviventris

[Vor. 74

ad
surface of the soil; I = under bark of stumps and stand-

ing trees, and inside fallen logs. Open symbol = Pseu-
doeurycea; half-closed symbol = Dendrotriton; closed
Sy — Bolitogl

mbers 2 through 10, and 12, are cloud forest in-
habitants E from Wake & Lynch (1976).

cia, B. engelhardti, B. franklini, Pseudoeurycea
sp., and B. resplendens (Fig. 10). All but the first,
which occurs at relatively low elevations, are
present in the cloud forest and make extensive
use of bromeliads. Bolitoglossa occidentalis is a
small, mainly lowland species that occurs in bro-
meliads, but it is most commonly found now
within the so-called "coffee zone," where it oc-
curs in agricultural plantings of bananas.

James F. Lynch and I are preparing a detailed
ecological account of our work in this transect,
and with his permission I present here some of
our data concerning use of bromeliads in cloud
forests by the above listed species. When we first
visited this area in 1969 primary forest extended
to roadside and bromeliads were abundant. When
we last visited the cloud forest region in 1980
the forest had been removed and pasture occu-
pied nearly the entire area between 1,500 m and

00 m. Below 1,500 m traditional coffee plan-
tations, which feature large shade trees and ex-
tensive plantings of bananas, had given way to
a near monoculture of coffee grown in hedgerows
without any suitable cover for arboreal salaman-
ders.

1987]

TABLE 2. Relative abundance of salamanders in bromeliads, San Marcos transect.

WAKE—ADAPTIVE RADIATION OF SALAMANDERS

Elevation Wet Season! Dry Season? Combined
Below 1,750 m — 3/25 = 0.123 3/25 = 0.12
1,750-2,000 m 6/39 = 0.15 41/75 = 0.55 47/114 = 0.41
2,000-2,250 m 48/239 — 0.20 29/64 — 0.45 71/303 — 0.25
2,250-2,500 m 89/121 = 0.74 140/191 — 0.73 229/312 = 0.73
2,500-2,750 m 81/25 = 0.72 27/30 — 0.90 45/55 — 0.82
Above 2,750 m 14/73 = 0.19 13/21 = 0.62 27/94 — 0.29

! May-September.
2 November- Fe ry.
3 Number of salamanders/number of bromeliads.

Salamanders are common inhabitants of bro-
meliads along the San Marcos transect (Table 2).
We found salamanders in approximately every
second bromeliad we opened. These bromeliads,
primarily members of the genera Tillandsia and
Vriesia, were located relatively low in the trees.

Salamanders in southwestern Guatemala are
most abundant in bromeliads at elevations be-
tween 2,250 and 2,750 m. From these elevations
down to approximately 1,700 m, bromeliads re-
main relatively common, and there are brome-
liads present at elevations up to approximately
3,000 m. All of the bromeliad specialists occur
in the cloud forest (roughly 1,500-2,750 m), even
though bromeliads are found both above and
below that formation. Above the cloud forest
those species that use bromeliads at lower ele-
vations (e.g., Bolitoglossa rostrata, Pseudoeury-
cea rex) shift almost entirely to terrestrial mi-
crohabitats. Salamanders are consistently more
abundant in bromeliads during the dry season
than during the wet season, except in the heart
of the cloud forest (2,250-2,500 m), where there
is less seasonality than elsewhere.

ABLE 3. Distribution of salamanders in brome-
liads, San Marcos transect, 16 January 1972, 2,400 m
elevation.!

Number of Salamanders
r Bromeliad

Frequency

0 15
l 10
2 8
3

4 1
5 l
6 1
8 1

Martin Feder accompanied us on one trip to
our transect and studied the thermal ecology of
some of the cloud forest salamanders pies
1982). He found that bromeliads, even in the
cloud forest, afford cooler and more tans tem-
peratures than microhabitats in immediately
surrounding areas. Bromeliad-dwelling salaman-
ders appear not to thermoregulate behaviorally
or k ario because thermal diversity in
their microhabitats is so low as to offer little
opportunity for such behavior. As is usual for
salamanders, there is a high correlation between
body temperatures of salamanders and prevail-
ing microenvironmental conditions (Feder &
Lynch, 1982), so the more stable the microen-
vironment, the less variable will be the temper-
ature of the salamander. The tropical salaman-
ders contrast sharply with more northern
plethodontids in having very limited ability to

undergo thermal acclimation (Feder, 1978, 1982).
Ibis may be either us Fuse Or ins eflect ar the
high fidelity t
displayed by many of these species (Feder, 1983).

The data and analyses in Tables 3-5 indicate
that the distribution of salamanders per bro-

TABLE 4. Distribution of salamanders in brome-
liads, San Marcos transect, 18 January 1972, 2,300-
2,350 m

Number of Salamanders

er Bromeliad Frequency
0 12
l 8
2 10
3
4

7
l
5 2

! These data are for 55 Dendrotriton bromeliacia and
three Bolitoglossa franklini taken from a sample o
bromeliads (X — 1.45 salamanders per bromeliad).

! These data are for 59 Dendrotriton bromeliacia and
four Bolitoglossa franklini collected in a sample of 40
bromeliads (X = 1.58 salamanders per bromeliad).

252

TABLE 5. Test for randomness, combined data from
Tables 3 and 4.'

Number of
Salamanders Expected
per Observed Frequency
Bromeliad Frequency (Poisson)
0 27 17.5
l 18 26.6
2 18 20.2
3 10 10.3
4 2 3.9
r fH. 2
6 7 0.3 5.5
7

1 >
8 l J

Chi-square (goodness of fit) = 8.50 with 4 df
0.05 > P > 0.1

ANNALS OF THE MISSOURI BOTANICAL GARDEN

' Data for 121 salamanders collected from 80 bro-
meliads (X = 1.52 salamanders per bromeliad).

.

meliadisnotsi different from random.
— p there is at least a suggestion that there
t be an excess of bromeliads that lack sal-
de as well as a deficiency of bromeliads
containing single salamanders. Thus, there might
be a tendency toward clumping under conditions
of high salamander abundance
romeliads might seem to be a near perfect
microhabitat for salamanders, in terms of food
availability, thermal stability, and constancy of
humidity. But concentration of salamanders in
bromeliads might attract predators. Spiders and
salamanders are the top resident carnivores with-
in bromeliads, and they do not prey on each other
very extensively. Some arboreal snakes forage
widely and may be important, although infre-
quent, predatory visitors to bromeliads; birds
might also be important predators. All tropical
salamanders have a specialized autotomy zone
at the base of the tail, and Shaffer (1978) ex-
amined tail loss frequency as a rough index of
relative predation pressure on 10 species of sal-
amanders from the San Marcos transect (parts
of tails may be lost in intraspecific aggressive

lation between rates of tail loss and elevation,
and we know that snake densities also decrease
with elevation. Two of the three cloud forest
species commonly found in bromeliads (Den-
drotriton bromeliacia, Bolitoglossa franklini) had
the second and third highest tail loss percentages
of the species studied (the highest was the ar-

[Vor. 74

Outlines of the left hind foot of two

FIGURE 11.

(this ne is 67.5 mm, snout-vent length) with
extensive webbing and is capable of generating suction
in arboreal ano Bolitoglossa occidentalis is a di-

minutive species (this specimen is 38.7 mm, snout-
vent length) with feet that superficially appear to be
webbed, but in reality are just incompletely developed
(paedomorphic) as suggested by the strong gradient in
phalangeal structure within each digit.

boreal lowland species B. occidentalis). Although
exposure to predation may be a relative cost for
living in bromeliads, the cost is apparently out-
weighed by Sa advantages, such as those men-
tioned abov
Although Diss dwellers in cloud forest
habitats differ greatly in morphology, ecology,
and behavior from even the most arboreal North
American plethodontids (e.g., Aneides), they are
not the most extremely specialized species on the
San Marcos transect. At elevations below about
1,400 m the cloud forests, and the lower cloud
forest salamander fauna, are absent. A new sal-
amander community appears at about 1,000 m
composed of three extreme morphological and
ecological specialists. This community includes
a relatively large and a relatively small arboreal
species of the genus Bolitoglossa and an elongate
fossorial species of the genus Oedipina.
olitoglossa salvinii, a large species, is an ac-

tive, climbing animal with a long prehensile tail
and large hands and feet with extensive inter-
digital webbing (Fig. 11). These animals, which
frequent surfaces of Heliconia and other large-
leafed plants on moist evenings, are capable of
producing suction with their large hands and feet
(Alberch, 1981).

Bolitoglossa occidentalis, the small species, has
small hands and feet that appear to be fully

ebbed. In reality the hands and feet manifest
incomplete development, a phenomenon known

f

1987]

as paedomorphosis that affects a number of fea-
tures of the organism (Wake, 1966; Wake &
Brame, 1969; Alberch et al., 1979; Alberch &
Alberch, 1981). Such similarly affected features

with digits that show a strong gradient of devel-
opment (Fig. 11). Although these animals do not
generate suction with their hands and feet (AI-
berch, 1981), they are very agile, partly as a result
of their small size. They have an extensive ven-
tral surface area (body, limb, and tail) relative to
their mass, so they "stick" to moist plant surfaces
by surface tension. Apparently these salaman-
ders are virtually restricted to leaf axil retreats.
They especially favor Heliconia and both culti-
vated and feral banana plants (Musa spp.). Smith
(1945) reported finding Bolitoglossa rufescens

ery similar in morphology and ecology to its
close relative B. occidentalis) to be abundant in
red bananas (he found about 250 per hour) at a
Veracruzian site. These animals are adept at
climbing small tendrils, stems, and strands of
moss.

Members of the small species of Bolitoglossa
are encountered only rarely in terrestrial sites,
and the large species, while occasionally found
on the ground (e.g., crossing roads on rainy
nights), also are basically arboreal. (An exception
may be the very large species B. dofleini, which
can be very common in terrestrial situations.)
But another group of lowland species, the very
elongate, short-legged genus Oedipina, is found
only at and beneath the surface of the ground.
Species of Oedipina are elongated as a result of
the addition of trunk and especially tail verte-
brae, and they have bizarre long tails (Fig. 12).
In addition to their extremely short legs and tiny
hands and feet, they have heads and bodies of
very small diameters; all of these features facil-
itate use of root channels and underground bur-
rows.

The San Marcos transect is special because
more species of salamanders occur there than in
any other area of the Pacific Versant in Middle
America. Volcan Tajumulco (4,200 m) is at-
tached to the Guatemalan Plateau at about the
3,000 m level, so on both sides of the volcano
there are substantial areas of moderate elevation
which trap moisture and thereby create favorable
salamander habitat. To the south and east the
Plateau gradually lowers and rainfall declines. In
the vicinity of Guatemala City, Volcan Agua (over

WAKE—ADAPTIVE RADIATION OF SALAMANDERS

FIGURE 12. eg d bove (top), from near
o de las Flores, Ver z, Mexico, and Oedipina
Le (bottom), from Fis a Julia, near San Rafael
de la Cuesta, San Marcos, Guatemala. The scale

bul r is 25 mm. These extremely elongated species are
semi-fossorial to fossorial in habit, and have evolved
convergently.

3,700 m) towers over the Plateau, to which it is
attached along only its northern flank at a low
elevation (about 1,000 m). The contrast in sal-
amander faunas between Volcán Tajumulco and
Volcán Agua is great. Only three species have
been collected on the slopes of Agua. There is a
single high elevation member of the genus Pseu-
doeurycea (P. goebeli), one low elevation species
of the genus Bolitoglossa (B. salvinii, a large, fully
webbed species), and a middle elevation gener-
alized species of the genus Bolitoglossa (B. mo-
rio). All three of these species also are present on
the San Marcos transect. There is a well-devel-
oped cloud forest with abundant bromeliads on
Volcán Agua, but the forest is localized and it is
isolated from similar habitats to the north and
west by low elevations covered by drier vege-
tation types. In 1969 we opened about 600 bro-
meliads on Volcán Agua, but found only two
salamanders! This contrasts sharply with the data

areas of the Guatemalan Pl e
Marcos transect it is found only at the top of the
cloud forest and in drier broadleaf forest above,
where the edge of the Plateau contacts Volcan
Tajumulco. Here the species occurs occasionally
in bromeliads (Fig. 10). On Volcan Agua, al-

ough B. morio remains uncommon in bro-
ae and apparently has not modified its mi-
crohabitat utilization patterns in any dramatic
way, its elevational range is 1,300-2,500 m. At
this site, where B. morio is the only salamander

e
a

Q oS "mA S
T f J 4
= NM
J } f nm ghe

COSTA RICAN TRANSECTS
o Q 25 p 100 i50

eM

FiGURE 13. Map of Costa Rica indicating the lo-
cation of the Irazü and Tapanti transects illustrated in
Figures 14 and 15.

present, it occupies an elevational range that
accommodates ten species on the San Marcos
transect. The patterns of species distribution in
the tropics (this example, but see below also) are
probably determined by combinations of
physiological constraints (in relation to physical
factors in the environment) as well as such in-
terspecific interactions as predation and com-
petition

COMPARATIVE ASPECTS OF
COMMUNITY ORGANIZATION

Until recently most fieldwork by my group has
been in Mexico and Guatemala (for general sum-
maries see Wake & Lynch, 1976; Wake et al.,
1987). In general, the results of our detailed stud-
ies of the San Marcos transect have been mir-
rored in other areas (cf. Fig. 9). Typically only
one or two species occur at elevations above 3,500
m; as one moves lower the number of species
present in a given habitat increases dramatically
at about 3,000 m and continues to be relatively
high until the lower limit of the cloud forest is
reached. At elevations below 1,000 m the num-
ber of species present typically declines, and at
sea level the largest number of species definitely
known to be present is four on the Osa Peninsula
of Costa Rica, where there are two species o
Bolitoglossa and two species of Oedipina. Pos-
sibly five species occur together at sea level in
the region of Bocas del Toro, Panama (Wake et
al., 1973 and unpubl. data), and in northeastern

M

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Costa Rica seven species are known from ele-
vations below 1,000 m, but not in local sympatry
(see below). The proportion of the total sala-
mander fauna present in the lowlands increases
at lower latitudes within Middle America (see
Table 5 in Wake & Lynch, 1976). There is evi-
dence of increased *'tropicality" (that is, of closer
packing of species in communities and increased
numbers of species present locally) as one moves
into the deep tropics. Finally, however, in north-
western South America this trend stops, perhaps
because plethodontids are not thought to have
dispersed into South America until late Pliocene
times (Wake & Lynch, 1976; Hanken & Wake,
1982). Except in the Chocó and the flanks of the
northern Cordilleran regions of Colombia, the
number of salamander sp tinlowland
sites in South America is not known to exceed
two, even in such biotically rich 1m as those
of the Río Palenque area of Ecuador.
Throughout Middle America, ai cadens are
found in cloud forests. In Mexico and in Nuclear
Central America, cloud forest salamanders make
extensive use of bromeliads as microhabitats,
even at the extreme northeastern limits of the
range of the supergenus Bolitoglossa in the Gó-
mez Farias region of Tamaulipas (where cloud
forests also reach their northern limit; Martin,
1958). But the manner in which cloud forest mi-
crohabitats are utilized changes in Costa Rica.

the tropics, thanks primarily to the work of Tay-
ie d 1954). However, very little has been

utilization and

patterns of co-occurrence of Costa Rican species.
Only four species are reported to occur in bro-
meliads (Robinson, 1977), although many more
species are known to inhabit cloud forests.
Recently I have been investigating the system-
atics and distribution of Costa Rican salaman-
ders in some detail. I have focused attention on
two general transects (Figs. 13-15). Results for
Costa Rica are preliminary, because major sec-
tions of these generalized transects have yet to
be searched thoroughly. Nevertheless, certain
marked contrasts with more northerly transects
are apparent.
dominant theme in the history of studies of
tropical salamanders has been that the species
which are the most conservative ecologically and
the most primitive phylogenetically occur in
Mexico, and that there is both increased spe-
cialization and a decline in the number of species

1987] WAKE—ADAPTIVE RADIATION OF SALAMANDERS
Volcan Irazu FR
Montane
30004 $ Wet Forest_
Ù m
S °
£ °
s > £
Š S Hs
4 = 8 &
S 3
` D
e `
` $ Qr
< X
2000 Q Š Š Š
Š S Q S S
Š SOR 3
S S Q à
o Ş ° >
S :
9 Š q
Š Š | Forest
x D t.
S S A, =
Å 3 Lr
10004 S n € da
N X ts f Premontane Rain
S 9 : Forest
Ss
500- = = EE MAN
&
o9
o Tropical Wet and
° O COE:
= i! | ce ERAN root Premontane Wet Forest
O
= V ee RA TNE T t
o 4
20 Ki O
FIGURE 14. Vertical distribution of pula salamanders along the Irazü transect, extending about 55
km from Finca La Selva to Volcán Irazü, Costa Ric

to the south (Dunn, 1926; Taylor, 1944, 1952;
Brame & Wake, 1963; Wake, 1966; Wake &
Lynch, 1976). Even though Wake & Lynch (1976)
documented the presence of a large number of
species below 10? latitude, we continued to es-
pouse the traditional view of relics in the north
(e.g., Chiropterotriton priscus of Coahuila and
Nuevo León, Mexico) and increasingly derived
forms to the south. The recent discovery of the
most primitive known genus of tropical pletho-
dontid in Guatemala (Elias & Wake, 1983) and
a fresh analysis of relationships of new and ex-
isting groups (Wake & Elias, 1983) have forced
me to re-evaluate my earlier views. It now seems
likely that salamanders have been in the tropics
of present-day Middle America for a very long
time, possibly throughout the Tertiary (see above
section on Ecological Geography and System-
atics). Middle elevations doubtless have been
important areas of both survival and radiation
in the group, and we have discovered that pat-

terns of microhabitat utilization differ dramati-
cally between Mexico and Nuclear Central
America on one hand, and Talamancan Central
America on the other.

Perhaps the most striking difference between

lier (Table 2), salamanders are sometime
abundant in cloud forests of northern Middle
America, where many species utilize bromeliads
(Wake & Lynch, 1976). In Veracruz, Mexico, and
in Nuclear Central America, elongate, fossorial
species of Lineatriton and Oedipina occur only
at elevations below the lower limit of cloud for-
est. In contrast, elongate members of the genus
Oedipina are well represented in Costa Rican and
Panamanian cloud forests, extending upwards to
elevations in excess of 2,000 m (Figs. 14, 15).
Here they utilize moss mats covering soil banks,
downed logs, and stumps. Furthermore, living

256 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

E &
Š Cerro de
Š la Muerte y) Y 2 à
E È Ad a
X. f
Š $ ae
3000 s XS
S
a s o <,
S I
Q Š ° q Montane
X 3 £5 Rain
4 © ë Š Á Forest
E E Qs Um -
$Š Saan $
Bs S D
š ES
Š & Lower
20004 S Montane
— ES š y ain
i S Š y ^
& S Sk x 3
NEUEN 82 _ 88 3s8h 8 Í Forest
š Sei] Š Se Fes ech s Lm
š ] $£| Š š Š ° š Š Š ° à p
= Š Š š Š SQ Š m Qi £
2 8 $ S ü SFA 73
Š Qj E] € Premontane
10004 : S a 4 p i 1
BS S
~ 5 Y & Premontane »,
š :
S š S ü Rain Forest Ma | OR
SP £ act
5004 $ Š i
8S 9$ Ar Tropical | Moist Forest
mo Š y^ Wet Forest
S 2
200 e Ü s |
a [| — Sines Tropical Moist and Premontane Wet Forest
T T T T T
40 60

O 20

FIGURE 15. Vertical distribution of PN aig p ege med a vepres Tapantí transect, which
a Ric 6

extends between Siquirres and Cerro de la Muert

a. This re EE 0 km long, contains more

species of plethodontid salamanders than any bou area in al ann

in moss mats in these cloud forests are species

is even an undescribed Costa Rican species of
Bolitoglossa that has been taken only in moss
surrounding twigs on trees and shrubs in cloud
forests. This substantial fauna represents an eco-
logical component that is rare (e.g., Dendrotriton
cuchumatanus occurs in moss mats in Guate-
mala, Lynch & Wake, 1975) or is missing in
cloud forests of Nuclear Central America.

he two Costa Rican transects offer some in-
teresting contrasts. Both extend up the Caribbean

about 55 km to the peak of Volcán Irazü. Sala-
mander distributions along this transect are
poorly known except for the region between 1,000
and 2,500 m. For example, Scott et al. (1983)
listed two species of Oedipina from Finca La
Selva, but study of specimens from the area sug-
gests that three occur there. The taxonomy of the
group is difficult, but it may be that none of the
three are the species listed. (Note that only two
species are indicated on Fig. 14.) There are pres-

ently 11 species of salamanders (in three genera)
known from this transect. As with the more
northerly transects, only one species (Bolitoglos-
sa subpalmata) is likely to occur at elevations o

7 3,000 m (and I cannot document its presence
at that elevation as yet, although I expect it may
be found) However, in contrast with more
northerly transects (cf. Wake et al., 1987), there
are at least five species present at elevations of

species of Oedipina), and two Bolitoglossa (B.
snags a nee that I have found in moss
er microhabitats ds = moe
"es sa P p )
and B. alvaradoi use bromeliads:
I expect that more species will be found on the
ds transect, in part because of the unusually
umbers of species present on the Tapantí
iue (Fig. 15). The latter transect 1s far more
eneralized in its boundaries than the first, and
ie essentially a broad (ca. 20 km) swath of ter-

1987]

ritory extending about 60 km from the vicinity
of Siquirres to the summit of Cerro de la Muerte.
The Tapanti transect is not a straight line, but
twists somewhat to encompass sites where some
of the rarer species are known to occur. There is
extensive habitat disturbance along this transect
(for example, in the vicinity of Turrialba), and I
justify weaving together an indirect transect as
an attempt to demonstrate the potential number
of species one might reasonably expect to find
on a continuous altitudinal transect under pris-
tine conditions. The Tapanti transect thus offers
a greater diversity of habitats than does the Irazü
transect and one might expect more species to
be present. To my surprise, the Tapanti transect
is the richest that I have found in the tropics,
with 21 species. There are only 26 species of
plethodontids known from Costa Rica (Scott et
al., 1983; their list and mine differ slightly but
we obtain the same total number of species), and
that about 80% of them occur along this transect
attests to its richness

The Tapanti anadi has a higher number of
species (seven) occurring below 500 m than does
the Irazú transect, and in both Panama and Nic-
aragua another species (Oedipina collaris) occurs
at elevations of «500 m (Brame, 1968), so there
is the likelihood that an eighth species eventually
will be found. Bolitoglossa alvaradoi apparently
occurs at elevations of < 500 m elsewhere in Cos-
ta Rica. Thus, as many as nine species might be
expected in the lowlands in the area crossed by
this transect.

Thirteen species occur in the cloud forest of
the Tapanti transect, if we accept 750-2,000 m
as its elevational bounds. In fact, cloud forest
conditions exist almost to 3,000 m, although in
general cloud forests are less well defined in Costa
Rica and Panama than farther to the north
(Myers, 1969). This transect contains the richest
cloud forest salamander fauna found anywhere
in the tropics. Here, too, I have probably under-
estimated the number of species present. At least
one more species of Bolitoglossa may be present,
and the two poorly known highland species of
Oedipina may well extend to lower elevations,
since the genus as a whole is strongly concen-
trated at lower elevations. Two species of Boli-
toglossa (an undescribed species and B. dimi-
nuta, Robinson, 1976) use mats of vegetation
including mosses and liverworts that surround
twigs and branches of trees. Several other species
use bromeliads and moss mats, but no quanti-
tative data are available. In the forests of Refugio

WAKE—ADAPTIVE RADIATION OF SALAMANDERS

257

Tapanti there is an especially rich epiphyte fau-
na, and this is the locality where the largest num-
ber of species are found. However, in marked
contrast to the situation in the cloud forest on
our San Marcos transect, the density of individ-
ual species is uniformly low on this transect. This
situation of high species diversity and low den-
sity of individual species is one more indication
of the increased “‘tropicality” of the Costa Rican
salamander fauna.

The contrast between the cloud forest that oc-
curs at around 1,000 m and the forest around
3,000 m on this transect is sharp. At high ele-
vations the density of Bolitoglossa subpalmata
is extraordinarily great, on the order of 9,000/ha
(Vial, 1968). Four species occur in sympatry at
around 2,500 m; in my experience B. subpalmata
is about 100 times more common than B. cer-
roensis, about 1,000 times more common than
B. sooyorum, and about 10,000 times more com-
mon than B. nigrescens! Doubtless there is a col-
lecting artifact involved, but the first species is
remarkably abundant and the last has been, at
the very least, elusive. The high density of a sin-
gle species at high elevation is a common theme
in tropical salamander biology.

Bolitoglossa subpalmata, which can be ex-
ceedingly abundant at high elevations in Costa
Rica, displays a marked shift in microbabitat

tlower

1

utilization and a reduction in

elevations. The species is primarily ground-
dwelling at high elevation, although it also uses
bromeliads. I have collected B. subpalmata from
bromeliads 30 m above ground level in an oak
tree at 3,000 m elevation. At elevations of «2,000
m the species becomes increasingly common in
bromeliads and is encountered only infrequently
in terrestrial situations. This species can be ex-
traordinarily persistent in the face of even drastic
habitat change, so long as bromeliads remain.
An anecdote illustrates this point. An old col-
lecting locality in the Montes de Aguacate, west
of San José, was visited recently. Only tiny frag-
ments of forest remain at this site, at about 1,500
m. Salamanders were common residents of bro-
meliads in one forest fragment where trees were
being felled. We opened 130 bromeliads and
found 55 salamanders, including 14 adults and
a subadult in a single bromeliad. As many as
three adults were found in the axil of a single
leaf. This is a graphic demonstration of the suit-
ability of bromeliads as microhabitat for sala-

anders, especially under conditions of great
habitat modification. I have had similar expe-

258

riences with Chiropterotriton lavae in fragments
of cloud forest above Jalapa, Veracruz, Mexico,
and with Bolitoglossa morio in an area devas-
tated by volcanic activity on Volcan Pacaya,
Guatemala

It is still too early to generalize very extensively
from our comparative transect studies.We have
come to be suspicious of species with broad el-
evational ranges (such as O. uniformis, Figs. 14,
15), and these deserve careful taxonomic re-eval-
uation, for most species occur within rather nar-
row elevational limits. We also have come to
expect few extreme highland or lowland species,
but there are more lowland species at low lati-
tudes. Cloud forests and salamanders are most
abundant at mid-elevations.

FACTORS INFLUENCING SALAMANDER
SPECIATION AND RADIATION

Species of plethodontid salamanders charac-
teristically display a high degree of genetic frag-
mentation (Larson et al., 1984; Larson, 1984).
Larson (1984) has argued that a general pattern
for the history of population structure in pleth-
odontids is that following origination they ex-
pand gradually and contiguously into regions to
which they have ecological access. Later, as a
result of climatic E which may result from
many p causes, their populations become
fragmented into ands among which there is
little or no genetic exchange. Subsequent climatic
changes may lead to re-establishment of ecolog-
ical access to areas separating isolated popula-
tions. This may lead either to renewed genetic
exchange or, depending on the level of genetic
divergence and its effect on isolating mechanisms
or mate recognition systems, to a variety of in-
teractions. There may be a hybrid swarm, a nar-
row hybrid zone, a narrow zone of overlap with
occasional hybridization, partial sympatry with
boundaries set by competitive interactions, or co-
existence with different ecological requirements.
All of these interactions have been documented
among plethodontids. The many studies of pop-
ulation structure based on electrophoretic anal-
ysis of proteins, together with studies of distri-
bution, phylogenetic history, and biogeography
(reviewed by Larson, 1984), suggest that geo-
graphic or allopatric speciation by subdivision is
the common mode in plethodontids. There are
similarities with the concept of “taxon cycles”
(Wilson, 1961; Ricklefs & Cox, 1972), which
usually have involved examples from island

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

chains. These postulated cycles of expansion and
contraction of geographic ranges and population
densities would establish the setting for the kind
of speciational processes I envision having taken
place among the tropical salamanders.

o primary factors in the history of the

tropical salamander fauna probably have been
the combination ofadaptation of species to cloud
forest environments and the long, complex his-
tory of tectonic activity in Middle America. There
are some relatively stable areas, such as the an-
cient core of Nuclear Central America, where the
upper and lower boundaries of the cloud forest
have shifted, but where cloud forest in a broad
sense probably has been present for much of the
Tertiary and Quaternary periods (see discussion
in Wake & Lynch, 1976). However, there are
many other areas where tectoni ts and
associated volcanic activity have been so great
and so persistent that cloud forests have been
only an ephemeral presence, shifting almost con-
stantly.
The three foci for salamander radiation in the
tropics (Fig. 2) all have stable, ancient tectonic
core areas surrounded by regions of great tectonic
activity. (For summary of tectonic history, see
Hendrickson, 1986.) These areas have been cited
by other workers as having phylogenetically dis-
tinctive faunal components. For example, Sav-
age (1982) stated:

I now believe that the distinctive montane her-
petofaunas of the southern Sierras of Mexico, Nu-

more or
less similar sea of widely distributed ancestors.

Nuclear Central America (Fig. 16) illustrates
the ideas outlined above. That region has as its
core the ancient Sierra de los Cuchumatanes, lo-
cated on the southern part of the North d
plate. The area has been characterized as th
Middle American Megathrust by Plafker (1976),
because it is the conjunction of three major plates.
The Cocos plate, to the southwest, is being sub-
ducted near the intersection of the North Amer-
ican and Caribbean plates (for evidence of the
widespread influence of this phenomenon see
Singh et al., 1985). The latter is being forced
eastward by the combined plate movements, and
a small western tip of the plate is effectively
"caught" between the North American and the
Cocos plates. The zone between the Caribbean

1987]

WAKE—ADAPTIVE RADIATION OF SALAMANDERS

259

Isthmus
of
| Tehuantepec

MEXICO

HONDURAS

Inspired by and based largely on Plafker (1976).

and North American plates is outlined by the
Motagua and Polochic fault zone. This has been
an area of intense tectonic activity for millions
of years and Plafker (1976) has estimated that at
least 200 km of lateral movement of plates along
this fault zone has occurred since the Miocene.
The western tip of the Caribbean plate, trapped
between the other plates, is being ripped, or de-
coupled (Plafker, 1976). Grabens have formed,
with small volcanic cones rising within them.

volcanoes are lined up. Volcan Tajumulco and
Volcan Tacana lie at the northwestern corner of
the Caribbean plate, where the three plates meet.
In this topographically complex zone of maximal
geological turbulence, the largest number of co-

ins of the core region are areas

in large part to the fragmentation and reassembly of areas of cloud forest.

occurring species of salamanders in Middle
America is found, along our San Marcos transect.

The relatively stable upland of the Sierra de
los Cuchumatanes is in many ways an even more
interesting area than the Pacific volcanic belt. We
began fieldwork in this area in 1974, at a time
when only two species of salamanders were
known from the Caribbean slopes of the Cuchu-
matanes, despite a number of brief LUIS trips
by different herpetologists. Results of our inves-
tigations have been summarized by Elias (1984),
who found 13 species of salamanders in this re-
gion (see also Wake et al., 1987). As elsewhere
in Middle America, the cloud forest is of special
interest, for six species with narrow elevational
ranges occur just above the cloud line, here lo-
cated at about 1,300 m. Two new genera of sal-
amanders were discovered in this cloud forest

260

(Elias & Wake, 1983; Wake & Elias, 1983), in-
cluding the exceptional Nyctanolis, a morpho-
logically primitive genus which appears to be the
sister group of all other tropical salamanders.
Thus, on the one hand, the ephemeral cloud for-
ests of the tectonically active and topographically
complex margins of the Cuchumatan uplands
have contributed to speciation and led to the
highly disjunct distributional patterns illustrated
previously for Dendrotriton and Nototriton (Figs.
). On the other hand, the more stable cloud
forests on the northeastern slopes of the core of
the Cuchumatan region have served as refugia
for what must be extremely ancient lineages.

CONCLUSIONS

We still have much to learn about the sala-
manders of the New World tropics, and even the
best known areas of Middle America have yield-
ed many recent surprises. Earlier misconceptions
concerning the probable history and ecology of
tropical salamanders have led to underestimates
of the age and diversity of the group and have
contributed to our relative ignorance of the ecol-
ogy of the cloud forest and lowland species, es-
pecially the arboreal and fossorial forms. New
species are being discovered more rapidly than
they can be described, for many species are known
from small series. Our knowledge of the com-
parative osteology and of molecular evolution of
this group, while still fragmentary, is sufficient
to demonstrate that parallelism and convergence
are rampant.

This, in turn, implies both that there may be
only a limited number of ecological roles avail-
able to tropical salamanders, and that there may
be functional and developmental-historical con-
straints which impose limits on the evolutionary
potential of the group. An especially clear case
of convergence is the elongation associated with
fossorial life in the genera Oedipina (from south
and east of the Isthmus of Tehuantepec) and Li-
neatriton (from north of the Isthmus) (Fig. 12).
The former has become elongate by increasing
the numbers of vertebrae; the latter by increasing
the length of the individual vertebrae, which are
identical in number (in the trunk) to all tropical
genera except Oedipina (Tanner, 1950; Wake &
Lynch, 1976). Earlier in this paper I highlighted
the convergence in the Chiropterotriton-Dendro-
triton-Nototriton assemblage. Within Bolitoglos-
sa webbing of hands and feet has evolved both
convergently (Alberch & Alberch, 1981) and in

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

parallel (Wake, 1966; Wake & Brame, 1969; Lar-
son, 1983; Elias, 1984). These phenomena have
made phylogenetic analysis very difficult, for any
phylogenetic hypothesis requires extensive con-
vergence, parallelism, or evolutionary reversal
(Wake & Elias, 1983).

Epiphytes appear to have been significant fac-
tors in the evolution oftropical salamanders. The
convergence in morphology and behavior of
bromeliad-dwelling salamanders is one indica-
tion of this importance, and the extent to which
bromeliads and moss mats are utilized as mi-
crohabitats is another. It is also significant that
salamanders persist in the face of great environ-
mental change so long as fragments of forest with
the preferred microhabitats remain.

The tropical salamanders, which originated
from a Laurasian ancestral group (Savage, 1973),
are a marked exception to a common pattern of
tropical origin and subsequent temperate inva-
sion. Late in the history of
but nevertheless a very long time ago (perhaps
at the beginning of Tertiary), they invaded the
area that has become modern Middle America.
Compared with salamanders generally, the trop-
ical salamanders have been phenomenally suc-
cessful. But now their survival and, indeed, the
survival of much of the diversity of tropical eco-
systems is at risk, for the lowland forests and the
middle elevation cloud forests that harbor most
tropical salamanders are being cleared at rates
that almost defy belief. Not a tree is standing
over extensive parts of our San Marcos transect,
which was in an almost pristine state as recently
as 1969. In a single human lifetime the results
of perhaps a hundred million years of evolution
will have been dramatically changed, if not ex-
tinguished.

group,

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[Vor. 74

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1987] WAKE—ADAPTIVE RADIATION OF SALAMANDERS 263

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193.

APPENDIX I

Use of bromeliads and moss mats by neotropical salamanders. All species known to occur = either
of these microhabitats are listed below. Literature references do not include all observations for a
given species, but either the first or the best documented example. Where no eive Aia Mad field
notes or notes from museum collections made by the author are cited.

> š

B. yucatana
Chiropterotriton arboreus
C. chiropterus

C. chondrostega
C. lavae

C. multidentatus
Dendrotriton bromeliacia

D. megarhinus
D. rabbi
D. xolocalcae

Nototriton barbouri

Bromeliad Occurrence:
roni tase pete DBW notes
B. arborescandens Taylor Lay
B. Me Trapido (1942)
B. cuchumatana Stuart Ev Elias (1984)
B. dunni Schmidt (1942
B. engelhardti Schmidt (1936a), Wake & Lynch (1976)
B. flavimembris Wn
B. franklini Mid (1941), Wake & Lynch (1976)
B. hartwegi e & Brame (1969), Elias (1984)
B. helmrichi uds t (1936a
B. hermosa diae et al. (1983)
B. jacksoni Elias (19 ay
B. lignicolor Dunn (19
lincolni Wake & im EA. Elias (1984)
B. meliana Wake & Lynch (1982)

)
Wake & Lynch (1976), Elias (1984)
Stuart (1943)

Brame & Wake (1963)

Shannon & Werler (19552)

Stuart (* "probably, " 1948)

Schmidt (19362), Taylor & Smith (1945)

Ruthven (1922)

Dunn (1937)

Wake et al. (1970)
Brame & Wake (1963)
Brame & Wake (1972)
DBW notes

Rabb (1955)

DBW notes

Martin (1958)

958
Schmidt (1936a), Wake & Lynch (1976)
dei 960)
as & ie (1975), Elias (1984)
or (194
s EH

264 ANNALS OF THE MISSOURI BOTANICAL GARDEN

APPENDIX I. Continued

N. nasalis

N. picadoi

N. veraepacis

oeil ea bellii
unnata

piel
firscheini

goebeli
leprosa
. nigromaculata

DVT BWR
2 3

. smithi
Thorius dubitus
Moss Mat Occurrence:
Bolitoglossa diminuta
; is dias
B. subpalm
n cuchumatana,
Nototriton ns
N. richar
Myetancls perni x
Oedipina poelzi
O. pseudouniformis
. uniformis
Pseudoeurycea rex
P. scandens
P. werleri
Thorius dubitus

Dunn (1926), Schmidt (1942)

Picado (1913

Lynch & Mrs (1978)

DBW notes from T. J. Papenfus

Wake & Seh TUA ie DBW notes
Stuart (1954)

Werler & Smith (1952), Shannon & Werler

Schmidt (1936a), and DBW notes
DBW notes

DBW notes

Walker (1955), Martin (1958)

DBW notes from T. J. Papenfuss
DBW notes

Robinson (1976 — in a liverwort mat)
Wake et al. (1973)

Taylor (1952)

Lynch & Wake (1975)

Taylor (1954

notes
Elias & Wake (1983)
)

58)
Shannon & Werler (19552)
Taylor (1941)

[VoL. 74

REVISION OF EREMANTHUS (COMPOSITAE: VERNONIEAE)!

NANDA F. F. MACLEISH?

ABSTRACT

Historica d

1 £ dias & T hi d

ly I
traits of s de dus aggregation of heads into a glomerule), pluriseriate involucres, and

achenes

with a persistent pappus. Syncephaly i is characteristic of the Lychnophorinae, which is con-

In the present treatment Eremanthus includes

sidere

Vanillosmopsis, but excludes Albertinia, Chresta, Glaziovianthus, Prestelia, Pycnocephalum, Paralych-

mopsis taxa) are presented.

Recent studies have shown that Eremanthus
Less. (Compositae: Vernonieae) includes both
Eremanthus Less. (sensu Schultz-Bip.) and Vanil-
Josmopsis Schultz-Bip., but excludes Albertinia
Sprengel, Chresta Vell. Conc., Glaziovianthus G.
Barroso, Prestelia Schultz-Bip., oo
(Less.) DC., Sphaerophora Schultz-Bip., and
Stachyanthus DC. (MacLeish, eo pv
1984c, 1985a, 1985b, and MacLeish & Schu-
macher, 1984). Thus, Eremanthus comprises 18
species of syncephalous trees and shrubs largely
restricted to cerrado (sensu Eiten, 1978, 1983)
of the arid Central Plateau of Brazil. This is a
regibn dominated by woody Compositae; geo-
ly rare in the family

4

Heywood et al., 1977).

The name Eremanthus is derived from the
Greek eremos (solitary) and anthos (flower bear-
ing) in reference to the heads, which bear single
flowers. Vanillosmopsis refers to the vanillalike
odor characteristic of this group of plants. Vanil-
losmopsis is locally called candeia, candle, whic
refers to its ability to burn luminously. The close
relationship between Vanillosmopsis and mem-
bers of Eremanthus sensu Schultz-Bip. is fre-

De Candolle (1836) is the only other daha
to combine Eremanthus and Vanillosmopsis, al-
though he mistakenly placed them in Albertinia.
He correctly noted that E. incanus and E. elaeag-

and Stachyanthus (now Argyrovernon nia).

Eremanthus thus com-

nus approach Vanillosmopsis in degree of syn-
cephaly, number of florets per head, and achene
and pappus characters; and that V. erythropappa
and V. polycephala resemble Eremanthus in their
high degree of syncephaly and, in the latter, re-
duced number of florets per head. In the course
ofthe present study, numerous specimens of nat-
ural hybrids between these two genera were iden-
tified. It soon became apparent that the taxo-
nomic distinction between Eremanthus and
Vanillosmopsis was largely artificial.
Host/parasite relationships and terpenoid
chemistry support the close relationship of these

mopsidia H. S. Jackson & Holway, has been found
on E. mattogrossensis and on V. erythropappa.
Because rusts (Uredinales) have evolved with
their vascular plant hosts, their taxonomy may
suggest relationships among vascular plant taxa
(Thorne, 1979; McCain & Hennen, 1982). Urban
(1973) demonstrated the systematic utility of
Puccinia and Vernonia coevolution in determin-
ing subgeneric categories. Hence, a close rela-
tionship between Eremanthus and Vanillosmop-
sis is supported by the presence of a single rust
species on members of both genera. In addition,
terpenoid profiles appear to be very similar in
members of Eremanthus and Vanillosmopsis (H.
Schumacher, pers. comm.). Also, many authors
have noted the presence of identical terpenoid

! Work at the University of Georgia was supported partially by National Science Foundation Grants DEB
t for

81-13522 and ee 79-09831. I am grateful to the Brazilian governmen
o members of the Rio de Janeiro Jardim Botánico for generous assistance. In addition, I

and, in particular,

the opportunity to collect in Brazil

thank the E of the ppu herbaria who loaned specimens ae pe project rie ll Baker, Graziela M.

Barroso, Christopher Meach
Depart

m, and Heiko Schumacher, who we

especial assis

ment of Library Science and Information cee aan [ceni tei. ym Georgia 30322,

ANN. Missouni Bor. GARD. 74: 265-290. 1987.

266

yav sisqa in the two groups, one further sup-
(Bake ., 1972; Gar
cia et al., 1976; Vichnewsid et P 1977; Har-
borne & Williams, 1977; Robinson et al., 1980).
The systematic utility of terpenoids in members
of Vernonieae has been discussed by Abdel-Baset
et al. (1971), Mabry et al. (1975), Harborne &
Williams (1977), and Robinson et al. (1980).
Eremanthus can be distinguished from other
syncephalous genera by a combination of char-
acteristics (MacLeish, 1984a, 1984c). These in-
clude woody habit; indument composed pri-
marily of stellate hairs; terminal compound cymes
(rarely reduced to a cyme) of glomerules; 1-4
florets per head; 10-ribbed (rarely 20-ribbed)
achenes with 3—5-seriate pappus and prominent
nectary; and advanced pollen of type A (sensu
Keeley & Jones, 1977). Members of Eremanthus
have complex glomerules derived from reduc-
tion of internodes between subglomerules ar-
ranged in a compound cyme. A complete dis-
cussion of glomerulescence arrangement and
derivation for Eremanthus and related genera
can be found in chapter 1 of MacLeish (1984c).
The only other syncephalous Vernonieae that ap-
proach the combination of characteristics evi-
dent in Eremanthus are Paralychnophora
MacLeish and Lychnophora Martius (MacLeish,
1984c). However, both of these have a biseriate
pappus and glomerules derived from the inter-
node reduction of corymbose subglomerules. In
addition, Paralychnophora is characterized by
axillary, solitary glomerules and angled achenes;
and Lychnophora by ericoid leaves and promi-
nently twisted inner pappus series.
mong nonsyncephalous Vernonieae, Ere-
manthus appears to be related to Vernonia sub-
sect. Eremosis (DC.) S. B. Jones (MacLeish,
1984c). This subsection also has a woody habit,
congested terminal capitulescences, reduced
numbers of florets per head, ribbed achenes, and
pollen of type A. However, this group is distrib-
uted primarily in Central America and northern
South America and has a biseriate pappus. If
Eremanthus is derived from Eremosis stock, there
are two Brazilian refugia that may have served
as centers of diversification and dispersal: Vea-
deiros and Rondônia (also called “Araguaia” and
* Airpuana" by Prance, 1982). Veadeiros is the
current site of several locally endemic members
of sect. Nectaridium (subg. Vanillosmopsis) that
are virtually indistinguishable from many mem-
bers of Vernonia subsect. Eremosis. Rondónia
contains a single Eremanthus species, E. ron-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

doniensis, which is restricted to that area and is
the most unspecialized member of subg. Ere-
manthus.

TAXONOMIC HISTORY

Historically, A/bertinia Sprengel, Chresta Vell.
Conc., Glaziovianthus G. Barroso, Pithecoseris
DC., Prestelia Schultz-Bip., Pycnocephalum
(Less.) DC., Sphaerophora Schultz-Bip., Stach-
yanthus DC., and Vanillosmopsis Schultz-Bip.
have been placed adjacent to or in synonymy
with Eremanthus. These genera are restricted to
the arid Central Plateau of Brazil and have not
been evaluated comprehensively since Baker’s
(1873) treatment of Compositae in Martius’s
Flora Brasiliensis.

In 1829, Lessing established Eremanthus to
include syncephalous trees or shrubs with one
floret per head and a paleaceous pappus. At the
same time he placed syncephalous trees and
shrubs with (presumably) numerous florets per
head, filiform pappus, and alveolate receptacles

Ibertini d syncephalous h ernonia
sect. Pycnocephalum. Also in 1829, two species
referable to Eremanthus s. lat. were described by
Velloso as Chresta: C. cordata and C. lanceolata.
However, the portion of Flora Fluminensis con-
taining Chresta was not effectively published un-
til 1881 (Carauta, 1973). Thus, de Candolle ac-
complished valid publication of Chresta in 1836
when he cited Velloso's figures, which had been
published in Florae Fluminensis Icones (Velloso,
1831). Of the two species described by Velloso,
only C. cordata was included by de Candolle in
Chresta (as C. spherocephala). This distinctive
taxon is widespread throughout Brazil. In con-
trast, the figure and description of C. /anceolata
could represent any one of several species. If one
takes into account the itineraries of Velloso and
contemporary collectors (Urban, 1906), C. /an-
ceolata is either Vanillosmopsis erythropappus,
Eremanthus incanus, E. glomerulatus, or E.
mattogrossensis. Kuntze (1898) believed this
taxon to be synonymous with V. erythropappus
but provided no account of his reasoning. In the
present treatment, C. /anceolata is excluded be-
cause of doubtful identity, and C. cordata is re-
tained in Chresta.

Two years later, Lessing (1831) dissolved A/-
bertinia when he transferred the type species into
Vernonia, and a second taxon, correctly consid-
ered to have one floret per head, into Ereman-
thus. In 1836, de Candolle treated syncephalous
taxa as five different genera. Syncephalous trees

=

1987]

and shrubs were included in Albertinia (he in-
correctly assumed that the type species had a
single floret per head), and syncephalous herbs
were placed variously in Chresta, Pycnocepha-
lum, Stachyanthus, and Pithecoseris. Schultz-Bi-
pontinus (1861, 1863) recognized the error im-
plicit in de Candolle's interpretation of the deeply
pitted receptacle A/bertinia as fused heads ho-
mologous with those found in many Eremanthus
species. In contrast, Schultz-Bipontinus consid-
ered the heads of Albertinia to contain many
florets, each of which is partially enclosed in a
receptacle cavity, and the genus to be monotypic
and closely related to Vernonia. However,
Schultz-Bipontinus placed taxa with one (rarely
three) floret(s) per head, high degrees of synceph-
aly, and achenes with a persistent pappus in Er-
emanthus or the newly-described Sphaerophora
(further delimited by achene surface features),
and those with three (to one) florets per head,
various degrees of fusion, and achenes with a
deciduous pappus in Vanillosmopsis, a second
new genus (Schultz-Bipontinus, 1861, 1863). In
1873, Baker followed Schultz-Bipontinus in his
delimitation of Vanillosmopsis; however, in his
classification, Eremanthus comprised not only
the species of Schultz-Bipontinus, but also those
of Sphaerophora, Chresta, Pycnocephalum, Pres-
telia, and Stachyanthus. Except for Barroso's
(1947) transfer of herbs with yellow and red co-
rollas to Glaziovianthus, and Robinson's (1980)
suggested recombination of all herbaceous taxa
into Chresta, the boundaries of Eremanthus and
related genera have remained largely as Baker
delineated in 1873

Following Baker, curators endeavoring to
identify Brazilian specimens have placed most
syncephalous Vernonieae with achenes bearing
a persistent pappus in Eremanthus. However,
syncephalous heads are characteristic of Lych-
nophorinae, which is currently considered to be
an artificial assemblage (Jones, 1977; MacLeish,
1984c). Traditionally, the distinction between
Eremanthus and Vanillosmopsis has been based
on the presence or absence of syncephalous heads,
one versus three pue r head, and paleaceons
and persiste
pus. In fact, ihe characteristic of lacking sineephe
aly i used w exclude Vanillosmopsis from
Ly aced)
despite generic descriptions that clearly pee
Vanillosmopsis as including several highly syn-
cephalous taxa (Baker, 1873; Bentham, 1873).
De Candolle is the only author until now to com-

MACLEISH — EREMANTHUS REVISION

267

bine Eremanthus and Vanillosmopsis, although
erroneously, in Albertinia. Currently, following
de Candolle, Eremanthus is considered to in-
clude trees and shrubs with reddish-purple co-
rollas, compound cymes (rarely reduced to a
cyme) of glomerules, 1—4 florets per head, ribbed
achenes with persistent or deciduous pappus of
3-5 intergrading series, and advanced pollen of
type A (subechinolophate with elongated ger-
minal furrows often barely separated at the poles,
sensu Keeley & Jones, 1977).

e Candolle and Schultz-Bipontinus used the
number of florets per head, degree of syncephaly,
number of pappus series, and various achene sur-
face features to delineate subdivisions of genera.
These are good characters that readily indicate
natural groups of related species. Thus, subg. Er-
emanthus includes sect. Eremanthus and sect.
Synglomerulus. Eremanthus sect. Eremanthus is
a group of closely related taxa (E. glomerulatus,

7. goyazensis, E. mattogrossensis, and E. ron-
doniensis) that may have arisen along an east-
west gradient across southern portions of the
Central Plateau (Fig. 1). This section is charac-
terized by coriaceous leaves, a compound cyme
of 10-60 glomerules, 5-100 heads per glomerule,
head fusion via the interwoven pubescence of
phyllaries, one floret per head, and sericeous
achenes with a stramineous, coroniform, persis-
tent, paleaceous pappus. Section Synglomerulus
comprises Eremanthus argenteus, E. auricula-
tus, and E. cinctus. They differ from sect. “Ere.
manthus primarily in reduction of the glomer-
ulescence to a cyme (not compound) of 2-10
glomerules
E omaniku subg. Vanillosmopsis comprises
two sections and is characterized by membra-
naceous to subcoriaceous leaves, compound
cymes of more than 100 glomerules, 1-12 heads
per glomerule, 1—4 florets per head, and glabrous
achenes with a purple or white (rarely stramin-
eous), not coroniform, deciduous, filiform pap-
pus. Eremanthus pohlii, E. graciellae, and E.
brasiliensis form sect. Nectaridium, which is
characterized by near absence of syncephaly
(heads solitary or slightly appressed basally in
pairs) and is restricted to the northwestern arm
of the Central Plateau. Eremanthus capitatus, E.
arboreus, and E. uniflorus are members of sect.
'anillosmopsis and have 2-9 heads per glomer-
ule, with bases slightly appressed and fused by
the interwoven pubescence of the phyllaries. The
first two of these species range south to north
along the northeastern arm of the Plateau, and

268

À sect. Eremanthus
subg. Eremanthus
@ sect. Synglomerulus
* subg. Isotrichia
X subg. P d h
A sect. Vanillosmopsis
subg. Vanillosmopsis
© sect. Nectaridium

A

FIGURE 1.

the last is restricted to the dde sa: arm
Eremanthus erythropappus and E. polyc

have glomerules with 6-12 heads that are closely
appressed and fused by the concrescence of phyl-
lary and receptacle tissues. These two species are
found only in the southern part of the Central
Plateau.

In contrast, three species (E. incanus, E.
elaeagnus, E. seidelii), historically assigned to
Eremanthus s. str., exhibit characteristics inter-
mediate to those of both subg. Eremanthus and
subg. Vanillosmopsis. Not surprisingly, these
species are largely restricted to the southern part
of the Plateau, an area where the two major com-
plexes are sympatric. This is also the region where
a majority of hybrids are found, and it is possible
that subg. Pseuderemanthus (E. elaeagnus and
E. seidelii) and subg. Isotrichia (E. incanus) rep-
resent established hybrids between members of
the major complexes.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Distribution of Eremanthus in the Central Plateau of Brazil.

TAXONOMIC TREATMENT

Eremanthus Less., Linnaea 4: 317. 1829. TYPE:

Eremanthus glomerulatus Less.

a S e Schultz-Bip., Jahresber. Pollichia 18—

16 61. TYPE: Vanillosmopsis glomerata

EU Bip. = Eremanthus erythropappus (DC.)
MacLeish.

Trees or shrubs to 10 m tall; stems reddish-
brown lanate-tomentose or gray lepidote-tomen-
tulose, often darkened basally by fire; branches
few to many, often tortuous. Leaves bna al-
ternate, strongly coriaceous to membranaceous,
largely restricted to ends of branches, subsessile
to petiolate; blades 2-20 cm long,
wide, narrowly elliptic to obovate, the bases acu-
minate to obtuse (occasionally auriculate, pan-
duriform, or retuse), the apices acuminate to ob-
tuse, the margins entire, adaxially glabrate to
Jepidare: abaxially silvery-canescent to gray-
brown, lanate-tomentulose to lepidote-tomen-

1987] MACLEISH — EREM.

tulose; trichomes stellate, the stalk with 1—2 cells;
primary venation pinnate and abaxially promi-
nent, the midrib furrowed; petioles 1-20 mm
long, lanate or lepidote, base slightly expanded.
Peduncles slender, to 35 cm tall, terete, reddish-
brown to gray, lanate-tomentose to lepidote-to-
mentulose, expanded at apex; bracts foliaceous,
both intra- and extraglomerular. merules-
cence a terminal cyme of 2-20 glomerules or a
compound cyme of 8-60(-100) glomerules; mat-
uration of glomerules centrifugal. Glomerules 5-
15 mm tall, 4-12 mm diam., hemispherical to
spherical; head maturation concurrent to cen-
trifugal within a glomerule. Heads homogamous,
1-100 per glomerule, closely to slightly ap-
pressed and free, or coherent by pubescence of
phyllaries, or connate by concrescence of recep-
tacle and phyllary tissues. Involucres obconic or
cylindrical, 2-12 mm tall, 0.3-4 mm diam.;
phyllaries in 4-7 imbricate, graduated (rarely
subequal) series; innermost phyllaries deciduous
at maturity: mare ins subscarious, entire to
o

ten
with purple apex, glabrate to lanate-tomentulose

REVISION 269

or lepidote-tomentulose, the inner surfaces stra-
mineous and vernicose; single vein (when pres-
ent) often with mucronate extension. Florets 1—
3(4) per head; corollas purple to white, 5-9 mm
tall, the tube and lobes subequal; tube 2-4.5 mm
tall; throat 0.5-1 mm tall; lobes 2.5-3.5 mm tall.
Anthers 2-4 mm long. Pollen tricolporate with
a continuous micropunctate tectum, short echi-
nate (Vernonia type A). Style 4-6 mm long, 0. 1 5-
0.3 mm wide, noticeably flattened, the branches
short! to mediuer bine l- 2. J mm long. Achenes

1
Lyla J

1-4 mm n tall, glandular, glabrous or sericeous,
10-ribbed (often obscurely or rarely to 20), apex
constricted and dark; carpopodium absent or mi-
nute; nectary 0.2-0.45 mm tall, persistent or tar-
dily deciduous, occasionally pubescent. Pappus
3-5-seriate, of stramineous, white, or purple,
strongly coroniform to not coroniform, often ba-
sally fused, persistent to promptly deciduous, pa-
leaceous to filiform, strigose bristles; outermost
series 0.4-3 mm long, innermost series 3-8 mm
long. Chromosome number: n = 1

tur binate),

KEY TO SPECIES OF EREMANTHUS

la. Glomerulescence a compound cyme of 3-60 glomerules or a cyme of 2-20 glomerules; heads (1-)20-

E per seen with phyllaries in 3—6 series; florets
ericeous with stramineous (rarely white or purple), strongly to subcoroniform, persistent
ou

cylindri

spher

1(2-4) per head; achenes cylindric-turbinate

3a. js om cylindric-turbinate (rarely re 10-ribbed; pappus stramineous (rarely purple),

coroniform, persistent, paleaceous; heads

onic (rarely cylindric), coherent by interwoven

pubescence of phyllaries (subg. Eremanthus =
4a. Glomerulescence a compound cyme of (3-)10—60 glomerules (sect. Eremanthus).

5a. Stem pubescence reddish-brown lanate-to

mentose; heads 20-100 per glomerule.

re a l4—V» length; leaves S AU Bi Sa rarely E peas -to-

lomerulatus

6b. Heads ed entire length; leaves strongly coriaceous, . abaxially esie to-
2.

i
g

mentose
. Stem renin gray lepidote- tomentulose; heads 5- -45 per ‘glomerule.

E. goyazensis

7a. Heads coherent '^ length; leaf blades 6-16 cm long, 2-10 cm wide uu

7b. Heads coherent Ys length; ‘leaf blade 2-7 cm n long, 0.5-1. 5 cm n wide .
.4.

E. mattogrossensis

“rondoniensis

4b. Glomerulescence a a |cyme (not compound) of 5- -20(-30) glomerules (sect. Synglomerulus.
8a

em pubescence gray lepidote-tomentulose; leaf base acute to attenua

argenteus

8b. Stem pubescence reddish- brown lanate- tomentose; leaf base variously lobed res

acute).

9a. Leaf bases auriculate, the blades 5-8 cm long, appressed- ascending; heads
r glomerule, Rai 12 lengt 6.

ute, the blades 6-19 long, not appressed-ascending; heads

7. E.

3b. Achenes cylindric, ; papp

us off-white to purple,

3
E. auriculatus

cinctus
subcoroniform, tardily decid-

uous, E heads nde connate by concrescence of tissues (subg. /sotrichia) ...
8.

E. incanus

270

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

2b. Florets 3-4 per head; heads slightly appressed, free; achenes cylindric; glomerule hemispherical
s).

(subg. Pseuderemanthu

a E E. seidelii

phyllaries in 5-7 series; y pene 1-4 adis

und cyme of pus than 100 glomerules; heads 1-12 per glomerule,
ead; achenes cylindric (rarely cylindric- b pores

sei
E. elaeagnus
with

with a purple or white (rarely stramineous
pappus; leaves membranaceous to su

(rarel

ubcoriaceous (subg. Vanlpsmopsis

lla. Heads solitary or in pairs slightly appressed basally, cylindric; pappus never twisted (sect. Nec-

taridium).

12a. Florets 2-3 per head; involucre with prominent truncate base, phyllaries purple.
ad

13a. Florets 3 per he
13b. Florets 2 per he

. ll. E. pohlii
. 12. E. graciellae

Florets 1 per head;

15b. Florets 34 per

16b. n 3-4 per head.

a. Heads 2-5 per glomerule |...
17b. Heads 6-9 per glomerule ...........

Eremanthus Less. subg. Eremanthus. Ereman-
thus subg. Eueremanthus Schultz-Bip., Jah-
resber. Pollichia 20-21: 393. 1863

Leaves strongly coriaceous to subcoriaceous.
Glomerulescence a compound cyme of 3-60
glomerules or a cyme of 2-20 glomerules. Glom-
erules subspherical to hemispherical. Heads (1-)
20-100 per glomerule, obconic; with phyllaries
in 3-6 series, closely appressed, coherent by in-
terwoven pubescence of phyllaries. Florets | per
head. Achenes cylindric-turbinate, 10-ribbed, se-
riceous. Pappus stramineous (rarely purple), co-
roniform, persistent, paleaceous, strigose bris-
tles.

Eremanthus Less. sect. Eremanthus

d cyme of (3-)10-
60 glomerules. Glomerules 5-10(-15) mm tall,
5-15 mm diam

1 1 (^ mÈ

—

. Eremanthus glomerulatus Less., Linnaea 4:
317. 1829. Albertinia glomerulata (Less.)
DC., Prodr. 5: 82. 1836. TYPE: Brazil: no
other data, Sellow s.n. (holotype, P; isotype,
TEX)

Albertinia rufiseta DC., Prodr. 5: 81. 1836. TYPE: Brazil.
a erais: siccis apricis montosis Serro Frio
Martius s.n. (holotype, M).
a e re DC., Prodr. 5: 81. 1836. Ere-
nthus pallidisetus (DC.) Schultz-Bip., Jahres-
Bee Pollichia 18-19: 165. 1861. TYPE: Brazil. Mi-
So hun campis editis ad Calumbi Praed. Serro
artius s.n. (holotype, M).
es biis Gardner, London J. Bot. 5: 235.

ad
l involucre without truncate base, phyllaries brown ...... b
11b. Heads (2-)5-12 per glomerule, cylindric or obconic; pappus often twisted (sect. Fanilosmops
14a. Heads un a Bae and connate by tissue concrescence, 6-12 per glo
15a. Floret r head; heads coherent '^ length
r head; heads coherent nearly entire length . 15.
14b. r = iy slightly appressed and coherent by pubescence of phyllaries, 2- 9 per glomerule.
l6a. l per head 6.

E. brasiliensis

l. E polycephalus
E. erythropappus

16. E. uniflorus

AE ena c | T; E, capitatus
"————— ÉOÉ Inr 18. E. arboreus

1846. Eremanthus stellatus (Gardner) Schultz-
Bip., Jahresber. Pollichia 18-19: 164. 1861. Er-
emanthus stellatus (Gardner) Schultz-Bip. var.
pud Schultz-Bip., Mota Pollichia 20-
21: 1863, n das Brazil. "Per-
nam dre (now B a Rosa, district of
se Rio Preto, Sept. 1839, Canine 2896 (holo-
type, BM; isotypes, 2 in F (fragment and photo
of W), G, , 2 in NY
Eremanthus stellatus (Gardner) Schultz- Bip. var. poh-
ultz-Bip., Jahresber. Pollichia rii 394,
1863. TYPE: rs in summitate montiu e
. S. Anna, piu d eig
W; isotypes, "F (fragment), 2in N

Slender to robust tree, 2-10 m tall, to 20 cm
diam.; stems reddish-brown lanate-tomentose;
branches few to many. Leaves subcoriaceous,
sessile to petiolate; petioles to 10 mm long, la-
nate-tomentulose; blades 4—14(-20) cm long, 1-
5.5(-7.5) cm wide, elliptic to ovate, the bases
acute to obtuse, the apices obtuse (rarely rounded
or retuse), margins entire; adaxially tomentulose
to glabrate, abaxially gray-brown lanate-tomen-
tulose (rarely lepidote-tomentulose). Peduncle
slender, to 20 cm tall, ribbed, reddish-brown la-
nate-tomentose to gray lepidote-tomentulose.
Glomerulescence a compound cyme of 20-60
glomerules. Glomerules 5-15 mm tall, 9215 mm
diam., hemispherical. Heads 20-90 per glom-
erule, closely appressed basally, fused “4 to !^
length by pubescence of phyllaries. Involucres
obconic, 3.5-5.5 mm tall, 1.3-2.5 mm diam.;
phyllaries in 4—6 series; outermost phyllaries nar-
rowly obtrullate, 1.2-3 mm long, 0.4-1.6 mm
wide, the apices acute; innermost phyllaries ob-
lanceolate, 3.1-5.2 mm long, 0.6-1.4 mm wide,

1987]

1
200 km

0 100
Elevation:..... 500 meters
„1000

MACLEISH — EREMANTHUS REVISION

€ E. glomerulatus a E.

A E. goyazensis

mattogrossensis

am E. cinctus

Figure 2. Distribution of Eremanthus subg. Eremanthus in Minas Gerais.

the apices acute; margins entire; abaxial surfaces
stramineous with purple apex, glabrate to lanate-
tomentulose. Corollas purple to white, 4-7.8 mm
tall, the lobes acuminate. Anthers 2-3.4 mm long,
the apical appendage acuminate to rounded, the
bases obtuse to rounded. Achenes 2.5-4.1 mm
tall, glandular, sericeous between the 10 ribs, the
apex slightly constricted; nectary 0.15-0.25 mm
tall, tardily deciduous. Pappus 4-5-seriate, of
strongly coroniform (at maturity) strigose bris-
tles; outermost series 1-2.2 mm long; innermost
series 4—6.2 mm long.

Flowering and fruiting occur from March to
October.

Eremanthus glomerulatus is distributed
throughout Minas Gerais, Goiás, and adjacent

regions in Bahia and Sao Paulo (Figs. 2, 3, 6), at
altitudes of 700 to 1,500 meters. It is found com-
monly in large colonies and may dominate cer-
rado and campo rupestre habitats. In Minas Ge-
rais, this species often is used for fence posts
because of its abundance.

Eremanthus glomerulatus is the eastern mem-

ber of a group of closely related taxa (sect. Ere-
manthus) found throughout the Central Plateau.
Eremanthus goyazensis is slightly more robust
with a greater fusion of heads and is found in the
northcentral branch of the Plateau. In contrast,
E. mattogrossensis is less robust and has lepi-
dote-tomentose (rather than lanate-tomentose)
stems and leaves; it extends throughout the west-
ern arm and into the southwestern side. Ere-
manthus rondoniensis is closely related to E.
mattogrossensis and is restricted to the far west-
ern portions of the western arm. Eremanthus
glomerulatus appears to be the member of this

complex most closely related to the taxa histor-
ically considered to compose Vanillosmopsis. The

and occasionally purple pappus of E
latus are reminiscent of characteristics found in
the transitional taxa, E. incanus, E. elaeagnus,
and E. seidelii. In addition, it is the only member
of this complex that is sympatric with both the
transitional taxa and the majority of the Vanil-
losmopsis complex.

he many epithets ascribed to E. glomerulatus

272

represent variations in pappus color (purple or
white) and number of series, branch position (an-
gular or not), and leaf shape. Eremanthus glo-
merulatus can be found throughout most of the
Central Plateau, and the observed variation can
be attributed to maturity and habitat differences.
These epithets were applied by earlier authors
who were attempting to identify Brazilian col-
lections in the absence of adequate documenta-
tion. Baker (1873) was the first author to rec-
ognize E. glomerulatus as a widespread and
slightly variable species

Paris specimen of E. glomerulatus is clear-
ly the holotype. It is annotated appropriately by
both Lessing and Schultz-Bipontinus and match-
es the description given by Lessing in 1829. Orig-
inally both the Paris specimen and the Texas
specimen were housed in Berlin because both are
stamped “hb. berol."

Additional specimens examined. BRAZIL. BAHIA: Rio
das Contas, 22 Jul. 1979, King et al. 8099 (US). DISTRITO
FEDERAL: Gama, 28 Aug. 1965, Irwin et al. 7926 (MO,

NY, US). GorAs: Cristalina, 3 Apr. 1973, Anderson
8004 (F, MO, NY, UB, US). MINAS GERAIS: Nova Lima,
15 Apr. 1945, Williams & Assis 6626 (F, GH, MO,
NY, S).

Possible hybrids. x E. elaeagnus (C. Martius ex
DC.) Schultz-Bip. MINAS GERAIS: Serra do Cipó, : Oct.
1980, ni. et al. 715 , GH
MO, NY, S, US), 7 Oct. 1980, et al.
716 (GA, Ra. 30 Jan. 1980, King & Almeda 8354
(US). x E. mattogrossensis Kuntze. MINAS GERAIS: Sac-
ramento, 22 Jul. 1980, Schumacher B10 Sea x E.
seidelii MacLeish & Schumacher. MINAS GE
tween Furnas and Piui, 25 Aug. 1981, Schumacher
1008 (GA).

2. Eremanthus goyazensis (Gardner) Schultz-
Bip., Jahresber. Pollichia 18-19: 165. 1861.
Albertinia goyazensis Gardner, London J.
Bot. 6: 425. 1847. TYPE: Brazil. Goias: hilly
campos near Villa de Arrayas, Apr. 1840,
Gardner 3804 (holotype, BM; isotypes, 2 in
F (fragment and photo of B), 3 in G, GH
(photo of B), 3 in NY, TEX (photo of B),
W)

Eremanthus weddellii Schultz- is Jahresber. ee
lichia 18-19: 165. 1861. TYPE: Brazil, Goiás, e
in Salinas, May 1844, Weddell 2032 Fiori
P).

Small tree, 1—3 m tall, the trunk often gnarled;
stems reddish-brown to gray-black lanate-to-
mentose, to 8 cm diam.; branches few. Leaves
strongly coriaceous, sessile to petiolate; petioles
to 10 mm long, lanate-tomentulose; blades 7.5—

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

18 cm long, 3.5-10 cm wide, elliptic to ovate,
the bases obtuse, the apices obtuse to rounded
(rarely retuse), me margins entire; adaxially
sparsely
lepidote- tomentulose. Peduncle nde. to 20 cm
tall, ribbed, gray-brown lepidote-tomentulose.
Glomerulescence a compound cyme of 10—40
glomerules. Glomerules 5-15 mm tall, 7215 mm
diam., hemispherical. Heads 25-100 per glom-
erule, ee appressed, pee entire length

y lanose pubescence o yllaries. Involucres
one 4.6 mm tall, 0. 4 1.5 mm diam.; phyl-
laries in 3—4 series; outermost phyllaries narrow-
ly obtrullate, 1.8-3.2 mm long, 0.2-0.8 mm wide,
the apices acute to acuminate; innermost phyl-
laries oblanceolate, 2.9—4.8 mm long, 0.3-0.8 mm
wide, the apices acute; margins entire; abaxial
surfaces stramineous, glandular, glabrate to la-
nose-tomentose. Corollas pale purple to white
basally, 5.2-8.2 mm tall, the lobes acuminate.
Anthers 2.4—4.4 mm long, the apical appendage
acuminate; bases obtuse to rounded. Achenes 2-
2.8(-3.8) mm tall, glandular, sericeous between
10 ribs, often obscurely ribbed; apex slightly con-
stricted; nectary 0.25-0.35 mm tall, tomentose,
tardily deciduous. Pappus 4-5-seriate, of strong-
ly coroniform (at maturity) strigose bristles; out-
ermost series 0.8-2.3 mm long, the innermost
series 4.6-6 mm long. Chromosome number: n =
15.

Flowering and fruiting occur from January to
September.

! ^d si ? Hd E LP e. |

d through-
out southern Goiás and nearby Minas Gerais
(Figs. 2, 3). It is infrequent in cerrado and campo
rupestre at altitudes of 700 to 1,600 meters.

Eremanthus goyazensis is the northcentral ex-
tension of the glomerulatus complex. It is dis-
tinguished from E. glomerulatus by its strongly
coriaceous leaves with abaxial lepidote-tomen-
tose surfaces and heads entirely coherent by in-
terwoven pubescence. In addition, it can be easily
distinguished from the mattogrossensis-rondon-
iensis complex by its lanate stem pubescence,

5-100 heads per glomerule, and high degree of
cohesion.

Additional specimens examined. BRAZIL. DISTRITO
FEDERAL: Chapada da Contagem, 18 Aug. 1964, Irwin
& Soderstrom 5287 (NY, TEX, UB). Goiás: Terezina,
18 Mar. 1973, Anderson 7354 (F, MO, NY, UB, US).

3. Eremanthus mattogrossensis Kuntze, Revis.
Gen. PI. 3(2): 145. 1898. TYPE: Brazil: “Mat-

1987]

4

5,

. argenteus

. auriculatus

. goyazensis

E

E

E. glomerulatus
E

E. mattogrossensis
E

pares

. cinctus

Elevation:
m IT 500 meters
21000 ”

a
ome LN 2

FIGURE 3.

togrosso" (now Mato Grosso): Jul. 1892,
Kuntze s.n. (holotype, NY; isotypes, photos
of B, F, GH, TEX).

Shrub or slender tree, 1-4 m tall; stems gray-
brown lepidote-tomentulose; branches few.
Leaves coriaceous, sessile to petiolate; petioles
to 15 mm long, lepidote; blades 6-16 cm long,
2-10 cm wide, elliptic to page 4 pu the
bases acute to acuminate, the ap to
rounded, the margins entire; adaxially sparsely
lepidote to glabrate, abaxially gray lepidote-to-
mentulose. Peduncle slender, to cm tall,
ribbed, gray-brown lepidote-tomentulose.
Glomerulescence a compound cyme of 6-35
(-80) glomerules. Glomerules 4.5-10 mm tall,
5.5-12 mm diam., subspherical. Heads 10—45
per glomerule, closely appressed basally, coher-
ent '2 length by pubescence of phyllaries. Invo-
lucres cylindric, 2.3-4 mm tall, 1.1-2.4 mm
diam.; phyllaries in 5—6 series; outermost phyl-
laries narrowly obtrullate, 1.3-2.3 mm long, 0.2-
0.6 mm wide, the apices acute; innermost phyl-
laries oblanceolate, 1.8—3.4 mm long, 0.5—0.9 mm
wide, the apices acute; margins entire; abaxial
surfaces stramineous, glabrate to lepidote-to-
mentulose. Corollas pale purple to white, 4.4—

.6 mm tall, the lobes acuminate. Anthers 2.1-
3.4 mm long, the apical appendage acuminate to
acute, the bases obtuse to rounded. Achenes 2-
3 mm tall, glandular, sericeous between ribs, 10-

MACLEISH — EREMANTHUS REVISION

273

I
wy

E rida

T eo ELS
o NEFKEE s OUS i
J MS ` FI + phe Lu -

/ SS d. LO USER eee aes

pcr ae
200

kilometers

Distribution of Eremanthus subg. Eremanthus in Goias.

ribbed (often obscurely), the apex slightly con-
stricted; nectary 0.15-0.25 mm tall, tomentose,
tardily deciduous. Pappus 3-5-seriate, of strong-
ly coroniform (at maturity) strigose bristles; out-
ermost series 0.8-3.1 mm long; innermost series
3.3-6 mm long.

Flowering and fruiting occur from April to Au-
gust.

Eremanthus mattogrossensis is widely distrib-
uted throughout the western section of the Cen-
tral Plateau of Brazil (Figs. 2-4) at elevations of
500 to 1,000 meters in cerrado. It is particularly
common in Mato Grosso, where it is called **vel-
ludo do cerrado," velvet ofthe cerrado, or **casca
freta," cleft bark.

Eremanthus mattogrossensis is transitional
between b. goyazensi and E. rondoniensis and

l location. All three
are found within the western arm of the Central
Plateau. Eremanthus rondoniensis is restricted
to the far western portion of the Serra dos Parecis
near the border of Mato Grosso and Rondônia,
and E. goyazensis is restricted to eastern portions
of the western arm (principally southern Goiás).

The type specimen, bearing only the citation
“Mattogrosso,” must have been collected in
western Mato Grosso, near Rondônia. Zanoni
(1980), in his excellent survey of Kuntze’s col-
lecting trips, pointed out that Kuntze entered
Brazil through Bolivia and explored only a small

/
mattogrossensis /

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

m E. 57° `
f N
æ E. cinctus / -—- 6? )
[4 z3
` r4
9 E. rondoniensis > e "d
\ /
[|
)
|]
z
`Y /
/
“semasa REND /
ERE: seed!

A. Melasi

re Elevation:
. 500 meters
:. 1000 i

FIGURE 4. Distribution of Eremanthus subg. Eremanthus in western Central Plateau.

area of Mato Grosso near the border. Zanoni also
noted that the bulk of Kuntze's personal her-
barium, including many type specimens, is at the

w York Botanical Garden. Thus, their spec-
imen of E. mattogrossensis, which bears the ap-
propriate collecting information in Kuntze's
handwriting and which matches his type descrip-
tion, is likely the holotype.

Additional specimens examined. BRAZIL. DISTRITO
FEDERAL: Brasília, 15 Jun. 1981, Heringer et al. 7057
(MO). GorAs: 75 km from Aragarças, near Piranhas,
23 Jun. 1966, Hunt & Ramos 6160 (MO, NY, UB).
MATO GROSSO: Barra do Garças, 6 May 1973, Anderson
9801 (G, MO, NY, UB). MINAS GERAIS: Furnas, 5 Jul.
1982, Schumacher 2008, 2009, 2010, 2012, 2014 (GA,
je n Sao Felix do Xingu, 12 Jul. 1978, Rosario

Y). SAO PAULO: inter Canna Verde et S. José,
mdi at 670 (BR, S).

Possible hybrids. x E. glomerulatus Less. MINAS
GERAIS: Sacramento, 22 Jul. 1980, Schumacher B10
(GA). x E. seidelii MacLeish & Schumacher. MINAS
GERAIS: between Furnas and Piuí, 25 Puig 1981, Schu-
macher 1003, 1004 (GA).

4. Eremanthus rondoniensis MacLeish & Schu-
macher, Syst. Bot. 9: 89. 1984. TYPE: Brazil.
Rondônia: Vilhena, 13?16'S, 58*52'W, 18
Apr. 1977, Bantel & Silva s.n. (holotype,
RB).

Small tree, ca. 1 m tall; stems gray-brown lep-
idote-tomentulose, often basally darkened by fire.
Leaves coriaceous, sessile to petiolate; petioles
to 5 mm long, lepidote; blades 2-7 cm long, 0.5-

ces obtuse entire; adaxially
deca lepidote to ioc abaxially gray-brown
epidote-tomentulose. Peduncle slender, height
unknown, terete, gray-brown lepidote-tomen-
tulose. Glomerulescence a compound cyme of
3-20 glomerules. Glomerules 5-7 mm tall, 7-10
mm diam., hemispherical. Heads 5-15 per glom-
erule, closely appressed, coherent '4 length by
pubescence of phyllaries. Involucres obconic, 3.5-
5 mm tall, 1-2.2 mm diam.; phyllaries in 4—6
imbricate graduated series; outermost phyllaries
widely trullate, 1.4-2.1 mm long, 0.5-0.7 mm
wide, the apices obtuse; innermost phyllaries ob-
lanceolate, 2.4-4 mm long, 0.8-1 mm wide, the
apices acute; margins entire; abaxial surfaces
stramineous with purple apex, glabrate to lepi-
dote-tomentulose. Corollas white, 5.1—6 mm tall,
the lobes acuminate. Anthers 2.6-3 mm long,
the apical appendage acuminate, the bases
rounded. Achenes turbinate, 1.9-2.2 mm tall,
glandular, obscurely-ribbed, the base sericeous,
the apex constricted with prominent dark collar;

1987]

nectary 0.2-0.5 mm tall. Pappus 3-5-seriate, of
strongly coroniform (at maturity) strigose bris-
tles; outermost series 0.6-1.8 mm long; inner-
most series 5—6.8 mm long.

Flowering and fruiting occur from April to July.

Eremanthus rondoniensis is distributed along
the Rondónia-Mato Grosso border in the west-
ern portion of the Serra dos Parecis (Fig. 4). It
is found in periodically burned cerrado. Ere-
manthus rondoniensis is closely related to E.
mattogrossensis and represents a western exten-
sion of the g/omerulatus-goyazensis- mattogros-
sensis complex.

Additional specimens examin ned. BRAZIL. MATO
te Rondônia):

RONDÔNIA (latitude and longitude indicate Mato Gros-
so): Vilhena, 13?46'S, 59°50'W, 17 Apr. 1977, Bantel
& Silva s.n. (RB)

Eremanthus sect. Synglomerulus MacLeish, sect.
nov. TYPE: Eremanthus argenteus MacLeish
& Schumacher.

Glomerulescentiae cymosae. Glomerulis 5-10(-30);
(5-)10-25 mm longis, 10-25 mm diam

WG

. Eremanthus argenteus MacLeish & Schu-
macher, Syst. Bot. 9: 84. 1984. TYPE: Brazil.
Goiás: 33 km N of Alto Paráiso towards
Cavalcante, 1,370 m, 14 . 1980,
MacLeish, Martinelli, Smith & Stutts 734
(holotype, RB; isotypes, F, G, GA, GH, MO,
NY, UB, US).

Slender tree, 2-3 m tall; stems ash gray lepi-
dote-tomentulose; branches few. Leaves coria-
ceous, sessile to petiolate; petioles to 5 mm long,
lepidote-tomentulose; blades 7.5-14(-18) cm
long, 2.5-6 cm wide, elliptic, the bases acute to
attenuate, the apices rounded to obtuse, the mar-
gins entire; adaxially silvery lepidote; abaxially
silvery lepidote-tomentulose. Peduncle slender,
to 30 cm tall, terete, ash gray lepidote-tomen-
tulose. Glomerulescence a cyme of 6-20 glom-
erules. Glomerules 5-20 mm tall, 10-25 mm
diam., subspherical. Heads 10—80 per glomerule,
closely appressed, coherent for ?^ their lengths
by pubescence of phyllaries. Involucres obconic,
3.5-5 mm tall, 1-2.5 mm diam.; phyllaries in 4—
5 series; outermost phyllaries narrowly obtrul-
late, 1.7-2.9 mm long, 0.4 m wide, the
apices acute; innermost phyllaries narrowly ob-
trullate, 3.6-5.1 mm long, 0.5-1.1 mm wide, the

MACLEISH — EREMANTHUS REVISION

245

apices acuminate, margins entire to subfim-
briate; abaxial surfaces stramineous with purple
apex, glandular, glabrate to tomentose. Corollas
purple, 5.2-7.4 mm tall, the lobes acuminate.
Anthers 3-3.6 mm long, the apical appendage
acuminate to acute, the bases acute to obtuse.
Achenes 2.5—4 mm tall, glandular, sericeous, ob-
scurely ribbed, the apex slightly constricted; nec-
tary 0.2-0.5 mm tall. Pappus 3-5-seriate, of co-
roniform (at maturity; strigose bristles; outermost
series 1-2.5 mm long; innermost series 5-7 mm
long.

Flowering and fruiting occur from July to Oc-
tober

Eremanthus argenteus is found only in the

hapada dos Veadeiros of southcentral Goiás
(Fig. 3). Although common, it is restricted to the
granite outcrops of campo rupestre at high alti-
tudes.

This species resembles E. cinctus and E. au-
riculatus which also show glomerulescence re-
duction to a cyme of glomerules from the com-
pound cyme, which is more typical of sect.
Eremanthus. Eremanthus argenteus is easily
separated from both species by its silvery lepi-
dote surfaces and absence of auriculate or retuse
leaf bases and apiculate phyllaries. At present, it
does not appear to be related to any particular
species of Eremanthus.

Additional specimens examined. BRAZIL. GOIÁS:
Brasilia Richtung Campos Belos, 22 km nach Alto Pa-
raiso, 1,500 m, 28 Aug. 1981, Schumacher 1035 (GA,

M, MB, RB, W)

Possible hybrids. x E. uniflorus MacLeish & Schu-
macher. GorAs: Alto Paraiso, 28 Aug. 1981, Schu-
macher 1038, 1039, 1041 (GA).

6. Eremanthus auriculatus MacLeish & Schu-
macher, Syst. Bot. 9: 86. 1984. TYPE: Brazil.
Goiás: Brasilia Richtung Campos Belos, 22
km nach Alto Paráiso, 1,500 m, 28 Aug.
1981, Schumacher 1037 (holotype, RB;
isotypes, GA, MB).

Shrub to small tree, 0.8—1.5 m tall; stems red-

Leaves coriaceous,
cending-appressed; blades 5-8 cm
wide, trullate to ovate, the bases iia the
apices acute, the margins entire; adaxially to-
mentulose, abaxially gray-brown lanate-tomen-
tose. Peduncle slender, to 20 cm tall, ribbed, red-
dish-brown lanate-tomentose. Glomerulescence

276

a cyme (rarely compound) of 9-20 glomerules.
Glomerules 5-15 mm tall, 15-20 mm diam.,
subspherical. Heads 30-80 per glomerule, closely
appressed basally, coherent !^ length by lanate
pubescence of phyllaries. Involucres obconic, 4—
5.5 mm tall, 1.5-2.5 mm diam.; phyllaries in 4—
5 series; outermost phyllaries narrowly obtrul-
late, 2.3-4 mm long, 0.4-0 wide, the api-
ces acute; innermost phyllaries narrowly obtrul-
late, 4.2-4.6 mm long, 0.5-0.8 mm wide, the
apices apiculate; margins entire to subfimbriate;
abaxial surfaces stramineous with purple apex,
often axially green, tomentulose to lanate. Co-
rollas purple, 5.5-7 mm tall, the lobes acuminate
to acute, sparsely glandular. Anthers 3.3-3.6

sericeous, obscurely-ribbed, apex slightly con-
stricted; nectary 0.2-0.5 mm tall. Pappus 3-5-
seriate, of coroniform (at maturity) strigose
bristles; outermost series 1.2-2.2 mm long; in-
nermost series 5.5-7 mm long.

Flowering and fruiting occur from July to Sep-
tember

Eremanthus auriculatus is known only from
the Chapada dos Veadeiros in southcentral Goiás
(Fig. 3). It occurs on campo rupestre adjacent to
the cerrado characteristic of this region.

This species superficially resembles E. cinctus;
however, E. auriculatus is distinguished by as-

represents a further reduction in glomerulescence
internodes from a compound cyme to form a
cyme of glomerules. It is easily separated from
the glomerulatus-complex by the cyme of glom-
erules (vs. compound cyme), by auriculate leaf
bases (vs. acute to obtuse), and by apiculate phyl-
laries (vs. acute to acuminate).

Additional aeg examined. BRAZIL. GOIÁS: Alto
Paráiso, 4 Jul. 1983, Schumacher 3045 (GA, MB).

7. Eremanthus cinctus Baker in C. Martius, Fl.
Bras. 6(2): 162. 1873. TYPE: Brazil. Mato
Grosso: *Cuyaba" (now Cuiabá), 1834, Sil-
va Manso 55 (holotype, BR).

Eremanthus ated piae Baker in C. Martius, Fl. Bras.

y! 1873. TYPE: Brazil, no other data, Tam-
berlik s.n. ADS. W; photos of W, F, TEX).

Shrub or small tree, 0.5—4 m tall; stems red-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

dish-brown lanate-tomentose; branches few.
Leaves subcoriaceous, sessile; blades 6-19 cm
long, 2-6 cm wide, elliptic, narrowly elliptic, or
obovate, the bases retuse to acute, the apices
acute to obtuse, the margins entire; adaxially gla-
brate, abaxially tomentulose. Peduncle robust, to
15 cm tall, ribbed, reddish-brown lanate-tomen-
tose; 3-9 bracts with prominently enlarged bases
surrounding each glomerule. Glomerulescence a
cyme (rarely compound) of 5-30 glomerules.
Glomerules 10-25 mm tall, 10-25 mm diam.,
hemispherical. Heads 20-50 per glomerule,
closely appressed, coherent entire length by la-
nate pubescence of phyllaries. Involucres obcon-
ic, 3.5-5 mm tall, 0.7-2 mm diam.; phyllaries in
4—5 subequal series; outermost phyllaries oblan-
ceolate, 3.3-4.3 mm long, 0.5—0.7 mm wide, api-
ces obtuse; innermost phyllari ,3-
4.8 mm long, 0.7-1 mm wide, apices acute; mar-
gins entire; abaxial surfaces stramineous with

acuminate and glandular. Anthers 4—4.4 mm long,
apical appendage acuminate, bases acute.
Achenes 2.5-3.5 mm tall, glandular, sericeous
between 10 ribs, apex slightly constricted and
dark; nectary 0.1-0.2 mm tall. Pappus 3-4-se-
riate, of stramineous (occasionally rusty on dried
S mens), coroniform b maturity) strigose
de outermost series 1.5-2.5 mm long; in-
nermost series 5.5-7.5 mm long.

Flowering and fruiting occur from April to
September.

Eremanthus cinctus is known only from three
locations: western Minas Gerais, western Goias,
and near Cuiabá in central Mato Grosso (Figs.
2-4). Schumacher was able to locate this species
in southwestern Goiás, but reported (pers. comm.)
that the Minas Gerais locations cited by Good-
land on his 1967 collections have been com-
pletely logged for charcoal.

Baker (1873) separated E. pandurifolius from
E. cinctus on the basis of a solitary glomerule
versus a cyme of glomerules. This distinction is
artifactual — the type specimen of E. pandurifo-
lius is fragmented, making the glomerules appear
solitary. Eremanthus pandurifolius appears to
have been collected in western Minas Gerais,
because that is the region Tamberlik is known
to have explored (Urban, 1906). Likewise, Silva
Manso was known to have been practicing med-
icine in Cuiaba during the year recorded on the
label ofthe type specimen of E. cinctus. Although

1987] MACLEISH — EREM.

E. cinctus and E. pandurifolius were published
simultaneously, E. cinctus has been chosen be-
cause the type specimen is intact, whereas that
of E. pandurifolius is badly fragmented. Until
recently E. cinctus appeared to have a disjunct
distribution since there were no collections from
intervening mountainous regions, particularly
Serra do Verdinho, Serra das Araras, and Serra
da Sào Jeronimo.

Eremanthus cinctus superficially resembles E.
auriculatus but is distinguished by subcoriaceous
leaves with acute to retuse bases, several prom-
inent extraglomerular bracts, and total fusion of
heads. By contrast, E. auriculatus has coriaceous
leaves with prominent auriculate bases, and heads
appressed only at the bases. Eremanthus cinctus,
like E. auriculatus, is clearly related to E. glo-
merulatus, and represents a further reduction in
glomerulescence internodes from a compound
cyme to form a cyme of glomerules.

Additional pe in examined. BRAZIL. MATO
GROSSO: known onl

GERAIS:
Goodland 3766 (UB). Prata, 29 Sep. 1967, Goodland
1 (MO). Go1Ás: Montividiu towards Amorinopolis,
21 km after Montividiu, 800 m, 28 Apr. 1984, Schu-
II 4004 (GA, MB, UNICAMP).

subg. Isotrichia (DC.)

acLeis . nov. Albertinia Sprengel
sect. gn ia DC., Prodr. 5: 82. 1836.
Vanillosmopsis Schultz-Bip. subg. [sotrichia
(DC.) Schultz-Bip., Jahresber. Pollichia 18-
19: 168. 1861. LECTOTYPE, here designated:
E. incanus (Less.) Less.

Eremanthus Less.
M

Leaves coriaceous. Glomerulescence a com-
ound cyme of 8-50 glomerules. Glomerules
Su pherical Heads per glomerule 30-100, cy-
lindric, connate by concrescence of tissues, with
phyllaries in 4-6 series. Florets 1 per head.
Achenes abe 1 5-20-ribbed, sericeous. Pap-
pus off-white to purple, subcoroniform, tardily
deciduous, subpaleaceous.

8. Eremanthus incanus (Less.) Less., Linnaea 6:

682. 1831. Albertinia incanus Less., Linnaea

4: 342. 1829. Cacalia incana (Less.) Kuntze,

Revis. Gen. Pl. 2: 970. 1891. LECTOTYPE,

here designated; Brazil, no other data, Sel-

low s.n. (B; isolectotypes, BR, F (fragment
from P))

Slender to robust tree, 2-10 m tall, to 15 cm

diam.; stems gray-brown lepidote; branches few.

Leaves coriaceous, petiolate; petioles 4-17 mm

REVISION 277

long, lepidote; blades 5.5214 cm long, 2-6 cm
wide, elliptic to ovate, the bases acute, the apices
rounded to obtuse, the margins entire; adaxially
glabrate, abaxially gray lepidote. Peduncle slen-
der, to 27 cm tall, terete, gray lepidote. Glomer-
ulescence a compound cyme of 8-50 glomerules.
Glomerules 5-15 mm tall, 7-15 mm diam.,
spherical. Heads 30-100 per glomerule, closely

crescence of p
dric, 2.4-5 mm tall, 0.3-1.5 m ; phy
ries in 4—5 series; outermost phyllaries irianaulét:
1.7-3 mm long, 0.1-0.5 mm wide, the apices
acute to obtuse; innermost phyllaries narrowly
obtrullate, 2.6-4 mm long, 0.3-0.6 mm wide,
the apices acuminate, the margins entire; abaxial
surfaces stramineous with purple apex, glabrate
to lepidote-tomentulose. Corollas pale purple to
white, 4.2—6.6 mm tall, the lobes acuminate. An-
thers 2.2-2.7 mm long, the apical appendage acu-
minate, the bases acute. Achenes cylindric, 2.2-
2.6 mm tall, glandular, sparsely sericeous, 15-
20-ribbed, the apex slightly constricted; nectary
0.15-0.25 mm tall. Pappus 3—4-seriate, off-white
urple, subcoroniform (at maturity), tardily
Pied Soiree naqa strigose bristles; out-
ermost series 1-1.5 mm long; innermost series
5-5.8 mm ee

2

Flowering and fruiting occur from July to Oc-
tober

Eremanthus incanus is distributed throughout
the southern section and northeastern arm (Figs.
5, 6) of the Central Plateau of Brazil at elevations
of 800 to 1,850 meters in cerrado, secondary
forest, or caatinga. It is particularly common in
Minas Gerais where it is called “pão candeia"
Scone and or “‘candeia da serra" (mountain

dle), names Mer d attributed to subg. Va-
UNE
Eremanthus incanus is transitional between

gle floret per head, large sericeous achenes, an
subpaleaceous pappus. In contrast, in its lepidote
tomentum, relatively small glomerules, phyllary
tissue fusion, cylindrical achenes, and off-white
to purp ]
E. pine net of the Vanillosmopsis complex.
Lessing’s specimens were deposited at Char-
kow. Because Lessing’s original material was un-
available, a Sellow collection from B herbarium
that bears appropriate annotations by Lessing

4 ED ly
e, tardi

itapproaches

—

Elevation
jee vane, Oy meters
. 1000

T T T
0 100 (

FIGURE 5.

was chosen as the lectotype. Historically, the
name Albertinia bicolor (now Paralychnophora
bicolor) has been confused often with E. incanus
due to the prominent spherical glomerules in both
taxa. Paralychnophora bicolor is characterized by
solitary, relatively large, axillary glomerules, 2-
4 or 8-12 florets per head, and angled, glabrous
achenes with a biseriate pappus.

Additional specimens examined. BRAZIL. BAHIA:

n. de das Contas, Pico das Almas at 18 km
e Rio das Contas, 1,600-1,850 m, 24 Jul. 1979
King et al. 8136 (CEPEC, U INAS GERAIS
between Paràopeba and Diamantina, 3 Oct. 1980
MacLeish et al. 687 (C, G,
, US), between Mendanh

a and Diamantina, 4 Oct.
1980, MacLeish et al. 703 (BR, CEPEC, F, GA, M,
MO, RB, TEX).

Possible hybrids. x E. elaeagnus (C. Martius ex
DC.) Schultz-Bip. MINAS GERAIS: between Paraopeba
and Diamantina, 3 Oct. 1980, MacLeish et al. 688 (F,
G, GA, GH i
(DC ) MacLeish. MINAS
Barreto 47 x E. u

MINAS GERAIS: Itamarandiba, 6 Sep
1116a (GA, MB).

.) MacLeish.
. 1981, Schumacher

Eremanthus Less. subg. Pseuderemanthus
Schultz-Bip., Jahresber. Pollichia 20-21:

ANNALS OF THE MISSOURI BOTANICAL GARDEN

deis:
Sere

[Vor. 74

n mealet

Distribution of Eremanthus subg. Isotrichia and subg. Pseuderemanthus in Minas Gerais.

395. 1863. Eremanthus Less. subg. Pseu-
deremanthus Schultz-Bip. “A. Elaeagnus”
Schultz-Bip., Jahresber. Pollichia 20-21:
395. 1863. LECTOTYPE: here designated; Er-
emanthus elaeagnus (C. Martius ex DC.)
Schultz-Bip.

Leaves coriaceous. Glomerulescence a com-
ispheric g lomerules.

rets 3-4 per head. Achenes cylindric, "10- ribbed,
sericeous. Pappus stramineous or purple, coro-
niform, persistent, subpaleaceous.

9. Eremanthus seidelii MacLeish & Schumach-
er, Syst. Bot. 9: 89. 1984. TYPE: Brazil. Minas
Gerais: Furnas in Richtung Piui, kurz von
Staumauer, 800 m, 25 Aug. 1981, Schu-
macher 1006 (holotype, RB; isotypes, 2 in
GA, K, M, MB

Tree, to 4 m tall; stems gray lepidote-tomen-
tulose; branches many. Leaves coriaceous, sessile
to petiolate; petioles to 10 mm long, densely lep-
idote-tomentose; blades 4-9 cm long, 1.2-3 cm
wide, elliptic, the bases acute, the apices acu-

1987]

~

E. arboreus

_

. brasiliensis

c<
44
\

E. capitatus

e Ó © >

E. glomerulatus

@ E. graciellae
* E. incanus

O E. pohlii

x E. uniflorus

rr

Elevation:

100 200 km

FIGURE 6.
oiás

minate to acute, the margins entire; adaxially
sparsely lepidote, abaxially densely gray lepi-
dote-tomentose. Peduncle slender, to 15 cm tall,
terete, gray lepidote-tomentulose. Glomerules-
cence a compound cyme of 10-60 glomerules.
Glomerules 5-10 mm tall, 6-15 mm diam.,

hemispherical. Heads 1-7 per glomerule, closely
ai Bis basally, free. Involucres cylindric, 5.5-
7.2 mm tall, 2.5-4 mm diam.; phyllaries in 4—6
scies: outermost phyllaries trullate, 1.3-4 mm
long, 1-1.5 mm wide, the apices acute; innermost
phyllaries narrowly rhombic to obtrullate, 4.7—
7 mm long, 0.5-1.2 mm wide, the apices acute;
margins entire to subfimbriate; abaxial surfaces
stramineous with purple apex, lepidote-tomen-
tulose. Florets 3 per head; corollas purple to white,
5.4—6.5 mm tall, the lobes acuminate. Anthers
2.5-3 mm long, the apical appendage acuminate,
the bases acuminate. Achenes 2.2-3.1 mm tall,
glandular, sparsely sericeous between 10 ribs,
apex slightly constricted; nectary 0.1—0.2 mm tall.
Pappus 3-4-seriate, of stramineous, subcoroni-
form (at maturity) strigose bristles; outermost

MACLEISH — EREMANTHUS REVISION

279

LR Nis "f
z hus eer s o A
i A Ó `
Y $$;

Distribution of Eremanthus subg. Vanillosmopsis, E. glomerulatus, and E. incanus in Bahia and

series 0.8-1.2 mm long; innermost series 4.5—6.2
mm long.

Flowering and fruiting occur from June to Au-
gust.

Eremanthus seidelii is restricted to the cerrado
surrounding the Furnas reservoir in southwest-
ern Minas Gerais (Fig. 5). This species is closely
related to E. elaeagnus, which is restricted to the
Serra da Espinhaco of northeastern Minas Ge-
rais. However, E. seidelii is distinguished from
E. elaeagnus by having fewer heads per glomer-
ule (1-7 vs. 9-20), straw vs. mostly purple pap-
pus color, elliptic vs. narrowly elliptic leaves, and
flowering June-July vs. August-September

Eremanthus seidelii and E. elaeagnus are tran-
sitional between subg. Eremanthus and subg.
Vanillosmopsis. They approach E. glomerulatus
in their coriaceous leaves, only 10-60 glomerules
per glomerulescence, large sericeous achenes, and
persistent subpaleaceous pappus. In contrast, in
their lepidote tomentum, 1-20 heads per glom-
erule, relative lack of head fusion, 3-4 florets per

280

head, and occasionally purple pappus they ap-
proach many members of subg. Vanillosmopsis.

Additional specimens examined. BRA MINAS
GERAIS: vor Alpinopolis, 12 Jun. 1977, Seidel s n. (RB);
bei Capitolio, 800 m, 25 Aug. 1981, Schumacher n
(GA, MB); Furnas in Richtung Capitolio, 18 Oct. 1980,
Sc humacher s.n. (GA, MB).

Possible hybrids. x E. erythropappus (DC.)
MacLeish. MINAS GERAIS: Furnas towards Piui, 25 Aug.
1981, Schumacher 1005 (GA); Alpinopolis, 12 Sep.
bel Seidel s.n. (RB). x E. glomerulatus Less. MINAS

AIS: between Furnas and Piui, 25 Aug. 1981, Schu-
mac che 1008 (GA). x E. mattogrossensis Kuntze. MI-
NAS GERAIS: Furnas towards Piuí, 25 Aug. 1981, Schu-
Mule 1003, 1004 (GA).

10. Eremanthus elaeagnus (C. Martius ex DC.)
Schultz-Bip., Jahresber. Pollichia 20-21:
395. 1863. Albertinia elaeagnus C. Martius
ex DC., Prodr. 5: 81. 1836. Vernonia elaeag-
nus (C. Martius ex DC.) Schultz-Bip., Jah-
resber. Pollichia 18-19: 166. 1861. TYPE
Brazil, Minas Gerais, altis lapidosis Serro
Frio prope Tejuco (now Diamantina), Mar-
tius s.n. (holotype, M).

Small tree or shrub, 2-5 m tall; stems gray
lepidote-tomentulose; branches many. Leaves
coriaceous, sessile to petiolate; petioles to 10 mm
long, densely lepidote-tomentose; blades 5-12 cm
long, 1.5-3 cm wide, narrowly elliptic, the bases
acute to acuminate, the apices acute to rounded,
the margins entire; adaxially sparsely lepidote,
abaxially densely gray lepidote-tomentose. Pe-
duncle slender, to 14 cm tall, terete, gray lepi-
dote-tomentulose. Glomerulescence a com-
pound cyme of 10-60 glomerules, these 5-10
mm tall, 10-15 mm diam., hemispherical. Heads
9-20 per glomerule, closely appressed basally,
free. Involucres cylindric, 5-10 mm tall, 2.5-4
mm diam.; phyllaries in 4—5 series; outermost
phyllaries trullate, 1.5-3 mm long, 0.6-1.2 mm
wide, the apices acute; innermost phyllaries nar-
rowly rhombic to obtrullate, 3.5-6.5 mm long,
0.7-1. ak = wide, fue pieces sente: mangas en-
tiret
with purple apex, lepidote-tomentulose. pa
3-4 per head; corollas purple to white, 5.5-7 m
tali, the lobes acuminate, AE is 5- 3 mm s

ate,the bas-
es acuminate. Achenes 2.5-3.5 mm tall, glan-
dular, sparsely sericeous between 10 ribs, the apex
slightly constricted; nectary 0.1-0.15 mm tall.
Pappus 3-4-seriate, of purple (rarely stramin-
eous), subcoroniform (at maturity), strigose bris-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

tles; outermost series 1-1.5 mm long; innermost
series 6-6.5 mm long.

Flowering and fruiting occur from August to
September.

Eremanthus elaeagnus is restricted to the Ser-
ra do Espinhaco in northeastern Minas Gerais at
elevations of 900 to 1,370 meters (Fig. 5). It is
locally common and dominant along with mem-
bers of Lychnophora in campo rupestre. Because
ofthe gray tomentum covering most of this plant's
surface and its general resemblance to members
ofthe Vanillosmopsis complex, it is locally known
as "candeia branca,” or white candle. This taxon
is closely related to E. seidelii and, as noted under
that species, both represent intermediates be-
tween subgenera Eremanthus and Vanillosmop-

Additional Edda examined. BRAZIL. MINAS
GERAIS: between Paraopeba and Diamantina. 3 Oct.
1980, MacLeish et al. 689 (F, GA, GH, MO, NY, RB);

Possible hybrids. x E. glomerulatus Less. MINAS
GERAIS: Serra do Cipó, 7 Oct. rien icr et al.
715 (BR, F, GA, GH, K M MO, NY, RB, S, US),
MacLeish et al. 716 (GA, RB), 30 m 1980, King &
Almeda seis S). x E. incanus (Less.) Less. MINAS
GERAIS: between Paraopeba and Diamantina 3 Oct
ee MacLeish et al. 6 , GA MO, NY,

US). x E. GERA (DC.) Ma yis MINAS
GERAIS: Serra do Cipó. 7 Oct. 1980, MacLeish et al.
718 (GA).

Eremanthus Less. subg. Vanillosmopsis (Schultz-
Bip.) MacLeish, comb. nov. Vanillosmopsis
Schultz-Bip., Jahresber. Pollichia pa 19:
166. 1861. Vanillosmopsis subg. Euvanil-
losmopsis Schultz-Bip., Jahresber. Pollichia
20-21: 398. 1863. TYPE: Vanillosmopsis
glomerata Schultz-Bip. = Eremanthus
erythropappus (DC.) MacLeish.

eaves membranaceous to subcoriaceous.
Glomerulescence a compound cyme of 100 or
more hemispheric glomerules. Heads per glom-
erule 1-12, cylindric or obconic, solitary or
slightly to closely coherent by interwoven pu-
bescence of phyllaries or connate by tissue con-
crescence, phyllaries in 5-7 series. Florets per
head 1-4. Achenes cylindric (rarely cylindric-
turbinate), 10-ribbed, glabrate. Pappus purple or
white (rarely stramineous), not coroniform,
promptly (rarely tardily) deciduous, filiform.

1987]

gii suem Less. sect. Nectaridium (Schultz-

MacLeish, comb. nov. Vanillosmopsis

Ede: Bip. subg. Nectaridium Schultz-Bip.,

Jahresber. Pollichia 20-21: 400. 1863. TYPE:

Vanillosmopsis brasiliensis (Gardner)

Schultz-Bip. = Eremanthus brasiliensis
(Gardner) MacLeish.

Heads cylindric, solitary or in pairs slightly
appressed basally. Pappus never twisted.

11. Eremanthus pohlii (Baker in C. Martius)
MacLeish, comb. nov. Vanillosmopsis poh-
lii Baker in C. Martius, Fl. Bras. 6(2): 18.
1873. TYPE: Brazil: foz do Viera, Pohl 556
(holotype, K, not seen; isotypes, 2 in F (one
a photo of B), 2 in GH (one a photo of B),
NY)

Small tree, to 3 m tall; stems gray-brown lep-
idote; branches many. Leaves subcoriaceous,
petiolate; petioles 7-10 mm long, lepidote; blades
6.4—7.6 cm long, 1.8-2.4 cm wide, lanceolate,
the bases attenuate, the apices obtuse, the mar-
gins entire; adaxially glabrate, abaxially gray lep-
idote. Peduncle slender, to 25 cm tall, terete, gray
lepidote. Glomerulescence a compound cyme of
numerous (> 100) glomerules. Glomerules 6-9
mm tall, 5-12 mm diam., hemispherical. Heads
1-2 per glomerule, slightly w free. In-
volucres cylindric, 5-7.5 m 2-3.5 m
diam. with prominent truncate ie patera
in 6—7 series; outermost phyllaries widely deltate,
0.5-1.2 mm long, 0.5-1.0 mm wide, the apices
obtuse; innermost phyllaries lanceolate, 4—6.2
mm long, 0.6-2 mm wide, the apices acute to
acuminate; margins entire to subfimbriate; abax-
ial surfaces stramineous with purple apex, gla-
brate to lepidote-tomentulose. Florets 3 per head;
corollas purple, 5-6 mm tall, the lobes acumi-
nate. Anthers 2-3 mm long, the apical appendage
acuminate, the bases acute to rounded. Achenes
2.5-4 mm tall, sparsely glandular, rarely pilose,
10-ribbed, the apex slightly constricted and dark;
nectary 0.2-0.5 mm tall. Pappus 3-4-seriate, of
stramineous (rarely purple), tardily deciduous,
strigose bristles; outermost series 1-2 mm long;
innermost series 5.5-7.5 mm long.

Flowering and fruiting occur from July to Sep-
tember

Eremanthus pohlii is restricted to the Chapada
dos Veadeiros, which runs north to south in south
central Goiás, and nearby parts of Minas Gerais

MACLEISH — EREMANTHUS REVISION

281

(Figs. 6, 7). It occurs in colonies on the cerrado
at high elevations (ca. 1,400 m).

Eremanthus pohlii is one ofthe more primitive
(i.e., approaching Vernonia) members of the subg.
Vanillosmopsis. It is closely related to E. gra-
ciellae and E. brasiliensis, both of which are re-
stricted to the northwestern arm of the Central
Plateau. Eremanthus pohlii can be separated from
E. graciellae and E. brasiliensis by its three flo-
rets per head and relatively long leaves (6.5-7.5
vs. 2.5-6.5 cm long). The only other member of
the Vanillosmopsis complex that occurs sym-
patrically with these taxa is E. uniflorus. Ere-
manthus uniflorus is most closely related to E.
capitatus and is distinguished (among other less
discriminating characters) by a single floret per
head and 3-9 heads per glomerule.

Additional specimens examined. BRAZIL. GOIAS:
Brasilia towards Campos Belos, 15 km from Alto Par-
áiso, 1,400 m, 28 Aug. 1981, Schumacher 1032, 1033
(GA, MB), Schumacher 1034 (MB, NY). MINAS GERAIS:
S Pinheiro, 15 Aug. 1967, Heringer 11531 (NY,

RB).

12. Eremanthus graciellae MacLeish & Schu-
macher, Syst. Bot. 9: 87. 1984. TYPE: Brazil.
Bahia: BR 020 Brasília Richtung Barreiras,
15 km weiter in Richtung Barreiras von Fa-
zenda Prainha, km 374, 800 m, 28 Aug.
1981, Schumacher 1048 (holotype, RB; iso-
types, GA, K, M, MB)

Small tree, to 2.5 m tall; stems gray-brown
lepidote; branches many. Leaves subcoriaceous,
petiolate; petioles 8-15 mm long, lepidote; blades
4.5-6.5 cm long, 1.5-3 cm wide, elliptic to nar-
rowly elliptic, the bases acute, the apices acute
to obtuse, the margins entire; adaxially glabrate,
abaxially gray lepidote. Peduncle slender, to 25
cm tall, terete, gray lepidote. Glomerulescence a
compound cyme of numerous (more than a
hundred) glomerules. Glomerules 6-9 mm tall,
4-9 mm diam., obconic. Heads 1-2 per glomer-
ule, slightly appressed, free. Involucres cylindric,
5.2-7.5 mm tall, 2.1-2.8 mm diam., with prom-
inent truncate base; phyllaries in 6-7 series; out-
ermost phyllaries widely deltate, 0.6-1.2 mm
long, 0.7-1.2 mm wide, the apices acute; inner-
most phyllaries lanceolate, 3.8—6. Ë mm long, 0.6—
1.4 mm wide, the apices acute acuminate;
margins entire to subfimbriate; iud ee ee
stramineous with purple apex, glabrate to lepi-
dote-tomentulose. Florets 2 per head; corollas
purple to white, 4.2-7 mm tall, the lobes acu-

282
© E. capitatus Sani “ae
\
Jes
A E. erythropappus 1 ox J
e EA
O E. pohlii
O E. polycephalus

Q:

x `
ROME y
M

"X ES p

A. Wes ades

FIGURE 7.

minate. Anthers 2.1-2.8 mm long, the apical ap-
pendage acuminate, the bases acuminate.
Achenes 2.2-3.5 mm tall, sparsely glandular, 10-
ribbed, the apex slightly constricted and dark;
nectary 0.2-0.5 mm tall. Pappus 3-5-seriate, of
stramineous to white, deciduous, strigose bris-
tles; outermost series 1-2 mm long; innermost
series 5-7.2 mm long

Flowering and fruiting occur from July to Au-
gust

Eremanthus graciellae is distributed along the
Serra Geral de Goiás, which runs north to south
along the border of Goiás and Bahia (Fig. 6). It
occurs in colonies on the quartzite grasslands
characteristic of this plateau.

Eremanthus graciellae is also one of the prim-
itive (approaching Vernonia) members of Ere-
manthus. It can be separated from close relatives,
E. graciellae and E. brasiliensis, by its two florets
per head and intermediate length leaves.

Additional specimens examined. BRAZIL. BAHIA: BR

eis Brasília Richtung Barreiras, 15 km weiter in Rich-
ung Barreiras von Fazenda Prainha, 800 m, 28 Aug.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Elevation:
500 meters
1000 i

Me AME A.

o 200

kilometers

Distribution of Eremanthus subg. Vanillosmopsis in Minas Gerais.

1981, as 1046 (GH, MB), Schumacher 1047
(GA, MB).

13. Eremanthus brasiliensis (Gardner) Mac-
eish, comb. nov. Monosis brasiliensis
Gardner, London J. Bot. 5: 230. 1846. Ver-
nonia brasiliensis (Gardner) Schultz-Bip.,
Jahresber. Pollichia 18-19: 161. 1861, nom.
illeg. (nom. rej. pro Vernonia brasiliensis
(Sprengel) Less., Linnaea 6: 681-682. 1831.

= Albertinia brasiliensis). Vanillos osmopsis
brasiliensis (Gardner) Schult

ia 0.

F (fragments from G and P, photos of G and
; )

Small tree, to 4 m tall; stems gray-brown lep-
idote; branches many. Leaves subcoriaceous,
petiolate; petioles 5-10 mm long, lepidote; blades
2.4-4.2 cm long, 0.8-2.6 cm wide, elliptic to
narrowly elliptic, the bases acute to attenuate,
the apices obtuse, the margins entire; adaxially

1987]

glabrate and glandular, abaxially gray lepidote.
Peduncle slender, terete, gray lepidote. Glomer-
ulescence a compound cyme of 100 or more
glomerules. Glomerules 6-8 mm tall, 4-8 mm
diam., obconic. Heads 1-2 per glomerule, slight-
ly appressed, ie Involucres cylindric, 5-6.5 m
tall, 2-2. iam.; phyllaries in 6-7 series;
outermost uires widely deltate, 0.6-1 mm
long, 0.5—1 mm wide, the apices acute; innermost
phyllaries lanceolate, 3.5-6 mm long, 0.5-1 mm
wide, the apices acute to acuminate; margins en-
tire to subfimbriate; abaxial surfaces stramin-
eous, glabrate to lepidote-tomentulose. Florets 1

er head; corollas not seen. Achenes 2-3.5 mm
tall, sparsely glandular, obscurely-ribbed, the apex
slightly constricted and dark; nectary 0.3-0.4 mm
tall. Pappus 3-4-seriate, of stramineous, tardily
deciduous, strigose bristles; outermost series 1—
2 mm long; innermost series 5-7 mm long.

Flowering and fruiting probably occur from
August to September.

Eremanthus brasiliensis is distributed in the
northern part of the Serra Geral de Goias (Fig.
6), which runs north to south along the border
of Goiás and Bahia. This species is known only
from the type specimens. Schumacher (1982) un-
successfully attempted to locate it at the same
time that he rediscovered E. arboreus in southern
Ceará.

Like E. graciellae and E. pohlii, E. brasiliensis
is one of the more primitive members of the
Vanillosmopsis complex. It is distinguished by
one floret per head, small leaves (2.5—4.5 vs. 4.5-
7.5 cm long), and heads which lack the promi-
nent truncate bases of its closest relatives.

Eremanthus Less. sect. Vanillosmopsis (Schultz-
Bip.) MacLeish, comb. nov. Vanillosmopsis
subg. Euvanillosmopsis Schultz-Bip., Jah-
resber. Pollichia 20-21: 398. 1863. LEC-
TOTYPE: Eremanthus erythropappus (DC.)
MacLeish.

Heads cylindric or obconic. Heads per glom-
erule (2-)5—12. Pappus often twisted.

14. Eremanthus polycephalus (DC.) MacLeish,
comb. nov. Albertinia polycephala DC.,
Prodr. 5: 82. 1836. Vanillosmopsis poly-
cephala (DC.) Schultz-Bip., Jahresber. Pol-
lichia 18-19: 168. 1861. TYPE: Brazil. Minas
Gerais: planitie alta ad Piedade Villam,
Martius s.n. (holotype, M; isotype, M).

MACLEISH — EREMANTHUS REVISION

283

Albertinia pcd: C. Martius ex DG., Prodr. 5: 82.
1836. Vanillosmopsis saligna (DC. ) Schultz-Bip.,
eed iai Pollichia 18-19: 168. 1861. TYPE: Bra-
zil. Minas Gerais: in editis siccis rupestribus mon-
ium Serro Frio, Martius s.n. (holotype, M).

Shrub or small tree, to 3.5 m tall; stems gray-
brown lepidote; branches many. Leaves mem-
branaceous to subcoriaceous, sessile to petiolate;
petioles to 8 mm long, lepidote; blades 4.5-7 cm
long, 1-1.8 cm wide, lanceolate, the bases acu-
minate to attenuate, the apices acute to obtuse,
the margins entire; adaxially glabrate, abaxially
gray lepidote. Peduncle slender, to 23 cm tall,
terete, gray lepidote. Glomerulescence a com-
pound cyme of 100 or more glomerules, these
5—7 mm tall, 4-8 mm diam., hemispherical.
Heads 6-12 per glomerule, closely appressed,
connate !'^ lengt
tissues. Involucres cylindric, 4—5.
2 mm diam.; phyllaries in 6—7 series; outermost
phyllaries widely deltate, 0.6-1.2 mm long, 0.7-
1.2 mm wide, the apices obtuse; innermost phyl-
laries lanceolate, 3-5 mm long, 0.5—1.2 mm wide,
the apices acute as Svar the margins entire
purple apex, lepidote-tomentose. Florets 1 per
head; corollas purple, 4—6.5 mm tall, the lobes
acuminate to acute. Anthers ae : mm long, the
apical appendage acuminate, ases acute.
Achenes 1.5-2.8 mm tall, ee 10-ribbed,
the apex slightly constricted and dark; nectary
0.2-0.4 mm tall. Pappus 3—4-seriate, of stramin-
eous, white or purple, tardily deciduous, strigose
bristles; outermost series 0.2-1 mm long, the in-
nermost series 3-5.5 mm long.

Flowering and fruiting occur from June to No-
vember

Eremanthus polycephalus 1s distributed along
the Serra do Espinhaço, which runs north to south
through the center of Minas Gerais (Fig. 7). It
occurs in colonies on the campo rupestre habitats
characteristic of high elevations (700-1,370 me-
ters). The common name of “‘candeia,”’ or can-
dle, is applied locally.

Eremanthus polycephalus is most closely re-
lated to E. erythropappus and is sympatric with
it in the northernmost portion of its range.
Uniquely, in subg. Vanillosmopsis, both of these
taxa exhibit concrescence of phyllary and recep-
tacle tissues. However, E. polycephalus is easily
distinguished by its single floret per head (vs. 3-
4) and partial fusion of heads (vs. nearly total
fusion). This species, because of its single floret

284

per head and various characteristics associated
with the Vanillosmopsis complex, superficially
resembles E. uniflorus and E. brasiliensis, taxa
that are restricted to Goiás and nearby Bahia and
which are distinguished by their lack of tissue
fusion within glomerules.

Additional dodge examined. BRAZIL. MINAS
GERAIS: 14 km NE o mantina towards Mendanha
on Estrada 367, 1,370 A p Oct. 1980, MacLeish et al.
702 (GA, GH, K, LE, NY, RB, UB, US
Possible hybrids. x E. elaeagnus (C. Martius ex
DC.) Schultz-Bip. MINAS GERAIS: Serra do Cipó, 7 Oct.
35$ MacLeish et al. 718 (GA). x E. incanus (Less.)
MINAS GERAIS: Itamarandiba, 6 Sep. 1981, Schu-
LU 1116a (GA).

15. Eremanthus erythropappus (DC.) Mac-
Leish, comb. nov. Albertinia erythropappa
DC., Prodr. 5: 82. 1836. Vanillosmopsis
erythropappa (DC.) Schultz-Bip., HUE er.
Pollichia 18-19: 167.
Minas Gerais: Marianne, Vauthier 334 (ho-
lotype, G-DC (as IDC microfiche); isotypes,
G, 2 in GH).

Albertinia p lay d London J. Bot. 5: 235.
l : Brazil. Minas Gerais: near Villa do
Principe, nei 1840, Gardner 48 12 (holotype, BM;

s, F, 2 in G, 2 in GH, 3 in NY, S).

Vanillosmopsis srera Schultz- Bip., Jahresber.

Pollichia 18-19: 167. . Vernonia glomerata
Schultz-Bip., Bot. Zeitung. (Berlin) 3: 155. 1845.
nom. nud
syntypes; Brazil. Minas Gerais: Aug.-Apr. 1840,
Claussen 2063 (G; isolectotypes, G, 2 in GH, M,
MO). Syntypes: Brazil. (Rio de Janeiro): in col-
libus siccis calidisque inter Mandioca et Serra Es-
trella, Riedel s.n. (577) (not found); no other data,
Claussen 863 (853) (not found); no other data,
Schucht 75 (W, not found)

Shrub or tree, to 10 m tall, to 10 cm diam.;
stems gray-brown lepidote; branches many.
Leaves membranaceous, petiolate; petioles 3-15
mm long, lepidote; blades 5—10.2 cm long, 1.6—
3.6 cm wide, oblanceolate to elliptic, the bases
acute to acuminate, the apices acute to attenuate,
the margins entire; adaxially glabrate to lepidote,
abaxially gray lepidote. Peduncle slender, to 23
cm tall, terete, gray lepidote. Glomerulescence a
compound cyme of 100 or more oe
Glomerules 6-15 mm tall, 4-12 m m.,

per cole closely
appressed, connate nearly entire length by con-
crescence of phyllary tissues and pubescence. In-
Moin obconic, 5-7.2 mm tall, 2.1-4.2 m
diam.; phyllaries in 5—6 series; outermost p
alice widely deltate, 0.6-1.5 mm long, 0.7-1.3

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

mm wide, the apices acute; innermost phyllaries
lanceolate, 3.6-6.8 mm long, 0.5-1.2 mm wide,
the apices acute to acuminate; margins entire to
subfimbriate; abaxial surfaces stramineous with

all, the lobes acu-
minate. Anthers 2.1—2.7 mm long, the apical ap-
pendage acuminate, the bases acute. Achenes
cylindric-turbinate, 1.2—2.5 mm tall, glandular,
10-ribbed, the apex slightly constricted and dark;
nectary 0.15-0.25 mm ls Pappus 3-4-seriate,
of stramineous, white or purple, often curling,
promptly deciduous, ida. bristles; outermost

eries 1.2-3 mm long; innermost series 3-6.5
mm long.

Flowering and fruiting occur from June to No-
vember.

remanthus erythropappus is distributed
throughout southeastern portions of the Central
Plateau (Fig. 7) at 700 to 2,400 meters. It is ex-
tremely common in colonies amidst secondary
forest of the coastal range and cerrado of the
interior plateau. It is often used as a living fence
around fields because the high terpenoid content
of its leaves deters insects. Also, it is commonly
known as “‘candeia,” candle, or “pau de can-
deia," candlestick.
Because of its extensive range, E. erythropap-

manthus erythropappus is further distinguished
by its 3-4 florets per head (vs. 1), 6-12 heads
per glomerule (vs. 30-100 in E. incanus), and
nearly total connation of heads (vs. partial con-
nation in E. polycephalus). The epithets assigned
by Gardner (1846) and Schultz-Bip. (1861) to
this species are indicative of the variation as-
cribable to age and habitat differences.

Additional specimens examined. BRAZIL. ESPÍRITO
SANTO: Mun. de Cachoeus do Itapemirim, Vargem Alta
Morro do Sal, 700 m, 3 Aug. 1980, Ferreira & Borges
1882 (GA), 21 es 1948, Brade 19340 (RB). MiNAS

GERAIS: Serra rral, Nova Lima, 1,300 m, 6 Aug.
1945, Williams & Assis 8029 (BM, BR, C, F, G, GH.

MO, N ). RIO DE JANEIRO: Teresopolis,
Serra das B 24 Sep. 1980, MacLeish et al. 670

(C, GA, nid LE, MO, NY, RB, B, W). são
PAULO: Serr: A Ae uy: km m Sao José do
Barreiro idle k of Bocaina, 1,640 m, 30 Sep.

1980, PAS dei et al 679 (B, BM, CEPEC, GA, RB,
S, TEX, W).

1987]

Possible hybrids. x E. capitatus (Sprengel) Mac-
Leish. MINAS GERAIS: Carai, 6 Sep. 1981, Schumacher
1083, 1085, 1089 (GA). x E. incanus (Less.) Less.
MINAS GERAIS: Tiradente, 10 Sep. 1936, Barreto 4792
(F). x E. seidelii MacLeish & Schumacher. MINAS
GERAIS: Furnas towards Piui, 25
macher 1005 (GA), Alpinopolis, 12 Sep. 1977, Seidel

. (RB).

16. Eremanthus uniflorus MacLeish & Schu-
aa Syst. Bot. 9: 93. 1984. TYPE: Brazil.
Goiás: 33 km N of Alto Paráiso towards
P IE 1,370 m, 14 Oct. 1980,
MacLeish, Martinelli, Smith & Stutts 736
(holotype, RB; isotypes, F, G, GA, K, M,
NY, P, S, UB, US)

Shrubby tree, to 2.5 m tall; stems gray-brown
lepidote; branches many. Leaves membrana-
ceous to subcoriaceous, petiolate; petioles 5-20
mm long, lepidote; blades 3.8-8 cm long, 1.2-4
cm wide, elliptic, the bases acute, the apices acute,
the margins entire; adaxially glabrate, abaxially
gray lepidote. Peduncle slender, to 25 cm tall,
terete, gray lepidote. Glomerulescence a com-
pound cyme of 100 or more glomerules, these
3-7 mm tall, 3-10 mm diam., hemispherical.
Heads 3-9 per glomerule, slightly appressed and

adherent basally by pubescence of phyllaries. In-
volucres cylindric, 4-7 mm tall, 1-2 mm diam.;
phyllaries in 5-7 series; outermost phyllaries
trullate, 0.7-2 mm long, m wide, the
apices acute; innermost phyllaries m 3.5-
5.5 mm long, 0.5-0.9 mm wide, the apices acu-
minate; margins entire to subfimbriate; abaxial
surfaces stramineous with purple apex and mid-
vein, glabrate to lepidote-tomentulose. Florets |
per head; corollas purple (to white with age), 5-
7 mm tall, the lobes acuminate. Anthers 2.7—3.8
mm long, the apical appendage acuminate, the
bases acuminate. Achenes 2.4—3.2 mm tall, glan-
dular, 10-ribbed, the apex slightly constricted and
dark, occasionally pilose at base; nectary 0.05-
0.1 mm tall. Pappus 3—5-seriate, of purple to
stramineous, deciduous, strigose bristles; outer-
most series 0.8-1.8 mm long; innermost series
5.5-7 mm long.

Flowering and fruiting occur from August to
October.

Eremanthus uniflorus is restricted to the Cha-
pada dos Veadeiros in central Goiás (Fig. 6).
Although locally abundant, it is found only on
campo rupestre outcrops

A single floret per head. the relatively weak
coherence of heads, and 3-9 heads per glomerule

MACLEISH — EREMANTHUS REVISION

285

distinguish E. uniflorus from other members of
sect. Vanillosmopsis. This taxon is closely related
to E. capitatus, which is primarily restricted to
the northeast arm of the Central Plateau and
differs in that E. capitatus has 3-4 florets per
head and 2-5 heads per glomerule.

Additional specimens examined. BRAZIL. GOIAS:
Brasilia Richtung Alto Paráiso de Goiás, 50 km nach
Joà

m of Alto Paráiso, 1,220 m, 24 Jan
1980, King & Almeda 8291 (US).

Possible hybrids.
macher. Goás: Alto Paraiso, 28
macher 1038, 1039, 1041 (GA)

x E. argenteus MacLeish & Schu-
Aug. 1981, Schu-

17. Eremanthus capitatus (Sprengel) MacLeish,
comb. nov. Conyza capitata Sprengel, Syst.
Veg. 3: 507.1826. Vernonia capitata (Spren-
gel) Less., Linnaea 4: 270. 1829. Albertinia
capitata (Sprengel) DC., Prodr. 5: 82. 1836.
Vanillosmopsis capitata (Sprengel) Schultz-
Bip., Jahresber. Pollichia 18-19: 167. 1861.
LECTOTYPE: here designated; Brazil. Bahia:
inter Victoria et Bahia (now Salvador), Sel-
low s.n. (K; isolectotype, GH).

Polypappus discolor DC., Prodr. 7: 281. 1838. Vanil-
losm opsis rere (DC. ) Baker in C. Martius, Fl.
Bras. 7.1873. TvPE: Brazil. Bahia: Jacobina,
Jan. 1843, Bianche el (holotype, G; vile
BM, 4 in F (one photo of C), 2 in

GH,
k albertinioides ppa Bip., Jahresber.
Pollichia 18— 168. TYPE: Brazil: no other
da s ented Ere B, not found).

Shrub or tree, to 6 m tall, to 8 cm diam.; stems
gray-brown lepidote; branches many. Leaves
membranaceous to subcoriaceous, sessile to pet-
iolate; petioles 1-12 mm long, lepidote; blades
4.6-10.5 cm long, 2.4—4.2 cm wide, elliptic to
obovate, the bases acute, the apices acute to acu-
minate, the margins entire; adaxially glabrate to
lepidote, abaxially gray lepidote. Peduncle slen-
der, to 30 cm tall, terete, gray lepidote. Glomer-
ulescence a compound cyme of 100 or more
glomerules. Glomerules 4-10 mm tall, 3-12 mm
diam., hemispherical. Heads 2-5 per glomerule,
slightly appressed and coherent basally by pu-
bescence of phyllaries. Involucres cylindric to
obconic with age, 5-8 mm tall, 2-4 mm diam.;
phyllaries in 5-6 series; outermost phyllaries
widely deltate, 0.5-1.1 mm long. 0 2 mm
wide, the apices acute; innermost phyllaries lan-
ceolate, 3.4-7.6 mm long, 0.6-1.4 mm wide, the
apices acuminate; margins entire to subfim-

286

briate; abaxial surf: t to green, gla-
brate to lepidote-tomentulose. Florets 3(-4) per
head; corollas purple with white tube and be-
coming white with age, 4.2-7.2 mm tall, the lobes
acuminate. Anthers 2.2-2.8 mm long, the apical
appendage acuminate, the bases acute. Achenes
2-2.5 mm tall, glandular, 10-ribbed, the apex
slightly constricted and dark; nectary 0.3-0.4 mm
tall. Pappus 3—4-seriate, of purple, white or stra-
mineous, often curling, promptly deciduous, stri-
gose bristles; outermost series 2.5-3.5 mm long;
innermost series 4—6.5 mm long.

Flowering and fruiting occur from July to No-
vember.

Eremanthus capitatus is distributed through-
out the northeastern arm of the Central Plateau
(Figs. 6, 7) at elevations of 100 to 1,000 meters.
It occurs in large colonies in cerrado or on the
border of secondary forest and is known as “‘can-
deia,” or candle, in Bahia.

Eremanthus capitatus is closely related to E.
uniflorus, E. arboreus, and E. erythropappus. It
is distinguished by its 3—4 florets per head and
2-5 heads per glomerule. Eremanthus uniflorus
is likely derived from an isolated population of
E. capitatus in the northwestern arm of the Cen-
tral Plateau whose number of florets per head
has decreased from three to one. In contrast, E.
arboreus represents a population found in the
extreme northeastern part of the plateau whose
number of heads per glomerule has increased
from 2-5 to 6-9. Eremanthus capitatus and E.
erythropappus have the same number of florets
per head, but E. capitatus lacks the fusion of
receptacle and phyllary tissues observed in E.
erythropappus.

Flowering or immature specimens of this tax-
on are frequently misidentified as E. pohlii. When
young, each head is noticeably cylindric like those
of E. pohlii; however, as fruits mature, the in-
nermost phyllaries drop off and the outermost
spread out so that heads appear to be cylindric-
turbinate.

Sprengel's types are generally considered to
have been deposited at B, BM, BR, or P. Un-
fortunately, a holotype was not located in any of
these herbaria. However, a specimen was located
at K bearing the citation “ex herb. B" and the
correct citation in Sprengel's handwriting. There-
fore, the Kew specimen has been chosen as lec-
totype.

Additional specimens examined. BRAZIL. BAHIA: 33
km from BR 101 on road S to Canavieiras, 23 Oct.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

1980, MacLeish & Soares ind 758 (BR, C, CEPEC,
F, GA, GH, MO, NY, RB, UC, US). GorAs: 5-10 km
N of Veadeiros, Valley of the Rio Paraná, 19 Jul. 1964,
Prance & Silva 58251 (F, NY, RB, S, UB). MINAS GERAIS:
BR 116 Téofilo Otóni towards Bahia, 2 km from Padre
Paraiso, 8 Sep. 1981, 600 m, Schumacher 1082 (GA,
MB).

Possible hybrids.
MacLeish. MINAS GERAIS: Carai, 6 Sep.
macher 1083, 1085, 1089 (GA).

x E. erythropappus (DC.)
1981, Schu-

18. pisi arboreus (Gardner) MacLeish,
comb. Albertinia arborea Gardner,
Idea. Bot 5: 236. 1846. Vanillosmopsis
arborea (Gardner) Baker in C. Martius, Fl.
Bras. 6(2): 16. 1873. TYPE: Brazil. Ceara:
Serra de Araripe, Nov. 1838, Gardner 1713
(holotype, BM; isotypes, BR, 4 in F (includ-
ing fragment from P and photo of B), 2 in

G, 3 in GH (one is photo of B), 2 in NY).

Tree, to 6 m tall; stems gray-brown lepidote;
branches many. Leaves membranaceous to sub-
coriaceous, petiolate; petioles 5-10 mm long,
lepidote; blades 5—6.5 cm long, 1.6-2 cm wide,
narrowly elliptic to obovate, the bases attenuate,
the apices acute to acuminate, the margins entire;
adaxially glabrate to lepidote, abaxially gray lep-

tall, 8-15 mm diam., hemispherical. Heads 6-9
per glomerule, slightly appressed and coherent
basally by pubescence of phyllaries. Involucres
obconic, 5-7.5 mm tall, 2-3.5 mm diam.; phyl-
laries in 5-6 series; outermost phyllaries widely
deltate, 0.6-1 mm long, 0.5-1.2 mm wide, the
apices acute; innermost phyllaries lanceolate, 4—
6 mm long, 0.5-1.2 mm wide, the apices acute
to acuminate; margins entire to subfimbriate;
abaxial surfaces stramineous to green, glabrate
to lepidote-tomentulose. Florets 3(-4) per head;
corollas white or purple, 4-7 mm tall, the lobes
acuminate. Anthers 2-3 mm long, the apical ap-
pendage acute, the bases acute. Achenes 1.5-2.5
mm tall, sparsely glandular, 10-ribbed, the apex
slightly constricted and dark; nectary 0.25-0.35
mm tall. Pappus 3-5-seriate, of stramineous to
off-white (occasionally purple basally), often
curling, promptly deciduous, strigose bristles;
outermost series 2.5-3 mm long; innermost se-
ries 4—6 mm long.

debui. and fruiting occur from August to
September

1987]

Eremanthus arboreus is restricted to the north-
eastern slope of the Chapada do Araripe (Fig. 6),
near Crato, which runs along the border of Ceara
and Pernambuco at elevations of 700 meters. It
occurs in colonies on wooded slopes near the top
of the plateau which are less dry than the sur-
rounding caatinga.

Eremanthus arboreus is closely related to E.
capitatus and is distinguished by its 6-9 heads
per glomerule. This species probably represents
an extreme northeastern population of E. capi-
tatus whose number of heads per glomerule has
increased from 2-5 to 6-9. Until recently, this
species was known only from the type collection.
However, Schumacher (1982) was able to collect
it again from what was probably Gardner’s orig-
inal site during several expeditions designed to
rediscover members of Eremanthus that were
known only from type specimens.

Additional specimens examined. BRAZIL. CEARA:
Chapada do Araripe, hillside on oe g of Crato, 25
Aug. 1980, Schumacher s.n. (G RB), Crato to-
wards Nova Olinda, at slope near Ps ‘of plateau, 700
m, 31 Aug. 1981, Schumacher 1049 (B, GA, K, MB
RB).

EXCLUDED TAXA

1. Eremanthus Less. sect. Chresta (Vell. Conc.
ex DC.) Baker in C. Martius, Fl. Bras. 6(2):
166.1873. Chresta Vell. Conc. ex DC., Prodr.
5:85. 1836. TYPE: (Robinson, 1980): Chresta
sphaerocephala DC. = Chresta Vell. Conc.

2. Eremanthus Less. sect. Pycnocephalum (Less.)
Baker in C. Martius, Fl. Bras. 6(2): 168. 1873.
Vernonia Schreber sect. Pycnocephalum

ss., Linnaea 6: 629. 1831. TYPE: (Mac-
Leish, 1985a): Pycnocephalum plantagini-
Kori (Less.) DC. = Pycnocephalum (Less.)

3. boc Less. sect. Sphaerophora
(Schultz-Bip.) Baker in C. Martius, Fl. Bras.
6(2): 165. 1873. Sphaerophora Schultz-Bip.,
Jahresber. Pollichia 20-21: 402. 1863, nom.
illeg. (non Sphaerophora Blume, 1850, Ru-
biaceae). Paralychnophora MacLeish, Tax-
on 33: 106. 1984. TYPE: Paralychnophora
bicolor (DC.) MacLeish = Paralychnophora
MacLeish.

4. Eremanthus Less. sect. Stachyanthus (DC.)
Baker in C. Martius, Fl. Bras. 6(2): 167. 1873.
Stachyanthus DC., Prodr. 5: 84. 1836, nom.
rej. (vs. Stachyanthus Engler, 1897, Icaci-
naceae). Argyrovernonia MacLeish, Taxon

MACLEISH — EREMANTHUS REVISION

5.

6.

N

oo

9.

12.

287

33: 106-107. 1984. TYPE: Argyrovernonia
martii (DC.) MacLeish — Argyrovernonia
MacLeish.

Eremanthus Less. subg. Pseuderemanthus
Schultz-Bip. “B. Jodopappus" Schultz-Bip.,
Jahresber. Pollichia 20-21: 396. 1863. TYPE:
Vernonia crotonoides (DC.) Schultz-Bip. —
Vernonia Schreber subg. Vernonia sect. Ver-

n

Eremanthus angustifolius (Gardner) Baker in
C. Martius, Fl. Bras. 6(2): 170. 1873. Chres-
ta angustifolia Gardner, London J. Bot. 1:
240, tab. 8. 1842. TYPE: Brazil. Goiás: shady
places at Arrayas, Gardner 3802 (holotype,
BM); isotypes, 3 in F (includes fragment from
P and photo of B), 2 in G, GH (photo of B),
NY, S, W) = Pycnocephalum angustifolium
(Gardner) MacLeish (19852).

. Eremanthus bicolor (DC.) Baker in C. Mar-

tius, Fl. Bras. 6(2): 165. 1873. Albertinia bi-
color DC., Prodr. 5: 81. 1836. TYPE: Brazil.
Minas Gerais: altis, Martius s.n. (holotype,
M; isotype, M) = Paralychnophora bicolor
(DC.) MacLeish (1984a).

. Eremanthus crotonoides (DC.) Schultz-Bip.,

Jahresber. Pollichia 20-21: 396. 1863. Al-
bertinia crotonoides DC., Prodr. 5: 81. 1836.
TYPE: Brazil. Minas Gerais: montium sepi-
bus, Martius s.n. (holotype, M) = Vernonia
crotonoides (DC.) Schultz-Bip.

Eremanthus curumbensus Philipson, Kew
Bull. 1938: 298. 1938. TYPE: Brazil. Goias:
valley of the Rio Curumbo in the plains be-
tween the mountains, Glaziou 21645 (ho-
lotype, K; isotypes, BR, C, 4 in G, NY) =
Glaziovianthus curumbensis (Philipson)
MacLeish (1985b).

. Eremanthus descampsii Klatt ex De Wild.

& T. Durand, Ann. Mus. Congo, Ser. 1, Bot.
1: 99. 1898, non Vernonia descampsii De
Wild., Bull. Jard. Bot. État. 5: 97. 1915. TYPE:
Rep. of the Congo, Katanga, Vallee de la
Liula, 1891, Descamps s.n. (holotype, BR
not seen; fragment and drawing of BR, GH)
— Vernonia klattii MacLeish (1984b).

. Eremanthus eriopus (Schultz-Bip.) Baker in

C. Martius, Fl. Bras. 6(2): 169. 1873. Pres-
telia eriopus Schultz-Bip., Naturf. Ges. Em-
den 1864: 73. 1864. rvPE: Brazil. Minas
Gerais: in glareosis S. da Lapa, Riedel 1127
(holotype, LE; isotypes, 2 in F (fragment and
photo of P), P, TEX (photo of P)) = Prestelia
eriopus Schultz-Bip.

Eremanthus exsuccus (DC.) Baker in C.

—
Ww

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—
WN

—
a

—
-

—

[re]

—
o

N
e

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Martius, Fl. Bras. 6(2): 166. 1873. Chresta
exsucca DC., Prodr. 5: 85. 1836. TYPE: Bra-
zil. Minas Gerais: montosis Minarum, Mar-
tius s.n. (holotype, M; photos of M, F, TEX)
— Chresta exsucca D

. Eremanthus harmsianus Taub., Bot. Jahrb.

Syst. 21: 453. 1896. TYPE: Brazil. Goiás: Ser-
ra dos Pyreneos, Ule 2984 (holotype, HBG,
not seen; isotype, P) = Glaziovianthus spe-
ciosus (Gardner) MacLeish (1985b).
Eremanthus imbricatus G. Barroso, Rodri-
guésia 23-24: 6. 1962. TYPE: Brazil. Distrito
Federal: Brasilándia, Macedo 4 (holotype,
RB; isotype RB) = Chresta exsucca DC

. Eremanthus jelskii Hieron., Bot. Jahrb. Syst.

36: 462. 1905, non Vernonia jelskii Hieron.,
Bot. Jahrb. Syst. 36: 459. 1905. TYPE: Peru,
crescit prope Shanyn, Je/skii 776 (lectotype,
US (MacLeish, 1984b); isolectotypes, pho-
tos of B, F, GH, NY, TEX) - Vernonia
shanynensis MacLeish (1984b).

. Eremanthus labordeii Glaz., Mem. Soc. Bot.

France 3: 380. 1909. TYPE: Brazil. Goiás:
entre Paranaua et Rajadinho, 31 Jun. 1895,
Glaziou 21675 (lectotype, P (MacLeish,
1985a); isolectotypes, BR, C, 2 in F, 2 in G,
GH) = Chresta exsucca DC.

Eremanthus leucodendron Mattf., Bot. Gart.
Notizbl. 9: 378. 1925. TYPE: Brazil. Bahia:
Rio de Contas, Serra das Almas, Carrasco,
Luetzelburg 242 (holotype, M; isotypes, F
(photo of B), 2 in GH (includes photo of B),
M, TEX (photo of B)) — Vernonia leuco-
dendron (Mattf.) MacLeish (1984b)

. Eremanthus martii (DC.) Baker in C. Mar-

tius, Fl. Bras. 6(2): 167. 1873. Stachyanthus
martii DC., Prodr. 5: 84. 1836. TYPE: Brazil.
Bahia: siccis sylvis aestu aphyllis, ad Ju-
azeiro, Martius s.n. (holotype, M; isotypes,
3 in M) - Argyrovernonia martii (DC.)
MacLeish (19842).

. Eremanthus mollis Schultz-Bip., Jahresber.

Pollichia 18-19: 166. 1861, non Vernonia
mollis Kunth, Nov. Gen. & Sp. 4: 36. 1820.
TYPE: Brazil. Goiás: Montes Claros et Ponte
Alto, ante Bomfim, Pohl 171 (lectotype, W
(MacLeish, 1984b); isolectotype, 2 in F
(fragment and photo of B), GH (photo of B),
2 in NY, TEX (photo of B)) = Vernonia
pannosus (Baker in C. Martius) MacLeish
(1984b).

Eremanthus pabstii G. Barroso, Sellowia 16:
173. 1964. TYPE: Brazil. Goiás: Cristalina,
Heringer 9229/1442 (holotype, HB; iso-

N

22,

N
[9v]

2

A

N
ws

2

nN

2

~

2

co

t3
e

[VoL. 74

types, RB, UB) = Vernonia pabstii (G. Bar-
roso) MacLeish (1984b)

. Eremanthus pannosus Baker in C. Martius,

Fl. Bras. 6(2): 164. 1873. TYPE: Brazil. Goiás:
Curralinho, Manso 1 (holotype, BR) = Ver-
nonia pannosus (Baker in C. Martius)
MacLeish (1984b).

Eremanthus pinnatifidus Philipson, Kew
Bull. 1938: 299, 1938. TYPE: Brazil. Rio de
Janeiro: env. de Rio de Janeiro, Glaziou
14033 (lectotype, C (MacLeish, 1985a)) =
Pycnocephalum pinnatifidum (Philipson)
MacLeish (19852)

. Eremanthus plantaginifolius (Less.) Baker

in C. Martius, Fl. Bras. 6(2): 168. 1873. Ver-
nonia plantaginifolius Less., Linnaea 4: 251.
1829. TYPE: Brazil: no other data, Se//ow s.n.
(lectotype, P (MacLeish, 1985a)) = Pycno-
cephalum plantaginifolium (Less.) DC.

. Eremanthus purpurascens Glaz. ex Oliver,

Hooker's Icon. Pl. 4(3): plate 2282. 1894.
TYPE: Brazil. Minas Gerais: Serra do Cipó,
Glaziou 19464 (holotype, P; isotypes, BR,
C) = Prestelia eriopus Schultz-Bip.

. Eremanthus pycnocephalus (DC.) Baker in

C. Martius, Fl. Bras. 6(2): 166. 1873. Chres-
ta pycnocephala DC., Prodr. 5: 85. 1836.
TYPE: Brazil. Minas Gerais: campis deserti
inter Min. Nov. et f. S. Francisci, Martius
s.n. (holotype, M) = Chresta pycnocephala
DC

; Eremanihins reflexo-auriculatus G. Barroso,

6. 1962. TYPE: Brazil.

— Paralychnoph-

ora reflexoauriculata (G. Barroso) MacLeish
)

(1984a

. Eremanthus rivularis Taubert, Bot. Jahrb.

Syst. 21: 453. 1896. TYPE: Brazil. Goiás: re-
giao de Maranhào superior, Ule 26 (2962)
(holotype, HBG, not seen; isotypes, 2 in F
(fragment and photo of P), GH (photo of P),
P) — Pycnocephalum angustifolium (Gard-
ner) MacLeish (19852)

. Eremanthus scapigerus (Less.) Baker in C.

Martius, Fl. Bras. 6(2): 168. 1873. Vernonia
scapigera Less., Linnaea 4: 250. 1829. TvPE:
Brazil: no other data, Sellow s.n. (lectotype,
B (MacLeish, 19852); isolectotypes, K, P) —
Chresta scapigera (Less.) Gardner.

Eremanthus schwackei Glaz., Bull. Soc. Bot.
France 3: 380. 1909. TYPE: Brazil. Minas
Gerias: Biribiry, Glaziou 19562 (lectotype,
here designated, P; isolectotypes, BR, C, G)

1987]

o2
©

wə
—

Ww
N

[9v]
UJ

o2
A

W
CA

ww
nN

o2
N

MacLEISH— EREM.

= Paralychnophora schwackei (Glaz.)

MacLeish (1984a).

. Eremanthus speciosus ee Baker in C.

Martius, Fl. Bras. 6(2): 169. 1873. Pide.
speciosa Gardner, aa J. Bot 240.
1842. TYPE: Brazil. Goiás: dry campos near
Villa de Arrayas, Gardner 3801 (holotype,
BM; isotype, BM) = Glaziovianthus specio-
sus (Gardner) MacLeish (1985b).

. Eremanthus sphaerocephalus (DC.) Baker in

C. Martius, Fl. Bras. 6(2): 167. 1873. Chres-
ta spherocephala DC., Prodr. 5: 85. 1836.
TYPE: Brazil. Minas Gerais: Tejuco (now
Diamantina), Vauthier 294 (lectotype, G-D
(as IDC microfiche, MacLeish, 19852);
isolectotypes, G, GH) = Chresta sphaero-
cephala D

QE nasus sphaerocephalus (DC.) Baker in

C. Martius var. intermedia (Gardner) Baker
in C. Martius, Fl. Bras. 6(2): 167. 1873.
Chresta intermedia Gardner London J. Bot.
5: . 1 . TYPE: Brazil. Minas Gerais:
near Formigas (now Montes Claros), Gard-
ner 4818 (holotype, BM) = Chresta sphaero-
cephala DC

. Eremanthus veadeiroensis H. Robinson,

Phytologia 45: 94. 1980. TYPE: Brazil. Goiás:
Chapada dos Veadeiros, ca. 20 km N of Alto
Paraiso, /rwin et al. 32752 (holotype, UB;
isotype, US) = Vernonia oo (H.
Robinson) MacLeish (1984

. Eremanthus verbascifolius pa Martius ex

DC.) Schultz-Bip., Jahresber. Pollichi 20-
21: 397. 1863. TYPE: Brazil. Minas Gerais:
ferruginosis Serra do Ant. Pereira, Martius
s.n. (holotype, M) = Vernonia crotonoides
(DC.) Schultz-Bip.

. Vanillosmopsis bicolor (DC.) Schultz-Bip.,

Jahresber. Pollichia 18-19: 168. 1861. Al-
bertinia bicolor DC., Prodr. 5: 81. 1836. TYPE:
Brazil. Minas Gerais: altis, Martius s.n. (ho-
otype, M; isotype, M) — Paralychnophora
bicolor (DC.) ipn (1984a).
Vanillosmopsis lanceolata (Vell. Conc.)
untze, Re en. Pl. 3(2): 183. ^ dd
chius ) PM Vell. Conc., Arq,
Nac. Rio de Janeiro 8: 350. 1881. TYPE: isnt
zil: mediterraneis transalpinus prope pagum
Cunha, Velloso s.n. (lectotype, as figure, Fl.
Flum. Ic. 8: tab. 151. 1831 (MacLeish,
1985a)) = Identity doubtful.

. Vanillosmopsis syncephala Schultz-Bip.,

Jahresber. Pollichia 18-19: 168. 1861, non
Vernonia syncephala Schultz-Bip. ex Baker

REVISION 289

in C. Martius, Fl. Bras. 6(2): 64. 1873. TYPE:
Brazil: no other data, Sellow 948 (holotype,
B (destroyed); photos of B, F, GH) =
nonia crispa (Mattf.) MacLeish (1984b).

354. 19
in via a Sandia ad Chunchusmayo, Weber-
bauer 1324 (lectotype, NY, photo of B
(MacLeish, 1984b); isolectotypes, photos of
B, GH, TEX) = Vernonia ramospatana
MacLeish (1984b).

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ANNALS OF THE MISSOURI BOTANICAL GARDEN

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pragmatic chemists? Symb. Bot. Upsal. 22: 200-
209.

1906. Vitae itineraque collectorum botani-
In C. Martius (editor), Flora Brasiliensis 1(1):
1-154.
URBAN, Z. 1973. The autoecious species of Puccinia
on Vernonieae in North America. Acta Univ. Car-
ol. Biol. 1971: 1-84.
VELLOso, J. M. C. 1829. Pp. 1-352 in Florae Flu-
minensis. Rio de Janeir
831. Florae fluminensis icones 8: tab. 150-
151.

URBAN, I.

. 1881. Florae fluminensis. Arq. Mus. Nac.

Rio de Janeiro p 349-350.

is ipa W., . L. MACHADO, J. A. RAB
MURARI & W. Pus 1977. Eregoyazin ind er-
o two new guaiangolides from Ereman-
thus goyazensis. J. p Chem. 42: dia

ZANONI, T. A. o Kuntze, botanist. I. Bi-
Ripe ‘bibliography, dd travels. midi 32:
551-57

BIOSYSTEMATICS OF TETRAPLOID EUCHARIS
(AMARYLLIDACEAE)!

ALAN W. MEEROW?

ABSTRACT

Eucha

rainforest understory from Guatemala to Bolivia. The tw

ris is a genus of 16 species of petiolate-leaved, p Amaryllidaceae da lai to
. bonpla

northernmost species, ii and

E. bouchei, are the only two tetraploid (2n = 92) species so far known in ack genus. E ucharis bonplandii

entral

series for both ‘tetraploid species. On the basis of 17 floral characters, the three varieties of E

e tetraploid representatives of Eucharis exhibit

a a wide degree of “karyotypic heteromorphism. , E ucharis bouchei var. dressleri i is an unstable tetraploid.

ami

no ff f polyploidy

in some individuals. cm phenotypes of iach bouchei var. bouchei are quite variable and
tl
F

var iety I

Lines

and a bouchei may b

°
°

read tet traploid

Asas desp
x. The entry of Eucharis Into Central America was probably a “eka recent ons. It is

he Colomb

of heterozygosity may have been important in facilitating the migration of Eucharis across the Isthmus
of Panama.

The genus Eucharis Planchon & Linden
di pa aged 2 family" Pancratioidinae
sensu Traub (1957, 1963)] consists of 16 species
ofi rare, petiolate- ms bulbous geophytes in-

MAaAVIL

Gliatemala to Bolivia (Meerow, 1986). Most of

of pr imar y rainfor est from

de
Eucharis bouchei Woodson & Allen is a highly
polymorphic tetraploid (2n = 92) complex of
Central America (Fig. 1). The species is concen-
trated in Panama (Fig. 2), but has also been re-
corded from Costa Pies and Guatemala. Eucha-
ris bouchei is th t species of Eucharis
and the only one found north of the Darién Gap.
It is also the most variable species in the genus,
in characteristics that elsewhere justify specific

delimitation. Patterns of variation in floral size
and tube and limb habit form a complete mosaic
throughout the range of E. pen showing little
or no geographic consistenc

In my recent mono mid of Eucharis (Mee-
row, 1986), three varieties are recognized chiefly
on the basis of staminal cup morphology (Fig.
3): E. bouchei var. bouchei, var. darienensis Mee-
row, and var. dressleri Meerow.

Variety bouchei, most common around El Valle
de Antón in Coclé Province (Fig. 2), is recognized
by its largely edentate staminal cup in which the
trapezoidal free filament is not markedly con-
stricted distally into a narrow subulate portion

Figs. 1C, 3A, B). It is the most variable of the
three varieties, both in flower size and staminal
cup morphology. The staminal cup of variety

! I thank Robert Dressler, Mark Elliot, Mark Whitten, ce on Botanical Garden for providing living

material of tetraploid Eucharis species. Bijan D

ehgan p

ovided the supportive environment w is work

was accomplished. Charles Guy gave freely of his time xm E materials for the electrophoretic analyses.
Pete

applications. Brent

Mishle

r Goldblatt,

George K. Rogers, and two anonymous reviewers provided Teila B of an earlier version of this paper.
S

Part of this work was supported by NSF
America/World Wildlife Fund Fellowship in pis al B

cu
Gainesville, Florida 32611, U.S

ANN. Missouni Bor. GARD. 74: 291—309. 1987.

Doctoral pecu ion
P

R-8401208 and a Garden Club of

292

— A. Flowers. —

_Fic GURE l. Eucharis bouchei.

ouchei.—ii. Meerow 1125, FLAS. —iii. |. Mee-
FLAS.—B. Tepals, variety bouchei. i, ii.
Meerow 1125. —i. Outer tepal.— ii. Inner tepal. iii, iv.
Meerow 1157.—iii. Outer tepal.—iv. Inner tepal.—C.
Staminal cups, variety bouchei. —i. Meerow 1125.—ii.
Meerow 1157. —D. Ovaries, variety bouchei, longitu-
dinal section. —i. Meerow 1125.—ii. Meerow 1157.

darienensis, found in Panamá and Darién prov-
inces, is Dun pene or lobed (Fig. 3D).
The tally into a narrow
(<2 mm wide) subulate portion. These two va-
rieties occur in close proximity in one location
near Cerro Campana in Panama Province. The
rare var. dresslerii (Fig. 1Ai), with its acutely
toothed staminal cup (Fig. 3C) and non-trigo-
nous Ovary, occurs close to populations of var.
bouchei near El Valle

Northwesternmost populations in Panama
representing var. bouchei have the most derived
androecial morphology (Fig. 3A, B) relative to

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

more southeastern populations (var. darienensis,
Fig. 3D). The latter have staminal cups similar
to the generalized morphology characteristic of
Andean and Amazonian species of subg. Eucha-
ris. This may indicate that general movement of
E. bouchei in Central America has been away
from the Colombian border.

The only other naturally occurring tetraploid
Eucharis species known is the rare Colombian
E. bonplandii (Kunth) Traub. It is separated from
E. bouchei by its slightly glaucous leaves (all oth-
er Eucharis species have nonglaucous foliage),
shorter petioles, and longer pedicels. Eucharis
bonplandii is the northernmost species of Eu-
charis subg. Eucharis in South America.

Results of phenetic, karyotype, and prelimi-
nary electrophoretic analyses of E. bouchei and
E. bonplandii are presented in this paper. These
data offer insight into the evolutionary history
of Central American Eucharis and the origins of
tetraploidy in the genus, and provide a basis for
understanding the enormous degree of pheno-
typic variation present within E. bouchei.

The use of electrophoretic analyses of isozyme
variation in plant systematics has been extensive
in recent years. The subject has been reviewed
by Gottlieb (1971, 1977, 1981a, 1981b, 1982,
1984) and Crawford (1983, 1985). Unlike many
morphological characters, which may be influ-
enced by a great deal of environmental or de-
velopmental plasticity, the electrophoretic phe-

lectrophoretic studies of tropical plants are
few (Hamrick & Loveless, 1986; Heywood &
Fleming, 1986; Sytsma & Schaal, 1985). Neither
have plants of limited or rare distribution been
widely investigated (Babbel & Selander, 1974).
Members of Eucharis are tropical monocots
exclusive to rainforest understory. They are rare
and widely dispersed in the wild. Studies of iso-
zyme variation of any plant group fitting any one
of these characteristics are very limited. Isozyme
analyses of polyploid taxa are also not abundant
(Crawford, 1985; Soltis & Rieseberg, 1986). Thus,
an attempt to explore isozyme variation in nat-
ural polyploid Eucharis seemed a worthy avenue
of investigation.

MATERIALS AND METHODS
PHENETIC ANALYSES

Principal component and hierarchical cluster
analyses of 20 herbarium specimens of Eucharis

MEEROW — EUCHARIS 293

1987]

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294 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

3. Staminal cup variation in Eucharis
Mieres y B. Variety bouchei. — A. Lewis et al. 2617
(MO). — B. Allen 120 (US).—C. Variety dressleri (Mee-
row 1107, FLAS).—D. Variety darienensis (Gentry &
Mori 13945, MO).

e — 1) were conducted with CLUS-
. 2.1 (University of St. Andrews,
anne da on eai North Florida Regional Data
Center (NERDC) system of the University of
Florida. Three-dimensional scattergrams were
constructed from P factor scores utilizing
PC T, a program written by Bart Schutz-
man at the University of Florida. The small
number of OTUS (operational taxonomic units)
underscores the relative rarity with which Eu-
charis is encountered in the field. These 20 spec-
imens represent the only specimens from which
the full character set could be recorded. A num-
ber of additional specimens examined were col-
lected in fruit and therefore were useless for these
analyses.
Twenty-seven characters were used initially.
The results suggested that some of these char-

acters (e.g., all foliage characters, scape height,
ovary length) were unreliable due to environ-
mental plasticity, developmental variation, or
specimen preparation. Although living material
provides additional characters of potential utility
(e.g., leaf surface texture, pigmentation pattern
of the staminal cup), the inability to consistently
determine these characters in dried specimens
precluded their inclusion. Where any two char-
acters were highly correlated (more than 80%
correlation), which can result in data redundancy
(Sneath & Sokal, 1973), one of the two was re-
moved from the data matrix. In the final anal-
yses, 17 floral characters (Tables 2, 3) were se-
lected as the basic data set, of which 14 were
continuous, quantitative Characters. The re-
maining tł e characters were treated
by assigning a über cd value for each Sartor
state. Since CLUSTAN would treat these values
as continuous, the character states were num-
bered in a progressive transformation series, such
that any two successive numbers would reflect
putative character state relationship. These
Hansiormaton series were constructed by study
of d trends in the genus
E ucharis and by comparative study with closely
related genera of Amaryllidaceae.

Raw data were standardized using the “z-score”
method (Sneath & Sokal, 1973) by which initial
values for each character were replaced by stan-

TABLE 1. Operational taxonomic units sa for multivariate analysis of the Eucharis bouchei complex.

CR = Costa Rica, G = Guatemala, P = Pan

No Collection and Herbarium Variety Origin

l Alston 8727 (BM) dressleri P, Coclé

2 Meerow 1107 (FLAS) dressleri P, Coclé

3 Wendland 207 (GOET) darienensis G

4 Sullivan 718 (M darienensis P, Darién

5 Folsom 4402 (MO) darienensis P, Darién

6 Folsom et al. 6582 (MO) ouchei P, Panamá

7 Allen 5347 (US) bouchei CR

8 Kirkbride & Hayden 305 (MO chei P, Panamá

9 Witherspoon & Witherspoon 8372 (MO) darienensis P, Panamá
10 Duke & Elias 3661 (GH) darienensis P, Darién
11 Gentry & s 13945 (MO) darienensis P, Darién
12 Stern et al. 49 H) darienensis P, Darién
13 Skutch 1585 n ch
14 Seibert 466 (MO) bouchei P, Coclé
15 Lewis 2617 (MO) bouchei P, Coclé
16 Witherspoon & Witherspoon 8736 bouchei P, Coclé
17 Allen 1228 (GH) bouchei P, Coclé
18 Mori & Kallunki 2014 (AAU) bouchei P, Colón
19 Mori et al. 6586 (AAU) bouchei P, Colón
20 Meerow 1158 (FLAS) bouchei P, Colón

1987]

dard deviations from the mean value for that
character. A distance matrix was then calculated
using squared euclidean distance (Cormack,
1971). In addition to PCA, cluster analysis using
average linkage (unweighted pair group method
(UPGMA) of Sneath & Sokal, 1973) was also
applied to the 20 OTUs as a further test of phe-
netic relationship.

CHROMOSOME CYTOLOGY

Root tips were collected from living collec-
tions, pretreated for 2-3 hours at room temper-
ature in 10 ppm solution of o-isopropyl-N-phen-
ylcarbamate (Storey & Mann, 1967), rinsed in
distilled water, fixed in 3: 1 mixture of 95% EtOH
and chloroform at 18°C for 24 hours, then stored
after fixation in 70% EtOH at 18°C. Root tips
were hydrolyzed in 1 N HCI at 50°C for 2-3
minutes, squashed, and stained with iron aceto-
tuine, Only Min iin slides - were made.
Metap d on
a Nikon LLabenhot EA with AFX-
II camera attachment; haploid idiograms were
constructed from photomicrographs.

As absolute chromosome length can vary ap-
preciably from cell to cell due to differential ef-
fects of pretreatment (Tjio & Hagberg, 1951;
Schlarbaum & Tsuchiya, 1984), relative length
based on a value of 100 for the haploid comple-
ment was used to designate size class. Relative
size igiene are based on 30% or ' greater corre-
lations between al d from
Battagla, 1955) and relative leneth (RL) of mi-
totic metaphase preparations of various species
of Eucharis, Eucrosia, Phaedranassa, and other
Amaryllidaceae with 27 — 46, all of which have
similar relative length ranges: large, RL = 7.0
^s gangs length (AL): > 10 um]; moderately ane:

5.0-7.0 (AL: 7-10 um); medium,

Ii. (AL: 5-7 um); small, RL < 3.5 (AL: 2-
5 um). For tetraploid karyotypes, diploid RL val-
ues were halved to assign size class. Chromo-
some morphology, modified from Battaglia
(1955), is defined as follows: metacentric, Arm
Ratio (AR; long arm/short arm) — 1 0-1.10;
near-metacentric, AR = 1.10—1.50; submetacen-
tric, AR = 1.50-3.00; subtelocentric, AR =
> 3.00.

ELECTROPHORETIC ANALYSES
Population selection and sample size. Five
populations were included in these analyses (Fig.
2: Table 4), representing all living collections of

MEEROW — EUCHARIS

295

E2. Characters used for multivariate analysis
of E ned species

Flower number

Limb spread (mm)

Length of free filament (mm)
Width of free filament (mm)

Width of tube at throat (mm)
Length of outer tepal (mm)
Length of inner tepal (mm)

. Width of outer tepal (mm)

. Width of inner tepal (mm)

. Staminal cup length (mm)

LM — E E eel
ROOIOTIO9u9tuPtwmr

3: Irregularly toothed
4: Quadrate
5: Lobe
6: Edentate
. Cleft of staminal cup:

—
un

1: <% length of cup
2: 14—!⁄ length of cup
3: Vi—'^ length of cup
4: > length of cup
. Relative length of teeth:
0: Edentate
« length of filament
2: ^ length of filament
3: = length of filament
4: > length of filament
. No. ovules per locule

_
o

—
-l

E. bouchei and E. bonplandii in cultivation at
the University of Florida. These included three
populations of E. bouchei var. bouchei, one from
El Valle de Antón in Coclé Province of Panama,
and one each from Cerro Brujo and Río Iguanita,
respectively, in Colón Province; and one popu-
ation of var. dressleri, also from El Valle. The
fifth population represented E. bonplandii, a rare
species from Colombia, also tetraploid. Sample
size was one or three for each population (Table

Unfortunately, the small sample size does not
allow statistically significant exploration of ge-
netic variation among these populations. Most
electrophoretic studies in plant systematics have
involved taxa of characteristically large popu-
lation size in nature (see Crawford, 1983; Gott-
lieb, 1981a). Populations of Eucharis, however,
are characteristically small. Many species of Eu-

296 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

TABLE 3. Data matrix for PCA and cluster analyses of the Eucharis bouchei complex.

Character

OTU 1 2 3 4 5 6 7 8 9 10 ll 12 13 14 15 16 17
1 05 43 3.5 18 2.5 330 06.5 262 25.8 05.5 080 094 095 4 2 1 3
2 06 53 42 148 50 410 10.5 320 280 100 135 160 11.45 3 2 2 3
3 06 42 3.5 18 3.7 360 06.5 25.0 230 09.5 127 11.0 120 3 2 1 2
4 04 46 3.1 20 46 25.5 09.5 20.88 200 080 11.1 060 132 6 2 1| 2
5 05 43 2.5 22 43 2363 071 218 210 105 15.5 070 145 6 3 0 2
6 06 55 36 28 352 460 122 314 2300 138 152 106 174 6 3 0 3
7 06 69 35 28 S2 440 12.0 35.0 31.5 155 175 11.0 182 6 3 0 2
8 04 45 28 2.5 49 31.0 10.5 277 255 11.4 145 100 160 5 4 0 2
9 04 38 30 20 43 330 100 200 190 09.5 125 092 140 3 3 1 2
10 05 42 21 17 42 271 06.0 238 220 100 120 090 128 2 2 2 2
11 06 55 20 20 45 43.4 12.0 27.0 250 102 165 140 182 3 2 2 4
12 05 38 18 L8 5.0 30.0 09.8 220 21.0 11.5 150 090 123 2 3 2 2
13 04 41 50 51 51 35.0 10.5 242 21.1 09.8 145 120 154 4 3 0 2
ld 05 45 5.3 4.3 5.3 36.0 08.7 280 260 120 140 110 140 4 3 02
15 07 50 69 5.0 69 43.7 10.8 268 25.0 140 146 167 160 4 3 0 2
16 05 49 45 3.5 45 43.0 10.6 21.0 200 09.0 11.0 080 125 4 3 0 2
17 03 50 65 4.7 65 45.0 120 23.5 220 11.5 15.5 128 145 4 3 02
18 05 34 59 3.7 3.7 34.0 08.3 180 160 09.0 11.5 11.6 11.5 4 3 0 2
19 05 35 37 40 40 33.0 07.0 21.6 19.5 080 100 100 124 4 2 0 2
20 05 40 5.3 38 3.8 40.0 080 260 240 112 137 11.8 097 4 3 0 2

charis frequently occur as single, widely dis- for sample sizes as low as one. Nei stressed, how-
persed clumps (Meerow, 1986). Herbarium spec- ever, that with a limited population sample, a
imens of Eucharis regularly includ tati large number of loci must be analyzed. The num-
indicating the rarity of the plants encountered. ber of enzyme systems assayed in the present
Yet, if most Eucharis are primarily visited by study is not sufficient for this purpose. Instead,
trap-lining insects (sensu Janzen, 1971) flyinglong individuals were scored for the presence or ab-
distances, as may be the case (Meerow, 1986), sence of putatively identical bands (Table 5).
population size from the perspective of potential These scores were used to generate distance coef-
gene exchange may in fact be greater than oth- ficients by the unweighted pair group method
erwise expected from known population densi- (Sneath & Sokal, 1973). Bands showing the same
ties. Few workers have addressed the problem mobility, as determined by their position in the
of how to apply electrophoretic data to rare plants gel, were considered to be identical. The resulting
of characteristically small population size. Nei dendrogram could then be compared with the
(1978) presented modified formulas forunbiased results of phenetic analysis of morphological
genetic identities and distances that could be used variation, as well as with the data derived from

TABLE 4. Eucharis bouchei and E. bonplandii populations examined electrophoretically.

Desig-

Taxon nation N Collection Information Voucher?

Eucharis bouchei var. EBD 3 Panama, Coclé, El Valle de Antón Meerow 1107
dressleri
E. bouchei var. bouchei EBBI 3 Panama, Coclé, El Valle de Antón Meerow 1125
E. bouchei var. bouchei EBB2 l Panama, Colón, Río Guanche, Cerro Meerow 1157
Brujo

E. bouchei var. bouchei EBB3 l Panama, Colón, Rio Iguanita Meerow
E. bonplandii EBN l Colombia, Cundinamarca, vicinity of | Bauml i BUMP

Bogotá

* All vouchers deposited at FLAS unless otherwise indicated.

1987]

MEEROW — EUCHARIS

297

TABLE 5. Presence-absence data matrix for electrophoretic analysis of tetraploid Eucharis. Refer to Table 4
for population designations, Figure 14 for bands. (*) — used only for cladistic analysis. Blank space indicates
le

data unavailable.

Band

+
wr

Population l

"19

—
—
—
N
—
w
—
+
—
wa
—
nN
—
N

10 *18

ooco--ooooc|u
oo--—-—-ooo|o
Or Ke Omer eee

co—-—-o-—-—--—o|o
Ore ee eH ooo] A
e@ — — — — ° ° ° ° | %

tr!

=

ux
— — O — — O — — =
ooooorocoo-

—ooooooococ!|o
—oooooooo
—oooooooo
—oooooooo
—o—-oco----
—o-oo0----
eee — HK OOO
—o-oo-ooo
oO-oc--o---

—o Om
oe = P=

comparative chromosome morphology. A sim-
ilar method for analyzing isozyme data was used
by Ashton et al. (1984) for Shorea (Dipterocar-
paceae), and Chou et al. (1986) for several genera
of bambusoid grasses. The data matrix was ad-
ditionally subjected to cladistic analysis using
PAUP by David Swofford (Illinois Natural His-
tory Survey). The “Wagner method” of simple
parsimony (Farris, 1970; Kluge & Farris, 1969)
was applied in constructing the cladogram, and
E. bonplandii was designated as the outgroup for
polarization of character states.

Of course, without the benefits of formal ge-
netic analysis, there is no guarantee of genetic

omology between any two bands of seemingly
identical mobility. Future studies may allow more
precise analysis of genetic variation within and
among populations of tetraploid E ucharis.

Isozyme ext Crude ex-
tracts for isozyme electrophoresis were prepared
by grinding ten 5 mm diameter leaf discs in 1
ml of extraction buffer [100 mM Tris-HCl, 10
mM DTT, 20% glycerol, and 1 mM PMSF ad-
justed to pH 6.8 (Hames & Rickwood, 1981)].

Extracts were centrifuged twice, for ten minutes

and two minutes, and the supernatant was de-
canted by pipette after cach inde. ise:

BIO-RAD
Protean I exlvacrsilunide gel ee Gel rec-
ipes were adopted from Hames & Rickwood
(1981). Running gels were 0.75 mm thick and
7.5% acrylamide (10 ml 30% acrylamide-bis ac-
rylamide, 10 ml 1.5 Tris-HCl at pH 8.8, 19.85
ml H,O, 100 41 10% ammonium persulfate, and
15 wl TEMED). A 2.5% acrylamide stacking gel
(1 ml 3096 acrylamide-bis acrylamide, 1.92 ml
0.5 M Tris-HCl at pH 6.8, 9 ml H,O, 20 ul

ammonium persulfate, and 7.5 ul TEMED) was
employed. Running buffer was 25 mM Tris-gly-
cine at pH 8.3 (Hames & Rickwood, 1981). A
20 ul sample of the supernatant was loaded into
each stacking gel column. Gels were electro-
phoresed at a constant current of 50 mA until
a blue indicator line (40 ul of bromophenol blue
added to cathodal buffer) migrated off the anodal
end of the gel, generally four to five hours

Five enzyme systems were assayed: aspartate
amino-transferase (AAT), glutathione reductase
(GSSGR), malate dehydrogenase (MDH), phos-
phoglucoisomerase (PGI), and shikimate dehy-
drogenase DH). Staining recipes of Vallejos
(1983) were followed for AAT, PGI, and SKDH.
The staining system for MDH was that of Shaw
& Prasad (1970), and that of Kaplan (1968) was
used for GSSGR.

Resolution of additional enzyme systems (ga-
lactose dehydrogenase, glutamate dehydroge-
nase, hexokinase, and isocitrate dehydrogenase)
using the same buffer system were unsuccessful.
Extracts of Eucharis leaf tissue are characteris-
tically mucilaginous, which may impede electro-
phoretic separation or contribute to degradation
of some enzymes after extraction. Also, cath-
odally migrating isozymes cannot be resolved in
the same vertical acrylamide gel as anodally mi-
grating isozymes.

RESULTS
PHENETIC ANALYSES

Principal Component Analysis (Figs. 4, 5; Ta-
ble 6). Cumulative variance of 71.996 across 20
OTUs was resolved in the first three principal
components (PCs). Characters 5 (stamen width),

298

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Eucharis bouchel

@ var. bouche
Æ ver. dressieri
X var. darienensis

FIGURE 4. PCA scattergram based on variance across 17 floral characters in 20 OTUs representing Eucharis

bouchei

6 (tube length), 9 (inner tepal length), and 14
(toothing) contributed the greatest magnitude of
variance to PC1, especially character 6. PC2 is
largely a measure of outer tepal length (character
8), inner tepal width (11), staminal cup widt
(13), and toothing (14). Characters 1, 7, and 14
also substantially contributed to the variance re-
flected in PC2. Characters 2 (limb spread), 3
(length of free filament), 13 (staminal cup width),
and 15 (toothing) v
of variance in PC3.

e three varieties of E. bouchei do not clearly
resolve into three phenetic groups in Figure 4.
Although var. bouchei shows a tendency to as-
semble along PC2 (21.196 total variance), this
variety is still widely distributed along PCI
(38.7% total variance). One OTU each of var.

VCIC the most

FIGUR
across 17 flor
Eucharis bouchei, with PC2&3 emphasized.

PCA s scattergram based on variance
l characters in 20 OTUs representing

darienensis (no. 1 1) and var. dressleri (no. 2) form
an outlying group, as do 7, and 15 of
var. bouchei. Variety darienensis shows a mea-
sure of phenetic congruence, but intergrades with
var. bouchei.

If the scattergram for the E. bouchei complex
is rotated so that PC2 and PC3 are visually ac-
centuated (Fig. 5), grouping of OTUs becomes
largely a measure of androecial Yonne. In this
scattergram, the three
clearly, particularly var. bouchei. Variety dari-
enensis, however, still intergrades with several
OTUS of var. bouchei, but one of these OTUs
(8) was collected from Cerro Campana in Pan-
amá Province, an area of sympatry between these
two varieties. The third (no. 7) is a Costa Rican
collection.

Cluster analysis (Fig. 6). Two major clusters
are resolved in the UPMGA dendrogram, each
fairly heterogeneous. The first clusters at a dis-
tance coefficient (DC) of 1.356. An outlying OTU
(one of two representing var. dressleri) fuses
with this cluster at DC 1.921. Within this first
cluster, two subgroups emerge at DCs 1.207 and
1.213, respectively. The former is made up en-
tirely of OTUs representing var. darienensis. The
second represents var. bouchei, with the single
exception of OTU 5 (var. darienensis). OTU 5
forms together with OTU 8 (var. bouchei) an
outlying cluster to this second subgroup.

The second major cluster is formed at a DC
of 2.502, near where all clusters finally merge
(DC 2.779). This smaller cluster is more hetero-

d more

1987]

TABLE 6. First three principal components for
multivariate analysis of the Eucharis bouchei complex.

Component Number

Character
Number l 2 3
l 0.151 0.317 0.120
2 —0.265 —0.189 0.340
3 —0.244 0.112 —0.440
4 —0.197 —0.141 —0.147
5 —0.375 0.129 0.059
6 —0.615 0.099 0.133
7 0.032 —0.302 —0.116
8 —0.100 —0.336 — 0.106
9 — 0.404 0.069 — 0.061
10 0.017 0.168 —0.239
ll —0.030 0.386 0.137
12 0.156 —0.062 —0.045
13 0.013 —0.353 —0.524
14 —0.232 —0.381 0.216
15 —0.184 0.322 —0.436
16 — 0.007 0.056 —0.121
17 — 0.046 —0.195 0.020
Percent of
Variance 38.71 21.12 12.03

geneous than the first, but four OTUs of var.
bouchei cluster at a DC of 2.112. As in PCA,
OTUs 2 and 11 (var. dressleri and darienensis
respectively) form a phenetic group.

KARYOTYPE ANALYSIS

Tetraploidy in Eucharis is known so far to
characterize only E. bonplandii and E. bouchei.
Karyotypically, the tetraploid Eucharis species
are strongly heteromorphic (Figs. 7-12; Table 7).
Karyotypes of two geographically isolated and
morphologically distinct populations of E.
bouchei var. bouchei, from Coclé and Colón
provinces of Panama (Figs. 9, 10, 12A, B; Table
7) are quite different. The second largest chro-
mosome pair is submetacentric in the Cerro Bru-
jo (Colón) population of E. bouchei (Figs. 10,
12B) and also in E. bonplandii (Figs. 11, 12C).
Eucharis bouchei var. dressleri is an unstable tet-
raploid (Figs. 7, 8, 12D). Fifty percent of all root
cells from which metaphase counts were ob-
tained had 46 chromosomes.

ELECTROPHORETIC ANALYSES

Of the five enzyme systems assayed, only
SKDH was monomorphic across all populations
of E. bouchei and E. bonplandii. Only polymor-

MEEROW — EUCHARIS

499

2.635 4

2.108 +

1.580 +

DISTANCE

1.317 +

1.053 + -4

0.789 +

Ë |

" 9 na u i u n 16 ou M, NU ) 1 ",
* x | 0 #* x e

FicunE 6. Cluster analysis dendrogram based on
variance across 17 floral characters in 20 OTUs rep-
resenting Eucharis bouchei. Refer to Table 1 for iden-
tification of OTUs.

u

phic loci are discussed below and diagrammed
in Figure 14.

AAT (Figs. 13A, 14). Two well-separated re-
gions of activity were resolved for AAT, one rap-
idly migrating anodally (AAT-1) and the other
(AAT-2) considerably slower; these probably
represent two different loci of this dimeric en-
zyme. Electromorphs at both loci were consid-
erably more complex than in diploid species of
Eucharis (Meerow, 1986). Two putative alleles
are inferred from the phenotypes of AAT-1 in
the E. bouchei complex. Each “allele” of AAT-1
in all Eucharis characteristically resolves as two
very closely spaced bands. This may be the result
of breakdown products forming after extraction
(see Fig. 1 in Shields et al., 1983). Electromorphs
of pollen of diploid (2n = 46) Eucharis (Meerow,
unpubl.) also showed this banding pattern. Were
each component band of the doublet a distinct
allele, pollen would be expected to show only
one of the two (Gottlieb, 1982, 1984). Alterna-
tively, if the high diploid chromosome number

derived (Meerow, 1987a), the doublet banding
pattern may reflect duplication of the genome
and would show up in pollen.

300

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FiGURES 7-11.

tetraploid Eucharis species. 7, 8. E. bouchei

Root-tip ce fi
id cell. se i cell.—9. E. deren var. bouchei from Colón Province in Pana-
. E. bonplandii. Two small chromosomes

var. dressleri. i Diplo
ma. — 10.

ei var. bouchei from Coclé aped in Panama.— 11

are outside rad figure frame. All scales = 10 u

Band a was the most common “allele” of
AAT-1, found in all individuals analyzed except
for two putative homozygotes for "allele" c (the

o Brujo population and one individual of
the El Valle population of var. bouchei). Variety
dressleri and E. bonplandii are homozygous for
"allele" a. The Río Iguanito individual of var.

bouchei is homozygous for “allele” b, while two
individuals of the El Valle population show a
putatively heterozygous phenotype with appar-
ent heterodimerization

Four E. bouchei individuals resolved a four-
banded electromorph at the putative AAT-2 lo-
cus. Bands f and g were found only in E. bon-

1987] MEEROW — EUCHARIS

5 units

1234526 7 8 9 101 12 134 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46

RELATIVE LENGTH

ITTE] )
OTE Le
D

FiGurE 12. Haploid idiograms of tetraploid Eucharis karyotypes.—A. E. bouchei var. bouchei from Coclé

Province in Panama.—B. E. bouc.
diploid cell.— D. E. bonplandii.

plandii. Only two bands were observed in var.
dressleri (an unstable tetraploid), representing
“alleles” a and b or b and c, and one individual
of var. bouchei from EI Valle (*alleles" c and d).
All other individuals of E. bouchei resolved a
four-banded electromorph for AAT-2. Band e
was found in one of the three individuals of var.
bouchei from El Valle. As all diploid species of
Eucharis species resolve only a two-banded elec-
tromorph for this isozyme (Meerow, 1986), it
was inferred that the proliferation of bands with-
in E. bouchei represented the additive effects of
tetraploidy (Crawford, 1983, 1985; Gottlieb,

MDH (Figs. 13B, 14). Malate dehydrogenase
characteristically forms complex banding pat-
terns that require genetic analysis to decipher
(Kirkpatrick et al., 1985; Torres & Mau-Lasto-
vicka, 1982). Consequently, no attempt is made
to infer genotypes in any detail from the banding
pattern. However, the phenotype of the most an-
odal bands in E. bonplandii suggests the presence
of two alleles and their heterodimer [pollen of

hei var. bouchei from Colón Province in Panama. —C.

E. bouchei var. dressleri,

this species resolved only a single band at this
locus in a repetitive run (Meerow, unpubl.), sup-
porting this interpretation]. The intensity of the
bands in the two most cathodal regions of several
individuals suggests dosage effects in putative
homozygotes (two individuals of EBBI, and

Two bands were observed
in the single locus resolved for GSSGR, all in-
dividuals manifesting one or the other

PGI (Fig. 14). Onlyasingle region of activity
was resolved for PGI. Two bands were observed,
but the more anodal one was found only in the
putative heterozygotes (two individuals of E.
bouchei var. dressleri, and one of var. bouchei
from EI Valle), and in E. bonplandii.

Isozyme relationships. The nine individuals
for which electrophoretic phenotypes were re-
solved were coded for presence or absence of the
numbered bands in Figure 14, creating a data set
of 19 characters. Phenotypes for PGI were not
included in the cluster analysis, as data were not
available for all nine individuals. The resulting

302

TABLE 7.
otherwise stated.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Karyotype data, Eucharis bouchei and E. bonplandii. All vouchers deposited at FLAS unless

Chromo-
Chromo- so
PRI jii I Chromosome Size^ Chromosome‘
Taxon, Voucher, m Range (relative Groups Morphology
& Fig. No Number (um) ngt L ML M S L M M S
Eucharis bouchei 46,92 2.4-11.9 2.0-9.7 4 10 14 18 m: 4 2
dressleris nm: 6 10
(Meerow 1107) sm: 4 6 8
Figs. 7, 8, 1 st: 6
E. bouchei v 92 1.6-15.5 0.5-5.3 14 16 14 48 m: 2
bouchei Tom nm: 2 2 18
1157) Figs. 9 sm: 10 6 6 28
12B st: 2 10 6
E. bouchei var. 92 2.0-10.4 0.8-4.5 10 18 l6 48 m: 2 2 4 6
bouchei (Meerow nm: 2 6 28
1125) Figs. 10, sm: 6 8 4 14
12A st: 8 2
E. bonplandii 92 1.9-9.7 1.0-4.9 14 12 24 42 m: 6
(Bauml 686, nm: 6 6 18
HUNT) Figs. 11, sm 2 4 16 18
12D st 6 8 2

* Based on a value of 100 for the ig iio UP ges
, ML = moderately long, = , S =

e L = lon

mall.

S
“m= metacentric, nm = ki asin dpi sm — submetacentric, st = subtelocentric.

* Diploid cell analyzed.

UPGMA dendrogram (Fig. 15) and cladogram
(Fig. 16) illustrate the isozyme relationships
among these individual plants. As might be ex-
pected, both trees are similar in topology. All
three individuals of E. bouchei var. dressleri
(EBD) cluster at a distance coefficient (DC) of
only 0.118, indicating their close isozyme rela-
tionship. The El Valle (Coclé Province) popu-
lation of var. bouchei (EBB1) is rather diverse in
its patterns of isozyme variation. Two individ-
uals are similar, clustering at a DC of 0.118. This
cluster then fuses with the Río Iguanita (Colón
Province) pasa of var. bouchei (EBB3) ata
DC of 0. . The remaining individual of El
Valle var. be EBBI shows greater isozyme
relationship to Cerro Brujo (Colón Province) var.
bouchei EBB2 than other convarietal individuals
from El Valle. This heterogeneous cluster then
fuses with var. dressleri (EBD) at a DC of 0.431,
followed by the other El Valle individuals (EBB1)
and the single Río Iquanita (EBB3) individual
(DC 0.443). The single individual representing
E. bonplandii (EBN) remains a distant outlyer
from all plants of E. bouchei, joining the latter
species at a DC of 0.625

The cladogram based on isozyme data was 27
steps long with a consistency index (Kluge &
Farris, 1969; CI — total length minus homopla-
sies, divided by total length) of 0.704. The clado-
gram supports a monophyletic origin of E.
bouchei var. dressleri (EBD) from var. bouchei
(EBBI) in Coclé Province, but suggests that var.
bouchei may be polyphyletic. The Río Iguanita
(Colón Province, EBB3) individual of var. bouchei
forms a monophyletic group with two individ-
uals of El Valle (Coclé province, EBBI) var.
bouchei, while the other Colón individual (Cerro
Brujo, EBB2) forms a monophyletic group with
the third El Valle individual.

DISCUSSION

The Central American £. bouchei complex does
not resolve oe discrete penenie groups. Stam-
inal c logy, however te va-
rieties to a fair degree (Fig. 5). Floral s size char-
acters in this group (Figs. 4, 6) do not succeed
as well in resolving phenetic groups.

Chromosome number is very stable in Eucha-
ris, and polyploidy is infrequent. The origins of
the polyploids (i.e., whether auto- or alloploid)

EBD

EBN EBD EBB) EBD EBB-3 EBB2 EBB) EBB !

13. Representative gels for borus vois
i complex

-
c
—
-
E

re e ru
putative Mh discussed in text. Refer to Table 4 for
population designations.

are inconclusive (attempts to secure meiotic fig-
ures have been unsuccessful), but the high levels
of morphological diversity in E. bouchei might
suggest that they are alloploids. Differences in
chromosome morphology among the popula-
tions of E. bouchei examined (Table 7) suggest
that structural changes in the chromosomes may
have been important in interpopulational diver-
gence. Eucharis bouchei var. dressleri is an un-
stable tetraploid. Somatic cells of b» E tips
e both t

nte

PH iN J TIJ Vu

hav
Snoad (1955) a karyotype MA a: in

as polyploid counts were observed in the cells of
the latter species.

Polyploid species of Eucharis do not show any
marked effects of increased chromosome number
beyond an increase in size of root cells and sto-
mata, and slight thickening of the leaf blades.
Eucharis bonplandii, in addition, develops a
glaucous bloom on the leaves in strong light, a
novel characteristic for the genus.

MEEROW — EUCHARIS

| |!

EBD EBD EBD EBB-1

EBD EBD EBD

— ló
— 17

EBB-1 FBB-1 EBB-I EBB-2 EBB-3 EBN

EBD EBD EBD

i complex. Anoda

a à — aspartate
transferase, MDH = malate dehydrogenase, GSSGR =
glutathione reductase, PGI = phosphoglucoisomerase.

Eucharis bouchei var. bouchei, as presently
Sea Meera, 1986), is diverse in floral
] nd pat-

terns of i isozyme polymorphisms. Xon re-
lationships based on the isozyme data indicate
that this polymorphic variety may even be poly-
gically EBB3 is similar to

a O
ouchei (Fig. 1Aiii), they are not discontinuous

0.7004

0.600—

DISTANCE
>
[^]
°
1

0.2004
0.100
0.000
EBD EBD E8D
FiGURE 15.
designations.

enough to warrant a clear differentiation of a
fourth variety in the species. The two individuals
representing different populations from Colón
Province each have closer isozyme relationship
to Coclé var. bouchei than they have to eac
other, while the Coclé population itself appears
isozymically diverse. Cladistic relationships based
on this data imply that Coclé populations of var.
bouchei may be of heterogeneous ancestry. The
cladogram (Fig. 16) also points to Colón Prov-
ince as the likely origin of those ancestors. Di-
vergence between and among the Coclé popu-
lations and those in Colón Province, presumably
mediated by geographic isolation, may thus
an s process.

the basis of staminal cup morphology, I
i petens that Eucharis bouchei has been

). The Cerro Brujo pop-
ulation of E. bouchei var. bouchei may represent
an intermediate point in the divergence of a new
geographical race of E. bouchei. The best test of
this hypothesis would be the results of isozyme
analysis of E. bouchei var. darienensis, the one
variety for which material is not presently avail-
able. Variety darienensis occurs closer to the Co-
lombian border than any other population of E.
bouchei and has the most generalized staminal
cup morphology relative to Eucharis as a whole.
If my hypothesis is correct, var. darienensis

ANNALS OF THE MISSOURI BOTANICAL GARDEN

EBB1
UPGMA dendrogram oftetraploid Eucharis isozyme phenotypes. Refer to Table 4 for population

[VoL. 74

EBB2 EBBI EBBI EBB3 EBN

should have the lowest genetic identity with pop-
ulations of either var. bouchei or var. dressleri
from Coclé Province, and higher identity with
populations of var. bouchei from Colón Prov-
ince.

Colón populations of E. bouchei are geograph-
ically intermediate between most populations of
var. darienensis and the Coclé populations of
var. bouchei (Meerow, 1986). The two varieties
come into close proximity in the Cerro Campana
area in Panamá Province. Colón populations may
therefore also be genetically intermediate be-
tween the two varieties. Segregating genotypes in
such a case could produce populations exhibiting
a mosaic of varying genetic identity, some close
to Coclé var. bouchei, others perhaps closer to
var. darienensis. Further testing of this hypoth-

the first step in sympatric speciation. This variety
shows greatest isozyme relationship with certain
individuals of the Coclé population of var.
bouchei. In this regard, the presence of a second
band for PGI in certain individuals of var. dres-
sleri and El Valle var. bouchei may be significant
(Fig. 14), but a larger number of individuals must
be assayed to confirm this observation. This va-

1987]

EBD

MEEROW — EUCHARIS

EBBI

EBB3 EBBI EBB2

—: reversal
æ : homoplasy

16. Cladogram based on isozyme Ee of tetraploid Eucharis. Refer to Table 4 for population

P aypuy asi Broken line indicates zero-length branch

riety is also an unstable tetraploid. Fifty percent
of all chromosome counts of root tip cell mitotic
metaphase configurations have 2n = 46, the typ-
ical diploid chromosome number in Eucharis.
Variety dressleri lacks the additive banding pat-
terns or heterozygote phenotypes observed in
both loci of AAT in all other populations of E.
bo uchei, a factor, perhaps, of this karyotypic in-

Eun (CE et al., 1972), Nicotiana (Red-
dy & Garber, 1971; Sheen, 1972; Smith et al.,
1970), Triticum aestivum (Hart, 1970, 1979;
Jaaska, 1978; Torres & Hart, 1976), and Steph-
po (Gottlieb, 1973), and are usually in-
rpreted as indicative of allopolyploid origins
= ach 1983; Gottlieb, 1983; Soltis & Rie-
seberg, 1986). Pollen stainability of var. dressleri
is 10096 with Alexander's (1969) stain, suggesting
that gamete formation is not impaired by the
chromosome number instability. Nonetheless, I
have not successfully crossed this variety with El
Valle populations of var. bouchei
The rare Colombian ietraploid E. bonplandii
also lacks either heterozygote or additive band-
ing patterns for AAT. This may indicate an au-
topolyploid origin for this species (Crawford,
1985; Soltis & Rieseberg, 1986), or at least a
lower degree of heterozygosity than in E. bouch-
ei. The rarity of this species, in relative contrast
to E. bouchei, fits Stebbins's (1980) model of the
“unsuccessful” autopolyploid. The difficulty in
obtaining successful meiotic figures from bulbs

pletely inside the bulb)
this question. Eucharis bonplandii exhibits large

distance on the basis of isozyme phenotypes from
all individuals of E. bouchei. The question of
whether these two species represent a monophy-
letic group on the basis of their tetraploid origin
is not conclusive.

Increased heterozygosity is an expected con-
sequence of allopolyploidy (Crawford, 1983,
1985; Gottlieb, 198 1a; Soltis & Rieseberg, 1986).
A certain degree of fixed heterozygosity would
also be expected in an allopolyploid (Gottlieb,
198 1a; Soltis & Rieseberg, 1986), due to the pres-
ence of two genomes in the allotetraploid. Al-
though it is premature to assess the degree of
heterozygosity present in tetraploid Eucharis,
their morphological diversity and i dim poly-
morphisms indicate that it may

There is insufficient information on ne breed-
ing system and pollination biology of Eucharis
to support more than ad hoc hypotheses of the
origin of most species in the genus. The char-
acteristically small population sizes that are en-
countered throughout its range may indicate that
ounder effects (Mayr, 1954; Templeton, 1980a,
1980b) have played an important role in the
movement of E. bouchei across the Isthmus of
Panama, with subsequent isolation restricting
gene flow between localized populations. The pu-
tatively allotetraploid genotype and karyotypic
polymorphy of ouchei would favor the “‘hy-
brid recombination" type of "genetic transili-
ence," a mode of speciation hypothesized by
Templeton (1980a, 1980b). This is consistent with
the distribution of E. bouchei, most populations
of which are geographically isolated from each
other (Meerow, 1986). Additional support comes
from the morphological novelties expressed

306

within each geographical variety (or race, in the
case of var. bouchei). Templeton’s model of spe-
ciation requires a period of successful inbreeding
after the initial founding event. Though Eucharis
are primarily out-crossing, evidence from green-
house studies indicates only partial self-incom-
patibility (Meerow, 1986). At least one Ama-
zonian species of subg. Eucharis [E. castelanaena
(Baillon) Macbride] is autogamous (Meerow,
1986)

EUCHARIS IN CENTRAL AMERICA

Gentry (1982) suggested that two major op-
portunities, widely spaced in time, existed for
floristic interchange between Central and South
America. The first, occurring during the Late
Cretaceous, was limited to a series of volcanic
islands (the proto-Antilles; Dengo, 1975; Lille-
graven et al., 1979). The degree to which this
island arc remained above water is unknown. At
the beginning of the Tertiary, however, this link
between the continents was disrupted as the pro-
to-Antilles began a northward displacement. It
was not until the late Tertiary that the second
opportunity for floristic interchange began to co-
alesce, as formation of the Central American
trench and new volcanic activity gave rise to a
new series of islands. These islands eventually

(1982) concluded that only very well-established
Cretaceous taxa would have been able to take
advantage of the earlier connection via island-
hopping. Entries into Central America dating
from this earlier connection would be expected
to show strong taxonomic differentiation in Cen-
tral America. Gentry (1982) cited tribe Crescen-
tieae of the Bignoniaceae as a putative example
of early colonization of Central America by is-
land-hopping, followed by taxonomic differen-
tiation. On the contrary, any migration dating
from the Pliocene or Pleistocene would not be
expected to show much differentiation at the spe-
cific or at the generic level. I have characterized
the Eucharis bouchei complex as a semispecies
complex of geographically isolated races or va-
rieties not yet strongly differentiated. Patterns of
isozyme variation, chromosome cytology, and
morphological variation in this group suggest that
entry of Eucharis into Central America was fairly
recent.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

The species of subg. Eucharis geographically
closest to E. bouchei is E. bonplandii, a rare
species of central Colombia, and also tetraploid.
It is inconclusive whether these two species rep-
resent a monophyletic tetraploid group. None-
theless, the congruence of phytogeography with
chromosome number in these two species sug-
gests that this may indeed be the case. It is tempt-
ing to wonder if tetraploid Eucharis were at one
time more common in northern Colombia, and
if E. bu and E. bonplandii represent the
remnant populations of a once more widespread,
d tetraploid complex. Prance's (1982)
most recent distribution of Pleistocene refugia
based on phytogeographic patterns includes both
a Río Magdalena refuge in northern Colombia
(most collections of E. bonplandii are from the
Rio Magdalena valley south of Prance's pro-
posed refuge), and a Darién refi

enensis is most common in the area ofthe Darién
refuge and is putatively the least derived variety
of the species. The absence of collections of Eu-
charis subg. Eucharis from northern Colombia
is something of a mystery but may indicate that
extinction of intervening populations between E.
bonplandii and E. bouchei was widespread in the
recent geological past.

CONCLUSIONS

The Central American E. bouchei complex is
a tetraploid, putatively alloploid, possibly highly

1981) still actively evolving. Discrete patterns of
isozyme divergence have not yet solidified be-
tween morphologically distinct and geographi-
cally isolated populations of E. bouchei var.
bouchei. Founder effects and geographic isola-
tion probably were, and still are, important forces
influencing the continued evolution of E. bouch-
ei. In one case (E. bouchei var. dressleri) sym-
patric divergence may be in process
unprecedented degree of variation in E.
bouchei is thus likely the result of two main fac-
tors: (1) tetraploidy, accompanied or followed by
structural rearrangement of chromosomes, and
(2) a geologically recent colonization of Central
America by this primarily northern Andean and
Amazonian genus. The wide variation present in
E. bouchei likely represents the segregating phe-
notypes of a richly diverse genetic base. On the
basis of known distributions, it appears that sub-
stantial geographic barriers exist between groups

1987]

of populations, probably restricting gene flow be-
tween them. Left undisturbed, as is not the case
in the Neotropics today, these aggregates could
conceivably one day each justify specific recog-
nition.

The fact that E. bonplandii is the northernmost
species of Eucharis subg. Eucharis in South
America, and is also tetraploid, lends at least
circumstantial credence to the hypothesis that E.
bouchei and E. bonplandii diverged from a com-
mon tetraploid ancestor. The rare occurrence of
polyploidy in Eucharis strengthens this possi-
bility.
Stebbins (1985), in a recent review of poly-
ploidy, found a correlation between high fre-
quency of polyploidy and patchy g } (or
ecological) distributions, coupled with the oc-
currence of secondary contact between these dif-
ferentiated populations. Levin (1983) discussed
how chromosome doubling may “ ‘propel’ a pop-
ulation into a new adaptive sphere.” Though E.
bouchei does not exhibit any noticeably novel
ecological adaptations, its success in colonizing
the Isthmus of Panama may have been aided by
its polyploid-related genetic diversity. The het-
erogeneous isozyme patterns characteristic of El
Valle var. bouchei may indicate either multiple
ancestry for this population or that secondary
contact has occurred between it and populations
in Colón or Panama provinces to the east.

Eucharis bouchei offers an excellent opportu-
nity for detailed study of the evolution of a trop-
ical rainforest organism. Future work should seek
to quantify in greater detail the genetic variation
present within and among populations of this
actively evolving complex. Meiotic pairing fig-
ures from dissection of bulbs may help confirm
the nature of the polyploid origins of this species.

Paleotropical genera of “‘infrafamily”’ Pancra-
tioidinae characteristically have 2n — 22 or 20
chromosomes (Ponnamma, 1978; Zaman &
Chakraborty, 1974), while almost all neotropical
genera have 2n = 46 (Di Fulvio, 1973; Flory
1977; Meerow, 1987a, 1987b; Williams, 1981).
The latter number is likely derived through frag-
mentation or duplication of a single chromo-
some, followed by doubling of the genome
(Lakshmi, 1978; Sato, 1938). Increased hetero-
zygosity may therefore have accompanied a tet-
raploid origin of the neotropical tribes of the
Pancratioidinae from an ancestor with 2n = 22
" e somatic number characteristic of Pancra-

m L., the largest paleotropical genus of the
PEL (Ponnamma, 1978)]. The high generic

MEEROW —

EUCHARIS 307

diversity of neotropical pancratioids (ca. 15 gen-
era) in comparison with the paleotropical taxa
(4 genera) itself may be partially a consequence
of greater genetic variability. Comparative anal-
ysis of isozyme phenotypes between paleotrop-
ical and neotropical genera is planned and may
provide insight into the evolution of the Pan-
cratioidinae.

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58.

THE SHRUBBY GENTIAN GENUS MACROCARPAEA
IN PANAMA

KENNETH J. SYTSMA!

ABSTRACT

A single species of the lisianthoid genus Macrocarpaea had been known from Panama. Recent
m

The shrubby gentians of the Neotropics are
some of the most conspicuous elements of higher
elevation tropical forests. This complex of about

6 genera comprised Grisebach’s (1838) tribe
Lisyantheae. These lisianthoid genera are noto-
riously difficult to separate taxonomically with
the consequence that they have often been treat-
ed as synonyms of Lisianthius (or its orthograph-
ic variant "Lisianthus"). The taxonomic diffi-
culties are compounded by their remote montane
habitat which makes them poorly collected. Only
Lisianthius sensu stricto (Weaver, 1972a) and
Macrocarpaea (Ewan, 1948) have been ade-
quately monographed. Other genera of the tribe
Lisyantheae are now being taxonomically re-
vised in a multidisciplinary study (Maas et al.,
1984; Maas, 1985).

Macrocarpaea (Griseb.) Gilg is one of four lis-
ianthoid genera in Panama (Elias & Robyns,
1975). Symbolanthus pulcherrimus, Irlbachia
alata subsp. alata (formerly Chelonanthus ala-
tus), and seven species of Lisianthius (Sytsma,
1987) also occur in Panama. These four genera
can be separated in Panama by the following
characteristics:

la. Main stem terete; stigma capitate; old pla-
centae visible as whitish bands along margins
of mature capsules; pollen grains as monads
. Lisianthius
Main stem usually deii dri stigma bi-
furcate; old placentae not visible on mature
capsules; pollen grains as Min or tetrads.
6-10 cm long; corolla funnel-
orm; more or less distinct, membrana-
ceous; corona scalelike; pollen grains as
tetrads Symbolanthus
. Flowers to 5 cm long; corolla usually

=

N
[^

as monads or tetra
3a. Leaves Bee "bracteoles never

wallioides, M. anco: and M. subcaudata. Evi-
dence is provided to merge the Costa Rican M. jer ll

into M. macrophylla.

leafy; calyx z mm long; pollen
grains as tetra Irlbachia

3b. Leaves petiolate: bracteoles. leafy;
calyx 6-20 mm long; pollen grains
as monads u. Macrocarpaea

The genus Macrocarpaea is centered in the
Andes of northern South America but extends
into the Amazon Basin and the Guayana High-
land. Of the approximately 30—50 species in the
genus, only eight are known from Central Amer-
ica and adjacent West Indies. Macrocarpaea
domingensis Urban & E. Ekman and M. tham-
noides (Griseb.) Gilg are restricted to the Do-
minican Republic and Jamaica, respectively.
Cuba has two endemic species, M. pinetorum
Alain and M. pauciflora Alain. Three species have
been described from Costa Rica (Weaver, 1972b):

m
Nilsson, has been known from Panama near the
border of Costa Rica (Elias & Robyns, 1975).
The type specimen had represented the only col-
lection for this apparently epiphytic shrub prior
to 1975

Recent explorations of cloud forest habitats in
central regions of the Cordillera Talamanca, a
ridge extending from the border of Costa Rica
to near Panama City, and in the Cordilleras of

of the Missouri Botanical Garden
through the Flora of Panama Project (funded by
NSF grant BSR-8305425). Macrocarpaea sub-
caudata, an epiphytic shrub known previously
from only two collections in one region of Costa

! Botany Department, University of Wisconsin, Madison, Wisconsin 53706, U.S.A.

ANN. MiSSOURI Bor. GARD. 74: 310-313. 1987.

1987]

FIGURE 1.
square = M. macrophylla; triangle = M. subcaudata

Rica, occurs in one cloud forest region in the
mountains of central Panama. Recent additional
collections of M. browallioides now extend the
range of this distinctive epiphytic shrub. Several
populations similar to both the Costa Rican M.
valerii and the Colombian M. macrophylla
(Kunth) Gilg are now known from the Darién.
Analysis of morphological variation among the
Costa Rican, Panamanian, and Colombian pop-
ulations provides a basis for merging M. valerii
with the now more widespread M. macrophylla.

KEY TO MACROCARPAEA IN PANAMA

la. Calyx Ped mm long, lobes ovate-oblong,
rounded a
2a. seme viny subshrub, Ment narrow-
ly lanceolate, to 18 cm long
wide pa
. Terrestrial shrub to 4 m, leaves s broadly
elliptic, to 45 cm long and 26
wide M. macrophylla
. Calyx 15-18 mm long, lobes triangular, a
to cuspidate at apex 3. M. pese eee

N
c

ox

1. Macrocarpaea subcaudata Ewan, Contr. U.S.
Natl. Herb. 29: 224. 1948. TYPE: Costa Rica:
[San José] La Palma, Wercklé 16492 (holo-
type, US; isotype, NY).

SYTSMA — MACROCARPAEA

N

Geographical distribution of the genus Macrocarpaea in Panama. Circle = M. browallioides;

Epiphytic, viny subshrub. Leaves essentially
glabrous except for small scattered hairs, some-
what thickened, narrow-lanceolate, to 18 cm long
and 5 cm wide; petioles to 23 mm long. Inflo-
rescence terminal or axillary from upper nodes,
dichasium bi- or tri-ternately compound, often
long-stalked. Calyx campanulate, greenish, gla-
brescent, 6-9 mm long; lobes slightly unequal,
ovate-oblong, rounded or ciliolate at tips, 5-7
mm long, 3-4 mm wide. Corolla greenish yellow,
to 3 cm long; tube to 2.3 cm long; lobes slightly
recurved (incurved when dried), narrowly tri-
angular, to 9 mm long, to 6 mm wide. Stamens
inserted near middle of corolla tube; filaments
to 17 mm long, just surpassing corolla lobes;
anthers yellow, 4-5 mm long. Style just surpass-
ing anthers. Capsules woody, 10-12 mm long ex.
persistent beak of 3 mm. Flowering period at
least mid-April through May.

Distribution. 1,500 m in mountains east of
San José, Costa Rica and between 1,150-1,260
m near continental divide between Chiriquí and
Bocas del Toro provinces in central Panama (Fig.

lk

Additional idis examined. PANAMA. BOCAS
DEL TORO: Con tal divide on carretera del oleo-
ducto ca. 1 km x of Quebrada Arena, IRHE Hydro-

electric ia s Knapp 5089 (MO), MUR Se
(MO). CHIR

Hydroelectric bout 10.1 mi. NW of Los "umi x
Hornito, Antonio 4190 (MO).

Macrocarpaea subcaudata occurs only in cen-
tral Costa Rica and northcentral Panama. De-
termining whether this disjunction is real or not
must await further collecting in as yet inacces-
sible cloud forest regions of both countries. Re-
lationships of M. subcaudata to other species of
Macrocarpaea are unclear. Macrocarpaea bro-
wallioides of Panama shares the epiphytic habit
with M. subcaudata, but the two are clearly un-
related. Ewan (1948) considered M. cerronis Ewan
and M. salicifolia Ewan from the tepuis of the
Guayana Highland to be the closest relatives of
M. subcaudata. The former two species (with
related M. arborea (Britton) Ewan, M. quelchii
(N.E. Br.) Ewan, and M. tepuiensis (Gleason)
Steyerm.), however, differ from all other species
of Macrocarpaea in having pollen in tetrads
(Nilsson, 1968, 1970). Maas (1985) combined
these six Macrocarpaea species with pollen in
tetrads into /r/bachia quelchii (N.E. Br.) Maas.
Weaver (1972b) cited M. acuminata Weaver and
two West Indian species as the closest relatives
of M. subcaudata. The Costa Rican M. acumi-
nata is almost certainly related to, if not con-
specific with, M. macrophylla (including M. va-
lerii). Macrocarpaea acuminata and M.
macrophylla share with M. subcaudata similar
calyx features and might be close relatives of M.
subcaudata.

2. M hylla (Kunth) Gilg, Nat.
Pflanzenfam. 4(2): 94. 1895. Sap
macrophyllus Kunth, Nov. Gen. & S
183. 1819. TYPE: Colombia: trail over = "£
millo to Almaguer between Pansitara and
Rio Ruiz, 8,400 ft., Humboldt & Bonpland
(Willd. Herb. 3561 fide Grisebach; MO pho-
to 37455, of collection in Humboldt Her-
barium at Paris).

Macrocarpaea valerii Standley, Publ. Field Mus. Bot.
18: 928. 8. TYPE: Costa Rica: La Hondura de
San José, Valerio 692 (F).

Erect shrub or subshrub to 4 m tall. Leaves
essentially glabrous, except for scattered small
hairs, venation strongly prominent, broad-ellip-
tic, to 45 cm long and to 26 cm wide, blade acute
or abruptly acuminate; petioles to 3 cm long.
Inflorescence terminal or axillary from upper
nodes, dichasium simple or bi- or tri-ternately

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

compound, long-stalked. Calyx campanulate,
fleshy, 8-14 mm long; lobes strongly unequal,
ovate-oblong, hyaline-margined, 3-6 mm long
and wide. Corolla greenish-white or cream, nar-
rowly to openly campanulate, to 3.5 cm long;
tube 2.5-3.0 cm long; lobes recurved, broadly
triangular, 6-8 mm long and wide. Filaments to
18 mm long, included in or surpassing corolla
tube; anthers yellow, 3-5 mm long. Style to 15
mm long; stigma bilobed, the lobes 2 mm long.
Capsules woody, to 20 mm long, persistent-
beaked. Flowering period at least May through
July.

Distribution. 1,000-1,800 m in mountains of
Central Costa Rica; 1,000-1,400 m in Serranía
del Sapo and Serranía del Darién of Darién Prov-
ince, Panama (Fig. 1); and common at 1,500-
2,500 m in the western and central Cordilleras
of Colombia.

Additional specimens examined. PANAMA. DARIEN:
S

o
a expedition, Gentry & Mori 13655 (MO), CEU:
P ‘al 16975 (MO).

Macrocarpaea macrophylla is distinctive as a
tall erect shrub with large round leaves and
prominent venation. As here defined, M. macro-
phylla ranges from Colombia to Costa Rica.
Standley originally described the Costa Rican M.
valerii based only on the type specimen. Ewan
(1948) reiterated the differences between the two
species based on only two Costa Rican collec-
tions. These differences were confined to size and
shape of leaves and to corolla shape. Both sets
of characters are subject to sampling error de-
pending on how the plants were collected and
pressed. Subsequently, additional collections of
" M. valerii" in Costa Rica indicated that these
features were not consistent, causing Weaver
(1972b) to state that “the two are virtually iden-
tical, except the calyx of M. valerii is glabrous,
while that of M. macrophylla is spiculate." Co-
lombian, Panamanian, and Costa Rican speci-
mens of these two taxa at MO and WIS showed
considerable variation in the degree of surface
ornamentation on the calyx. No consistent calyx
differences or other foliar and floral differences
were seen among specimens described as M.
macrophylla and M. valerii. This necessitates the
inclusion of these Colombian, Panamanian, and
Costa Rican specimens into M. macrophylla
(Kunth) Gilg, the name with priority.

1987]

3. Macrocarpaea browallioides (Ewan) Robyns
& S. Nilsson, Bull. Jard. Bot. Natl. Belgique
40: 13. 1970. Lisianth(i)us browallioides
Ewan, Proc. Biol. Soc. Washington 64: 132.
1951. TYPE: Panama. Bocas del Toro: north-
ern slopes of Cerro Horqueta, Robalo Trail,
Allen 4932 (MO).

Epiphytic shrub to 1.5 m tall. Leaves glabrous,
+ coriaceous, dark green and black-punctate
above, gray-green below, elliptic to ovate-ellip-
tic, to 11 cm long and to 5 cm wide; petioles to
25 mm long. Inflorescence a compound dicha-
sium, + long-stalked. Calyx campanulate, light
green, 15-20 mm long; tube short; lobes ovate-
triangular, strongly overlapping, 12-16 mm long,
acute to long-acuminate at apex. Corolla fleshy,
yellow-green to cream yellow, infundibuliform,
3.0-4.0 cm long; lobes subequal, ovate-trian-
gular, acute at apex, to 1.2 cm long. Stamens
inserted ca. 6 mm above base of corolla tube, +
exserted; anthers yellow, 5-6 mm long. Style just
surpassing anthers. Capsule woody, to 14 mm
long, excluding persistent beak of 6-8 mm. Flow-
ering in February, May, August, and late No-
vember.

Distribution. 1,690-2,200 m, cloud forests of
cerros Horqueta, Pate Macho, and Colorado,
provinces of Chiriquí and Bocas del Toro.

Addi tional A Li examined. PANAMA. BOCAS
st along trail from end of Rio Palo Alto
road to Chiriqui/ Bocas del Toro border near peak of
Cerro Pate Mac v Hammel 5781 (MO). CHIRIQUI:
Cerro Pate Macho, ca. 5 mi. NE of Boquete, trail to
continental divide ‘leading to Finca Serrano, Antonio
2654 (MO); Cerro Colorado, along mining road 31. 6
km beyond bridge over Rio San Felix, 10.6 km beyond
turnoff to Escopeta, Croat 37155 (MO); Cerro Pate

a e of Pate Macho, Hammel 6112 (MO); NW

Finca OVIIO

ridge
no, | eiua et al. 4900 (MO).

Macrocarpaea browallioides is easily distin-
guished from the other Central American species
of Macrocarpaea by its large calyx and long-acu-
minate calyx lobes. Like M. subcaudata it is epi-
phytic, but unlike M. subcaudata it is erect rather

SYTSMA — MACROCARPAEA

313

than viny. Several of the collectors do not men-
tion an epiphytic habit. However, the distinction
between an epiphytic and terrestrial habit is ten-
uous on an extremely wet, peatlike soil as is
encountered on Cerro Pate M i
phytic as well as free-standing ‘although on mats
of living roots or trunks) individuals of M. bro-
wallioides were seen on this cloud forest Dead
The relationship of this species to other members
of Macrocarpaea is unclear. Neither Ewan, who
first described the species as a Lisianthius, nor
Robyns and Elias, who transferred it to Macro-
carpaea, gave any indication as to its likely rel-
atives. Only three known species approach M.
browallioides with its distinctive short calyx tube
and long, overlapping, and acuminate-tipped
lobes: M. cochabambensis Gilg of Bolivia; M.
glabra (L.f.) Gilg endemic to paramo vegetation
above Bogotá, Colombia; and M. guttifera Ewan
of Amazonia, Brazil.

LITERATURE CITED

Euas, T. S. & A. Ropyns. 1975. Family 160. Gen-
tianaceae. In Flora of Panama. Ann. Missouri Bot.
Gard. 62: 61-

EwAN, J. 1948. A revision of Macrocarpaea, a neo-
tropical genus of shrubby gentians. Contr. U.S.
Natl. Herb. 29: 209-249.

GRISEBACH, A. H. R. 1838. Genera et species Gen-
tianearum. Stuttgart & Tübingen.

MAAS, P. 1985. Nomenclatural notes on neo-
tropical Lisyantheae ( — uris Proc. Kon-
inkl. Nederl. Akad. Wetensch. C. 88: 405-412.

etal. 1984. wd Ves studies in n

Pike oup the Lisianthius complex. Acta Bot

eerl. 32: 371-374

Pollen morphology in the genus

significance. Svensk Bot. Tidskr. 62: 338-364.
. 1970. Pollen morphological contributions to
t. (Gentiana-

1-43.
. Biosystematics and evolution in
the Lisianthius skinneri (Gentianaceae) species
complex in Panama. Ann. Missouri Bot. Gard.

(MS).

Weaver, R. E., JR. 1972a. A revision of the neo-
tropical genus Lisianthius (Gentianaceae). J. Ar-
nold Arbor. 53: 76-100, 234-311.

2b. The genus Macrocarpaea (Gentiana-

ceae) i in Costa Rica. J. Arnold Arbor. 53: 553-557.

MONOECY AND SEX CHANGES IN FREYCINETIA
(PANDANACEAE)!

HANS-HELMUT POPPENDIECK2

ABSTRACT

In cultivation, did: sae of the s dioecious genus
sequentially and MM on dem plans

pistillate and s inflore ces

"D. 4 A

These eg are AM een) E with other cases of occasional deviations from dio

According to most textbooks and general ref-
erence works the genus Freycinetia is considered
to be dioecious (Warburg, 1900; Heywood, 1979;
Dahlgren et al., 1985; Stone, 1984), although
some exceptions have been noted (Stone, 1972;
Cox, 1981; Cox et al., 1984), drawing attention
to the significance of these presumably rare ex-
ceptions to the dioecious condition. In Freyci-
netia reineckei Warb., Cox has observed func-
tionally bisexual spikes, and both pistillate and
staminate inflorescences on the same branch in
Freycinetia scandens Gaudich., and Stone (1972)
has noted the same in Freycinetia negrosensis
Merr. and F. imbricata Blume

The following observations made in the trop-
ical greenhouses of the Hamburg Botanical Gar-
den confirm and extend these earlier observa-
tions and, by adding some new pieces to the
puzzle, seem to indicate that sex expression in
this genus is to a considerable extent under en-
vironmental control. Both Freycinetia funicula-
ris (Savigny in Lam.) Merr. and F. cumingiana
Gaudich. (synonymous with F. /uzonensis Presl,
according to Stone) have been observed to show
monoecy.

Both of the species mentioned above are easily
cultivated from cuttings, and the material in cul-
tivation probably represents single clones of each,
so that the observations made at different places
relate to genetically identical plants. Freycinetia
funicularis is comparatively widely distributed
in German botanic gardens, but its origin cannot
be ascertained; however, it is a native of Indo-
nesia, and probably was introduced from the Bo-
tanic Gardens in Bogor (formerly Buitenzorg),
where it has been in cultivation for perhaps a
century and still is common.

Freycinetia cumingiana (F. luzonensis) is a

Philippine species which came to Hamburg from
the Bonn Botanic Garden, and according to the
curator there, Dr. K. Kramer (pers. comm .), it
was introduced to Europe via Tübingen from a
source in the U.S.A., initially under the name
"F. cunninghamiana." It has been possible to
trace this clone back to its original source. Ap-
parently it had been sent by E. D. Merrill from
the Philippines to the New York Botanical Gar-
den, from there to Puerto Rico, and to the Fair-
child Tropical Garden in Miami, Florida, where
it caught the attention of Dr. R. A. Howard of
Harvard University, who reported his observa-
tions to Dr. B. C. Stone (see Table 1). Eventually
this species found its way to Europe, perhaps
from the Fairchild Tropical Garden

OBSERVATIONS ON GROWTH AND
SEX EXPRESSION

Observations on both species are summarized
in Table 1. It is remarkable that despite the pro-
ed diffe i hoot hit t i

í

S ee Fi
1), the two species have produced staminate and
pistillate inflorescences, both sequentially and si-
multaneously. Freycinetia cumingiana appar-
ently produces different kinds of shoots, prede-

M shoot diam
decreases with age, the initially pistillate shoots
convert to the production of staminate inflores-
cences. Evidently, shoot diameter and sex
expression are correlated in this species. The
qualitative “jump” may occur when the shoot
diameter decreases to less than 4 mm (Fig. 2;

able 2). Perhaps this is also the diameter range
wherein mixed i y form. In Fre rey-

' My thanks to Dr. B. C. Stone for encouragement, to Dr. K. Kramer for details on Freycinetia cumingiana,

J. Bogner on F. Led gen Dr. M. F
our greenhouse sta expert cultiv
? [nst

ANN. Missouni Bor. GARD. 74: 314—320. 1987.

eis dau Prof. Dr. K. Kubitzki for advice, and Mrs. H. Sch

wob-Tonn and

.f Alemana. Botanik u. rie Garten, Hesten 10, D 2000 Hamburg 52, West Germany.

1987] POPPENDIECK — FRE YCINETIA 315

TABLE 1. Comparison of growth and flowering of two species of Freycinetia in the Hamburg Botanic Gardens,
1982-1986

Freycinetia funicularis Freycinetia cumingiana

Large climber (“rooter”) with horizontal
clinging roots and vertical roots; inflores-
cences lateral on at least one-year-old, still
leafy shoots, often penetrating the leaf-bas-

Morphology cei “leaner” with perennial shoots
ich may flower in 2nd year; inflores-
cences ande in axis or on rather
PR lateral shoots; shoots dimorphic,
dia mm in staminate shoots and

PET mm in pistillate shoots.

Flowering particulars

1982 Abundant (ca. 50) inflorescences. Several vegetative shoots; one fertile, with
about 12 pistillate inflorescences
1983 Abundant but exclusively pistillate inflores- Only staminate inflorescences observed.
ces

1984 Numerous staminate inflorescences plus a Not recorded; allegedly staminate, but pres-
single pistillate inflorescence at the base of ence (or not) of pistillate inflorescences un-
the plant; viable seeds and offspring ob- certain
tained after hand-pollination.

1985 Twelve staminate inflorescences, most pro- Two ramified shoots with staminate inflores-

1986

Additional
observations

duced later than usual after severe pruning
had reduced the shade made by the plant
itself.

No inflorescences on the old plant. Young
plant (4 yr. old) with 12 staminate inflores-
cences mostly on vertical axes, 5 riu
inflorescences on subapical parts of hori-
zontally trained shoots, and one mixed in-
Vend on a vertical shoot. pores
yields fruit

Similar observations by J. Bogner in Munich
Bot. Gard. Younger plants from cuttings
first al pistillate then both pistillate
and staminate inflorescences in the next
and unde years. Encke's account
(1958) of “Freycinetia insignis" probably
refers to F. funicularis.

cences which had probably flowered the
year before; 3 pistillate shoots, at least 2
flowering for the first time (Fig. 1); several
fruits formed (no viable seed).

Staminate shoots continue to form staminate

apparently pistillate-determined shoots
have been formed. New shoot formed in
1985 (diam. 6.2 mm) is pistillate.

A somewhat different sequence of flowering
y Dr

as been observed
(Bonn Bot. Gard.): the first inflorescences
were staminate; later, also pistillate inflo-

res were p
the branches. Eventually also mixed inflo-

, when I found an inflorescence pro-
ducing pollen” (April 1969).

cinetia m this shoot differentiation is
lacking, s
ko seal pistillate inflorescences in 1983 is
unexplained. However, it seems to indicate that
an adjustment over the whole vegetative body
can take place, which may be hormonally (or

that the synchronized formation of

nutritionally) determined. Observations made in
the Munich Botanical Garden by Josef Bogner
(pers. comm.) suggest that larger and vigorous
plants of F. funicularis may develop inflores-
cences of both sexes.

It is difficult to assign these Freycinetia species

316

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FI f ah
black circles; stami nate a penance circles; vegetative shoots—

two species of Bicis Pistillate inflorescenc ces—
s. Le

laris, an approximately three-year-old | plant raised from a cutting, ribet for de first time; inflorescences,
at least one-year-old parts of the main shoots. X—plant t rained

oo from this point. * —pistillate inflorescences formed here in the
resent year, two pistillate shoots form

mingiana, showing (left to right) shoot of p

pet for ien first ES a two- dione pistillate shoot, and a (probably) three-year-old staminate shoot. Note

acrotonous ramifications, and ‘
influenced bo different yes of shadin

to one of the architectural models of Hallé et al.,
although some species have been so assigned by
Castro dos Santos (1981). This is because the
“opportunistic” growth of these lianas with eas-
ily formed accessory shoots makes it difficult to

plagiotropic
growth. In F. cumingiana, inflorescences are al-
ways terminal on the main shoot as well as on
the lateral shoots, so that branching in the floral
region is monopodial, except for occasional veg-
etative shoots. In F. funicularis, the inflores-
cences are lateral on short shoots.

When the staminate and pistillate inflores-
cences were produced simultaneously, hand-pol-
lination of the latter resulted in fruit formation;
in F. funicularis seeds from such fruits germi-
nated to produce viable seedlings. In F. cumin-
giana, attempts to pollinate by hand were un-
successful in both Hamburg and Bonn Botanic
Gardens. Although the pistillate heads enlarged
considerably and were not aborted, fertilization
apparently did not occur; at least the pollen was

a distribution of inflorescences and vegetative shoots probably

able to trigger development. It may be noted that
in the related genus Pandanus, no pollen stim-
ulus is necessary, at least in some species, and
parthenocarpy is normal. It must be remembered
that attempts to pollinate tropical plants in tem-
perate-zone greenhouses are chancy, for even if
the basic aspects of the breeding system and the
receptive period of the stigmas are understood,
it may be impossible to obtain fertilization. An
example of this difficulty is Cinnamomum verum
(C. zeylanicum), the true cinnamon, as recounted
by Kurz (pers. comm.). On the other hand, if
pollination is successful, the possibility of pseu-
dogamous agamospermy cannot be ruled out
(Gadella, 1983). These observations are casual
but may serve as the basis for some speculations,
if only for the purpose of stimulating further ex-
periments (which cannot be performed with
scarce material in cultivation).

The evolutionary significance of dioecy has re-

: 243-296.
1984) where a survey of the subject and further

1987]

TABLE 2. Branch diameter and sex expression in
r: wasa cumingian
was Visits ca. 3 cm below the inflores-
cence d iiie the expansion immediately below it).
The pistillate shoot measure is that shown in Figure 2.
e respective means are m, = 3.98 + 8 mm for
pistillate inflorescences, m, = 3.08 + 0.35 mm for
staminate inflorescences on pistillate shoots, and m, —
2.76 + 0.26 mm for staminate inflorescences on sta-
minate shoots. Branch diameter for pistillate inflores-
cences differs significantly from that for staminate in-
florescences.

aminate
Pistillate Inflorescences
Branch Diam. Inflores- 9 3
(in mm) cences Shoots Shoots
1.6-2.0 l
2.1-2.5 l 4
2.6-3.0 l 4 11
3.1-3.5 2 2 3
3.6-4.0 12 2
4.1-4.5 5 l
4.6-5.0 4

references may be found. Discussion centers
around the question of whether the so-called
benefits of dioecy (Willson, 1983) are EP PPE
or ecologically determined, i.e., whether the
motion of outcrossing or ecological aera
on reproduction are evolutionarily decisive. This
matter need not be resolved here, but our con-
cern is with occasional departures from dioecy,
a phenomenon that has been termed “leaky dioe-
cy” by Baker & Cox (1984). Such leaky dioecy
may be considered as one prerequisite for suc-
cessful colonization of remote islands, since
propagation by selfing becomes possible. Apo-
mixis is another kind of solution to this problem,
as is the parthenogenesis observed in some in-
sular animal populations (e.g., Cox in White,
1985; Mau, 8).

Rohwer & Kubitzki (1984) recently reported
an interesting example of leaky dioecy in the
neotropical riverine willow Salix martiana Leyb.,
which was found to produce regularly mixed cat-
kins with staminate, pistillate, and perfect flow-
ers. This was attributed to the difficulty of pro-
ducing offspring after a probable long-distance
dispersal and in an unstable environment. Mixe
catkins are not uncommon in Salix (Velenow-
sky, 1904; Toepffer, 1925) but the high incidence
in a tropical species, certainly much less studied
than its temperate congeners, suggests that here

POPPENDIECK — FREYCINETIA

317

FIGURE 2. Diagrammatic sketch of sex expression
on an initially pistillate shoot in two successive years.
oken line marks th

note that due to the straggling habit and the necessity
of training ui climber, some shoots may have been
removed. e further Ting the PIE inflores-

ences (open dise formed on the distal parts of the
Muse 1986.

it is a more or less regular phenomenon. Al-
though hermaphroditic flowers are unknown in
Freycinetia, the two genera may be taken as par-
allel cases for the evolution of monoecy (which
may be a reversal of a previous evolutionary
trend from monoecy to dioecy). Another aspect
of this possible parallelism will be discussed next.

Developmental plasticity is one of the char-
acteristics of liana species, as exemplified by their
*opportunistic" shoot architecture in compari-
son with trees (Etifier, 1981). This plasticity is
needed to adjust to a temporarily very hetero-
eneous habitat, where, for instance, light inten-
sity encountered by juvenile plants is drastically
different than that encountered by adults. In

go

ANNALS OF THE MISSOURI BOTANICAL GARDEN

\

FIGURE 3. Aspects of growth of two Freycinetia species. a, b. Freycinetia funicularis. —a. Staminate iud
rescence on a vertical shoot, ca. 18 cm diam.— b. Pistillate piece at anthesis (spikes 8 cm long). c

1987]

Freycinetia, at least in the two species observed
for this study, such plasticity extends to sex
expression. This is not surprising since environ-
mentally determined sex expression has several
selective advantages for species living in a
“patchy” environment (Charnov & Bull, 1977;
Willson, 1983), and sequential hermaphroditism
has been reported for many species. The factors
invoked include light intensity (e.g., Catasetum;
Dodson, 1962), hormonal and nutritional status
(e.g., Elaeis guineensis, Williams & Thomas,
1970), or disturbance of vegetative parts (e.g.,
Arisaema: Schaffner, 1921), all of which may
interact variously, and all of which may be ap-
plicable to the present cases. Thus, in F. funi-
cularis, horizontal training of the branches may
have altered the hormonal balance toward the
production of pistillate inflorescences; exhaus-
tion of nutrient reserves, self-shading of shoots,
or pruning may have been responsible for the
switch back to staminate inflorescence produc-
tion. Experimental proof for this would be dif-
ficult to obtain. At least, the : size- -correlated sex
expression in plants of F. will permit
further study. Cuttings have been made from the
differently determined axes and their fate will be
observed in coming years.

Clearly it should not be too difficult to find
plausible explanations for the adaptive values of
dioecy, monoecy, or plasticity in sex expression
in a given situation. However, the phenomenon
of leaky dioecy is still puzzling. Why is it re-
stricted to certain genera and absent in others?

hy, for instance, are no such cases known in
Populus or in Pandanus, which are the sister gen-
era of Salix and Freycinetia, respectively? In
Populus and Pandanus pollination is believed to
be chiefly by wind (Toepffer, 1925; Cox, 1982).
A possible explanation is that in anemophilous
plants, the staminate and pistillate functions must
suit quite different needs and, as a result, tend
to evolve divergently. Such divergence seems
improbable in zoophilous groups, where a high
degree of similarity is advantageous (in overall
form, size, color, odor, and perianth details,
though not in the details of staminal and gynoeci-
al form). The correlation of leaky dioecy with

POPPENDIECK —

FREYCINETIA 319

change from anemophily to zoophily is a pos-
sibility worth investigating.

To obtain evidence bearing on these questions,
many more observations are needed, preferably
on the same specimens over a number of years.
The statement by Cox et al. (1984) about the
value of living collections for this kind of study
may be emphasized again. Phenomena such as
these tend to escape the notice of both purely
field- and herbarium-oriented workers, but bo-
tanic gardens provide an excellent location in
which to study such things as “leaky dioecy."

VOUCHER MATERIALS

Herbarium material of both Freycinetia species
discussed is deposited in PH and HBG, and the
behavior of F. funicularis is also documented in
M (Bogner 1783). Additional prints for photo-
graphic documentation are on deposit in PH and
HBG

LITERATURE CITED

BAKER, H. G. & P. Cox. 1984. Further thoughts =
dioecism and islands. Ann. Missouri Bot. Gar
71: 230-239.

CASTRO DOS SANTOS, A. DE. 1981. L’appareil végétatif
des monocotyledons. Thèse, Acad. de Montpel-
lier, Univ. des Sciences et Techniques de Langue-
doc.

CHARNOV, E. L. & J. BULL. 1977. hen is sex en-
vironmentally determined? Nature (London) 266:
—830.
Cox, P. A. 1981. Bisexuality in the Pandanaceae: new
findings in the genus Freycinetia. Biotropica 13:
8.

. Vertebrate pollination and the main-
tenance of dioecism in the genus Freycinetia (Pan-
danaceae). — Naturalist 120: 65-80.

, B. WALLACE & I. BAKER. 1984. oecism

in the genus Fre reycinetia peat an L im
16: 313-314.

DAHLGREN, R. T. CLIFFORD & P. F. Yeo. 1985.
The Families of the Monocotyledons. Springer-
Verlag, Berlin, Heidelberg, aid
Dopson, C. H. 19 Pollination ie pa ariation in
the subtribe Catasetinae (Onehidaccae). pues Mis-
souri Bot. Gard. 49: 35-56.

ENcKE, F. 1958. Pareys Blumengartnerei. 2. Auflage.
Paul Parey Verlag, Berlin und Hamburg. (See p.
101.

ETIFIER, M. E. 1981. Données sur la strategie de

—
Freycinetia cumingiana. —c. Staminate inflorescence ca. 6 cm diam.— d. Pistillate inflorescence after anthesis,

spikes 1.5-2 cm long.—e. Bas
shoot with caducous bra

thinner).

al parts of shoots (from left to right, two old pistillate
acts, sais pistillate shoots interspaced with two staminate shoots, the latter much
—f. F. funicularis seedlings ca. six months old, their leaves about 2 cm long.

shoots, current pistillate

320
croissance de quelques lianes tropicales. Thése,
Acad. de Montpellier, Univ. des Sciences et Tech-

I 1983. Some notes on the deter-

mination of the mode of reproduction in higher

plants. Proc. Koninkl. Nederl. Akad. Wetensch.

Fowering Plants of the World.

Oxford Univ. Press, Oxford, London, Melbourne
MAU, K. G. 1978. Nachweis natürlicher Partheno-
gense bei Lepi

RETR acer (R ig Sauria, Gekkoni-
) mandra
ROHWER, KUBITZKI. nus Salix martiana, a

K.
regularly hermaphrodite willow. Pl. Syst. Evol. 144:

HAMM. J. H. 1921. Control of the sexual state
in Á š Pea 4 s d 45 A gs

Amer. J. Bot. 9: 72-78

SrowE, B. C. 1972. Materials for a monograph of
Freycinetia Gaud. (Pandanaceae). XV. The Su-
matran species. Federation Mus. J. n.s. (for 1970)
15: 203-207.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

1984. A guide to collecting Pandanaceae
(Pandanus, Freycinetia, and b raranga). Ann.
Missouri Bot. Gard. 70: 137-

TOEPFFER, A. 1925. Salicaceae. = 373- 377 in O. v.
Kirchner, E. Loew & C. Schroeter (editors), Le-
biu ie EA de Blütenpflanzen Mittel-Euro-
pas. Band II, 1. E. Ulmer, Stuttgart.

VELENOWSKY, J. Tur Vergleichende Studien über
die Salix-Blüte. Beih. Bot. Centralbl. 17: 123-128.

WARBURG, O. 1900. Pandanaceae. Pflanzenreich IV.9:
1-99. (Heft III.

WHITE, J. (editor).

1985. Plant Demography. Aca-

of Pandanus tectorius in Polyne
WILLIAMS, C. N . L. THOMAS. 1970. Observa-
tions on sex differentiation i in the oil palm, Elaeis
guin eue L. Ann. Bot. 34: 957-963
WILLSON, M. . Plant E ass Ecology.
John "is & Sons, New York. (See p. 79.)

A GUIDE TO COLLECTING LECYTHIDACEAE!

Scott A. MORI AND GHILLEAN T. PRANCE?

ABSTRACT

Because of their arborescent habit and unique flowers and fruits, Lecythidaceae present specialized
collection problems. Instructions for the preparation of more informative collections of Lecythidaceae
vided.

for use in taxonomy are pro

The Lecythidaceae sensu lato are a pantropical
family of small to very large trees. The family
includes four subfamilies: Planchonioideae with
55 species in six caps distributed through trop-
ical Asia, Malaysia, northern Australia, and the
Pacific Islands; Fie. with five species
in a single genus distributed in Madagascar, In-
dia, and Malaysia; Napoleonaeoideae, with 11
species in two genera distributed in West Africa
and one species in the upper Rio Negro of Ama-
zonia; and the Lecythidoideae, with about 206
species in ten genera distributed through tropical
merica from Veracruz, Mexico to southern
Brazil (Kowal et al., 1977; Prance & Mori, 1979).
Because our collecting experience has been most-
ly in the Neotropics, this essay emphasizes the
Lecythidoideae. However, the methods de-
scribed are applicable not only to other subfam-
ilies of Lecythidaceae but to tropical trees in gen-

ral.

General collections of plants provide the data
upon which the monographic and floristic treat-
ments of plant taxonomists are based. A single
collection of a species usually does not provide
sufficient data for its taxonomic description.
Consequently, collections are needed that rep-
resent: (1) all parts of the plant used in classifi-

stages of the life cycle of the plant (flowers, fruits,
seeds, and seedlings), (3) individuals from
tinct habitats in which the plant grows, (3 in-
dividuals from throughout the geographic range
of the species, and (5) intra- and interindividual
variation.

Many recent collections of neotropical Lecy-
thidaceae have added little to what was already

about those features of the plant, such as habit

and bark, that are not preserved on the herbar-
ium sheet. Careful notes on habitat are usually
lacking and even flower color is often not re-
corded in a manner that can be interpreted by
taxonomists. Worst of all, flowers are often dried
under such extreme heat and pressure that they
become so carbonized that their structure is im-
possible to determine. Even fruits are collected
in such a way that their taxonomic features and
morphological variation are difficult to interpret.
Finally, few collectigns or weir and fruits are
from

ibe-s same ie.

Another problem is the continued collection
of herbarium material of the same taxa from the
same area by the same collectors. However, some
duplication is necessary to voucher ecological
studies of tropical forest structure and compo-
sition. Collections of some Lecythidaceae, such
as Gustavia augusta L. and G. hexapetala (Au-
blet) J. E. Smith, continue to late without
augmenting the information needed for under-
standing the species. These collections do not
merit the cost of their processing or the valuable
space they occupy in herbaria. This same lack of
selectivity in collecting occurs in almost all woody
tropical families, for example, the Chrysobala-
naceae (Prance & Campbell, in press).

If further progress is to be made in the tax-
onomy of tropical woody groups, more careful
and selective collections by more sophisticated
collectors are needed. The purpose of this article
is to provide the information needed for making
these collections. While we emphasize Lecythi-
daceae, similar needs pertain to almost all rain-
forest woody plant families.

TREE CLIMBING

Because of their large size, many species of
Lecythidaceae can be collected only by felling or

! We are grateful to Bobbi Angell, Bethia Brehmer, and H. M. Fukuda for the preparation of the original

s wks and to Carol Gracie for rearr

anging Figure 4 and for her comments on the manuscript.

The New York Botanical Garden, Bronx, New York 10458-5126, U.S.A.

ANN. Missouni Bor. GARD. 74: 321-330. 1987.

322

40cm

FIGURE |. war el pipes pet to collect Lec-
ythidaceae. — A. tree im spikes
(*griffes"). — B se s En used b. native climbers
("*peconha"). Reprinted with ission fr e

m
Memoirs of the New York Botanical Garden

climbing. As a general rule, trees should be felled

earing,
struction, etc. Forest clearing provides an excel-
lent opportunity for making collespons dom
felled trees. Thet

from a herbarium specimen often does not war-
rant the sacrifice of a tree. This is especially true
when collecting near Indian villages or settle-
ments where trees may have economic value or
cultural significance to natives. Boom (1985) has
shown that 82% of the species of trees surround-
ing a Chácobo Indian village in Bolivia are used
in one way or another by the Indians.

We have found climbing to be the most sat-
isfactory method of obtaining specimens of Lecy-
thidaceae. Although numerous methods are used
for gaining access to tropical trees, we have em-
ployed (1) native climbers, (2) “Swiss Tree Grip-
pers" (Mori, 1984; available from Forestry Sup-
pliers, Inc., 205 West Rankin St., P.O. Box 8397,
Jackson, Mississippi 39204-0397), and (3) French
tree SBEHDEDR spikes (Fig. 1).

N becoming i InCI easingly dif:

ficult to find. We have employed them most ef-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

fectively to collect Lecythidaceae in Brazil, es-

still common because of the custom of gathering
edible fruits from Euterpe and Oenocarpus palms.
The Brazilian climbers can efficiently scale trees
under 35 cm DBH by using an adjustable canvas
loop called a “‘peconha” (Fig. 1). A climber can
be hired for about $10 per day.

For trees from 50 to 72 cm DBH we have used
“Swiss Tree Grippers.” To use them, lianas and
epiphytes have to be cleared from the trunks
before the trees can be ascended, but the trees
themselves, in contrast to those scaled with spikes,
are not damaged. The weight, bulk, and high
price of the grippers often counterbalances the
advantage of being able to climb the few addi-
tional trees not climbable with spikes.

e have found French climbing spikes (Fig.
1) to be the most efficient method for collecting
Lecythidaceae. Smaller spikes (24 cm diam.) are
used for trees from 10 to 25 cm DBH, whereas
larger ones (35 cm diam.) are employed for trees
from 26 to 50 cm DBH. The advantage of these
spikes over the single spur spike preferred by
some climbers is the ease with which trees can
be climbed. The climber's weight and safety belt
secures him to the tree in such a fashion that his
hands are free for using clipper poles, capturing
insects for pollination studies, etc. The advan-
tage of the single spike apparatus is that trees of
all sizes can be ascended with one pair of spikes.
The French spikes are available from Ets La-
coste, 24160 Excideuil, France for about $50 a
pair, not including shippi

All climbs should be ads with the climber
secured to the tree with a safety belt. We prefer
the Klein nylon protective belt with two D rings
in combination with the Klein nylon adjustable
lanyard (both available from Forestry Suppliers,
Inc.). The use of two lanyards insures that the
climber is attached to the tree at all times. When
the climber reaches a branch, one lanyard re-
mains around the trunk below the branch while
the other is secured around the trunk above the
branch to be passed. The belt around the main
trunk beneath the branch can then be released
and the climber is free to move above the branch.

We have climbed several thousand trees using
the French spikes without any falls. Neverthe-
less, the climber should be extremely careful be-
fore and during any climb. Before climbing, the
tree should be carefully inspected to insure that
it is in good enough condition to support the
weight of the climber, and for the presence of

1987]

stinging and biting animals. Special attention
should be paid to swarms of bees, as they may
be made up of the particularly aggressive African
honey bee, which has been responsible for the
death of at least one biologist. The bases of trop-
ical trees often serve to house creatures that vary
from sand flies that carry the protozoan disease
of leishmaniasis to snakes that can inflict painful
and deadly bites. The tree itself may be occupied
by bees, wasps, biting ants, and the deadly snake
Bothrops bilineata (an arboreal fer-de-lance). The
last is more frequent in fruiting trees, where it
lies in wait to prey upon visiting frugivores. Ex-
treme care should be used when reaching over
branches or when climbing past epiphytes. Near-
by trees should also be examined so that wasp
and bee nests are not disturbed during manip-
ulation of the clipper pole. Finally, the condition
of the climbing gear should be constantly mon-
itored and any frayed or defective parts should
be immediately replaced. The recent fatal fall of
a young botanist in Venezuela is a shocking re-
minder of the danger involved in making tropical
tree collections

Collection m specimens from the tree's canopy
by a collector on the ground or in the tree itself
is greatly facilitated by the use of a tree pruner
attached to a series of aluminum poles. We have
found the poles developed by botanists of the
Missouri Botanical Garden staff to be the most
efficient. This system includes three sets of two
telescoping aluminum poles attached to one
another by a spring activated button that passes
from a hole in one pole into that of the next.
Each pole is 1.8 m long so that when all six poles
are joined nearly 11 meters plus the height ofthe
collector can be attained. However, when more
than four poles are joined it is very easy to bend
the poles. In order to avoid this, it is best to cut
with the poles in the most vertical position pos-
sible and to wrap the cutting rope around the
poles several times in order to insure that the
cutting force is directed along the poles. We find
the best tree pruner to be the Snap-cut model 33
(available from Forestry Suppliers, Inc.) which
fits exactly onto the end of one of the smaller
aluminum poles. This head is light enough to
allow maneuverability yet strong enough to cut
most branches to about 4 cm in diameter.

LEAF VARIATION

Normal collection for herbaria does not ade-
quately sample intra- and interindividual vari-
ation in leaf morphology. Collections which in-

MORI & PRANCE—LECYTHIDACEAE COLLECTING

323

clude shade and sun leaves, young and old leaves,
and extremes in leaf size are invaluable in helping
to solve taxonomic problems. In Couroupita, for
example, there was much taxonomic confusion
due to differences in leaves from the crown ver-
sus those that appear on the cauliflorous inflo-
rescence. These leaves differ in shape, texture,
and pilosity and were sometimes described as
different species, though they in fact represent
variation within the same species. Consequently,
it is important to document leaf variation, even
if more than one herbarium sheet is needed to
do so.

FLOWERS AND FRUITS

The remarkable variation in the floral and fruit
morphology of Lecythidaceae can often be re-
lated to pollinators and seed dispersers. The two
principal floral types in neotropical Lecythida-
ceae differ in the structure of the androecium.
One type is radially symmetrical (actinomor-
phic) and the other is bilaterally symmetrical (zy-
gomorphic, Fig. 2). The actinomorphic androe-
cium consists of a fused basal portion, the
staminal ring, which is surmounted by free sta-
mens (Fig. 2). In the zygomorphic androecium,
the staminal ring is prolonged on one side into
a stamen-free area called the ligule, which ter-
minates in an appendage-bearing hood (Fig. 2).
The hood may be open (with a space between it
and the summit of the ovary) or closed and tight-
ly appressed to the summit of the ovary. New
World genera with the former type of androe-
cium are Asteranthos, Gustavia, Grias, and Al-
antoma, whereas the latter type is found in
Couroupita, Corythophora, Bertholletia, Lecy-
this, Eschweilera, and Couratari (Fig. 3). The
flowers of Cariniana are intermediate between
these two types (Fig. 3

Pollinator d d ]
the androecium. In gotiinvanmanephin species Jd
reward is always pollen. Moreover, the pollen
collected by pollinators from ic onis i
flowers is the same kind of pollen that affects
fertilization, i.e., no nectar or specialized pollen
are produced. In the zygomorphic species either
pollen or nectar may serve as the pollinator re-
ward oevet, His pellen souaga by polli-

—

morphologically (e. g., Couroupita guianensis
Aublet: Mori & Boeke, 1987; Mori et al., 1980),
from “g which affects pollination. This spe-
cialized or fodder pollen is found in the anthers
ofthe hood or in a group of stamens of the stam-

324

ANTERIOR

DISTAL

POSTERIOR

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Staminal Ring

Ligule

[VoL. 74

PROXIMAL

Staminal Ring

ANG
IS W

IGURE 2. Androecial structure of a zygomorphic-flowered New World Lecythidaceae. — A. Side view of
flower with petals removed, showing terminology used in describing floral orientation and in describing the
androecium. — B. Medial section of the androecium.— C. Artificially opened androecium.— D. Stamen from
staminal ring. Reprinted with permission from Flora Neotropica.

inal ring adjacent to the ligule. Color differences
help to separate the two types of pollen. For ex-
ample, in Lecythis pisonis Cambess. the fodder
pollen, located in the hood, turns black after 24
hours, whereas the pollen in the staminal ring
remains yellow. In L. corrugata Poit. subsp. cor-
rugata the fodder pollen, located in a row of
stamens on the ligular side of the staminal ring,

is yellow in contrast to the white pollen of the
remainder of the stamens. Nectar is the principal
pollinator reward in those zygomorphic species
with coiled androecial hoods.

The aforementioned androecial structures as
well as other floral characteristics provide fea-
tures useful in the classification of Lecythidaceae.
Unfortunately, most collections do not ade-

1987] MORI & PRANCE—LECYTHIDACEAE COLLECTING 325

GUSTAVIA
HEXAPETALA

CARINIANA
PAUCIRAMOSA

outil FDE

LECYTHIS

COUROUPITA PISONIS

GUIANENSIS

=
ANANA
PRIDE EIE
BERR

LECYTHIS
TUYRANA

BERTHOLLETIA
EXCELSA

' var I f:
: rU ESCHWEILERA ESCHWEILERA COURATAR
1 COLLINA LONGIPES STELLATA

FIGURE 3. Variation in androecial structure of New World Lecythidaceae. In all cases the androecium has
been removed from the flower and cut in medial section. Gustavia, Grias, and Allantoma are actinomorphic,
whereas the remaining genera display varying degrees of zygomorphy. Reprinted with permission from Flora
Neotropica.

326

quately preserve or describe these features. The
most common error in the collection of flowers
of Lecythidaceae is the use of excessive heat and
pressure in the drying process. It is useful to dry
at least some of the flowers outside the press in
a paper bag over low heat and to preserve flowers
in F.A.A. or 70% ethanol. Structure of the an-
droecial hood can also be recorded by photo-
graphing a medial section of it. The photographs
should subsequently be attached to the herbar-
ium sheet.

Flower color and size are also useful features
that have not been adequately considered in the
classification of Lecythidaceae because they are
seldom properly recorded. After drying, it is very
difficult to determine flower size, and therefore
the maximum diameters of several flowers from
the same tree should be measured and stated on
the label. It is also important to record the color
of all the separate floral parts. It is not sufficient
to state that the flowers are white, yellow, or
some other color. If indeed they are entirely of
a single color, this is unusual and should be clear-
ly indicated. In many species of Lecythidaceae,
the petals and androecial hood are different colors
and this should be noted. In addition, the color
and position of differently colored anthers should
be recorded, as this indicates whether one or two
types of pollen are produced within the same
flower.

The fruits and seeds of neotropical Lecythi-
daceae have evolved a variety of forms in re-
sponse to pressures exerted by different seed dis-
persal agents, fruit and seed predators, and
environmental constraints. Major trends caused
by these selective pressures are (1) indehiscence
versus dehiscence; (2) retention of the fruit on
the tree until the seeds have been released versus
drop of fruits with the seeds inside; (3) fleshy
versus woody pericarps; (4) development of mu-
cilage within the pericarp; (5) lateral versus basal
fleshy arils; (6) development of arils that com-
pletely surround the seeds; (7) loss of arils; (8)
development of membranous, winglike arils; (9)
development of plano-convex cotyledons as in
Gustavia; (10) development of leaflike cotyle-

ons as in Cariniana, Couratari, and Courou-
pita; (11) lack of well-developed cotyledons as
in Allantoma, Bertholletia, Corythophora, Esch-
weilera, Grias, and Lecythis; and (12) terminal
versus lateral seed germination. These characters
have coevolved with animal and wind seed dis-
persers and environmental factors throughout a
long history under rainforest conditions (Prance

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

& Mori, 1978, 1979, 1983; Mori & Prance, 1981).
The terminology used to describe the fruits and
seeds of Lecythidaceae is illustrated in Figure 4.

A particularly difficult problem in the taxon-
omy of Lecythidaceae has been the determina-
tion of the extent of intraspecific fruit variation.
Unfortunately, earlier monographers of the fam-
ily (Miers, 1874; Knuth, 1939) did not have a
concept of intraspecific fruit variation. They sim-
ply named all fruit variations as separate species.
However, beginning with Dugand (1947), bota-
nists have become aware that the fruits of Lecy-
thidaceae can display considerable variation
within the same species (Mori & Prance, 1981).
In Allantoma, fruit morphs representing as many
as six of Miers’s species have been found on a
single tree (Prance & Mori, 1979).

Collectors can help in the description of fruit
and seed features of Lecythidaceae. They can also
gather material that adequately shows fruit vari-
ation within and between individuals. It is es-
pecially important that the developmental state
of the fruits is recorded. Under heat, young fruits
may dehisce and thereby appear to be mature.
Because it is often very difficult to open up dried
fruits, some should be cut lengthwise through the
middle and seeds removed from the fruit with
the arils and funicles intact. The features men-
tioned earlier should be recorded at the time of
collection.

It is extremely useful to have flowers and fruits

turning to the same trees to gather representative
material from all stages of the plant’s life cycle.
This is especially critical in Eschweilera, where
the similarity of the leaves of many species can
lead to falsely relating fruits of one species with
flowers of another. Collectors can also contribute
to the proper correlation of flowers and fruits by
always searching in the canopy and under flow-
ering trees for old fruits. Ifthe fruits are too rotten
to collect, a photograph should be made and sub-
sequently affixed to the herbarium collection.
However, care must be taken to avoid attributing
fruits from the ground to an incorrect tree. If any
doubt exists, the fruits should be given a separate
collection number.

Local collectors can contribute much to our
knowledge of Lecythidaceae by becoming ex-
perts on the family in their areas. On-site studies
of taxonomy, ecology, pollination biology, pop-
ulation dynamics, and ontogenetic development
in species-rich areas will add much to our knowl-

1987] MORI & PRANCE—LECYTHIDACEAE COLLECTING
proximal end pedicel scar
infracalycine
zone
sepal scor
supracalycine
zone I
. A. line of operculor
distal end dehiscence
calycine ring
operculor iie of pos
MA ring
aril
aril
C --— f unicle D E
F. 6 G H. L. é
FiGurE 4. Fruit, seed, and seedling features of New World Lecythidaceae. — A-C. Lecythis ampla Miers.

Note ic basal aril in C. —D. Eschweilera
Bonpl. Note that this species lacks an aril.—
wing entirely surrounds the seed.—G. Un

sp. S em with lateral aril. —
Winged seed of C pos stellata A. C. a Note that the
ilaterally winged seed of Cariniana micrantha D

E. Seed of Bertholletia excelsa Humb. &

e.—H. Lateral

germination of Eschweilera tenuifolia bad Miers.—I. Apical germination of E. pittieri R. Knuth. Modified

with permission from Flora Neotropic

edge of the family. These studies are needed be-
cause many features that allow recognition of
species in the field are not apparent in herbarium
specimens. For example, in French Guiana, where
the senior author has recently completed a de-
tailed study of the 27 species of Lecythidaceae
in the proposed national park surrounding Saül
(Mori & collaborators, 1987), we were able to
resolve a number of taxonomic problems that
could not be understood with herbarium mate-

rial alone. Similar studies would be especially
useful in central and western Amazonia and in
the Chocó of Colombia.

ADDITIONAL NOTES

Collectors generally do not provide enough in-
formation on habitat, habit, and bark to aid in
the identification and classification of Lecythi-
daceae. These features are valuable in separating

328

species of Lecythidaceae and should be recorded
by the collector.

Species of Lecythidaceae are primarily found
in lowland moist forests. Nevertheless, a few
species have invaded savanna habitats [e.g., Ca-
riniana rubra Gardner ex Miers and Eschweilera
nana (Berg) Miers of central Brazil and L. schom-
burgkii Berg of Roraima, Brazil], and 14 species
of Eschweilera are known to occur at elevations
above 1,000 m (Prance & Mori, 1979). Within
lowland habitats, some species of Lecythidaceae
are restricted to the periodically inundated vár-
zea habitat where they may be among the most
conspicuous elements of the vegetation [e.g., Al-
lantoma lineata (Mart. ex Berg) Miers, Couratari
oligantha A. C. Smith, C. tenuicarpa A. C. Smith,
Eschweilera ovalifolia (DC.) Niedenzu, E. par-
vifolia Mart. ex DC., and E. tenuifolia (Berg)
Miers]. However, Lecythidaceae are most di-
verse in the nonflooded, or rerra firme, habitat.
Even within ferra firme some species appear to
prefer ridge tops, others hillsides or valley bot-
toms (Mitchell & Mori, 1987). Consequently,
it is important that collectors make very careful
habitat notes, indicating the vegetation type, al-
titude, and slope from which collections are made.
It is especially useful to note whether the plant
grows near a stream or other body of water, and
if the water is white, black, or clear.

Although all species of Lecythidaceae are trees
(some may occasionally grow as shrubs), the col-
lector should be aware of and note differences in
habit. Some species (e.g., Gustavia monocaulis
Mori) are unbranched pachycauls, others [e.g.,
G. grandibracteata Croat & Mori and G. superba
(Kunth) Berg] are branched pachycauls, and oth-
ers [e.g., G. hexapetala (Aublet) J. E. Smith and
most other species] are leptocauls.

Species of Lecythidaceae are found as under-
story, canopy, and emergent trees. In order to
communicate this information, the collector
should record the height and DBH of collections
as well as note the stratum to which this and
other individuals of the species belong. Only re-
productive individuals should be considered
when determining the tree stratum in which the
species belongs. For detailed studies of Lecythi-
daceae, diameter versus height diagrams should
be prepared. The graphs will level out at the
stratum in which the species reaches reproduc-
tive maturity. In his study of Surinam forests,
Schulz (1960) has prepared such graphs for a
number of species of Lecythidaceae as well as
for species of other families.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

The base ofthe trunk should also be described.
Some species (e.g., Lecythis idatimon Aublet)
have cylindric trunks to the very base, others
(e.g., L. chartacea Berg) have basally swollen
trunks, and others (e.g., L. zabucaja Aublet) have
well-developed buttresses. The height, width, and
thickness of the largest buttress should be mea-
sured. An undescribed species from French
Guiana has pneumatophores.

An important but infrequently described fea-
ture of Lecythidaceae is the bark. There are four
types of external bark morphology in the family.
In the first, the bark is very deeply fissured (e.g.,
Bertholletia excelsa Humb. & Bonpl., Corytho-
phora rimosa W. Rodr., and Lecythis zabucaja
Aublet); in the second it is nearly smooth (e.g.,
Eschweilera collina Eyma); in the third it is more
or less smooth, but shallow vertical cracks and
lenticels may be present [e.g., Eschweilera ped-
icellata (Richard) Mori and L. corrugata Poit.];
and in the last it may be markedly scalloped or
dippled [e.g., E. micrantha (Berg) Miers and E.
apiculata (Miers) A. C. Smith]. The thickness of
the outer and inner barks as well as the color of
the inner bark should also be recorded. The bright
yellow inner bark of Lecythis poiteaui Berg or
the flesh pink inner bark of Eschweilera apiculata
are excellent aids in field identification. Heart-
wood color should also be recorded when pos-
sible.

One of the best ways to communicate habit
and external bark features is with photographs.
The photograph should include a scale and the
number of the collection and should eventually
be affixed to the herbarium sheet.

SPECIALIZED COLLECTIONS

Known chromosome counts of Lecythidaceae
are summarized in Kowal et al. (1977). Addi-
tional counts are needed for Allantoma, Carinia-
na, Corythophora, and Couratari of the New
World Lecythidoideae, for most of the Old World
genera, for Asteranthos brasiliensis Desf. (the only
member of Napoleonaeoideae in the New World),
and for species of Gustavia to determine the ex-
tent of polyploidy. Collections of buds of all sizes
should be fixed in either Farmer’s (3 ethanol: |
glacial acetic acid, v/v) or Jackson’s (4 ethanol:
2 methanol: 2 chloroform: | proprionic acid: 1
acetone, v/v). The best results have been ob-
tained with fixation in Jackson’s solution (Ko-
wal, pers. comm.). If possible, the buds should
be left on the inflorescence, as this facilitates lo-

1987]

cation of buds in the proper stage for counting.
In addition, because of their thickness, buds
should be slit open to allow penetration of the
fixative. After 12 hours of fixation the buds should
be transferred to 70% ethanol and stored in a
freezer when possible.

For anatomical studies, collections of leaves,
twigs of several sizes (newly flushing ones are
especially useful), and bark should be fixed in
F.A.A. (10 ethanol: 7 distilled water: 2 forma-
lin: 1 glacial acetic acid, v/v). They may be stored
in F.A.A. or in 7096 alcohol. The bark sample
should consist of a block ca. 2 cm square and
should include a portion of the outermost sap-
wood to insure that all layers of the bark are
included in the sample.

Wood samples may be chisled from standing
trees or cut from felled trees. In the former case,
the block should be about 10 cm square by 10
cm deep and the bark should be left attached to
the wood sample. Smaller blocks can be removed
but, because of splitting of the sample upon
drying, they are not as desirable. The wound left
by the removed block should be painted over
with a sealer to minimize infection. In the case
of felled trees, larger blocks of wood or segments
of trunk that can later be cut into specimens
about 10 cm long (parallel to the long axis of the
trunk) by 5 cm wide (tangentially) by 2 cm thick
(radially) should be collected. Specimens should

e taken from breast height and from the bole
itself rather than from large branches or from the
buttresses. The specimens should be dried over
low heat to insure that a minimal amount of
cracking occurs.

There have been no detailed studies of any
aspect of the chemistry of neotropical Lecythi-
daceae (Prance & Mori, 1987). If collecting ma-
terial for study of flavonoids, the collector should
remember that the preservatives used in tropical
fieldwork (i.e., aqueous mixtures of formalde-
hyde or don Be pentachlorophenol
in alcohol, or alcohol) may remove these and
other compounds and thus render the specimens
useless for chemical study. Therefore, if material
is collected for chemical study, it should be kept
free of field preservatives and air dried. More-
over, all field preserved specimens should have
the treatment indicated on the label so that the
specimens are not subsequently screened for
compounds soluble in the preservatives (Coradin
& Giannasi, 1980).

It is now known that bees play a major role in
the pollination of most neotropical Lecythida-

MORI & PRANCE—LECYTHIDACEAE COLLECTING

329

ceae. A recent review of pollination in neotrop-
ical members of the family is provided by Mori
& Boeke (1987). Nevertheless, in order to un-
derstand the complex interactions of bees and
Lecythidaceae, many carefully documented col-
lections of bees visiting Lecythidaceae are still
needed. Especially useful are detailed studies of
the pollination of individual species of Lecythi-
daceae. Collections should be made at all times
of the day when flowers are open. Fortunately,
most species of Lecythidaceae open their flowers
at daybreak and drop them late in the same after-
noon. However, some species either flower en-
tirely at night or open their flowers during the
night. For example, the bat-pollinated Lecythis
poiteaui Berg opens its flowers at dusk and drops
its petals and androecia at about 0300, and the
bee-pollinated Gustavia augusta L. opens its
flowers during the night and drops its petals and
androecia by late afternoon the next day. During
the day, a French Guianan individual of G. au-
gusta was visited mostly by trigonid bees, where-
as the night-flying bee Megalopta genalis visited
its flowers before daybreak. Consequently, it is
necessary to determine floral longevity before a
pollination study is begun, and once this is es-
tablished, the plant must be observed and pol-
linators collected at all times when flowers are
open. In addition, the species should be studied
and pollinators collected throughout its entire
flowering cycle. The pollinators at peak flowering
may not be the same as those visiting the plant
at the onset or the end of the flowering period.
For example, Mori & Boeke (1987) have col-
lected completely different species of Trigona
dominating the flowers of G. augusta during the
day at different times during its flowering cycle.

We have collected bees visiting Lecythidaceae
by simply climbing into the crown and waiting
by flowers until insects enter them. Random cap-
ture of all insects that approach flowers of Lecy-
thidaceae will often give erroneous ideas of the
pollinating species because many species of in-
sects, such as wasps, visit flowers of Lecythida-
ceae to prey upon other insects, not to collect
pollen or nectar. In species with zygomorphic
flowers, it is especially important to observe the
position and behavior of the bee in the flower,
since this will indicate what type of pollinator
reward the bee is after. Euglossine bees are best
captured after they have completely entered the
flower; otherwise, they often are quick to spot
the movement of the net and escape

We use the chlorocresol method described by

330

Tindale (1962) to preserve our insect collections
in the field. This method keeps the insects moist
and free of mold and allows them to be pinned
directly upon their removal from the storage con-
tainer. In this method, a teaspoonful of chloro-
cresol is placed on the bottom of a plastic con-
tainer. It is held in place by a layer of cotton or
tissue followed by a tightly fitting layer of card-
board or blotter paper. The bees are then placed
in layers separated by tissue paper. Labels with
the collection data are inserted in each layer. The
plastic container is tightly sealed so that the
moisture of the insects’ bodies keeps them sup-
ple. If the collections are too dry, a little water
may be sprinkled into the container before it is
sealed. The insects should be pinned soon after
removing from the container as they dry out rel-
atively quickly upon removal. If the container is
properly sealed, the insects will remain in good
condition for months. Chlorocresol can be pur-
chased from BioQuip Products, P.O. Box 61,
Santa Monica, California 90406.

CONCLUSIONS

complete understanding of the taxonomy
and biology of neotropical Lecythidaceae must
be based, to a large extent, on adequately pre-
pared and documented herbarium specimens. It
is not good practice to simply collect Lecythi-
daceae without giving consideration to the prop-
er preservation of the structures used in the clas-
sification of the family. Moreover, it is essential
that more detailed descriptions of structures not
preserved on herbarium sheets, such as flower
color and bark characteristics, be noted by the
collector. Better descriptions of the phenological
state of the plant and. its habitat are needed as

£

well. Sp the study of chro-

mosomes, anatomy, chemistry, and pollination

biology will add greatly to our knowledge of Lec-

ythidaceae. It is also important that local collec-

tors become more selective in the species p
a

a given locality, en

therefore, their continued collection adds no new

information to our herbaria and only causes un-

due expense and takes up valuable space in her-
aria.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

LITERATURE CITED

Boom, B. M. 1985. Amazonian Indians and the forest
environment. Nature 314: 324.

CORADIN, L. & D. E. GIANNASI. 1980. effects of
chemical preservatives on plant m edis to be
used in chemotaxonomic surveys. Taxon 29:
40

DUGAND, A. 1947. Observaciones taxonómicas sobre
M Lecythis del norte de Colombia. Caldasia 4:
1-426

TES R. 1939. Lecythidaceae. Pflanzenreich IV.
219a: 1- jt (Heft 105).
Kowa, R. R., S. A. Morr & J. A. KALLUNKI. 1977.

Chromosome numbers of Panamanian Lecythi-
daceae and their use in subfamilial classification.
Brittonia 29: 399-410.
Miers, J. 1874. On the Lecythidaceae. Trans. Linn.
Soc. London 30: 157-318.
S. A. Jep 1987. Chapter X.
collaborators, The Le-

Use ‘of “Swiss Tree Grippers” for
making botanical collections JA tropical trees. Bio-
tropica 16: 79-80.
. BOEKE. 1987. Chapter XII. Pollination.
In S. A. Mori and collaborators, The Lecythida-
ceae of a lowland neotropical forest: La Fumée
Mountain, French Guiana. Mem. New York Bot.
Gard. 44: ea
G. T. NCE. 1981. The "sapucaia" group
of PEH (Lecythidaceae) Brittonia 33: 70-80.
& co The Lecythidaceae
ofa eee d SR La Fumée Moun-
tain, French Guiana. Mem. New York Bot. Gard.
44

—— J. E. ORCHARD & G. T. Prance. 1980. In-
trafloral pollen differentiation in the New World
Lecythidaceae, subfamily Lecythidoideae. Science
209: 4 3.

PRANCE, G. T. & D. G. CAMPBELL. (In pi The
present state of tropical floristics. Tax

& S. A. Moni. 1978. salone dn ‘the fru
and seed of neotropical hie Daria
30: 21-33.

NIMES Part I. Fl.
270.

Neotrop. Monogr. 21:
& 1983. elec and distribution

of Lecythidaceae and Chrysobalanaceae. Sonderb.

Naturwiss. Ver. Hamburg 7: 163-186.

& . Chapter XIII. Future re-
search. In S. A. Mori and collaborators, The Le-
cythidaceae of a lowland oo n 2
Fumée Mountain, French Guiana. Mem
York Bot. Gard. 44: 156-163.

SCHULZ, J. P. 1960. Ecological studies on rain forest
: northern Suriname. Meded. Bot. Mus. Herb.
Rijks Univ. mie 163: 1-267.
ie, N. B. 62. The chlorocresol PTE for
field St J. Lepidop. Soc. 15: 195-1

CHROMOSOME CYTOLOGY OF OLDENBURGIA

(ASTERACEAE—

MUTISIEAE)'

PETER GOLDBLATT?

ABSTRACT

As determined from root tip mitoses, three a the four species of the endemic Cape genus Oldenburgia

hav et UIC oc

to metacentric ch romosomes. Base . In i thé genus IS MCI to be x =
is probably a paleotetraploid relict. It has no close relatives in Africa, a
arborescent South American genera of Gochnatiinae, perhaps the most primitive

related to a group o
subtribe of Mutisieae.

Oldenburgia is a distinctive and taxonomically
isolated, southern African genus of Asteraceae
tribe Mutisieae. It is currently being revised by
Pauline Bond, and this cytological study was un-
dertaken in conjunction with her work, the genus
being unknown cytologically until now. O/den-
burgia consists of four species, all of which occur
in the Cape Province of South Africa, between
Grahamstown in the east and the Tulbagh dis-
trict in the west. All species are restricted to the
nutrient-poor, sandstone-derived soils of the Cape
geological system. All species are woody and O.
grandis (syn. O. arbuscula) is a small tree. Mu-
tisieae are poorly represented in Africa, but rich-
ly developed in South America, where several
genera are arborescent.

MATERIALS AND METHODS

Counts were obtained from root tips harvested
from germinating seeds. The root tips were treat-
ed in 0.003 M hydroxyquinoline for six hours at
refrigerator temperatures, then fixed in 3:1 ab-
solute ethanol-glacial acetic acid for two to five
minutes, and stored in 7096 ethanol. The roots
were squashed in FLP orcein (Jackson, 1973)
after a six-minute hydrolysis in 10% HCl at 60°C.
Vouchers are cited in Table 1.

OBSERVATIONS

The three species examined have a diploid
number of 21 — 36 and similar karyotypes with
relatively small chromosomes, 1.5-3 um long.
The chromosomes are too small to be easily char-
acterized, but submetacentric pairs predominate
and there are a few distinct metacentrics.

and a similar karyotype of ag irre aad ric
Oldenburgia
nd it seems i jm most closely

DISCUSSION

Oldenburgia is a member of the basal tribe
Mutisieae of Asteraceae and is usually regarded
as belonging to Gochnatiinae, possibly the most
primitive of the four subtribes of Mutisieae (Ca-
brera, 1977). The genus is characterized by thick,
coriaceous, spirally arranged leaves, large radiate
capitula, and bilabiate florets with a well-devel-
oped pappus. Its affinities within Gochnatiinae
are at present obscure. It appears to have no close
relationships to other African members of the
subtribe (Bond, in prep.), which include Erythro-
cephalum, Achyrothalamus, Pasaccardoa, Di-
coma, and Ainsliaea. The last two genera lack
ray florets (unlike Oldenburgia), Erythrocepha-
lum and Achyrothalamus lack or have a reduced
pappus (well-developed in Oldenburgia). Pas-
accardoa, which comprises a few annual species,
comes closest to Oldenburgia in floral characters
but is unlike it in vegetative features. It seems
likely that O/denburgia is most closely related to
a group of largely arborescent South American
genera that includes Chimantaea, Cnicotham
nus, Pleiotaxis, Gongylolepis, and Wunderlichia,
as well as Gochnatia, which also occurs in south-
east Asia. There are no counts for any of these
genera. Counts for African genera of Gochna-
tiinae include n = 12 and 11 for Ainsliaea (Ar-
ano, 1957), and there is only a single count for
Dicoma, n = 11, for an Indian member of the
genus (Bhandari & Singh, 1977). These genera
appear distantly related to O/denburgia on cy-
tological as well as morphological grounds.

Basic chromosome number for Asteraceae is
probably x = 9 (Raven, 1975; Solbrig, 1977) and
the same number is most likely basic for Muti-

; P by grant DEB 81-19292 from the United States National Science Foundation.
? B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri

63166, USA.

ANN. MissouRi Bor. GARD. 74: 331-332. 1987.

332

TABLE 1. Chromosome numbers in Oldenburgia.
All counts are original in this paper. Localities are in
the Cape Province, South Africa.

O. grandis (Thunb.) Baillor (syn. O. arbuscula DC.):
2n = 36; cuit ivated at Kirstenbosch Botanic Gar-
dens bat Gal-
pin s.n. n. (National Botanic Gardens 984/13 in NBG).

O. papionum DC.: 2n = 36; Wangenheim farm, Raw-
sonville, rock crevices, Bond 1722 (NBG).

O. paradoxa Less.: 2n = 36; Robinsons Pass, Oute-
niqua Mts., Bond 1726 (NBG).

sieae and Gochnatiinae as well. Oldenburgia ap-
pears on present evidence to be a paleotetraploid
genus probably allied to a group of South Amer-
ican genera, mostly of the Guayana Highlands,
and geographically isolated in Africa. Its basic

romosome number is possibly the same as for
the family and tribe.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

LITERATURE CITED

ARANO, H. 1957
taxonomic considerations in the t
Japanese J. Genet. 32: 293-299.

BHANDARI, M. M. & D. M. SimcH. 1977. In IOBP
chromosome number reports LV. Taxon 26: 107-
109

. Karyotype analysis and its karyo-
ribe Mutisieae.

CABRERA, A. L. 1977. Mutisieae — systematic review.

V. H. Heywood, J. B. Harborne & B. L. Turner

(editors), The Biology and Chemistry of the Com-
positae 2: 1039-1066.

JACKSON, R. 1973. Chromosomal evolution in Hap-
lopappus gracilis: a centric transposition race.
Evolution 27: 243-256.

Raven, P. H. 1975. The bases of angiosperm phy-
logeny: cytology. Ann. Missouri Bot. Gard. 62:
724-764.

SOLBRIG, O. T. 1977. Mar aang ag evo-
lution in the family Compositae. /n V. H. He
wood, J. B. Harborn rner (editors)
The Biology and Chemistry of the Compositae

2

NOTES ON CIPURA (IRIDACEAE) IN SOUTH AND CENTRAL
AMERICA, AND A NEW SPECIES FROM VENEZUELA!

PETER GOLDBLATT? AND JAMES E. HENRICH?

ABSTRACT

Cipura is a small genus of Iridaceae — Tigridieae that is widespread in tropical South and Central

America, Mexico, and the West Indies. It

— of interior Brazil. The basic chromosome number in Cipura is x = 7
ave been established here for C. xanthomelas and C. dolre s both senate e 2n =

During preparation of treatments of Iridaceae
for three regional floras, Flora Mesoamericana,
Flora de Nicaragua, and Flora of the Venezuelan
Guayana, the genus Cipura Aublet posed several
problems in typification and delimitation of
species. As a result we studied this small genus
in some detail. Our conclusions relate to the ge-
nus as a whole and are presented in the form of
a review. A new species, C. rupicola, is described
from western Venezuela, and two more unde-
scribed species are included in a key to the genus
but are not published as new species here for
reasons given below. A complete revision of Cip-
ura is being prepared by P. Ravenna as part of
a treatment of Iridaceae for Flora Neotropica.

Cipura is one of a distinctive group of New
World Iridaceae comprising tribe Tigridieae
(Goldblatt, 1982), which is characterized by a
bulbous rootstock, plicate leaves, and a basic
chromosome number of x = 7. The first genus
in the alliance to be described (Aublet, 1775), it
was based on C. paludosa, from what is now
French Guiana. Cipura comprises five or prob-
ably a few more species, distributed from south-
ern Mexico in the north to Bolivia, southern Bra-
zil, and Paraguay in the south (Fig. 1). Cipura
paludosa occurs over almost the entire range of
the genus, but the other species have narrower
and sometimes very restricted ranges.

Cipura is closely related to the larger genus
Cypella Herbert from which it differs in having
erect inner tepals that partly conceal the stamens
and style-stigma apparatus and a large cauline

t compri
understood. The type species, C. paludosa Aublet, is often treated as including the similar C. c

ises at least five species and is at present poorly

am-

he differences between the two are detailed. A
l

hromosome

leafinserted at the flowering stem apex just below
the single or few and closely set rhipidia (spathe-
enclosed inflorescence units). As in Cypella and
several other genera of Tigridieae, the fugacious
flowers have broadly clawed outer tepals; inner
tepals with an adaxial nectariferous area on the
limb (usually concealed by a fold in the tepal
surface); free stamens with weak filaments; and
anthers adhering to the style branches.

The style divides above into three thickened
branches. In the less specialized species there are
one or two pairs of erect appendages (crests) which
exceed each transverse stigma lobe. Such elab-
orate style branches are probably basic in Cipura
and the apparently simpler structure in C. pa-
ludosa and C. campanulata, in which the style
branches are not developed and the crests are
absent or reduced, is derived.

The species of Cipura currently recognized are
C. xanthomelas Martius ex Klatt, of which C.
flava Ravenna is almost certainly a synonym;
the new C. rupicola, C. paludosa Aublet; and C.
campanulata Ravenna. In addition, there are two
undescribed species: Cipura sp. 1, a large and
violet-flowered species, some herbarium speci-
mens of which have been annotated C. formosa
Ravenna by P. Ravenna; and Cipura sp. 2, also
with violet flowers, from southwestern Venezue-
la and adjacent Colombia.

In the review we include the following: 1) typ-
ification of Cipura paludosa, which involves the
delimitation of C. campanulata, a largely Central
American species often confused with C. palu-

1 pu by Grant DEB 81-19292 from the United States National Science Foundatio

Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box

salee. T

299, St Louis, Missouri

3 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

ANN. Missouni Bor. GARD. 74: 333-340. 1987.

334

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

=

LJ 200 400 600 800 1000km
O 100 200 300 400 600 800 miles \

100

Geographical ee of the species of Cipura. The occurrence of C. paludosa in Nicaragua,

FIGURE 1.
iru El Salvador, Guat
we

dosa; 2) description of C. rupicola and a discus-
sion of its differences from the closely related C.
xanthomelas; 3) brief discussion of the two vi-
olet-flowered species; 4) key to the species; and
5) synonymy and description of C. paludosa.

TYPIFICATION AND DELIMITATION OF
IPURA PALUDOSA

Cipura paludosa, described by Aublet in 1775
from plants growing near Cayenne, is the most
widespread and common species in the genus. It
has pale bluish, or sometimes white, flowers with
a cylindric-fusiform ovary included in the spathes
and non-clawed, broad, closely imbricate inner
tepals about half as long as the outer tepals (Fig.

2A). Pen = except from above, the sta-
mens, stigma. The style branches are
hardly Í but the style is thickened
above and weakly three-lobed apically (Fig. 2A).
The stigmas are simply expanded lobes located
opposite and above the anthers which form a
tube enclosing the upper part of the style. Ob-
scure crestlike appendages extending above the
stigmas can sometimes be seen, but they are often
microscopic. The inner tepals of C. paludosa are
concave below and have a conspicuous central
zone of yellow nectariferous tissue outlined in
violet.

Identification of the type of Cipura paludosa
with living populations is not without problems.
The type collection was made by Aublet in wet

1987]

savannas at the foot of Mt. Kourou in what is
now French Guiana. The illustration accompa-
nying the description conforms best with blue-
flowered plants that are fairly widespread in the
Guianas and Brazil, which generally have short
stems, relatively broad leaves, and 2-3 flowers
in each rhipidium. The flowers are clearly illus-
trated by Aublet as having erect and imbricate
inner tepals and a style with short stigmatic lobes
(flowers are not present on the specimen that we
have designated lectotype in the Rousseau Col-
lection in the Paris Herbarium). In the type figure
the stigmatic lobes are also most unusual. They
are drawn as large, ascending and acute struc-
tures, quite unlike those in any living plants of
any species of Cipura. We assume that these
strange structures were included in error.

A species described by Kunth (1816) from
eastern Colombia, Cipura graminea, is regarded
here as conspecific with C. paludosa. The type
specimen lacks flowers but has comparatively
narrow leaves and elongate included capsules, of
which there are two in some rhipidia. The Bo-
livian species Cipura major, described by Rusby
(1910), and a plant from Peru, C. goodspeediana
(Vargas) R. Foster (1962), based on Cypella
goodspeediana Vargas (1945), with pale blue
flowers and short flowering stems 7-10 cm long,
also conform with C. paludosa. Following Foster
(1946), we regard both as synonyms of C. pa-
ludosa. Cipura major differs mainly in having
very narrow leaves, but similar leaves are known
on some specimens of C. paludosa.

Similar and closely related is a species com-
mon in Mesoamerica which almost always has
white flowers. When dried it is difficult to dis-
tinguish from the often violet- or pale blue-flow-
ered Cipura paludosa, with which it was included
or confused in the several local and national flo-
ras for Mesoamerica. Living plants that we ex-
amined display some important differences not
apparent in most dry specimens. The inner tepals
are initially more or less erect but soon become
semipatent and are held well away from the sta-
mens and style-stigma apparatus (Fig. 2B). Also,
the inner tepals lack nectariferous tissue and, at
least to the unaided eye, have no glands. In ad-
dition the Mesoamerican species almost always
has only one flower in each rhipidium (two or
three in C. paludosa), and the flower emerges
laterally from the spathes and is thus somewhat
secund. The leaves are often very narrow, and
the flowers have comparatively short pedicels 5—
7 mm long. Names in the literature that appear

GOLDBLATT & HENRICH — CIPURA

FIGURE 2. Dorsal views of the flowers (x2) and
detail of the stamens and style from the side (x 6), of
Cipura paludosa (A), and C. campanulata (B).

to apply to this species are C. campanulata Ra-
venna (1964), the type of which is from Yucatán,
and C. inornata Ravenna (1984) from Caracas,
Venezuela. We have been unable to see the types
of either of the species described by Ravenna.
Holotypes are in the private herbarium of P.
Ravenna who is unwilling to loan them to us.
An isotype of C. campanulata, cited as at the
Kew Herbarium (Ravenna, 1964), has not yet
been received there.

From the description, Cipura inornata appears
to differ hardly at all from C. campanulata and
is reduced to synonymy here. Ravenna distin-
guished it, although not expressly, by the absence
of style appendages but these are also lacking in
C. campanulata. Ravenna (1964) explicitly dis-
tinguished C. campanulata from C. paludosa by

336

its cernuous (i.e., secund) rather than erect flow-
ers and by the inner tepals being nearly equal to
the outer, his measurements being 19 mm long
for the inner tepals and 21 mm for the outer.
We have four living collections of the white-
flowered Mesoamerican Cipura and thus are able
to assess, to some extent at least, its floral vari-
ation. The size, shape, and orientation of the
inner tepals are most like the outer tepals of any
of the species of Cipura and are about three-
quarters as long (Fig. 2B), thus usually shorter
by some 4-5 mm and are reasonably close to the
dimensions given by Ravenna for C. campanu-
lata. Moreover, the flowers are nearly always dis-
placed to one side because they emerge laterally
from the spathes rather than apically. Thus, there
seems little to distinguish this plant from C. cam-
panulata and, despite not having seen the type
material, we feel that the name applies to the
widespread Mesoamerican species. We must note,
however, that several specimens from Yucatan
that we have seen are somewhat more robust and
have broad, strongly plicate leaves that are dis-
tinctly striated when dry. Nevertheless, we prefer
not to recognize this form as a distinct taxon.
Cipura cubensis Griseb., based on Wright s.n.,
anno 1865, is also white- flowered (Grisebach,
1866) andi d
C. paludosa. We have been unable to locate the
type, and it is impossible to tell from the pro-
tologue whether C. cubensis corresponds better
with C. campanulata or C. paludosa, the pres-
ence of only the latter being established in Cuba.
The name C. cubensis must for the present be
excluded. We note, incidentally, that C. paludosa
sensu Grisebach which is based on the cited col-
lection, Wright 3256, is a species of Cypella.
Cipura paludosa and C. campanulata are
probably the most specialized and least repre-
sentative species of Cipura. They are distinct in
their linear to fusiform ovaries, included in the
rhipidia, and inner tepals lacking distinct claws.
Both species are autogamous, unlike other mem-
bers of Cipura, and have broad inner tepals, style
branches represented by short stigmatic lobes,
and anthers coherent around the thickened upper
part of the style. The thickened part of the style
probably represents fused style branches, which
are present and free in other species of Cipura.
Cipura paludosa is apparently a diploid species
(Goldblatt, 1982), 2n — 14, although structural
heterozygosity and derived numbers of 13 and
12 have been recorded in plants of horticultural
origin. The only count for C. campanulata is

das separate from

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

tetraploid, 2” = 28 (reported by Goldblatt in
1982 as C. paludosa).

Cipura paludosa is the most widespread species
in the genus, extending from northern Bolivia in
the south to the Caribbean, Mesoamerica, and
the West Indies in the north (Fig. 1). It varies in
vegetative features. The stem is often quite short,
reaching less than 8 cm from the ground, but in
some plants may be up to 15 cm long. The leaves
are usually relatively broad, 4-10 mm wide, but
they may occasionally be only 1-2 mm wide. Its
most distinctive features seem to be the number
of flowers (and hence capsules) produced in each
rhipidium, usually two or three, rarely more or
fewer, and the closely imbricate inner tepals with
conspicuous nectar guides.

In contrast, Cipura campanulata has pure white
or rarely pale blue flowers without yellow nectar
guides, and the inner tepals are initially erect
(parallel to the axis of the flower) but later be-
come somewhat outcurving. The style branches
and stamens are similar to those of C. paludosa
but always. lack appendages. Cipura campanu-
lata seasonally wet or marshy sites from
Nayarit and Vera Cruz, Mexico through Meso-
america to northern Venezuela and Colombia
(Fig. 1).

YELLOW-FLOWERED SPECIES

A yellow-flowered species of Cipura, C. xan-
thomelas, was described by Klatt in 1882, based
on collections made by Martius and Regnell in
Brazil. It is characterized by globose, exserted
capsules and yellow flowers. The relatively large
yellow flowers have outer tepals 30-35 mm long.
The inner tepals are about half as long as the
outer, ca. 14 mm long, and have distinct claws
ca. 8 mm long with red striations on the inner
surface and dark markings on the knee and apex.
The style branches are long and have well-de-
veloped crests ca. 2.8 mm long. Cipura flava
(Ravenna, 1964) appears to be conspecific and
pending examination of the type, deposited in
the private herbarium of P. Ravenna, we are
provisionally regarding it as a synonym. Bulbs
of C. xanthomelas are quite large, usually 2-2.5
cm in diameter, with blackish resinous tunics,
the spathes are (2.5-)3-5 cm long, and the nearly
globose capsules are exserted and typically 12-
14 mm long. The species is known to us from
numerous herbarium specimens mostly with
poorly preserved flowers and from a living col-
lection grown at the Missouri Botanical Garden.
The diploid chromosome number in this collec-

1987]

tion (Plowman et al. 9306, MO) is 2n = 28, and
the species thus appears to be tetraploid. Details
of the karyotype of this and the following species
will be reported elsewhere. The flowers of our
living collection appeared to differ significantly
from other material in having partly united fil-
aments, a feature that we have not seen in other
specimens. Cipura xanthomelas is centered in
the Brazilian state of Goias (Fig. 1) and extends
into the neighboring states of Mato Grosso, Mi-
nas Gerais, Maranhao, and possibly Piaui, while
an unusual variant is recorded from western Per-
nambuco, adjacent to Piaui.

A similar yellow-flowered species, here de-
scribed as Cipura rupicola, has come to our at-
tention from Venezuela. It has pale yellow flow-
ers with unusual inner tepals having slender claws,
between which the anthers can be seen, but the
long style branches are partly concealed by the
erect tepal limbs (Fig. 3). Cipura rupicola differs
from C. xanthomelas in being less robust, in its
non-resinous bulbs 1.5-2.5 cm in diameter, and
shorter spathes 2.6-3.6 cm long. The flowers are
also smaller and lack the markings on the inner
tepals and the red striations on the claw char-
acteristic of C. xanthomelas. The outer tepals
are 25-28 mm long and the inner tepals have
claws ca. 9 mm long and limbs ca. 8 mm long.
The style branches lack crests. Like C. xantho-
melas, C. rupicola has exserted capsules, globose
in shape and usually 8-11 mm long, thus some-
what smaller than its relative. Chromosome
number in C. rupicola is 2n — 28 and it thus
appears to be tetraploid like C. xanthomelas.

Cipura "n Goldblatt & —L sp. nov.
TYPE: Venezuela. T.F.A., Dept. Atures: near
Puerto D Davidse 4 Miller 26437
(holotype, MO; isotypes, COL, K, NY,
VEN). Figure 3

Plantae 12-40 cm altae, tunicis nigribus non resi-
nosis, floribus flavis, tepalis exterioribus 25-28 mm
longis, interioribus ca. vies = unguibus angustis,
filamentis liberis 3 mm lon ntheris mm longis,
ramis styli 5 mm a sine ae capsulis obovoideis
8-11 mm longis exsertis

Plants 12-40 cm tall. Bulb 1.5—2.5 cm diam.;
tunics dark brown to blackish, brittle-papery but
not noticeably resinous. Leaves 3-5, all but one
basal, strongly plicate, narrowly lanceolate, about
as long to slightly longer than the flowering stem,
(6-)9-14 mm wide; subterminal cauline leaf sol-
itary (second smaller leaf occasionally present),
smaller than the basal, 7-15 cm long, about 12

GOLDBLATT & HENRICH — CIPURA

337

mm wide. Flowering stems 1—3 per plant, erect,
lateral to the basal leaves, 8-20(-35) cm long,
bearing 2-several crowded rhipidia either sessile
or on short branches. Inner spathes 2.6-3.5 cm
long; outer V»—^A as long. Flowers bright yellow,
eae a more or less distinct perianth tube ca. 2
m long; outer tepals 25-28 mm long, oblong,
eios slightly near the base, somewhat distally
wisted from horizontal, weakly divided into limb
and claw, the claw ascending, plane, the margins
becoming transparent, the limb horizontal, chan-
neled, ca. 20 mm long, to 10 mm wide; inner
tepals ca. 18 mm long, the claws ca. 2.5 mm
wide, narrower towards the base, ascending and
widely separated from one another and thus
forming broad windows making the anthers vis-
ible, curving inwards above to become horizon-
tal at the apex, the limbs erect, imbricate, the
apex rolled abaxially, ca. 8 mm long, ca. 8 mm
wide, bearing a small circular yellow zone at the
base, running on to the apex of the claw. Fila-
ments free, erect, thickened below, 3 mm long;
anthers ca. 3.5 mm long, free from the style,
latrorse, with a broad connective wider than the
anther lobes; pollen yellow. Ovary included in
the spathes at anthesis, ca. 4.5 mm long; style
ca. 4 mm long, dividing into 3 erect branches,
these hidden icai by the inner tepal limbs;
style branches 5 mm long, plane, truncate, with-
out apical appendages, cspanded apically into
bilobed papillose stigmas. d from
the spathes, obovoid, 8-11 mm long; seeds an-
gular, to 2 mm at the longest axis. Chromosome
number 2n = 28. Flowering time June-July.

Distribution. Venezuela, T.F.A., Dept.
Atures, in the vicinity of Puerto Ayacucho, grow-
ing on shallow soils of granite hills and outcrops
and on the surrounding dry stony savannas (Fig.

e

POES

Additional specimens examined. VENEZUELA.
TERRITORIO FEDERAL AMAZONAS: DEPT. AT

VEN); bet

de las r o Ayacucho, Williams 13099 (K,
US, VEN pris medio de las rocas, Puert
Ayacucho, Williams 12967 (F, VEN); medio y alto

m N of Puerto Ayacucho, Romero 1254 (MO)

VIOLET-FLOWERED SPECIES

There are two violet-flowered species known
to us. One, Cipura sp. 1, fairly well represented

338

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FIGURE 3. Cipura rupicola. Habit (x0. 5) lateral view of the flower (x ve and detail of the stamens and style
ein from the side including a dorsal view of the stigmatic region ( x

in some herbaria, is native to Brazil, occurring
in Bahia, Goiás, and Mato Grosso. It is easily
recognized by the unusually large bulbs with res-
inous tunics, relatively broad rigid leaves, a short
cauline leaf which is typically shorter than the
spathes, and only one or sometimes two rhipidia.
The flowers are unusual in their violet color and
large size. The outer tepals are ca. 40-45 mm

long and 30 mm wide, and the inner tepals are
ca. 22 mm long, with a long ascending claw to
ca. 18 mm long and a limb that curves forward
over the claw and then is rolled outwards distally.
The style branches are flattened and petaloid but
fairly narrow and have two pairs of crests, the
adaxial ca. 5 mm long and the abaxial about half
as long. The species is known to us only from

1987]

dry specimens and details of the inner tepals and
style branches are not completely clear.

The second violet-flowered species, Cipura sp.
2, of which we have seen only two specimens, is
from southwestern Venezuela (Huber 5711, VEN
and adjacent eastern Colombia (Vincelli 1230,
MO). It also has resinous bulb tunics, but is taller
than C. sp. 1, ca. 45 cm high, and has two to five

—

GOLDBLATT & HENRICH—CIPURA

339

rhipidia at the stem apex subtended by a cauline
leaf about three times as long as the spathes. The
flowers are much smaller than those of C. sp. 1
with outer tepals ca. 18 mm long. They are com-
parable with those of C. paludosa and C. cam-
panulata, and it is probably most closely related
to these two species. Floral details are obscure
owing to poor preservation.

KEY TO THE SPECIES OF CIPURA

la. Flowers yellow; capsules globose and exserted from the spathes.
2a. Inner tepals yellow with blackish markings on the knee and at the apex, less than half as long as
the outer tepals, to 14 mm long . xanthomelas
2b. Inner tepals entirely pale yellow; somewhat more than half as long as the outer, to 18 mm long
C. rupicola
. Flowers shades of blue to violet or white; capsules cylindric-fusiform or nearly globose; included in
the spathes
3a. Rhipidia solitary or two and subtended by a cauline leaf shorter than or about as long as the
C. sp. 1
3b. diem usually more than one and subtended by a cauline leaf usually at least twice as long as
the spathes.
4a. "Bulbs 15-20 mm in diameter, with tunics not or only slightly resinous.
5a. Flowers white, rarely pale blue; inner tepals without yellow nectar guides or a zone of
nectariferous tissue; dd solitary in each rhipidium; inner tepals about 3⁄4 the length
of the outer and not i cat campanulata
. Flowers usually pale bluish to violet, rarely white; inner tepals with a yellow Pius guide
and a zone of nectariferous tissue, often outlined in violet; flowers (1—)2-3 in each rhi-

—
c

tA
I"

; inner tepals closely naa about half as long as the outer „u C. paludosa
C. sp. 2

pidium
4b. Bulbs about 30 mm in diameter, with tunics heavily resinous

NOMENCLATURE AND DESCRIPTION
OF C. PALUDOSA

Cipura saa Aublet, Hist. Pl. Guiane 38-39.
1775. TYPE: French Guiana: near Mt. Kou-
rou (as a. Aublet s.n. (lectotype, P—
Herb. Rousseau, designated here). Figure 2.

E a d din Kunth, Nov. Gen. Sp. 1: ene 1816.
bem San-

Í m del Angostura, Humboldt & Bo npland

S.A. atin cad E d DE ted here; isolectotype,

P— Herb. Bon

Cipura major cenas "Bull. New York Bot. Gard. 6:
493. 1910. TYPE: Bolivia: Tamupasa, 1,800 ft.,
Neila 546 (lectotype, NY, designated here;

Cypella and — Revista Univ. Cuzco
71.

944. TYPE: Peru. Convencion: Hda. Po-
trero, Vargas 2509 due ype, CUZ, not seen; iso-
ue , GH); Cipura goodspeediana (Vargas) R

Foster, Rhodora 64: 31
Cipura paludosa subsp. mexicana Ravenna: Phytolo-
5-1 : Mexico. Sinaloa:
joe 278 (holotype, Herb. Ravenna, not seen).

Plants 16-27 cm tall. Bulb 12-20 mm diam .;
tunics dry and brittle-papery, rarely slightly res-
inous. Leaves finely plicate, linear-lanceolate, 1—
3 basal, 16-27 cm long, 2-5 mm wide; subter-
minal cauline leaf single, largest, clasping the

spathes below, 16-23 cm long. Flowering stem
3.5-8(-15) cm long, bearing several crowded
rhipidia, either sessile or on short branches. In-
ner spathes 30-35 mm long; outer ca. 73 as long
as the inner. Flowers pale to bright blue, or white,
the tepals connate into a distinct tube ca. 2 mm
long; outer tepals fading to white at the base,
oblanceolate, ascending to nearly horizontal, dis-
tally twisted ca. 30°, the margins rolled outward,
the right more so than the left, 2.5—2.8 cm long,
1.3-1.5 cm wide, with a white nectary in a central
zone ca. 1 cm above the base; inner tepals 1.5
cm long, 8-10 mm wide, light blue with a white
apex, darkest at the margins, more or less ob-
ovate, erect and imbricate, the apex rolled out-
wards, strongly concave, with a yellow nectary
in a median band extending ca. 2 mm from apex
to 2-3 mm from the base, broadening to 4 mm
at the widest, surrounding a white to light blue,
raised nonglandular area, sometimes with semi-
parallel dark blue bands extending at right angles
to the nectariferous zone. Filaments 2-3 mm long,
thickened and sometimes contiguous at the base
for up to 1 mm, free and threadlike above; an-
thers adhering to the style, 3.5-4 mm long, la-
trorse with a slender connective broadest at the
base and tapering to the apex, pollen white. Ovary

340

included in the spathes, 6-10 mm long; style 6—
8 mm long, cylindric, 3-lobed above, the lobes
usually emarginate, | mm long, 1 mm wide, dis-
tally papillose, with 1—2 obscure (often micro-
scopic) erect, crestlike appendages at the base.
Capsules oblong-cylindric, 12-18 mm long, in-
cluded in the spathes; seeds angular, ca. 1.3-1.8
mm long. Chromosome number 2n = 14.

Distribution. Bolivia and Paraguay, through

southern Brazil to Central America, southern
Mexico, and the West Indies (Fig. 1

LITERATURE CITED
AUBLET, F. 1775. Histoire des Plantes de la Guiane

? "Studies i in the flora of Bolivia —
r. Gray Herb. 161: 3-19.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

GOLDBLATT, P. 1982. Chromosome cytology in re-

lation to suprageneric eae of neotropical
ridaceae. Syst. Bot. 7: 1

GRISEBACH, A. 1866. Catalogus Plantarum Cubensis

ip
' Erganzungen und Berichtigungen

KLATT, F. W.
zu Baker’ s Systema Iridacearum. Abh. Naturf. Ges.
Halle 15: 337-404.

KuNTH, C. S. 1815 [1816]. Voyage m Humboldt et
Bonpland. Sixiéme Partie. Botanique. Nova Gen-
era et Species Plantarum (= N s Genera e
Species Plantarum quas . . .), Volume 1. Libraire
Grecque-Latine-Allemande, Paris. Gide Fils, Paris.

1964 a i l

. (Bu
otes on ligase IV. Phytologia 56:

193-195.

RusBv, H. H. 1910. New species from Bolivia col-
lected by R. S. Williams— 1. Bull. New York Bot.
Gard. 6: 487-517

ITHOMIINAE (LEPIDOPTERA: NYMPHALIDAE):
SUMMARY OF KNOWN LARVAL FOOD PLANTS!

Boyce A. DRUMMOND III? AND KEITH S. BROWN, JR.*

ABSTRACT

he known interactions between the larvae of ithomiine butterflies and their host plants (about 400,
90% in the Solanaceae) are described in a table, illustrated, and briefly discussed.

The widespread and diversified use of plants
in the family Solanaceae by man is reflected in
the large number of applied scientific papers pub-
lished on these plants (see the taxonomic index
of any issue of Biological Abstracts). The alka-
loidal nature of most plants in this family has
led to their extensive use in folk and proprietary
medicine, consciousness expansion, and recently
as a source of pharmaceutical intermediates.
Other important uses include fodder, fencing,
support, insecticide, ornament, and perfume.

In tropical America, the most important groups

inae. These insects overcome t
physical and chemical defenses of these plants
and turn them to their own use, at least as rec-
ognition cues, if not necessarily as protection
against predation (Brown, 1987). Useful Sola-
naceae frequently attacked by Ithomiinae in-
clude Lycopersicon, Cyphomandra, Solanum sect.
x el d and solasodine-producing or tobac-
tituting Solanum. Also regularly eaten are
E Eun tuberosum, S. melongena and relatives,
as well as Capsicum and Physalis. Many orna-
mental and medicinal Solanaceae (Brunfelsia,
Cestrum, Solandra, Markea s.l., Juanulloa
Brugmansia, Acnistus, Solanum pseudocapsicum
and Solanum sect. Jasminosolanum) are heavily
damaged by ithomiine larvae. Nicotiana and Pe-
tunia seem to be immune to these herbivores.
This paper lists the known i t ti (to mid-
1985)1 butterflies and their lar-

IÇ

! This paper was part of the Second I

val food plants, 90% in the Solanaceae (Table 1).
It is the data base for papers by Drummond
(1985), Brown (1985), and Brown & Drummond
(in prep.). The 40 butterfly genera for which food
plants are known or inferred are placed in phy-
logenetic order in the Table, grouped into tribes
as first proposed by Fox (1961), followed basi-
cally by Mielke & Brown (1979), and modified
by recent studies of early stages leading to a nu-
merical phylogeny by Brown (in prep.). Nomen-
clature for the butterflies follows Mielke & Brown
(1979) except in a few cases in which recent stud-
ies, especially of chromosomes, indicate changes
in status. Nomenclature of the plants follows the
thesis of Mary Fallen (1983, Hamburg) for Apo-
cynaceae, recent compendia of the Solanaceae,
and the Index Kewensis. We believe that iden-
tifications of both insect and plant taxa are ac-
curate to the generic level in all cases, to the
subgeneric level for Solanum in essentially all
cases, and to the species level where given in the
vast majority of cases. Many older records, either
not confirmed or regarded as unlikely in view of
broader recent studies, have been excluded from
the list; these include especially those in the ag-
ricultural literature of Brazil, compiled in D’Ar-
aujo e Silva et al. (1968) and continuing up to
the present day [such as a recent report of Me-
chanitis lysimnia nesaea Hübner as a pest of Pas-
siflora edulis Sims. in northeastern Brazil, clearly
a misidentification of Heliconius ethilla narcaea
(Godart) or Eueides isabella dianasa (Hübner),
common in the area]. Doubtful records are
marked with a question mark in parentheses,

on the Biology and Systematics of the Solanaceae

presented at the Missouri pedum Garden on 3-6 August 1983.

? We are grateful to W. H. H
J. R. Trigo, R. F. Monteiro, G. pia mas M., G. Small, W

er, J. Vasconcellos-Neto, L. E. Gilbert, S. Knapp, J. Mallet, Renata S. C. Dias,

. W. Benson, A. H. Watson, Condorcet Aranha, Lucia

F. d'A. Carvalho, F. Fernández Yépez, and W. G. D’ Arey a information on ithomiine host plants, discussion

of the patterns revealed here, and identification of organ

3 Pikes Peak Research Station, Colorado Outdoor “upan sN Center, Florissant, Colorado 80816, U.S.A.

Please request reprints from this address.

4 Departamento de Zoologia, Instituto de Biología, Universidade Estadual de Campinas, C.P. 6109, Campinas,

Sao Paulo 13.100, Brasil.
ANN. Missouri Bor. GARD. 74: 341-358.

1987.

342

whereas sure records with tentative (uncon-
firmed) plant identifications are indicated by a
simple question mark after the name. Localities
and sources are coded and given at the end of
the table. Numbers or letters in parentheses after
a plant name are voucher symbols for that species.

The following genera (with number of species
in parentheses) of Ithomiinae have yet to be ob-
served or suspected as larvae on any plants; from
preliminary field abana they are predict-
ed to use the sola s genera indicated in
each case: Raswelia tit apsicum UM. Pa-
tricia (2) (Dunalia), a new genus near Hyposcada
(1) (Lycianthes). Paititia (1) (Cyphomandra), Ar-
emfoxia (1) and Pagyris (1) (Witheringia, Du-
nalia, Brugmansia, and relatives), and Dygoris
(1) and Veladyris (1) (Solanum sect. Geminata,
Cestrum).

The following genera of Solanaceae, with one
or more species available to Ithomiinae in their
tropical or subtropical moist habitats (genera re-
stricted to dry habitats or temperate zones not
included), have not yet been seen to be used by
any species of Ithomiinae. Solanoideae: Jalto-
mata, Athenaea (expected for Epityches and Ith-

treae); Nicotlana. Petunia, Fabiana, Nierember-
gia, Bouchetia (Nicotianeae); Schwenkia,
Protoschwenkia, Melananthus (Schwenkieae);
Parabouchetia (Parabouchetieae); Leptoglossis,
Browallia, Streptosolen (Salpiglossideae), Het-
eranthia (tribal position unclear). While some of
these genera may be found to be used by Ith-
omiinae with more observation, members of
others have been watched for years within large
Ithomiinae communities and have not been seen
to support the larvae; in a few cases, ovipositions
were followed by larval death (e.g., in Campinas,
SP, Mechanitis polymnia casabranca Haensch
has oviposited on both Capsicum annuum and

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Nicotiana sp., but the larvae died without feeding
or developing).
The complete picture of the known food plant
e of the Ithomiinae, based on the data
n the table, is presented in Figure 1, in which
Dian: genera (from top to bottom) and butterfly
tribes (from left to right) are arranged in phylo-
genetic order based on currently accepted evo-
lutionary sequences (the vertical position of but-
rfly genera represents only convenience in
presentation of the figure; details are given in the
phylogeny of Brown, in prep.). The complexity
of the relationships illustrated in the figure in-
dicates that strong ecological influences may out-
weigh the presumed evolutionary history of the
interactions. See papers by Drummond (1986)
and Brown & Drummond (in prep.) for discus-
sion of these aspects.

c
O

LITERATURE CITED

Brown, K.S., JR. 1987. Chemistry at the Solanaceae/
Ithomiinae Interface. Ann. Missouri Bot. Garden
74: 359-397.

Phylogeny of sud Ithomiinae (Lepidoptera:
Nymphalidae). (In pre
& B. RUMMOND TIT. Biochemical and phy-
logenetic coevolution: evaluation of the Solana-
Miri one oo (In prep.

D'ARAUJO E SILV WG E D. M.
GALvAo, A. i L. GONÇALVES, b
SILVA & L. SIMONI. 1968, ieee aes ‘dos
Insetos que Vivem nas Plantas do Brasil—Seus
Parasitos e Predadores. Ministério da Agricultura,
Rio de Janeiro, 4 volumes.

DRUMMOND, B. A., III. 1986. Coevolution of ith-
omiine butterfli d sol pl t Pp. 307-
327 in W. G. D'Arcy (editor), The Solanaceae:
Biology and Systematics, Columbia Univ. Press,
New ;

Fox, R. M. 1961. A checklist of the "ergo "
Tribes Tithoreini and Melinaeini. J. Lepid

3.

MiELKE, O. H. H. & K. S. BRowN, JR. 1979. Suple-
mento ao “Catalogo dos Ithomiidae Americanos"
de Romualdo Ferreira D'Almeida (Lepidoptera:
Nymphalidae: Ithomiinae). Univ. Federal do Pa-
raná, Curitiba.

DRUMMOND & BROWN-ITHOMIINAE 343

1987]

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ag wry jeung (SIIN) Ws8anjuvy Dapuniuoydty (19p[94 X 1o9p[o 1) ou:
ANY uA ezjuNy (WUO) SINDI1SSb4I pipuptuuaot/díD
JeJA- VÀ uA Jag (PUNA) prjofis42aip Dapunwuoyday (&epo[qnoq) pisapap
Td HO 9zjunw (WHO) simnpoissp42 napurwuoudá;)
HA ‘AM t: 9) IgunçI (SIIN) 7122(42401| vapunwuoydíp (seg) OyjuDjau npisd pipu |.
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aa uir] SIIIN (1029[U9S) ¿pupoaməutu vojmuvnf (uosltA9H) snpydouaw snjudouaws
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ay ued uius `d `4 PYIUVINI] (snuiuvisajmu28) noo4njy lopng $7/2/]p4pd
fy] "2 ZEMS D4O]fipup48 DAPUDIOS
DT HA WO `SuI9H DYJUDINaU (wunipodou1uiua3jy) v2xapnjy.
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ay AS ‘ory SIJIN (SWIG) paojfipiuia (DIUUOLYIOSSAG) DAYID nippeow n(104Dd
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ay oferey "Qory 22412202 DaYADW JƏuIÁƏAA SiSuanpuu
8x ue (i) YOTA 22412202 vDayADW (snoeuurT) atuəutu ətuəutu D2DutJ2JV
(punxa po]]nupnr uo pasod
ax uA (¿) &31y,q (IEUNG) Dapunjosau (snuiupisaipnyog) v2xy40jy.— -140) ^^aH % 'Áepiqa piəsədáy
HM uo (i) ZEMS pn4ojfipuva8 DAPUDIOS "A[ES X "urpor) adoay) viaiadAy sisaanq
(pupo51iəutu po]]nupnr uo
ay uA (¿) &o1y,q (rTeunq) vépunjosau (snyjuvisaynyrg) vayan paseas) uOSIIMAH 2 "[QnOQT $141042
HM xO (i) ZUVMS paopfipura8 pipup]oç "A[ES 8 UPON 1428U1pnpISs SIVID42 SpA]
INISVNIT3]A
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DRUMMOND & BROWN-ITHOMIINAE 347

1987]

IeJA- V4
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ZUCMG naio] (unuoi9JS01d2T) wunupjosg
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Sne ^2 ajtqpiava (uunuowajsojdaT) unuvjos

‘ds (winuowajsojdaTJ) unuvjos

jeunqg wnsosdaj-osojnunss (unaau1uva24g) tunup]oç
‘doos tunupillunpua (wunaauupa2a4g) UNUDjOS

Sne ^2 uo1yoojp32i4 (UuNdaYyJUDAII_) UNUDIOS
Iojpusg XƏ 'P[U9S w71uu12u02 (wumna4211upa24g) tunup]oç
TUNE wunuwissiypajnop (n404doiuvoy) uunup]oç

Wey] wns (unuowajsojdaTy) unuvjos

"uie Mens (1unuow21s01d21) unuvjos

‘boef wnijofiruowrajs (pdivz01sv T) unudjos
90 Mpaadspoos (wnaau1uvaa4g) UNUDIOS
‘ds (uinsayjuDaaig) wunurjog

jeunq (SIIN) 1122040| Dapunwuoyday
JOYA 04n3u2jnos2 uo21s42d0204]

"ure asuaojinb (pdavooisv'T) uinuvjos
peung wnaopfijissas (pd4pooisvT) wunuvjos
‘ds (uinuvjos) wnurjog

‘ds (uinuowajsojdaT) unuvjos

peung wnsopfijissas (pdipootsp'1) wunuvjos
"qea unuy (pdupootsp']) unung

7] Duasuojau (uinuoulajsojdaT) unuvjos

‘Woy (peung) wuinauiSnasafoaov4yI0 (uinuowuasojdaT) unuvjos

"org 14n4adsp (unaau1upao4g) UNUDIOS
“dea wnunri]pppuan[2opnjos (unaau1upaa24g) wnunjog
"Word sdə2up (ni40sspg) wnunjog

qosuoeH VDIUDAGDSDI

(snaeuurT) viuwudjod

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yosusey aaipdina

1Jədurpne1S Sapissi4op
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saeg vnuyIsi piuwudjod sillupuo2]w
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saiseds pue (uonooag 10 snuesqns) snusy
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ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

348

sa elg uoAegq ?g zin] 1471704224D2U1 (uunuow1s01d2]) UNnUDjOS
do SoA `do5S wnupuunpiu (uinsayjudaaig) UNUDIOS
sa elg (eouviq ene) "ds (uuna42u1una24g) unuDjos
sd eig (oout4q ournj) “ds (un4211una24g) wnuvjog (sniouqe) vuus]
HM SIV ezjuNy (WUO) sgnpoisspa2 DApUDUOYdAD
ag wry yeund wnjouljoad (pd4v201sv T) wnunjos
HM Sly `SIƏd wuniupdia (wunaau1upa24g) UNUDIOS
aa wry (peel) “ds (unup]os) unuvjos
aa wry Jog 4n1u0]042D14 (unup]oçs) unuDnjos (UNINDO) VS1]ə DIUUISA]
HA *IO UPA Saplounspdls (pi1uv2D421JA) wunuvjog
HM ‘91 HO IN Wnjopyjaquin (uinsayjuvaasg) UNnUuDIOS
£y] HO jeund wnsosns (wmna4211uva24g) uunurjog sojeg snssiaop snss4uop
aa wry wey] asuaojinb (pdupootspT) wnuvjog
aa wry [eunqq 14njDpu1J22d (pdupootsp'T) wnuvjog
aa wry Teunqq wnusopfipissas (pdavooisv T) wnunjos (spuqáy 7) 1epnng $7:7d222p
ay STL peung wnusopfijissas (pdav2oisv T) unuvjos
aa uirT wnijJoftav2un] “IU (DiUD2DA21JA) wi nuv]og
aa wry (EEEL) “ds (unuowasoidaT) unuvjos
aa uir] boef (9TEL) ‘ds (uunuowd21s01d2T) ununjos UOS]IM9H $712DZDiu snəpznta
aa 24A 7] wmsowwnu (n40gdougupoy) uunup]oç
ag 24A wey asuaojinb (ndar2oisv T) unuvjos UOS]IA9H SnauljuDU
ol WO UINUISSI]DAINID “IU (DAOYdOYIUDIP) uunup]oç
HM HO ITV $apiooisdpo (paoydoyjunoy) unuvnjos
HA WO [eunq xo [duog 77 ‘quiny wnyofiasv (p401dotu1uvoy) unuDjos
HM dO wey] asuaojinb (pdupootsp'T) wnunjos
HM HO UPA Saplounipdis (Di[1UD2DA201A) uunup]oç
HM uo sJoq Unpidsiy (uunuow1so1d2T) uunup)oç "A[ES X "UIpor) n7D4njps sidpuawui
WN org Joujpuos sijAjsopo19s napurwoud;»)
ay duro Jəu1puəS (“YOOH) sun43p4f pipuputot/d4ə
WN org Ioujpuos nuinjaa DapuDWOoYdaD
ay duro JIA, tnzuə]noəsə uo21s42d02]
Zif ory jeund wnsopfijissas (pd4vooisp T) wnuvjos
ay duro UuQAEd X ZINY wumipupjo4adsp (umnuowas0)d2) ununjos
,90INOS ,Al[?207] saisedg pue (uorjoas 10 snudsqns) snuoar) sor»odsqns pue ‘saisedg 'snuar) *'oqu
5Sjue[d 1SOH :Əpuruuoull
'Penunuo) “| 18v]

DRUMMOND & BROWN-ITHOMIINAE 349

1987]

HM Ww ‘ysuRH XÁJDIOYIUOI DIuow«taqq
HM Wo UO]1O]JA p048 DaUUINIOD (¿1əƏpəəJ 9e2oeuousor))
HM HO jsueH DaUINsUDSUOI n2uwunjo?) qosuaeH saprupraa DUDIUIBAIA
(pupo21xətu po]lnuphnr uo Impe 01 paiva)
ay ue (¿) YOTA 72u12202 DAYAD IY (uositMaH) 7482 nprosodÁH
INITAAITO
aJ Teod "u]Ioer) (7T) Sapiojpsdyd DAPUDIIN
do SOU uitou1o8e (7T) 7240q4D DIsuDWsnig
€ 181 əz1uns (WUO) Sljnb31sspuo pipuptuot/dAO
WN ory Ioujpuog S1/41S0pv1s DApuDUoYyddy
aN duro 1Jəu1puəs (ooH) supupi pipuptuou d(O
sa elg Joujpuos 2u1nja4 v4purwuoud«)
ay ‘do elg JOA 04412uagn2s2 uo2is42d02(T
do elg “T wnso4aqni (2070104) uunup]osç
NAf ums jeung unan (p40doi]uvoy) uunup]oç
8M Teod Joujpuog tnso4ə22p (vaoydoyiuvoy) uunup]oç
8M ‘NAL OW ‘wns lI V Sapiooisdvo (paoudou1uvop) uinuvjos
NAL wins yuelyos wnaindindoajo (vaoydoyiunoy) unuvjos
NAL uing yas unyofiydosjol (0401douuvoy) uinuvjog
GAw ory “boef wunwuissipomov (vaoydoyiuvoy) wununjos
8x Ory ‘ds (uinuowajsojdaT) unuvjos
8x AS ‘Or ZLIEMS 7440] (uÓnuoui9]s0)d2'T) uunup]oç
an [sig ‘ds (uinuowajsojdaT) unuvjoy
NAL wins 7] Uunipbpnoiurd (unuowoa1so1d27T) unuvjoy
NAL wing SNIEN ^2 ajtqpiapa (uinuowajsojdaT) unuvjoy
8x AS ‘Td "PIM 2472218178 0f (uunu0121501d2T) uunup]os
8x duro Jəu1puəS tun]pno4p (unuouuəlso1də'T) UNUD] OS
NAL wns suMOq
3? unus ^g ueur&T osuonbsnuq (14nuoi21s01doT) uunup]oç
sa elg (osuew eof) ‘ds ((unuotuəlso1d2T) nuvos
Sa elg `IDUƏAA 24n7snqo4 (uunuowo)1so1de' T) uunup)oç
Sd elg Iou]puog u714D48D (uunuown91so1do' T) wnunrjog
82D SOU "ureT winiyofilaquidsis (uunuowim1sojd2T) wunupjog
c«99Jnog ,AM[?207] soioods pue (uonoos 10 snuasqns) snuar sorv»odsqns pue 'saroadg 'snuoar) ‘aqu
usjue|d 1S0H :eeuruiou
'penunuo) “| s18v[

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

350

ay ay ‘ds sayjuv19A 7

8x puo ‘ds (uunupjosounusp[) ununjos eproui[y T] 2uipunu p32liqolupo psshpot/q
ay quq (ds s2uupiocT sojeg Dulinydjns

aa wry KIV q DUDIPADMOY SaYyUDIIAT yosusey 14D

ay 109 o ds sauupiotT [Bay W X04 MMosuyol pionuti

ay puoy ds sauy1uvioT eproui[v.(] 2uD242D

aq wry (STEL) i dS SayjuDIIAT [8931 ?? x04 sisuaDíronpvo siydjds

ay puoy (ds sauunio«T ppiƏuu[V.Gd Sisua, pssənp

HM t: ©) (¿) ‘ds wnuvjog

HM Wo ds sauuvio«T1

HM HO 191118 paojfimua “IU SaYIUDIIAT ue£urpor) vivu DSOJO] S2u2802dDNI
ay SOU Joujpuog DUuDISaaU si]ps(uqQ

ay dure) 'IPIU»S (7T) $uaosa4oquo snisiuop

8x SOU JayxIZUNH (J9UIPUIs) r40jf1424Q DIGOSsSD A

ay Ider Ioujpuog wnsonxalf tunoisdpƏ

WA ory Ioujpuag xo snrieJA `O ajiqvaiu wundisdvy (194215) adwodna sayotyidy

ININJOO3dVN

ay niod log ASUIOIDYI “IU (20]D10g) UNUD] OS (19gdoH) rjjiuidsi»

ay OW says V JSWIIOY WnuDIZUDMS (010pidoT) wnuvjog (12u142AA) DJIONbD

8x va ‘OLOW a jpu3nis (pdav2o0tsv T) unuvjos (anyu9qQ) 1212111

8x puo SNIEN ‘O wunsoipisui (uunuowuig]so1d2T) uunupioç (s1opues) 1/]21/24nq

ay puoy Jag 254202712 “IU (201010q) wnurjog “AOU “dss D24]$D

HM Wo UPA Sapiounapdis (DiJJup2DA421JA) UNUDIOS (nq X png) sua2saqna

AV Ho ueuiua21r) uwumijpofimajoaa (2010104) tunup]oç (nq) pspavd voijaz

MS nioqd ‘ds (viaosspg) unuvjos (uosltA9H) njjtapupt

aa wry] (OTEL) “ds (puuvovaotjg) wuinuvjos

aa wr] (61€L) “ds (plaosspg) wunuvjos

aa wr ueulru2217) tun1]ofi]jna]o42 (201010q) uunup]oç

aa wry AQ[PURIg 1UOXDU SaYyJUDIIAT (19P[24) 27514030

IeWw-VvA USA IIg UNEAZ] (piaAosspg) uunuvjog (uosjIAoH) DUaIYDOU
HA Ds) INg tun3Áz14] (piaosspg) UnuDjos
HA HO 191118 VLOMU souuniotT (utA[eS) 2u1214
HM Ho IMU VOM “IU SaYyJUDIIAT (J9WIAS A) DINDd 2u1401214 D142]()
8x nod (à) “ds sauuDiotT (UOSIIMƏH) vjjturo snu98 MƏN
;99Jnog ,Al[?207] saroods pue (uorjoos JO snuagsqns) snuay sor»odsqns pue ‘saidedg 'snuar) “qu L

;Sjue|d 1SOH

:eeurmuou

'Penunuo) [318v]

DRUMMOND & BROWN-ITHOMIINAE 351

1987]

aa WITT “dem wWnuviyppualyoajyos (wumnaau1unaa24g) WUNUDjOS (u5suəpH) n4nips pajnfiuəs
aa wr (TIEL) ds (nu1uv20421JA) tunup]oç (yosuse}) nu42aq pluon]/
SUMO(]
NAf wing % uiuis ^g UeWAT asuanbsnaq (uunuowi21s01d2T) uunup]oç
a ory ZEMS 440] (UiNUOWWaTSO]daT) uunup]oç
NAL wing 7T uinjpojnaiund (uinuowuajsojdaT) wnupjog
NAL wins '[PU9AA 14n7$nqo4 (uunuow21s01d2T) UNnUDIOS
NAL duo SNIEN ^2 aj1quiava (uunuowu2)s01do'T) unuvjos
8x duie;) snniejq ^O wnsoipisul (unuowuajsojdaT) unuvjos
Zif ory jeung wunĢojfiua204 (unu0w21s01d2T) uunup]oç
NAL uns doog wnupijunvu (wunaauupao4g) unung
NAL duro soj[nuog V IIWIIOY turtuDiz]4DMSs (n010p1də'T) uunup]oç
Gad ory 1o1tog xo [eun(q 1u7137u284D (DjopidaT) wnuvjog
NAL wins Sne ^2 4u0j120]D8214 (wunaauupa24g) unung
ay w Ioujpuog xo SNR `O tun]Jətuu92 (unaaiuupaa2ag) UNUDIOY
8M dure) IMPUI xo NOYIS wnuut2u02 (wumna2u]una24g) UNUDIOS
ay Da "ds (vapniqnpuy) wnuvjos
NAL duro JIoujpueg unuunu (plopidə'T) unuvjos
aA BON 'IISA unpnnyf (unupjosounusof) UNUDjOS (eanpsiog) vjavp
aN puoy g ‘ds (pjopidaT) wnunjos
ax puo v ‘ds (vjopidaT) wnunjos (KTA) 1£1412u
ay ue UINUDIDPUIIYIAJYIS (uunaouubvaa4g) UNUDjOY (yosuaeH) vajdu
8x S1, jeung wnusopfijissas (pd4p2o0isv T) uunup]oç (uexrz) vivmovwojdwuos viuoutu si«cuodépg
ay USA IUe A 17141 (vd4v201sv T) unuvnjos "^OU ‘dss
HM *IO U€£A3 Sapiounavdis (0u1uv20421JA) tunup]oç
HM HO UOLIOW 2% Ad[PURIS 5422524220 (1unuow2Js01d2'T) UNUD] os CATES 29 UPOD) pupuunəaəp vsj22xa
€x niod eow wnssa4ddp (uunaau1ura2a4g) uunup]oç
ay nd SNIEN `O 2f1qpi4va 716 (unuowas01dIT) wunuvjos (3ydoH) pju
€ oq-ues ‘ds ({winsayjunaaig) wnuvjos "^OU “dss
IeW-VA uA 'U£ A wn, (pdipootsb'1) uunup]oç (uosuaeH) nuo nino su4]pí H
ay [dry jeung 14712201400 (DYyJUDIDAN PY) UnuDjOS (yosuseH) p.otusəp
8M ueJA "boef wniyofiapoun) (pu1up2D421JA) wnur]os
ax ole, ‘ds (uinuowajsojdaT) unuvjos
an uel snie ^2 wnsoipisul (uunuow1so1d2T) uunup]oç (so1eg) DJOISOYIUDX DIOJSOYIUDX SI1ANVSADH
¢90INOS ,AM[?20'] sor»odg pue (uonoog 10 snuo3qns) snuay soroadsqns pue 'saroadg ‘snuay QUL

5Sjue[d 1SOH

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"penunuo) `p sigv L

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

352

ay Ider (5) 18u1puoS iunsojra UinoIsdDD
ay Ider Joujpuog wnsonxayf wunoisdp)
TA OW 9zjun* (pueouoJA) npon] unsisdv> eprouiy.q 14421]
HM ‘DI Ho IIHT Daopunjos nigu12111M
HM yD Ionig paojfiniu sauuniotT UOSIIMSH pJ/!]nd
HM xO “IIH. 222Durjos D1ISu1121/114A
HA wo Ioxrzung (Ao[puelS) 212147042180 D18u142U11M
Dy] Wo (9) ds sayjun19A 7 saeg $1u2420dd1 visvip vnuoyi]
IeJq- V4 uA 199AS (CPIM) SUajoaanns Disunwusnag
IeW-V4 USA `IPI[Uu5S (T) $ua2sa40qav snisiuop
ay USA uluny n22Duvjos nipunq
JejJA- VI uA JoxrzungH (UNY) purdi DISIAN (uosit^3H) 201101442 BOYIOWAD D14IJDAJN
INIINOHL]
WOd BON [eunqq wn5s04d9] -050puv48 (wuniayiuvaaig) uunuvjog
NAS q duo `do5S tunupnumnpua (unaauuva24g) unupjog (uosuoeH) pulu x pad]
aN yury peung wnsoésns (uun42u1uDa24g) unung
ay qurq yory unsadsp (wuniayiuvaaig) wununjos (&epo[qnoq) vuydvj
ay puoy boef wnyofiuowpass (pdav2oisv T) unuvjos
8x puo jeung wnsosns (un42(1uDA248) wu nupjog
ay puoy yory uniadsp (uinsaylunaaig) unuvjos (saeg) 114Dq “IU
ay on jeund wnsosna (uunaaijupa24g) uunup)oç
ay ‘NAL dwy ‘Uew yry 1naədsp (uinsayjuvaaig) uunup]oçs (saeg) z1upq
ay oq-ues Yory wnaadsp “IU (wna2uuva24g) UNUDjOS (qyosuseg) nuu]
aa wq SOI[NYIG P IIWIOY X9 "pIIUAA 40/021q (unaəyruvaəag) unung (1opng) vıpəwuaə1u1
JeNWN-VA uA (,,2100eq81,,) “ds (uuna2u1uDa24g) wununjos
IeJA- VÀ uA yry w742dsD (uinsayjuvagig) uunup]oç (uepop) vajona
gy (51 ueg AO IPN wumivjjaquan (uinsayjuvaaig) wununjos
AV ‘D1 Ds) peung winsoésns (uinsayjuvaaig) unung (seg) p!uponə) vojono s1adyjodA fy
ay puoy ‘boef wunijofiarour] (pyluDIDANIP) UNUDIOS
ay JOM [ePun(q 14n2201402 (DiJ1UD2DA21JA) UNnuUDjIOS ‘Aou ‘dss unada]
ay puoy SNIEN ‘O wnsoiprsui (unuouwo1soi1d2T) uunup]oç 'AOU ‘dss $7124214DU4
ax puoy jeung wnsosns (una4ou1upa24g) uunup]oç eplowlly.q siuydpp
8x Ie) jeund wnsosns (uinsayjuDaaig) unuvjos eproui|y (T Saplouydpp
ay JOW ‘ds ({uinsayjuDaaig) unuvjos
ay duy "Qory wm4adsp (uinsayjudaaig) uunup]oç UMOIg SISuUaDdDUD siuudpp
¢90INOSG m 9111227921 səd pue (uonoos JO snud8qns) snup saisadsqng pue ‘saisedg ‘snuan ‘QUL

;Sjue[d 1S0H

:oeurruou

'penunuo) ‘p sigv

DRUMMOND & BROWN-ITHOMIINAE 353

1987]

gl Ds fe) peung wnsosna ‘iu (wumaau1upaa4g) tunup]oç
ry] wo (cc) ds (uinsayuvdaaig) tunup]oç (194215) r13n]y DUUAIIG
ay puoy jeung wnusopfijissas (pd4poo1sv' T) tunup)oç
ay puoy ‘boef wnijofiruowr4is (pdipoolsp'1) ununjos
ay puoy snien 2 wnsoipisui (uunuowmo1so1d2T) uunup]oç
ay puoy UOAR_ V ZINY wnsopfipudis (UiNUOWAsSo]daT) wnurjog eproul[V.(] 24D242D 71240]
ay 5Sq-ues "qory tunuaədsp ‘su (1un421]upa24g) tuunup]oç (12uIA3 M) D2140]
ay uA JONG 7742zDW “IU (UindayJUDAIIg) tunup]oç (19p[94 ?9 19p[24) p31upta DUIpY DUUdILIG
SUAO(T ??
ay Ider Qus ‘Q ueul&T WnuDayIDMYIS '1U (UiNsaYyIUDAIIg) tuunup]oç (sneuos) pnoəspd puua]pÁH
ay og-d ‘ds s2uunioT (UOSHMƏH) p42fixn42 v1s(wuv]2A
HM ‘OT Ds (@) unq (SINN) 1222(3401| Dapuduoyddy (nq X png) 742u0] snua3 MON
ay BON ‘TRA tunpl522p]/ ((unup]osounupr) uunup]oç (19pI94) OYJUDx
ay NLW UOARY V ZINY wnsopfipunss (uinuowo1so)do'T) unung
8x puoy Joujpuog ajisuad (uunurjosounusp[) tuunup)oç ppi9uu[V.d /S0S$DADA] D2u2]
8^ uo ADIY q wmupijaupras `!u (Sp42204puy) UNUDIOS
HM Wo ‘UUO % UELEWA) 2D47D]2-2D]2UDS SAYJUDIIAT (UOSIIMƏH) 2iz2u DIZaY DIWOYIIIDD
ININNSO8I(]
aa ur "1 p)]pj)nSup syosdyd UOS}IMOH PidD]ps DsDAap
ay puoy 7] vipjnsun sipstud
aa wry ‘J suaosaqnd sippstuq
ay duro 'IPIU?S (7T) SUIISILOQAD SNISIUIP
8x duro Ioxrzung (1oujpuosS) 240j/1424q DIGOSSDA UOS]I^9H 2150u83D DISOUSD
ay duro `IPIUu39S (7T) suə2sə24oqap sn]siu5 F JəuqnH otu4up oudap
HA uo 'IPIU2S (T) $422s240q4n snjsiuop
HA wo JoyIZUNH (Ao[puelS) 27nauno p18u1421/1144
HM yo Ioxizung (YLUNY) Dud DIsadjOND (seg) souax
HM wo 'IPIU»S (7T) $422s240q4n snjsiusp
fYI'HAA HO Kosy qA 114014 DIBUL2IILM
HM >) IoxizunH (ulun%) Dud DISadJOND sajeg nroipjvaau
IeW-VA uA ‘IDH. VADUD] os niduu2u11M
JeW-VA USA JoyIzunyY (u1uny) india Dis24]0n;) “MOH ?9 "ÁpIqg Psspupndi psspupunjdi
HA wo Joxyrzung (PUNY) 24india 718241007) Inq P Ng vjoulsnjd n142]22
aa wry (LZEL) ds DisaajonD yosusey p7/714ptup
,90INOS ,AM[?20] satoadg pue (uorjoog IO snuasqns) snusy soroodsqns pue 'saroods 'snuar) ‘QUL

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ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

354

ado SOU (¿) “wey wnijofiaquaásis (uunuowm1s01d2) unuvjos
8x duo "7 wumoisdpoopnesd (voisdpoopnasq) tunup]oç
€x ory ¿ds (pipunuay) ununjos
ay Ider 'dureo `HƏA 2unanappo (vipuun) tunup]osç (zig) papuawtuy DaDUaWAY DIZMA
aq gi (0€€1) ‘ds (DIDUNUAD) wnuvjog (qu»suseg) uətu41
8x Sd Iou]pusgs wWnsO/fixD] (p3punua2)) uunup]oç ppi9ur[V.Gd ?? UMOIG DI4DUDO DDDOSHD42)
JeJA- V4 uA ‘ds (p;Dunuo22)) wununjos (UOSIIMSH) 217
HA WO ZINYIS zunup]]1lup (vinur?) uunup]oç (saeg) vjj140p nuni
aa wry z[nuoS iunapnjun (ppuru) wnuv]os (saeg) 71220d rj1220d
ay Pa ‘ds (uunuowajsojdaT) uunup]oç
8x puoy "boef wnijofituowmmais (pdapootsv T) uunuvjos
ay puoy sniwey ‘O wunmsoipisui (unuowajsojdaT) wnunjos eploWl[Y.q !upy1z OSaU D1u1]D42))
3gI4 MIN
do SOU wuniofiaquatsis (unuowa)jsoid2T) unuvjos
do SOU UOAR, V ZINY 14717D42324D2U1 (unuouuo]soldə'1) uunup]o ç
ay orgy ZURMS 1un440] (uunuow91so1d2T) uunup)oç
NAf uing TT umipnoiuvd (unuowajsojdaT) wnuvjog
NAL wins IPuƏAA LuNISNgOd (unuouw2s01d2T) wnuvjog
NAT ung SNIEN ^D 2jpqpiapa (uunuowm1soid2T) uunup)oç
NAf wins doos wnupijiinviu (uunaauiupa24g) uunup]oç
NAL wins snniejq ^D 4071/20]03214 (un421/1uDA248) tunuvjog Io]srounmg Du17J22
HA ‘91 Ds fe) u194 (Teunqq) wWnauisnisaf-OaIDAYIO (UNUOWAISO]daT) WnUDIOS (19p[94 ?9 19p[24) DujAyona 042p
ay uA INI muazpy (unaiəyiuvaəag) wunurjog (1343) Dual
HA Ds (0) UPA Sapiounapdis (DYyJUDIDAIIPY) tunup]oç (,.sisuanb14i1y),,) `Aou “dss pural
HM hi9) IN 1477jjJ2quun (uinsayjuDdaadig) wunuprjog
HM HO DOW @ 3SS9S 2$U2ADD402 (UUNAaYJUDAIIG) tunup]oç (19p[94 % 19p]24) $24(jo
AV HO Siəd uinpidsiy (uimnuowo]so1d2T) wnuvjog
HA wo Io] Wnipjjaquin (uinsayjudaaig) tunup)oç
OI uo jeung wnsosns “IU (uuna2u1upa24g) uunup]oç
HM xO “OW WZ 9SSƏS 25u2ADD402 (wn42u1uDa24g) tunup)oç sonuq X INNY 7770]24
OI uo "boef wnijofiar2un] (pu1uD27421JA) tunup]o ç
wa X9JA ZUBMG Unao] (ummnuowua)sold2T) uunup]oç
HA YO ‘Wray (PUNA) umoaui8na43/-02204420 (uunuow21s01d2) unuvjos
HM Wo ‘Slag unpidsiy (unuotuəlsold2) uunup]oç
HA uo IJN wmpjjequan (uinsayjuvaaig) uunup]oç
c«99Jnog ,ÁM[e207] so1sedg pue (uorjos JO snuagsqns) snup satoedsqng pue 'saioodg ‘snuay ‘aque

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DRUMMOND & BROWN-ITHOMIINAE 355

1987]

8M Idee ‘ds (DJpulluay) tunup]oç
gy duie;) "IPIUSS 1717021420] 14n4152)
ay ‘Wea IDY “duo SOINYIC W JIWIIOY WnuDYIZjUDMS (DjopidaT) unuvjoy
qo SOU Od xo [RUN WinajUuasiv (DJOpidaT) wnuvjos
ay duo 7] tunə1sdpəopnəsd (po21isdpoopnəsq) uunup]oç
gy duro ‘TPA Dpupuanappo (njmunua2)) uunup]o ç sneuos DIDI
AW HO jeung wnpjdydojpsau wuma1s27)
HA WO ‘ds (njbumu22)) tunup]oç
HA Wo ‘ds (ppuru) wnuvjog
HA WO AQ[PURIS X uo1iojN 11$2u24Q (DIDUIUAD) tunup]oç
HM WO jeung xo '[duog X `quinH 147240q4D (njbunua2)) wnunrjog
gT WO Jig asuajqod (DJDUIUAD) UnuDjIOS Inq X 1opng 2/j10u
HM WO ÁI PURVIS ?g UOLIOJ :71$2u24Q (D1Du1u22)) uunup]oç UIA[ES ?? URUIPOD 2/]D3D
HA WO Jeung xo '[duog ?? 'qwuny wnpnu (p)punuəD) UnUuDIOS (UMO) 0714102
HM dO (3) ‘ds (p;punu22)) wnuvjog
HM IO (A) ‘ds (vipunuan) wnurjog
HM HO (O) ‘ds (n;munu22)) tuunup]oç
HA WO (g) "ds (p;punuəD) wnuvjos
HM t: ©) (v) ‘ds (pjpunua2)) wnurjosg
HA WO Áo[pueig W UOLO 71$2u24Q (D1Du1122)) WNUDjOS
HM Wo jeung xo ‘|duog 7 '-quing 147240q4D (vipura) tunup]oç
HM WO z[nuoS wuna4pjjuup (pjmunua2)) wununrjog (Uta Jes) xə/duu1s xajduuis
IeJA-V.À USA (ds (DJDunu22)) wunupjog X04 12qaaq
HA Wo Jag (19ujpuog) r42iuru«s SAYJUDIIAT
HM WO ADIY q (C1[n0))) S1suajJINISa SAYIUDIIAT
HA Wo Ig Vof saujuniotT1 (UOSIIMƏH) DUILID puəlip DIMAUOAA]
gy ory `HƏA DUDANADD) (plputtuəD) wununjos qosuaeH sisodiajs DUISND]I
8x 109 (¿) ‘ds (pipuimay) wununjos
HM Wo jeung xo ‘[duog 77 'quing wnpnu (p1IpunuəD) tunup]oç
H^ wo z[nuoS wumapjjup (DIDUIUAD) wunurjos (saeg) rruiajvs ppposid:;4
ay duro `IPIU9S 1477081427] uani1səOƏ
ay duro UO(T ^£) NjppualyIajYyIS WNASID
ay duro SNIEN ^2 WUNUDIUƏIUIPUIS wunajs2))
do SOU 7] mbavd wnijsay
do SOU 7] wunuanjou winsjsay
¢90INOS ,AlM[?20] səroədç pue (uorjoog 10 snuasqns) snuar) səroədsqnç pue 'saroadg ‘snus ‘QUL

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'penunguo) “| s18v]

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

356

ay USA asuadu “IU (pjDunuox)) tunup]oç (uosjyIMoH) D2104pup

gl Wo (çI) "ds wnajsay (utA[ES) v4] DINUOAPUD

HA Wo [£5 Y SNIEN ‘O wnjoun] wun4Js2?)

HA wo ‘ds winijsay (UOS}IMIH) 970 2u034014

HM Wo ds wnuajsay

HA wo ds tn41$2)

HM HO TED ?? SNIEN 72 wunjpuv wunajs2?)

HA Wo Agouely umsojn8na winijsayD

HM HO 7] wunuanjoou wun4js2?)

HA wo jeung wnpyjdydojpsau winajsaD

HA *O Agouely 23/18D4f uinsjsaD (uositMoH) DUuassijod D1249
ay 109 "ds winajsay (GƏPPA X 19p[24) $14g72q1]

ay oq-ues ds wnajsay (UOSIIMSH) v20qis2udjp sijiuauodA H
8x OW ds 14n4/52)

8x idef "IPIUSS t7150quu«402 1041827)

ay ‘NAL duro `IPIUu3S tungp3Əl4əp] 1n4152))

8x duro uoq `D !IJDpuəll/22]1/9S wn4152)) (uosS)LAƏ9H) DINLA

aa wry (pce) “ds uinajsaD (UOS]IMOH) VUI rpposopnasqd
ay uA asuadi4 “IU (DIDUIWIAD) UunUDjOS (uosllA9H) Didapay vuəpəy

ay puoy (1uoquinoop) “ds wnajsaDd "AOU “dss

ay aly o ds (vipuun) ununjos AOU ‘dss

aa wry] IPIU9S 1417031420] wn4152)

aa ur] (PEEL) ds wn4j$2) (19U1Á9 A) S1/DUOLJDU Dj2[UADZ

AV WO AQ[PURIS ?? uon] MSauUadg (DIDUIWUAD) uunur]og (a[odot ND) 2721do:s2D2 Dssnuos

OT ao (£I) “ds wun4js27)
HM HO "I unuaniou wnajsay (utA[ES #@ URWIPOD) pi3z St4Apox)

INIGISAGO:L)

ay Sa 1J9u1puəS uunuo]fixp] (pipunua2)) uunupjos

8x S3 HIH 1S puinbopnəsd (p]punuəD) wnuvjog (IWI) 723714n2

ts va ¿ds (winsayjuvaasg) uunup]os CPA 2? PRA) Ə2i/Jupxruot

ag uirT unq xa '[duog X `quinH wnpnu ‘Ju (ppuru?) unuvjog qosuoeH vsunds vjjus24

8x USA asuadi JU (DIDunua2)) tuunup]oç (UOS}IMO}) 977110]
HA Ds) ZYNYIS uwunapjnuy “IU (vivui) WUNnUDjIOS soniq X "ning osupiuiajn{
H^ yə ÁƏIpue1S ?? uon] 7$2u24q (D]Dutu21)) wnuvjog "A[ES ?? "urpor) suaosaapnf

;99Jnog ,Al[9207] SƏI99dS pue (uorjoos 10 snuo3qns) snuar) saroodsqns pue “səroəds 'snuar) ‘QUL

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'ponunuo) ‘| slgy L

DRUMMOND & BROWN—ITHOMIINAE 357

1987]

`ur OOI-OS ‘zeig 'seuozeury ‘sneue JO YOU UOIsay = URW

‘WI Qc ‘zeig u1ojseo ‘OURS ojuidsq 'so1equrT = yur
"ul Qgc '1openoq uiojseo *euooououirT] = WIT

"WOOT I-O08 "Ttzeg 3S ‘ojneg ogg 'rerpunf ‘def op geus = Idee
"ui 000'7-000' I 'IIZe1g 3S 'o11rouef op ors ‘LIENEI ap IPuODEN ias im = eu

*'I£Ze1g ‘SBIOD jo IWS *OQ[2A SEION pue ‘BIURIOH ‘BADAOUDH O]UES = SIOD

'Ul Qc '[Ize1g ‘eieg U491SƏA ‘OILY = OILA
"ui 008-07 '[Izeg u1oj1seo ‘oues ojuidsq WIDYWON = Sq
"101098 u191SƏA ATJSOW *JOpEA[ES [3 = SIF

"eqn;) Jo puesi 3u1 Jo wed ujojseo Ajqeqoig = eqn?

"Ul Q09‘ 01 [9^9[ BAS ‘BOY LISON ut SILIO, SNOVA = YO

"ui OOS‘ 1—00p ‘LIN ‘Bole onune A[ISOW *erquio[o?) = [OD

"ui 004-00c ‘Zeig ‘Ied urouinos 'sefe1e;?) sop LG = IBD
‘wu 008—009 ‘zeig ‘Omeg oes 'seurdure;) jo uoigos = due)

"ut Q0C1—000'I ‘zeg ^e19po4 o1unsiq ‘eI[Iseig = [sig

"uaiseoqinos Ápisour DENT SnOLIEA ‘zeig = Eig

w OS zeig MS ‘DY ‘OouRIg Ory Ieou = 3V

:səpoo Áji[950[ JO IST] [£209q e ud[y z
‘MINUDIOS snuodqns ut 19010 Yoea 0) 1x9U
pooe[d Suroq SUIT} Əy} 10j 21e i isi d ais pue 2jopido'] 'ripbnqnpu ‘(£861
‘9¢9-SE9 :7E uoxe] ‘ddeuy `S 33S “INL uo4puaporo'] 10 NEL D4puapoioT =)
pibunuəDp suonoos pue :(e1ouo8 se pojeoar IY) SUOTIIAS Se SaYyJUDIIAT
pue uooir$42do2] 3pn[our osp[e p[noo [23] auos yorum “2010104 snuagqns UI o1e
suonoos : J| snuasqns əy} ul 3IE£

D40udouupoy pue Dduvootsv] 'DIJUDODADIAL suonoos 'unupjos snuog ou] UJ ,

ay OW "IPIUSS :/7g)104D. Uins]say Dssapa
ay puoy (auosquinoop) “ds 14714182)
aa wry (pce) "ds uinajsay (so1eg) ajaydau sipso42312H
ay idee "ds uinajsay
8M OW IIUIPUIG xo NOYIS tun40Jf1]ISSsaS 14n4182,) (uosi1MoH) DSDpY
ag uir] (ESEL) ‘ds wnaisa) (UOSHMƏH) 2u1J040
aw uu "ure 2uniJo[] wumnaisa?) (uoslta9H ?9 Áepo[qno(q) 73/D20
AV Wo ids (pipunuay) unuvnjos
HM yo ‘ds windjsayd
HA Wo jeung winjjAydojpsau winsjsay (saleg) suosspo DidajodA JT
NAL wns "IPIUoS 14717231427] wn4152)
ay duro uoq `O ropuaj2opj2s wuna1sa?)
NAL wns SNIEN ^O 1nupuiulpuəas UNASID (uosltA9H) DUIUOIDS pnuiuojps DISUN] IP
HM uo "DOIN ?9 9SSƏS 25u24Dp402 (UINAIYJUDAIIG) WUNUDIOS
HM wo ‘ds wn41$2))
HA WwW Agouel{ 2/18D4f n41$27)
AV Wo "ds winajsay (ULIOND) ə2JJəuup ajjauud
AV Wo Agouel{ MO]punis uinsjsay
HA Wo "ds winsjsay (UOSIIMS}{) 042u
or eqno “dS winajsay (1IPPLYIS-YILIH) pupqno
c«99Jnog ,AlI[E207] səioədç pue (uorjoos JO snuoSqns) snuar) saloadsqns pue 'səroədç ‘snuay ‘aqui L

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ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

358

‘BOTY VISO UIOJJ 19S ejep I31e] STY} JO sojepdn pue suorjeAJ12sqo 1U9001
sJOW pu? *ejosauurjA JO ÁPSIAUN ‘SISAL (qud “8161 ^A ‘QLH = HM
'uosuag ^A Jnupoow JO suonegA19sqo poausiqndu(] = g^
"niod pue
eueÁnr) APW `f pue ddeuy `S jo suorgA12sqo 1uəoər pousiqndu(] = MS
'9LI-SLI :£8] Ə2UƏIDS “SLEI `ƏlOOd ^A A P "fH PPPA = dA
Trzeg
‘dS 'seurdure;) op [enpeis3 AUN ‘SISIL “OS JAN `I861 ^4 ^W 'o112100JA = WA
'uongoguoa Suuinboi sp10221 Áueul ‘7/-¢¢ :p o[pned
OBS 7193 ‘OOZ “DIV “PH6I “POE-IBE :£€ ZMID Op[E^SQ "1su[ “WOW '8E6I
urag “sonbisojo1aidopiday] SZULI “7761 VINL ^ “epiotu[v.q = (HH
‘O6ITT -zz “90S “WY Uy "uo “2961 "JN ^W XOA = AA
"o[neq OLS [BUS ‘ENON JesaD O[ned = WOd
`ə[qe)sn.n A[qeqoud “ç [7] :p o31sue( op ors “SN “YTV "1881 ^N ‘EIRIN = WN
“617VIT ‘P6I-681 :9p 1SIšolouuoludq `£ 16] Os[e '(1o1srg [enieN])

SUOT]EAJ9SQO 1U92091 IJOW pue soureu jue[d JO SUONO 191e] YIM
*uoda1 SLO 8961 & ut Ansouu HəqrIO 73 7 jo uon eurnojur pəousilqndun = ry]
"e191dop
-ido T uerunuoary jo songo[e1e» snouea ut uonegunojur “Y ‘PIemÁeH = HM
'8tC-S€0€ :9 eordonoig
'FL61 (uosuog "A ^M UA) Li I :96 7908 Mj Hay 'suei[ '0L61 (eprou

-Ied yonu yim K[1u2001 * UMOIg YOY JO spjooo1 i im ÁNSOWN = ay
‘[Izeig ‘ojneg oes 'seurduie;)
3p [eNpelsy opeprs1oAtu(] “sisəu L S'IN ‘OS6I "f *O19N-SO[IƏ39uooseA = NAS
"urep1ojsury `səun[oA € 's1opuir[A ouosureeuungs '668I-8c8] "f ‘ddas = Sf
“8CC-STT :¿S UOPUOT “90g "ju 'suei]
‘POGI -PLI-OLI :Z ANID VEN PIPIA Pepruug `f `p681 “1 f ‘Addny = OTL
“BUBARH "?ueqn;) eIZOjOWO UY e[ e uorinquiuo) ‘1881 "f ‘YoR[puNyH = of
`əlqguonsənb Áuew ‘zesg 3S ut uexrz `A `f Jo sp1ooo1 pousrqndu(] = Z4f
‘8P9-9P9 :OST 9J1nIEN
`PL6I (Siolgioqello5 YIM) pue *'66£-68€ :961 uopuoT '[ooZ `[ 'Z86I ul
Ajjetoadsa *1e3p3 `y uyof Aq suoneasosqgo pəusriIqndun pue pousiqng = qv í
"PUIEUEd UI [EW ‘g uopiop JO suoneasssqg = SH
`< 81 (WY “Jed AY “9261 Ose
pue ‘Nd “eur “JA seureT open wo uoreuLiojur pausiqndu(] = 15
qes AKjeqonr $CC-1:I 'qugef OOZ *988] “ZW "19]njN = WA

poirsodop [euojeui uo xoy [UA V Áq S3}0U ƏAISU91XƏ Zurpnyout ° p|ənzəuə A ‘Abd
-EIE *erurouo18y op pene Əu1 JO uonoo[[oo aui ur uongutuioJul = JeJA-V.4
`uonguuiguoo gurpoou ‘INIIAI [eEorurouo.18e
WOJ} SpJ0221 Jop[o AYLIOMIsNIjUN Ápsoui ‘8961 “TE 19 BAIS 2 Ofnely.g = Sd
“6-€ :9 `dəT “xa “90g ju] Og 0861 `f ‘EZEN BI 9P = Wa

"uorneurnmuoo ambas 9194 pojuosaud əsou1 pue
[nJiqnoq *97] :d pue 9-1 :v "TIT INS Op spuein
ong "uro1u -biy ‘0961 pu? “ZçI[-I :9p oəptAƏluo],] “UOISY ‘ORY "Ao “(S101
-EJOQ*[[OO YIM) / C6] Á[[eroodso ‘suoneoygnd snouea JA ‘O ‘oyuezaig = go
‘yIzeig *o[neq oes op opepis13A
-IUN ‘SISIL "(qud `£¿61 `D 'seureT Aq ponrodai ory Usop ut noeg = Tg
'P861 u3nouug suoroauroo pue suongA1əsqo IIYUNY pue "aj[IAsauter)
“EPHOLY Jo ÁUSIAUN ‘SISAL "qrud 79/60 "III ^v ^g 'puoununiq = qg
';at
ZIT :9C “AN “Z "uq Ys 76161 NEPON ^A IA? "JN `V “SunoA = W/AV
"Áude18or[q!q 10} £9-/€ :6z eono1oouiorg BIW *0861 295 “W `V "SunoA = AV
'06-LL :c "do XIW “90g ^^23I 79/61 `V "Ipuousánjv = WY
:BJBP JO s221nos *s3poo 22u213J31 JO ISI] [£E2n9qeud[vy ç
"ur QOL 01 [249] Bas *10pen54 UJo1S3AA JO UIE] [eIseOD = JM
`ur OOO Z 01 [PAI] Bas 'soni[eoo| SnOLIBA *e[onzouaA UIIYWON = UA
'Sani[eoo[ snouea ‘Aeng = yA
"ur Qe zerg *eJeq ‘SUNULIOL Ory 'Ininong] = on]
"ui 008 01 [PAZ] VƏS *pepruu] = UUL
"ui 90£-0c ‘Izeg 3S 'o[neg oes [eISBOd “UNIA OBS = AS
"'|3^3| Bas *]S£02 9ureuuns = Ins
"ui 009-066 ‘Zeg 1s?2ugnos ‘ojneg oes *23ueuing = UNS
"ut OOP 1-006 '[!zeg 3S *o[neg oes ‘LIZIN LIIS = BIZINS
"ut 007—001 ‘zeig MN 'Seuozeury
*OI8ƏN ors 1oddn LIOMO ap PULY ues jo 1seaujiou 'soge] 319S = 21/
`1Openoq ujəunoçs = JS
"ul 008 '1openoq
isvo *(e10ure7 pue ezinbe[enr) 01 seoejA) urseq JOALI OSerjues Jodd() = 53-ueg
“WOOT ‘Izeg ‘gred ‘sodomy = [diy
"ur 00C-0S ‘zerg Uloy OU ourorjxo *eultejos[ = 103
`u 009
—00c 'seuronbury “guroiqosu L ‘nef *opeJo[o;) zeig MS *"eruopuos = puoy
“UL 008 01 [343] Bas zesg AS *Orrouef op ory Jo UOIZIY = ory
`ur Q00'T 01 [9^9] Bas '[rze1g UJZYINOS 3ut911xə ‘PNG op opueir) ory = SOY
“UW 009 ‘Izeg 3S 'o[neg oes ‘Oeo ory = [I
`r Q67-0S 'Izeg AN 'o»nquieuroq = quq
‘Nag [LNU = nag
"ur 008‘1—000‘1 ‘Zeg ‘stelayH seul ‘sepled op so5og = 9d
"ul QOO'T 01 [AA2] BAS 'sani[eoo[ SNOVA *eureueg = ueq
"ui 000'c-009 ‘Nag ujəulou 'soquin] pue 'e1eong ‘Uf = IAN

"ut 00€$-00c

‘zerg WINS ‘(IHON Op) OSSOID OJEW JO IWS dy} Jo Led WASIM = NIW
"ui OSS

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"ui 01

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CHEMISTRY AT THE SOLANACEAE/ITHOMIINAE
INTERFACE!”

KEITH S. BROWN, JR.?

ABSTRACT

The secondary chemical constituents of 42 species in sixteen genera of Solanaceae, of five species
in three genera of Apocynaceae- Ris amp p of 142 species in 45 genera of Ithomiinae (Lepi-
doptera: Nymphalidae) whose larvae feed o ese plants, have been analyzed and compared in a
standardized manner. Large orb spiders (Nephila clavipes), which cut field- caught adult Ithomiinae
out of their webs, were used to assay fractions present in or derived
from larval food plants; the fractions were lied externally to the palatable n uiii butterfly
Biblis hyperia. All extracts and fractions from Solanaceous plants were inactive, not defending Biblis
against predation by Nephila. The — use defensive compounds of adult Ithomiinae (also found in
s and occasionally in Apocynaceae u y some Ithomiinae larvae) are dehydropyrrolizidine
alkaloid (PA) monoesters and their N-oxides, stored in up to 2096 of dry weight. These compounds
are obtained from a variety of natural sources by the adults after emergence from the pupa (freshly
hatched individuals of both sexes in 26 genera were readily eaten by Nephila), especially from de-
composing Boraginaceae-Heliotropoideae and flowers of Compositae-Eupatorieae, for which they
serve as pollinators. onus Adenostemma, and many Eupatorium flowers were confirmed as
rich sources ofthe alkaloid
mutualism with the cim pollinators. The highly diversified and variable Solanaceae toxins. seem
not to be stored and used for defense by the Ithomiinae, but may be important in mediating larval
feeding preferences and oviposition in the butterflies, which show appreciable chemical specificity in
host plant preference. Patterns of ithomiine | larval use of oe Solanaceae at the generic level, however,
fthe oups. An

not p
to sequential adaptation by radiating lines of Jibi. to dnm classes of chemical toxins in the
already diversified Solanaceae. By obtaining their PAs, necessary for their defense and reproduction
(pheromone synthesis), from a source stabilized by mutualism, the Ithomiinae avoid dependence on
the Solanaceae chemicals, constantly destabilized by divergent selection in the antagonistic system.
Quantitative determination of PA concentrations in parts of different Ithomiinae and We: dian
permits the drawing of a diagram for the flow of these substances in natural ecosystems

(April 1983), at the 35th and 36th Annual Reunions of the Sociedade Brasileira para o Progresso da Ciéncia
(Belém, Para, July 1983 and Sao lees July 1984), and at the first meeting of the International Society of
Chemical Ecology (Austin, Texas, June 1984). For a preliminary s see Brown (1984); detailed
analysis of the insect side of the interface is presented in Brown (19 85).

2 I am especially grateful to Prof. Joao Vasconcellos-Net o for tł tal biological background

of this paper was read and criticized by T. M. Lewinsohn, W. W. Benson, L. E. Gilbert, M. Rothschild, and J.

A. Edgar. R. F. D’Almeida, O. H. H. Mielke, and Gerardo Lamas M. stimulated my interest in the Ithomiinae,

and special attention to the interface with ree was infused by W. W. Benson, L. E E. Gilbert, and - A.
e,

e
were most helpful with systematic aspects oe the "Solanaceae an nd H. F. Lei tao Fo with. the Compositae. E
Aparecida Henriques assisted greatly in extraction and fractionation (Table 3), and José Roberto Trigo and
Renata S. C. Dias helped in field study of the Ithomiinae/Solanaceae interaction. T. Eisner provided information
and reprints about the Utetheisa/PA/Nephila interaction first studied by him and his students in Florida. R.
Forster and D. A. Monteiro facilitated access to field sites in the Fazenda Santa Elisa, Instituto Agronómico de
Campinas. A. and C. Kascheres provided important services in NMR and mass spectra at the Instituto de
Química, UNICAMP, and I. Valio and L. Sodek permitted extensive use of the Micronal colorimeter in the
Departamento de Fisiologia Vegetal, UNICAMP. Chemistry of the Solanaceae was lucidly presented to me by
W. Evans, K. Schreiber ; D. Lavie „and J. G. Roddick. I especially thank L x. Brower, J. A. Edgar, and M.
Rothschild for helpin nd bioassay techniques. FAPE
contributed to the original installation of the Laboratorio de Ecologia Quimica (project 74/0544) and Rhodia
NCAMP to its reinstallation. The CNPq provided a research fello

3 Laborat torio de Ecologia Quimica, Departamento de Zoologia, Instituto s COE Universidade Estadual

de Campinas, C.P. 6109, Campinas, São Paulo 13.081, Brazil.

ANN. MIssouRI Bor. GARD. 74: 359-397. 1987.

360 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

CLAY 07

oO "o

A. R*H solamine
ReCOOEt solaurethine C " m ed ñ
R=COCH=CH(CH2) 12CH3 K. l-acetyl-B- z etaine

i carboline-3-
carboxylic

solapalmitenine

G. anatabine*

B. cuscohygrine

A083
N
LA,

L—- Nis.

C, tigoidine 8

ad

°

pesos un
N. capsiamide* ^
C
P.
E

C2
hyoscine d '
J

D. withasomine E. hopeanine?? B. lupanine M. capsaicin*
[ORNITHINE]
E bim LALIE j
FIGURE |. Representative alkaloids known from the Solanaceae, with probable biosynthetic pathways; see

Table 2 for occurrence in genera. Asterisked alkaloid types are not known yet in leaves of natural Ithomiinae
host plants.

Plants in the cosmopolitan family Solanaceae (Brown, 1979, 1987; Brown & Benson, 1974).
are widely known for their toxic qualities derived They even pull in other less abundant and per-
from an exceptionally diverse suite of alkaloids haps less protected, but typical aposematic but-
(Fig. 1) and steroidal bitter principles, terpenes, terflies such as Heliconiini (Brown, 1972a; Brown
saponins, and phenolic glycosides (Fig. 2; Evans, & Benson, 1974), Danainae-Itunini (Brown,
1979; Schreiber, 1979; Harborne & Swain, 1979; 1987), Acraeinae (Brown & Benson, 1974), and
Kirson & Glotter, 1982). Such a rich larder of Troidini (which are usually central models of
potential poisons could be expected to repel, de- their own mimicry rings), as well as myriad
ter, or intoxicate almost all herbivores while Batesian mimics (Satyrinae, Nymphalinae,
causing a few to become specialists; the special- Charaxinae, Riodininae, Pierinae, Dismorphiin-
ists might be predicted to sequesier the Solana- ae, Hesperiidae, Castniidae, Geometridae, Diop-
ceae poisons and use them in their own defense tidae, Arctiinae and Pericopinae— the last two
against predators. In fact, the herbivorous insects possibly distasteful Müllerian mimics— and
found on Solanaceae leaves in the Neotropicsare members of other insect orders such as Odonata
often restricted to the family and some of them and Homoptera). Adult Ithomiinae were in-
are brightly colored as ifto suggest unpalatability volved in the original proposals of Batesian and
(Table 1). These form ideal systems for the in- Müllerian mimicry and have been shown to be
vestigation of biochemical ecology and coevo- rejected by a variety of vertebrate predators
lution (Brown, 1980). (Bates, 1862; Belt, 1889; Brower & Brower, 1964;

One of the most important groups of Solana- Haber, 1978; Coimbra-Filho, 1981). One known
ceae herbivores in the Neotropics is butterfly lar- exception (Brown & Vasconcellos-Neto, 1976)
vae of the nymphalid subfamily Ithomiinae involves complex learning behavior by a single
(Drummond & Brown, 1987). The brightly col- tanager population (Pipraeidea melanonota),
ored adults are regarded as prime distasteful ^ which squeezes out fatty abdominal contents of
movers in regional insect mimicry complexes individuals in winter reproductive diapause dur-

1987]

STEROIDS

Q. BITTER
PRINCIPLES

Q4 withaphysalin A

SAPONIN OH

R. hispinin B

!
H Ò- Rha-Rha

U. solanesol |

BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY

S. perulactone

POLYPRENE

361

TERPENOIDS PHENOLIC GLYCOSIDES

. DITERPENES

š i
i OH
ë
H
W. cestric
acid

MI raimonol

c

M 4, I uo didum ad

uerci
rutinoside

`

has TRITERPENE

Z. PUNGENT
OILS
(unknown

o
T. lanosterol composition)

EN H,OH

IGURE 2. Examples of non-nitrogenous secondary compounds found in Solanaceae used as larval hosts by

Ithomiinae. Carote
See Table 2 for distributi on

ing the cold of early morning, treating them as
it does fruits with a bitter rind. Similar learned
behavior has been observed in jays (Corvidae)
eating ithomiines in Costa Rica (R. Hagen, pers.
comm.), as well as in orioles and grosbeaks at-
tacking wintering monarchs (Danaus plexippus)
in central Mexico (Calvert et al., 1979; Fink &
Brower, 1981). Like most aposematic insects,
Ithomiinae have tough and resilient bodies; sur-
viving individuals squeezed by the tanager were
often captured in flight one or more days after
attack. They often remain on the forest floor for
weeks after death, avoided by predatory and
scavenging ants. Even the giant tropical orb spi-
der Nephila clavipes, which often clutters the air-
space of ithomiine colonies with broad sticky
webs and takes most aposematic butterflies with
typical rapaciousness (Vasconcellos-Neto &
Lewinsohn, 1982, 1984), cuts out all Itfhomiinae
from its web rapidly (10 sec.-2 min.) after con-
tact with any part of the body or wings. Other
spiders, especially flower-frequenting Thomisi-
dae (crab spiders), may take Ithomiinae regular-
ly, however (Drummond, 1976; pers. obs.).

It has been suggested frequently that the pro-
tection of adult Ithomiinae against predators is

noids in other pigments are not included; for flavonoids, see Harborne & Swain (1979).

due to alkaloids or other toxic chemicals se-
questered and stored by the larvae from Sola-
naceae and passed on to the adult (Brower &
Brower, 1964, p. 154; Young, 1972, p. 291;
Drummond, 1976, p. 268, 1981, p. 63; Brown,
1980). A good precedent for this suggestion exists
in the storage of cardiac glycosides (Fig. 3A) by
larvae of Danainae—sister-group to the Itho-
miinae (Ackery & Vane-Wright, 1984)— which
are transmitted to adults and help protect them
against avian and other predators (Brower et al.,
1967; Brower, 1969; Brower & Glazier, 1975).
Nevertheless, no evidence has been obtained yet
for the presence of any Solanaceae secondary
chemical sequestered naturally into the tissues
of any herbivore. Indeed, all results reported by
Rothschild (1973) showed metabolism and ex-
cretion of Solanaceae alkaloids by specialist her-
aes a id experiment showed retention of

Manduca sexta (tobacco hornworm)
(e up et al., 1979), but this herbivore does
not normally encounter these insecticidal com-
pounds in nature. Since Solanaceae alkaloids and
steroids are relatively stable compounds, abun-
dantly available to Ithomiinae through the larval
food plant and toxic enough to be eminently suit-

[VoL. 74

ANNALS OF THE MISSOURI BOTANICAL GARDEN

362

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1987] BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 363

INSECT DEFENSIVE COMPOUNDS

MEC T

INSECT MALE PHEROMONES

i ead ^ x

CARDENOLIDES

"iaispencii
UR MM lactone"
alkaloid monoesters
A. Calotropi E. lycopsamine
T sclepias "i i . i F 9 CH3
F. (epi-e) intermedine
G. (7«-0H) echinatine a
OAc š : :
H. (7«-OH, epi-«) rinderine
I. (epi-«e,8) indicine li. Danaidone
H202
9 CHO
oleandrose-O (0.15%) “a H
I H MeOH u oc
glucose 100° 7 , z Ps
P n 6 N ¿
B. Urechitoxin (Urechites) n m H
Ac.0 : =
eroe 3 MN. Án
so qo ie J. Echinatine N-oxide
acetate : MS
NH, 100° K. (epi-7,0,@) Indicine N-oxide
^ APOCYNACEAE TOXIN
OO HO CHOR Pon
CO 2% BF lal Soe CHZ0H
NH
2 CO co
(CH) "Hac
3-hydroxy-L- 579. 5 min cH Ehrlich
C. kynurenine MATTOCKS- f =a 0. oqui
u BINGLEY " pro ins oss)
(Ithomiinae) ASSAY N is 562 n nm ——— asa

Danaine and ithomiine ED and defensive compounds, including cardiac glycosides (A,

able for protection against predators, their ab-
sence in Solanaceae-feeding insects was quite a
disappointment (Rochschild, 1973). The only
known “secondary chemicals" isolated from the
Ithomiinae before this work were the amino-acid
yellow pigment 3-hydroxy-L-kynurenine (Fig.
3C; Brown, 1967; Brown & Domingues, 1970)

and a Sermon lactone related to some es-
terfving acids of pyrrolizidine alkaloids (Fig. 3D),
as well as the presumed precursors of the latter
(Fig. 3E, F) (Edgar et al., 1976).

Especially interesting to find in adult Ithomi-
inae were these latter compounds (dehydropyr-
rolizidine alkaloids, or PAs, Fig. 3E-K), since
they have been shown to be very important in
both defense and reproduction of the butterflies
in the sister group Danainae (Edgar & Culvenor,
1974; Schneider et al., 1975; Schneider, 1977;
Rothschild & Marsh, 1978; Boppré, 1978; Edgar
et al., 1979; Edgar, 1982). Unlike the rather spo-

N), ware pen Men alkaloids (E-I) and their N-oxides (J,

assay, and deplete toxins (B, E,

radically stored cardenolides (Rothschild et al.,
1975; Dixonetal., 1978; Malcolm & Rothschild,
1983; Fig. 3A, B), PAs are universally present in
field-captured adult Danainae, which actively
seek them out and sequester them (Boppré, 1984)
from flower nectar, plant exudates, and even oth-
er insects (Bernays et al., 1977). Some danaines
(Euploea) also obtain PAs from apocynaceous
larval food plants (Parsonsieae: Parsonsia; Ed-
gar, 1982). Larvae of a primitive ithomiine iso-
lated in the New Guinea region, Tellervo zoilus,
feed on these same plants; adults contain PAs
presumably derived from Parsonsia (Edgar,
1982), as do adults of a primitive Central Amer-
ican ithomiine (Tithorea tarricina) whose larvae
feed on Prestonia (Apocynaceae: Parsonsieae)
containing the same PA found in the butterflies
(Fig. 3E; Edgar, 1982; Edgar & Harvey, in prep.).

The purpose of this paper is to examine in
detail, by chemical analysis and field bioassay,

364

three basic questions related to the specific in-
teraction between Neotropical Ithomiinae and
their Solanaceae larval host plants. These ques-
tions derive from conventional aspects of insect-
plant coevolutionary theory, especially as ap-
plied (with much success) to aposematic insects
on poisonous plants (Ehrlich & Raven, 1965;
Benson et al., 1976; Brown, .

(1) To the extent that host plant specificity
may exist in the Ithomiinae/Solanaceae relation-
ship at generic and specific levels, how does it
relate to chemical mediation of cues for ovipo-
sition and larval feeding?

(2) How might such chemical specificity re-
flect mutual interaction of these two groups over
long evolutionary time, leading to diversification
in both groups as a function of reciprocal selec-

tive pressures—that is, what is the evidence for
coherent patterns of biochemical or “‘parallel di-
versification” coevolution?

(3) To what extent do the Ithomiinae use the
poisonous secondary chemicals of their Solana-
ceae larval hosts for their own defense against
predators in any stage of their life cycle?

A preliminary phylogenetic (Brown, in prep.)
and chemical (Brown, 1984, 1985) survey of the
Solanaceae/Ithomiinae interface, accompanied
by an efficient spider bioassay for predator-de-
fense compounds, has indicated surprisingly that,
while the first question seems to merit an affirm-
ative response, the other two questions must be
answered by “little or no evidence in favor." The
Ithomiinae apparently colonized the Solanaceae
well after the generic diversification of this plant
family in the New World, tolerating and using
the diverse secondary chemicals (Figs. 1, 2) for
oviposition cues to regulate specificity, but not
for defense. Efficient chemical protection of adult
Ithomiinae is generally unrelated to toxins en-
countered by larvae but not stored; instead, the
adults seek out PAs in flower nectar and decom-
posing leaves, using them not only in pheromone
synthesis (Fig. 3D) but also in defense, exactly
as in the Danainae. The results of this survey are

plant specificity, and alternative
ecological and physiological factors in the rela-
ionship

MATERIALS AND METHODS
STUDY SITES

Field observations of interactions between So-
lanaceae and their herbivores were undertaken

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

principally in two very different artificial tropical
forest systems in the Fazenda Santa Elisa, Insti-
tuto Agronómico de Campinas, Sao Paulo (600
m elevation): Monjolinho (a forest garden of na-
tive plants) and Amarais (an old eucalyptus for-
est with natural under- and middlestory) (see
Brown et al., 1981, for photos, maps, and de-
scription). Supplementary observations were
made in Sumaré, SP (similar to Amarais, 550
m); Martinho Prado, SP (floodable riverine for-
est, 400 m); Serra do Japi, Jundiai, SP (native
seasonal and montane forest on rocky soils, 850-
1,150 m); Serra Negra, SP (humid montane for-
est, 1,000 m); Bertioga, São Vicente and Mon-
gagua, coastal Sào Paulo (perhumid tropical for-
est, near sea level); Pocos de Caldas, Minas Gerais
(montane forest, 1,200 m); coastal Rio de Janeiro
and Espírito Santo (disturbed tropical rain forest,
sea level to 800 m); Goiânia, Goiás (riverine
thicket, 800 m); various parts of Rondónia, SW
Brazil (seasonal forest on rich soils, 100—650 m);
various parts of NE and S Pará, Brazil (tropical
forest, sea level to 800 m); the region north of
Manaus, Amazonas, Brazil (disturbed tropical
rain forest on poor soils, 50 m); and in northern
Venezuela (deciduous and cloud forest, sea level
to 1,400 m). Some limited data have also been
obtained in Mexico, Panama, Colombia, Ecua-
dor, and Peru

JUVENILE BIOLOGY

Ecological observations in the field were con-
ducted with the aid of binoculars and suitable
recipients for living adults and larvae (for details
see Brown & Benson, 1974, and Brown, 1972b)
Identification is secure for adult ithomiines, at
least 9096 certain for juveniles, and secure for
Solanaceae host plants at the genus or section
level but still uncertain in a few cases at the species
level. Early stages of ithomiines were brought
into the laboratory when necessary and reared
to adults in glass or plastic dishes on separated
leaves of natural or experimental host plants,
kept subhumid. At least one fertile egg could
usually be expressed from any field-captured re-
productive female ithomiine by careful pressure
on the abdomen with thumb and forefinger,
working backwards slowly from the fourth seg-
ment. Such eggs were kept with expected or po-
tential food plants until hatching (3-9 days). A
large number of potential food plants could be
found in Monjolinho or other areas in Campinas;
many others were cultivated as available in my
garden.

1987]

LIVE INSECTS OR FRESH LEAVES

vigorous «d RIS ze
with

10

BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY

365

at room temperature

volume of MeOH:H20 (8:2 or 9:1)

filter and wash with another smaller portion

extract

solid residue

virgini to dpt. volume, under vacuum, homogenize
OPTIONAL 40? ving all methanol; acidify with 6% volume of conc. H2SD4 eh 2 x 10x
extract 3x with an equal volume of CHC13 extracts
lower layers
y unite,
make basic with conc. NH40H (to pH 10) unite, dry with evaporate chitin or
extract with 2 equal volumes of CHC13:Me0H 3:1 anhydrous Na2SD4 under fiber
and one larger volume of CHC13 vacuum
evaporate under vacuum
lower layers E
-
A
aqueous unite, dry with v i I (nonpolar material: mostly fats,
layer anhydrous Na2S04 moderate polar acid waxes, carotenes, hydrocarbons)
(basic) iris x I or — esl
des uec evaporate under vacuum pigme rin steroi
many u mE podere
extract residue B
with MeOH (2 cycles) ( 7 . analysis by TLC on
solid extracts moderately polar basic silica gel
erige alkaleids and
thei 5,
inorganic selt vaporate wir glycosides, Aminas) Further fractionation on
es) 3504 under vacuum analysis by TLC, columns of silica gel
and C silica gelH,
7 fractionation on

CHC14-MeOH +

(highly polar material:

alkaloid N-o pci Te
m e nal reducti opt
loop at this atage

and

zwitterions, some salts)

Nis Fu
deactivated basic alumina
or partition columns

PAEA by TLC
aper chromatography s> Further fres

ionation on Sephadex or

t
-exchange column

E4. Mild and rapid standardized fractionation scheme for fresh leaves, flowers or insects, adapted to
alkaloid- -containing materials, giving fractions for chemical and ecological analysis and assay.

CHEMICAL FRACTIONATION

A standard extraction and fractionation was
developed for all plant or animal material (Fig.
4). Leaves or flowers of plants observed to be
used by Ithomiinae larvae or adults were sepa-
rated from petioles or peduncles and maintained
fresh until extraction; juvenile and adult Ith-
omiinae were kept alive in dishes or envelopes
until the moment of extraction. Thus, all ex-
tractions were performed on fresh and recently
functioning living tissues. The method was
adapted to rapidly remove all organic substances
of moderate to high polarity and at the same time
deactivate enzymes in the living material; stable
nonpolar compounds were removed afterward
The initial use of heat, aeration, acids, bases, or
acetone (which can condense with primary
amines) was avoided. Exhaustive extraction was
sacrificed in favor of rapidity; the more labile
polar compounds (especially glycosides) were
immediately and efficiently dissolved by vigor-
ous maceration in the first extracting solvent
added to fresh material. The leaves, flowers, or
insects were divided if necessary into pieces of
no more than 2 cm maximum linear dimension
by scissors, already under or falling directly into
methanol-water 8 : 2 (usual for leaves to extract
less chlorophyll) or 9: 1 (for adult insects), using
about ten times the volume in milliliters as the
fresh weight of material to be extracted. The

”

extraction was immediately completed with scis-
sors (if less than 40 or with a kitchen
homogenizer (Braun **Mini-Pimer" with a two-
blade metal propellor) introduced into the sus-
pension from above. Within a minute, leaves
were reduced to tiny fragments and insects to
pieces of chitin, partly coated with a fatty layer.
The extract was homogenized for 1-3 minutes
and filtered under suction; the insoluble material
was washed and stirred with a further amount
of aqueous methanol, up to half the original vol-
ume. Nonpolar compounds (fats, waxes, caro-
tenes, some terpenes, and free flavonoids), 1
soluble in aqueous methanol, were Sewg end
extracted from the residue with two portions (each
5 x original fresh weight) of ethyl acetate, giving
a “fat fraction" (F) and leaving only polymeric
organic fiber or plaque (cellulose, lignin, chitin).
The filtered and united aqueous-methanolic
extracts were evaporated under vacuum at less
than 40°C until all the methanol was removed.
The resulting aqueous suspension (at times after
filtration, giving a chlorophyll or nonpolar frac-
tion F) was acidified with conc. sulfuric acid
(roughly 696 of the total volume, making it about
= 1 M in acid) and extracted three times
Wil: an sanal volume of chloroform. If occa-
t this or later stages
(especially in some plant extracts) did not sep-
arate after 6 hours, they were filtered or broken

366

with MeOH. The organic layers were united, dried
with anhydrous granular sodium sulfate, filtered,
and evaporated under vacuum to dryness to give
a relatively more polar but nonalkaloidal frac-
tion (A), often including some chlorophyll from
leaves (usually little soluble in aqueous metha-
nol) plus terpenoids and steroids (incl
cardiac glycosides, withanolides, and saponins),
flavonoids, coumarins, and organic acids (Fig.
2). Half or all of the aqueous phase was then
made strongly basic (pH above 10) with conc.
ammonium hydroxide and extracted with two
equal volumes of chloroform-methanol 3: 1 and
one to three volumes (1.5 x) of chloroform; this
gave effective partition between water-methanol
8:1 and chloroform-methanol 6:1, obviating
salting out of polar organic constituents from the
upper layer. The organic layers were combined,
dried, and evaporated as above to give a mod-
erately polar alkaloid fraction (B) containing gly-
coalkaloids, tropanes, pyrrolizidines, and sim-
pler bases (Figs. 1, 3). The remaining aqueous
phase retained alkaloid N-oxides, very polar gly-
cosides, more polar acids, and salts and sugars;
it was directly evaporated and extracted with
methanol (two cycles with filtration) to give a
polar extract (C) and leave insoluble organic salts,
mostly ammonium sulfate. If the alkaloids cor-
responding to the N-oxides were desired, the oth-
er half of the acidic solution (after CHCl, ex-
traction, 2 N in H,SO,) or fraction c after
dissolution in 2 N H,SO, was reduced by stirring
for 2 hours at room temperature with an excess
of zinc dust (usually in grams equal to one tenth
of solution volume), filtered (paper cone, gravity
funnel) alcalinized with an excess of conc.
NH,OH (until all zinc salts dissolved as the zinc-
ammonia complex), and extracted as above with
CHCl;-MeOH (2x) and CHCI,. If this gave a
much greater weight of alkaloid than in unred-
uced extracts, significant amounts of N-oxide
were indicated. The complete extraction and
fractionation scheme is illustrated in Figure 4.
Crude or purified fractions were analyzed by
NMR spectra in CDCI, + 1% TMS, on the Var-
ian T-60 of the Instituto de Quimica, UNI-
CAMP, or eventually by IR spectra (in CHCl,
or KBr) on a Perkin-Elmer Infracord. Mass spec-
tra were performed by Mrs. Concetta Kascheres
on the Varian MAT 311A instrument of the In-
stituto de Quimica, at 70 eV, probe temperature
60-105?C. Chromatographic analysis on TLC
plates (coated microscope slides) used silica gel
H (no binder) and varying amounts of chloro-

g most

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

form-methanol or benzene-ethyl acetate, plus 196
NH,OH for alkaloidal fractions; other systems
used ethanol, acetone, and amines. Separations
on adsorption columns (silica gel or alumina ac-
tivity III were usual) followed results of the TLC
analysis and used careful gradient elution. AI-
kaloids were purified by partition chromatog-
raphy with the system ethanol-chloroform-water
(usually 1 :9 : 1), stationary (upper) phase 0.75 x
(V/W) on Celite 545 and including phenol red
as indicator (Brown & Kupchan, 1962) or chlo-
roform and buffer solutions.

DETERMINATION OF PAS

Dehydropyrrolizidine alkaloids (PAs) were
quantitatively assayed in fresh plant or insect
material (they are destroyed after death or dry-
ing), cut up under absolute methanol (at least
10x), and, after at least one day, colorimetric
determination on aliquots (usually 1/20, prefer-
ably containing no more than 100 ug PA; 40 ug
gave initial absorbance off scale > 2.0) and the
Beer’s law curve became bove 150 ug)
with a modification of the Mattocks (1967, 1968)-
Bingley (1968) method. The assay is very sen-
sitive as modified (< 0.1 ug is easily seen), very
accurate for N-oxides (+ 1%), reasonably accu-
rate for total PA (+ 10%), and extremely specific
and applicable on total plant or insect extracts
with minimal interference by other components;
the final product of Ehrlich condensation shows
Amax 261.5 nm (in isoamyl acetate-ethanol-ace-
tone about |: 1:2 with 2% BF,) with e ~ 57,000
(Fig. 3L). For total PA determination, the aliquot
is treated with 0.5 ml of freshly prepared 0.15%
methanolic H,O, (from 200: 1 dilution of 30%
aqueous H,O, stabilized with 5 mg/ml of
Na,P,0,) and heated in a boiling water bath for
30 minutes, followed by drying with hot air for
5 minutes. This procedure destroys about 40 +
5% of the total alkaloid N-oxide formed, but
milder methods of oxidation give incomplete re-
action or difficulties in the following steps. The
resulting completely dry N-oxide (or an aliquot
of the original extract air-dried at 80°C) is taken
up in 1.0 ml isoamyl acetate, treated with 0.125
ml acetic anhydride, and heated 2 minutes in the
boiling water bath. After being cooled to room
temperature, the solution is treated with 1.0 ml
ofa solution of absolute ethanol : 2096 BF,-meth-
anol: p-dimethylaminobenzaldehyde (50:5:1)
and heated at 56-59°C for five minutes. Absor-
bance is read against a blank which passed
through all the reactions, at 561.5 nm, after di-

1987]

lution to 3.8 ml with anhydrous acetone; further
dilution with acetone gives a linear decrease in
absorbance. At room temperature absorbance
reaches a maximum several hours after the last
reaction, about 5% above the reading immedi-
ately after it, and then diminishes; it may be
maintained for many days at 4°C. Each micro-
gram of PA in the original aliquot contributes to
the absorbance 0.05 (N-oxide) or 0.03 (total base)
(standard curves with echinatine and lycopsa-
mine samples, Fig. 3G or E, provided by J. A.
Edgar). Determinations were routinely done in
racks of 40 13 x 100 mm glass test tubes, always
doing total base and N-oxide determinations in
the same batch to equalize the initial aliquot
sampling and give an internal control on abnor-
mal results. Separate pipets and bulbs should be
used for each reagent in the series; water must
be rigorously eliminated from the test tubes be-
fore the Polonovski reaction (Ac;O-isoamyl ace-
tate).

PAs with a carbonyl group conjugated with the
unsaturated ring (7-ketones, or 9-aldehydes or
carboxylates, such as Fig. 3M-O) or with no dou-
ble bond in the ring did not give any blue color
with this method. Dihydropyrrolizines (with the
nucleus of Fig. 3M, N) without carbonyl conju-
gation could be directly determined on aliquots
to an accuracy of about + 10% using the final
reaction in the series. They were usually de-
stroyed in the first reaction step (oxidation) if
present in the extract.

NEPHILA BIOASSAY

Fr apne were bioassayed for predator-de-
fense compounds, following observations of
a and Lewinsohn (1982, 1984),
with natural populations of the giant tropical orb
spider Nephila clavipes (L.). This predator, com-
mon from December through July in Campinas
(the same period as that of maximum ithomiine
density in the same places) and present in small
numbers year round, cuts out of its web, alive
and unharmed, any field-captured ithomiine that
is introduced naturally or experimentally, after
contact with any part of body or wings. If the
butterfly struggles excessively (ithomiines gen-
erally remain quiet, awaiting liberation), it may
be attacked or wrapped in silk, but the spider
does not suck it (Vasconcellos-Neto & Lewin-
sohn, 1982, 1984). Nephila spiders are among
the most abundant, effective, and aggressive po-
tential predators of flying insects in Campinas
and throughout the more seasonal Neotropics,

BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY

367

and can be safely regarded as very important in
the evolution of predator defense mechanisms
in the Ithomiinae. Vertebrate predators have not
been used yet in the bioassay, but they were shown
to reject an itl by Brower & Brower (1964),
to reject almost all ithomiines by Haber (1978),
to reject two ithomiines by Coimbra-Filho (1981),
and to accept only fatty abdominal contents of
ithomiines in Brazil by Brown & Vasconcellos-
Neto (1

Living panado of the palatable nympha-
line butterfly Biblis hyperia (Cr.), readily at-
tacked and eaten by the spider, were painted with
aqueous solutions, suspensions, or emulsions of
fractions or compounds (corresponding to 0.5-
1.5 butterfly, 2 g fresh weight of leaves, or 0.2 g
fresh weight of flowers), covering the entire body,
legs, antennae, and wing bases to at least half the
radius, and let dry. Still alive, they were thrown
into part of the web of an adult, non-satiated,
and healthy spider (as judged by web structure),
from a distance of 0.5-1.5 m on the side opposite
the spider, between 1200 hours and 1500 hours
on a warm sunny day. The spider normally ad-
vanced immediately to attack the butterfly (de-
lays of up to 5 minutes can occur if the spider is
“dozing” or distracted). If the Biblis was punc-
tured, sucked, and then cut out or wrapped in
silk by the spider, or (to still its struggles) was
bitten, wrapped in silk, cut out, and taken to
another part of the web and sucked, the test was
regarded as negative. All such tests were repeated
at least twice with different spiders and different
individuals of Biblis. A positive test consisted of
the spider’s drawing back from the animal, at
times inspecting various parts with its sensory
palps but not biting or sucking, and eventually
cutting out the living butterfly by breaking all
necessary strands (usually cutting them with the
third pair of legs), manipulating the entire insect
and letting it drop unharmed. This test, usually
repeated at least three times with at least two
Biblis, was regarded as strong evidence for effec-
tive predator-deterrent compounds in the ex-

e
-
°

t.
Biblis is common year round in most parts of
the Neotropics, not especially fast- or high-flying,
and readily attracted to fermenting baits. It is
brightly colored black and red, regarded as an
“incipient mimic" by the Browers (1964; Brow-
er, 1969; Brower et al., 1971). Its larva feeds
upon Tragia, an urticant euphorbiaceous vine,
and the adult possesses dorsal scent glands in the
abdomen that it displays in evident defensive

368

behavior when handled. Nevertheless, it is readi-
ly eaten by Nephila and thus is an ideal substrate
for the bioassay of chemical fractions. Alterna-
tive organisms used with similar success in the
bioassay include some pierids (Eurema) and
Heliconius erato, also eaten by the spider (Vas-
concellos-Neto & Lewinsohn, 1982, 1984).

CHEMICAL RM nena AND
COEVOLUTION A
SIRE INTERFACE

HOST PLANT UTILIZATION PATTERNS

In the principal field sites around Campinas,
SP, at least 42 species of Solanaceae are used (or
potentially should be used, by analogy with other
regions) by larval Ithomiinae (Figs. 5-8). A fur-
ther four species have been inspected frequently
but show no signs of usage: Capsicum praeter-
missum Heiser & P. G. Smith, Lycianthes ran-
tonnetii Carr. ex Lescuyer, Solanum wendlandii
Hook. f. and Solanum americanum Mill. Many
additional species occur in nearby field sites (at
least 15 more species of Solanum) or in gardens.

Six of the 25 species (in 17 genera and 11 of
the 13 neotropical tribes) of Ithomiinae known
from the Campinas region (Figs. 5, 9) can be
regarded as only occasional tors (the two Mel-
inaea, Episcada philoclea, Pseudoscada quadri-
fasciata, and Hypoleria adasa and goiana), these
possibly do not find enough adequate host plants
available to establish permanent populations. At
least 15 additional species in as many genera
occur in healthy populations within 200 km of

of the 19 species regularly present feed only on

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Apocynaceae-Parsonsieae (Fig. 5). Thirty-six
species of solanaceous food plants have been re-
corded for 14 of the remaining 17 species and
may be confidently predicted for the rest (Fig. 5);
no species is monophagous but only Mechanitis
(and possibly Pteronymia) could be called poly-
phagous (the rest are best regarded as quite nar-
rowly oligophagous, especially in the chemical
sense, as will be shown below)

This pattern is a microcosm of the general pic-
ture of food plant usage by the Ithomiinae (see
Drummond & Brown, 1987), comprising about
400 species X species interactions. Monophagy
or narrow oligophagy is the rule at the present
level of information, not only for local popula-
tions but also for whole genera. A summary of
all these data (Table 2) with chemical informa-
tion added from many sources in the literature
(with preliminary verification in this work, see
Table 3) shows a reasonable specificity in the
interactions between Ithomiinae and Solanaceae,
at least at the level of genera of ovipositing fe-
males and host plants. Most of the ithomiine
genera are known at present from only one or
two Solanaceae genera or subgenera of Solanum;
even generalists like the common and ubiquitous
Mechanitis show a strong epos for only two
subgenera of Solanum in most s

The records are still I. ices Ith-
omiine ovipositions are not commonly ob-
served, and 23 of the 52 genera are still repre-
sented by three or fewer confirmed food plant
records. The possibility cannot be discounted that
the specificity patterns shown in Table 2 are pri-
marily due to lack of adequate information. Con-
fident patterns of food plant specificity are seen
only in the well-documented interactions of the

—

Known and expected food plant relationships of Ithomiinae in the Campinas region, São Paulo.

FIGURE 5.
Heavier e: have all been verified in interbreeding populations occupyi

diameter circle Campinas and Barão

ng various habitats within a 5-km

Geraldo; note prevalence of oligophagy and some polyphagy

e
(Mechanitis, Prittwitzia, Pteronymia). Neither Ithomiine tribes nor Solanaceae genera are in evolutionary order
(see Table 2 for this), but have been arranged so as to simplify the figure

FiGURE 6. Solanaceae cal southeastern Brazil inc iir in Campinas, São Paulo unless otherwise indi-
itianum S. s

cated): s us m.—A. S. m
robustum. —F. S. LUE
authors' names, see Figure 5.

wartzia

E 7. Solanaceae of southeastern Brazil “ee nee
otanic o des Janeiro). —
ansia suaveolens. — N. Br

GUR
mansia candida (cultivated; Jardim B
Miers (Jardim Botánico, Rio de Janeiro). —

m. —C. S. capsicoides. — D. S. aculeatissimum. —E.

—G. S. an ei “H S. megalochiton. —]. S. pseudoquina St. Hil. For

—J. Solandra grandiflora (cultivated). —K. Brug
L. Markea (Dyssochroma) ehe e (Sims)
unfelsia australis. —O. Vassobia

breviflora. — P. Acnistus arborescens. — Q. micis ipee alum. —R. Cestrum intermedium (Joinvile, Santa ‘Ca a-
tarina).

1987] BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY 369

Known ( m; ) and presumed (——————-—- ) food-plant records
for Ithomiinae in the region of Campinas, Sao Paulo, southeast Brazil
(23* s, 48° w, 600-1,000 m)

ITHOMIINAE FOODPLANT

TITHOREINI:
TITHOREA harmonia pesudethra Butler PELTASTES peltatus (Vell.) Woodson
NADENIA vi Mi

(NEW TRIBE): TEMNAD violacea Miers

AERIA olena olena Weymer PRESTONIA coalita (Vell.) Woodson

APOCYNACEAE

PRESTONIA dusenii (Malme) Woodson

PRESTONIA acutifolia Schum.

METHONINI:

METHONA themisto (Hii BRUNFELSIA uniflora (Pohl) D. Don

BRUNFELSIA pauciflora (Cham. & Schldl.)
Benth.

(NEW TRIBE)
PLACIDULA euryanassa (Feld. & Feld. )'™ —
ku suwa BRUNFELSIA australis Benth.
MELINAEINI: SSDS LTT A
MELINAEA ethra (Godart) ~ "c ess DATURA stramonium L.
NI n. ~~ P
MELINAEA ludovica paraiya Reakirt -— “T NEL BRUGMANSIA suaveolens (Willd.) Sweet
SSI na T BRUGMANSIA candida Pers.
SS
ITHOMIINI TM. 7 SOLANDRA grandiflora Sweet
ITHOMIA d drymo Hübn TUS
iind y» ~ TY ie MARKEA (DYSSOCHROMA) longipes (Sendtner)
~~ Miers

ITHOMIA agnosia — Hewit
ikani D' Must.

nec PHYSALIS neesiana Sendtner
ee ACNISTUS arborescens (L.) Schldl.
PITYCHES eupompe (Geyer) ———— VASSOBIA breviflora (Sendtner) Hunziker

RIPOIHEBIS- eur Lea n Me reisen ee CAPSICUM flexuosum Sendtner
HYPOTHYRIS ninonia daeta (Boisduval) NICANDRA physaloides (L.) Gaertner
CYPHOMANDRA sciadostylis Sendtner

S A CYPHOMANDRA divaricata Sendtner
0-98 ff

CYPHOMANDRA fragrans (Hook.) Sendtner

MECHANITIN
sp T1 cetoides (Ros
alb.) pal

E

lida Godm. & de

X hippodamia (Fabr.) CYPHOMANDRA crassicaulis (Ortm.) Kuntze «

MECHANITIS polymnia casabranca Hnsch: 4 LYCOPERSICON esculentum Miller E
NS SX SOLANUM mauritianum Scop. ie

NN SOLANUM granuloso-leprosum Dunal u

MECHANITIS lysimnia lysimnia (Fabr.) SOLANUM concinnum Schott ex Sendtner Ds
aS AS SOLANUM megalochiton C. Martius °

`< n

SOLANUM murinum Sendtner
g gioi Jera (thoes Felde SOLANUM swartzianum Roemer & Schultes

na ae SOLANUM melongena L.

A NN KN.
NW `

UN SOLANUM paniculatum L.

(NEW TRIBE) E P "
PRITTWITZIA hymenaea hymenaea VM S SOLANUM asperolanatum Ruiz & Pavón
(Prittw.) SOLANUM robustum Wendl.
EPISCADA philoclea (Hewit ) m SOLANUM variabile C. Martius
N
SOLANUM insidiosum C, Martius

EPISCADA carcinia Schaus w \
MN
` SOLANUM brusquense Lyman B. Smith & Downs

PTERONYMIA carlia Schaus SOLANUM arcuatum Sendtner

SOLANUM viarum Dunal

GODYRIDINI: SOLANUM capsicoides All.
PSEUDOSCADA quadrifasciata Talbot wmm, SOLANUM atropurpureum Schrank

SOLANUM jatrophifolium Dunal

PSEUDOSCADA erruca (Hewitson)

HYPOLERIA adasa (Hewitson) wa. SOLANUM aculeatissimum Jacq.

DIAJ

HYPOLERIA plisthenes D
HYPOLERIA goiana D'Almeida — Í

SOLANUM pseudocapsicum L.

SOLANUM caavurana Vell.

CESTRUM schlechtendalii G. Don

MCCLUNGIA salonina salonina CESTRUM laevigatum Schldl.
ewteson) CESTRUM sendtnerianum C. Martius

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

>
x
E
z
=
=
T
Q
=
<
z
=
°
en
E
u)
<
=
Q
<
Z
<
o
O
N
|
Z.
z
°
x
m

ANNALS OF THE MISSOURI BOTANICAL GARDEN

“QS. U iii É
P mh AQ; :
lite.

4 (4
«6:14 Y Y ` 1 @ M h
" C» OCA aN ‘ | )
"3v >

= j

1987]

genera Tithorea, Elzunia, and Aeria (with Apoc-
ynaceae: Parsonsieae), Methona, Melinaea,
Thyridia, Mechanitis, Hypothyris, Ithomia, Dir-
cenna, Pteronymia, and Greta. At a higher level,
almost all members of the last two tribes show
collectively a strong specificity to Cestrum and
surprisingly also to the distant section Geminata
of Solanum. It is probable that more work will
provide more evidence of the restriction of Na-
peogenes to Lycianthes and confirm the nascent
patterns in Placidula, Scada, Epityches, Oleria,
and Callithomia, but this is mere speculation at
this time.

At the species level, Mechanitis females tend
to divide up the local Solanum, resulting in min-
imal overlapping of oviposition (see Drum-
mond, 1976; Haber, 1978; Vasconcellos-Neto,
1980, 1986), but the divisions do not coincide
with simple taxonomic lines in the plants. They
also vary from area to area and are not correlated
with larval choice or survivorship in experi-
ments, indicating an unstable ecological deter-
mination of the partition.

Though the butterflies may be able to recog-
nize their food plants, biologists have greater dif-
ficulty in Solanaceae identification, even to the
genus level, which may result in some spurious
patterns in Table 2. In 1969, a top Solanaceae
taxonomist identified for me a specimen from
near Rio de Janeiro as a Capsicum on the basis
of its cleft anthers, but both S. Knapp (pers.
comm.) and I regard this plant as Solanum (sect.
Geminata) caavurana. The “diagnostic” trait in
this case may have been an artifact of the drying

BROWN SOLANACEAE/ITHOMIINAE CHEMISTRY

373

method used for the specimen. If such evolu-
tionarily distant and chemically distinct taxa
could be potentially confusing to an experienced

expert, how will they appear to the average field
biologist who, having just observed a female ith-
omiine oviposit on a (presumably solanaceous)
bush with glabrous entire leaves and no flowers,
faces the maze of complex and confusing tax-
onomy in the family? This plant could be placed
preliminarily in over 30 genera, five with over a
hundred species and one with over a thousand.
Thus the “solid” data base for host plant usage
in Table 2, from which a number of completely
unlikely records have already been purged, may
yet suffer fundamental modifications with fur-
ther work.

COEVOLUTION AND PHYLOGENETIC
DIVERSIFICATION

An attempt to relate the phylogeny of the Ith-
omiinae with that of their larval food plants, such
as was done with reasonable confidence and re-
sults for the nymphaline tribe Heliconiini (Ben-
son et al., 1976), has met with very little success
(Drummond, 1986; Brown & Drummond,
prep.; Drummond & Brown, 1987). Only ie
broadest pattern can be seen in the progression
from Apocynaceae to Solanaceae (with Gesne-
riaceae in the middle to provoke the imagina-
tion). The most primitive ithomiine genera to
use Solanaceae (Athesis, Methona, Olyras, and
Melinaea) feed on the genera regarded as highly
advanced in this family. Two large tribes (Me-
chanitini and Napeogenini) run their food plant

—
FIGURE 8.
K-M, P, R. Second instar larvae, about 4-7 mm long.—
S-DD. Fourth or fifth instar larvae, an 20-35 mm long. —EE. Prepupa and p
pseudethra a ru sp.), Pirac
Napeogenes sulphurina Bates Hans Sp., a vine), Ipojuca, Pernamb —

apa.—F, DD
P eee eii canaria Brown & D'Alm

anjuba, Goiás.— B, U. Aeria olena (Prestonia pé teg Campinas,
CC. G

m sp., note simple ES poe Para.—E, |

Juveniles of Ithomiinae with food plant and locality. A-J, N-O. Eggs, about 0.6-1.0 mm high. —

. Adult ovipositing MA AA J. Vasconcellos-Neto). —

. A. Tithorea harmonia
SP. —C, AA.
arsauritis xanthostola

n Sendt.), Santa Teresa, Espirito

onymia hemixanthe (Feld. & Feld.) (Solanum SUPE Brev vantheru?) Ubata, Bahia.—I, R. Me-

i m 102 (Solanu
(Solanum asperum), Linhares, Espírito deii
originally reported as “Forsteronia sp."), Jar

(4 vir mop,

anf grandiflora D. Don), Barinitas, Vene

Bryk (Prestonia? sp.), Belém, Para.—
hona ME reip sus australis), Pinos on E —

E. H cidit eucled laphria
S ssp. Mone ulei (D m.) Cuatr.,

e V are from watercolors by Emily Fountaine,

uela.
d in the British Museum (Natural Histon. Solanologiss are requested to photograph and rear through

jn le adult or to preserve in alcohol any eggs (with ridges as

arvae of these types found on identifiable

plants and send them to B. Drummond or K. Brown for uec Ed and registry in the data bank for Table 2.

374

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

TABLE 2. Approximate number of known interactions (species x species) between Ithomiinae and their larval
food plants, with data on plant chemistry (Figs. 1, 2) and on attraction to pyrrolizidine alkaloid sources.

Plant Families: Genera*” (no. species)

. Solanaceae
Attraction
to: Solanum (1000)
= 8 Yo . § >= =
2 s g Se ERR Y Š a = @ Q2 X d^ =<
ii ša Gg Sa NEFS P t Setasacss SzS Š Š
sp Se sSssS SISESE ES S S š EQgES Ise, SSS Š g
Bese SESS DOE SOQ DOE E Š E š DP 22293923 $53 § Š
SEs gre PERSIS SS S $ S goggas ggs i $
Aš us ef83 GSS asa $85 O 5 FSFE aldàd 333 Š š
Alkaloids (Fig. 1)
Ste roids,
a PAs (3E-I) ? === ABFLH ---- BL ABCL BL BC?L C? ABC BC BCP C BC C? LC BCE
onstit
uents Other C C Fig. 2)
Ithomiinae: Cardiac
Tribes, Genera!“ glycosides,
(no. species) XYZ WY ~ RTUWXYZ -R? Z RWXY YQ? Q QSY QY QYZQR XY XY RWXYZ XY
A. Tellervini*
Tellervo (Ay l I 5
B ithoreini
Elzunia (2) l ? 4
dais dd l 1 16 1?
C. (N
shi pe 2 3 8
D. (New Tribe)
Athesis (1) l l 1
E. Methonini
Methona (7)" l l 12
F. (New Tribe
Placidula (1) 2 3 1? 6
G linaeini
Olyras (2) 2 2 1? 1
^ utresis (2 2 2 I? 1
Melinaea (10) 2 2 1 9
H nitini
Thyridia (1)" 2 3 6
(2) 1 3 l
la l 3 l 3
Forbestra ( l 3 l
Mechanitis (5)"' l 3 3 1556 1 7 2 1? |!
Ll. Oleriini
Hyposcada (8) 2 3 3 1
New genus (3) l 1 l
Oleria (30)" 2 3 148 3
J. peogenin
AKA 2 3 2 l 2
"es a (1) 2 3 1? 1
Napeogenes (18)" 2 3 1? 8
Garsauritis (1) 2 3 4
Hyalyris (16) 2 3 5
Hypothyris (16) 2 3 1 2413 1
K. Ithomiini
Miraleria (1) 2 3 l 2 1
Ithomia (21)" 2 3 2 2 11 3 5
L. Dircennini
Callithomia (3)^ 3 3 13 I
Velamysta ( 3 3 1
New Ge 2 ? 1
Dircenna (7)" l 3 14 17
Hyalenna (5) 3 3 1
M. (New Tribe)
Ceratinia (5) 3 3 3 3
Ceratiscada (3) 2 3 2
rittwitzia (1) 2 3 3 1? 5
Episcada (15) 2 3 4
Pteronymia (41)" 3 3 25 3 3 2

1987] BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY 375

TABLE 2. Continued.

Plant Families: Genera*” (no. species)

Solanaceae
Attraction
to: Solanum (1000)
a 5 doa S —
Ithomiinae: š E g is E š Š Rn T 9 a oe e "S cu e
Tribes, Genera** s oo g a Ss E 3 > S E S Ss A ago fsa & s
(no. species) Ups DugE Bog ESS ES 8 § — 3 "EF s» Vo ss Š s
ee be SESS CSR RSs ° vk Ë E 38 BS 8 SS39 Us S sss E 3
esse DID Es -1 20 SSE 2 Š BEE 232353 BID š $
= = = e I Fi =
S3 as BESS GSS ase § S55 Š § FSFE AgI S83 Š Š
N. Godyridini
Godyris (10)^ 3 3 5
Pseudoscada (6)" 2 3 5
Hypomenitis (6) p. 3 2
reta (10 2 3 l l 16
Hypoleria (1 1)" 2 3 1 6
Mcclungia (1) 2 3 3
Heterosais (3)" 3 3 3

a Families are in approximate order of advancement and genera of Solanaceae in order according to Hunziker
(1379), from primitive to advanced.
^ In addition to Nicotiana (see note i, below), important Solanaceae genera available to Ithomiinae in tropical
America for which no food plant records are yet known include J dicii Athenaea (but nid uc used by
Epityches and pos mia), e Saracha, Iochroma, Salpichroa, Jaborosa, Lycium, Grabowskia, Trianaea,
essea, Metternichia, Petu Fabiana, Nierembergia, oS Protoschwenkia, Schwenkia, Melananthus,
Ppbounedia, oet y Sin za Streptosolen, and Het
—weak attraction usually including both sexes, 2—males a attracted, 3—males strongly and females
regulary attracte
ribes and genera follow the order of Mielke and Brown (1979) as modified by Brown (in prep.), from
* os to advanced (including genera within each tribe).

* No food plants have yet been recorded for the following genera of Ithomiinae (number of species in paren-
theses). * = Andean cari usually of y qd us cloud forests, though Roswellia extends out into the foot-
hills.— Tribe D. *Roswellia (1), *Patricia (2).— Tribe G. Athyrtis (1).— Tribe H. Paititia (1).— Tribe I. *a new
genus near Hyposcadi i — Tribe J. 7 (1).— Tribe K. *Pagyris (1).— Tribe N. *Dygoris (1) and *Ve-
ladyris (1). Total, 9 genera with 10 s

f The recent divisions of these ina prod into a number of smaller Es (Hunziker, 1979), most of them
used by the same Ithomiinae, are here included under the collective older names.

e Restricted to the Old World tropics, regarded by some as closer to the Denials than the Ithomiinae (see
Ackery & Vane-Wright, 1984).

^ Food plant patterns are constant from the northern to Edd southern extremes of the range of the genus, usually
from Central America to southern sl Aes. of 18 gen

! Ovipositions of Mechanitis polym asabranca were B S ENE in Campinas, São Paulo, on "iaa ad
Capsicum annuum L. and feral Nicotiana (rosette only), but the ipe did not grow on these pla
j The published structure for *hopeanine" (1E) is biogenetica t

with the data presented in the einge phe: (Iyer, 1978); no alkaloids of ` Brunfelsia are lau under
investigation.

phylogen ds ( d dbutterflies does not lead to a fruitful parallel. The patterns
use Por due) veins plants). And although for the Ithomiinae are complex, and the arrange-
the most morphologically advanced ithomiine ment of the plant groups in Figure 5 so as to
genera concentrate on the relatively advanced produce a maximum parsimony scheme (lowest
Cestrum, they use equally well the supposedly number of crossing lines in the middle) has pro-
primitive section Geminata of Solanum (Table duced some strange and thought-provoking
2). That the neotropical Ithomiinae representa proximities between genera normally widely sep-
widely diversified butterfly group (51 genera in arated; note especially the positions of Brunfelsia
13 tribes) could help explain why a comparison and Cestrum.

of their host plant utilization patterns with those While the Solanaceae indeed may have rep-
of the 65 rather homogeneous species in no more resented a new field for adaptive radiation of the
than ten genera of the single tribe Heliconiini Ithomiinae in the New World, it is probable that

376

the plant family was already very diversified
chemically and taxonomically before the butter-
fly subfamily began to move onto it. A large
number of available genera (at least 24) are still
not known to be attacked by Ithomiinae (see
Table 2). Furthermore, the genus Brunfelsia, re-
garded as a relatively advanced member of the
Solanaceae, apparently entered the Caribbean
area and diversified greatly (as section Brunfel-
sia) before Methona, a relatively primitive ith-
omiine genus in a monotypic tribe, could evolve
as a specialist herbivore on it (Plowman, 1979,
p. 489). It is possible, however, that the extensive
speciation verified in Solanum and Cestrum
eventually may be related to their heavy use by
a variety of Ithomiinae (Table 2). This can be
investigated only by careful chemical analysis
and bioassay with larvae, Mines in parallel
twoi tly developin y lin
of species or populations Pie ‘show. specific in-
teractions.

urther considerations of the incongruities be-
tween phylogenies of genera of Solanaceae and
Ithomiinae are presented in Brown & Drum-
mond (in prep.).

1 1

O
n

BIOCHEMICAL COEVOLUTION

The taxonomic diversification of the Solana-
ceae has been accompanied by an impressive
diversification in secondary chemicals, leading
some to suggest polyphyly for the family as pres-
ently constituted. Indeed, no other plant family
can challenge the Solanaceae in having alkaloids
representing all four major biosynthetic path-
ways (lysine-ornithine, phenylalanine-tyrosine,
tryptophan, and steroid), an equal number of less
important pathways (nicotinic acid, anthranilic
acid, glycine, acetate), and a further set of aber-
rant or combined sources unique to the family
(leading to capsaicins, solanines, withasomnine,
and hopeanine) (Fig. 1), not to mention more
common amines such as choline, noradrenaline,
and hypoxanthine; some species also produce
very toxic peptides. As is often the case, these
alkaloid-rich plants are singularly poor in lower
(volatile) terpenes, but some Datura, Cypho-
mandra, and especially Cestrum and Solanum
section Geminata share a similar and abundant
pungent oil that may include terpenes. Diter-
penes and triterpenes abound in some Solana-
ceae, and nonalkaloidal steroids are represented
as saponins (such as diosgenin glycosides) and
withanolides/physalins, unusual steroidal lac-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

tones with marked biological activity (Fig. 2).
Flavonoids and coumarins also are documented
amply in diverse members of the family (Fig. 2;
Harborne & Swain, 1979), and further phenolics
and their glycosides (including tannins) are al-
most always found when sought in polar extracts.
The C,; alcohol solanesol is an unusual unsat-
urated linear nonaprenol (Fig. 2). Lacking are
reports of cyanogenic glycosides, cardenolides,
glucosinolates, nonprotein amino acids, and ir-
idoid glycosides, but their absence cannot be de-
finitively affirmed because few phytochemists
have apecttically sought them in the Solanaceae.

hus the p this family are well defended
chemically and attacked by rather few insect her-
bivores other than Ithomiinae (Table 1). The only
ones that seem to promote similar damage in
natural systems are generalist grasshoppers and
meloid beetles, perhaps molluscs in more cal-
cium-rich areas, and specialist chrysomelid bee-
tles. The plants also at times are defended ad-
mirably against vertebrate chewers by prickles
(Symon, 1986) but these have very little effect
on smaller invertebrates, who walk and chew
around them or build silken pads over them.
Other defenses (tough leaves, glandular tri-
chomes) are more effective against invertebrates
(Vasconcellos-Neto, 1980, 1986), as may be fur-
ther quantitative defenses (t amino-acids,
and resins) and a host of ecological strategies such
as phenology (see Vasconcellos-Neto, 1986) hab-
itat, growth form, seed dispersal by chiropte-
rochory, and encouragement of spiders and pred-
atory pentatomids on the leaves, among others
observed in Brazil.

When the generic preferences of the butterflies
are analyzed in terms of plant chemicals (Table
2), the associations seem to show patterns, though
the data are still chemically and biologically very
incomplete. There appears to be a certain ten-
dency toward chemical specificity, at times with-
out reference to taxonomic relationship (as in the
similarly potent-smelling Solanum sect. Gemi-
nata and Cestrum). It seems probable that female
Ithomiinae search for and find specific chemical
cues for oviposition on certain genera of Solan-
aceae and may be led to place eggs on unrelated
plants with similar chemicals. What else would
permit a Mechanitis polymnia female in Cam-
pinas in May 1981 to recognize a small Nicotiana
plant in Monjolinho as a potential Solanaceae
host, or to oviposit on Capsicum annuum in my
garden, especially when the larvae survived on
neither? (The first shares nicotine and the second

1987]

steroidal glycoalkaloids with a usual host, So/a-
num mauritianum.) Based on the patterns in Ta-
ble 2, some still unknown chemical may be pre-
dicted to set Solanum subgenera Bassovia,
Potatoe, and Lycianthes apart from all other So-
lanaceae (see Napeogenes, Oleria, and Calli-
thomia) and possibly associate Lycianthes with
the withanolide-elaborating genera (see Ithomia)
and even with Solanum sect. Geminata (Ptero-
nymia uses both).

Further support for some degree of chemical
mediation at the oviposition sector of the inter-
face comes from the broad geographical consis-
tency of the more specific relationships, at least
from Mexico through Costa Rica and Ecuador
to south Brazil for the 18 genera for which food
plants are known over this range (Table 2; Drum-
mond & Brown, 1987). This surely must have
been established over long evolutionary time in
developing phyletic lines of the butterflies. The
chemical proximity of disparate taxonomic
groups of plants, treated as interchangeable by
ovipositing females, also supports a chemical
mediation at the interface. In order to separate
this possible evolutionary component in food
plant usage from the ecological noise in the sys-
tem, careful comparative tests with free-flying
females must be performed under controlled
conditions, coupled with larval feeding prefer-
ences in multiple-choice tests, and thorough
chemical analysis of all potentially active com-
pounds in the food plants chosen or rejected.

The patterns broadly reflected in Table 2 and
supported on a local scale by Figure 5, however,
do not support any easily envisioned hypothesis
of mutual interaction and fine coevolutionary
adjustment between the two groups at a taxo-
nomic level. Although some regularity in chem-
ical cues for oviposition and chemical specificity
is suggested, the data are still very limited and
give little support for parallel diversification in
the two groups. Much more complete biological
and chemical information will be necessary be-
fore any claims of biochemical coevolution can
be advanced at this plant/herbivore interface. The
present information already seems to falsify many
attractive and conventional hypotheses.

In view ofthe fact that Ithomiinae populations

probably are controlled principally by certain
limiting adult resources (suitable humid open-
understory *pockets" and sources of PAs, see
below), they may exert very little, ifany, selective
pressure for chemical and phylogenetic diversi-
fication on their host plants. In any case, the

BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY

377

specific choices seem to vary appreciably with
local ecological conditions. Under a moderate
degree of Ithomiinae attack, some Solanaceae
even respond by vigorous new growth and flow-
ering (especially in Brunfelsia and in section Bre-
vantherum of Solanum). Under these conditions,
it is difficult to establish the most basic prereq-
uisites for “classical” coevolution between her-
bivores and host plants.

AN ALTERNATIVE HYPOTHESIS: SEQUENTIAL
COLONIZATION BY CHEMICAL ADAPTATION
(TABLE 4)

The results of chemical extraction, fractiona-
tion, and Biblis/ Nephila bioassay of 48 ithomiine
host plants are presented in Table 3. In the cases
investigated so far the Solanaceae have shown
in their leaves the compounds already reported
or presumed for each species, in good quantities.

If these chemical characteristics of the host
plants are combined with the utilization patterns
in Table 2, a most suggestive picture emerges, as
summarized in Table 4. Nothing is known of the
paleophytochemistry of these plants beyond the
global variation in secondary compounds seen
in widespread geographical populations of each
genus today, but it may be presumed that they
were diversified generically before Ithomiinae
began attacking them in the New World (since
the most advanced genera were attacked first)
and probably similarly protected chemically to
modern species.

It then becomes possible to trace a series of
small hypothetical advances in larval toleration
of food plant chemicals, each facilitated by pre-
adaptations on the existing host plants, which
lead to a complete picture of chemical specificity
as observed today (Tables 2, 4). The scheme is
clearly oversimplified but is attractive as a hy-
pothesis for the sequential colonization of plant
hosts, already diversified taxonomically and
chemically, by progressively adapting herbivores
(Futuyma, 1983). It is in excellent agreement with
the great diversity and apparent nonstorage by
herbivores of the Solanaceae chemicals and the
initial dependence of the Ithomiinae stock on
PAs (see below).

Thus, Apocynaceae-feeding Ithomiinae de-
pendent on PAs probably did not adapt sufh-
ciently to the great chemical variability in the
Parsonsieae (seen today also), which in some cases
repelled or poisoned larvae, in others left adults
unprotected against predators and deficient in

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

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380

PA-derived sex attractants and other phero-
mones (Fig. 3D). Adults increasingly exploited
PA sources (Boraginaceae leaves, Compositae
flowers) and may even have found some substi-
tutes for PA precursors in Solanaceae (Brunfel-
sia?). Since they were preadapted to bicyclic
8-carbon monoester alkaloids and their N-oxides
(both PAs and tropanes fall under this classifi-
cation), the generally well-protected, tropane-rich
Solanaceae represented an open niche. At least
two separate radiations could have moved onto
these plants (discussion of the move over onto
the Gesneriaceae must await further biological
and chemical study of Hyposcada and its food
plants; the move onto Brunfelsia, whose alka-
loids are still enigmatical, was probably a side-
track of one of the radiations). The line that col-
onized the tropane-containing Brugmansia
(represented by Placidula and Miraleria today)
also encountered in these plants the steroidal bit-
ter principles, to which they were also preadapted
through their experience with cardenolides in the
Apocynaceae. Tolerance of these bitter steroids
permitted further colonization of the many gen-
era defended by them, most of which also contain
tropane alkaloids, sometimes with N-oxides.
Utilization of Capsicum by this line also gave a
gateway, through its steroidal glycoalkaloids and
saponins (Table 2), to the immense and under-
utilized resource represented by Lycianthes and
Solanum, on which the more advanced genera
persist today.

The second radiation found tropanes initially
in Solandra and relatives, and thence in Cy-
phomandra, on which it adapted to tolerate the
steroidal glycoalkaloids also present there and

cada). As a final step, adapta-
tion to saponins and strong-smelling oils present
in some Solanum (especially section Geminata
and relatives) permitted exploitation of the
abundant Cestrum in the forest understory. In
all cases, tolerance of toxins encountered pre-
viously in the evolutionary history of each line
could be maintained as new enzymes were de-
veloped to detoxify progressively more effective
classes of plant defensive chemicals, in many
cases not correlated strictly with taxonomy and
evolution of the plants. Indeed, some of the more
primitive genera— possibly with a broader range
of biosynthetic capabilities, according to present
concepts of biochemical evolution in plants—
seem to have been the last to be colonized by
the most advanced Ithomiinae.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Persuasive evidence for this hypothetical sce-
nario (Table 4) can come only from extensive
studies of female oviposition preferences and lar-
val tolerance of food plants or chemical com-
pounds. Preliminary data (Table 2) and some
initial experiments are encouraging: Placidula
cannot eat Solanum, nor sun Aeria accept any
Solanaceae, but Mec/ n use tropane- and
withanolide-containing iim (Datura, Nican-
dra), and some Cestrum-feeders continue to use
Solanum and even Lycianthes. Experiments in
this area inevitably will encounter problems re-
lated to individual and local adaptation, lateral
radiation (as onto Brunfelsia), and loss of ances-
tral genes but should in the long run help to define
the viability of this proposed sequential expan-
sion of Ithomiinae through progressive chemical
tolerance to Solanaceae food plants.

HERBIVORES AS PHYTOCHEMISTS

The chemosensory apparatus of the average
insect is capable of detecting very small amounts
of specific chemicals, either through smell (an-
tennae) or taste (tarsal or buccal chemorecep-
tors). This highly specialized analytical instru-
ment transforms chemical structures into
recognizable electrical potentials through me-
chanical contact between compound and recep-
tor and uses these potentials to stimulate a va-
riety of response behavior sequences. It can be
fooled at the receptor end by substances of dif-
ferent chemical composition but similar confor-
mation and spatial relations among active bind-
ing sites. It is far more sensitive and accurate,
however, than the majority of the instruments
in the modern phytochemist’s laboratory. Its
specialization is selected over many generations
of advantages accrued by those who could rec-
ognize a few chemical compounds and thereby
compete better for higher quality food, defense,
or mates.

To the extent that ovipositing female Itho-
miinae seem t
ificity in host plant recognition (Table 2), which
involves use of chemoreceptors on the forelegs
("drumming" on leaves) and antennae, they can
serve as precise analytical tools for the phyto-
chemist. If some chemical patterns are already
evident on a very rough scale in the food plant
choices of the Ithomiinae (Table 2), these could
be used to suggest analytical methods for still-
uninvestigated Solanaceae and predict chemicals
in others that have undergone only preliminary
analysis.

1987]

BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY

381

E 4. Possible scheme for sequential colonization of Solanaceae by Ithomiinae through progressive

TA
preadaptive tolerance of secondary chemical classes.

Substances Tolerated

Step Ithomiines (genera) Plants Colonized

l Tellervo, Elzunia, Apocynaceae: Parsonsieae 3E-K: PAs and N-oxides
Tithorea, Aeria (Parsonsia, Prestonia, etc.) 3A-B: Cardenolides

2a Athesis, Placidula, Solanaceae: — 1C: Tropane alkaloids and

'raleria ‘apsicum N-oxides

2b Olyras, Melinaea Fearing a 2Q: Bitter steroids
Hyposcada Solandra PA precursors?

2c Methona Brunfelsia

3a Epityches, Ithomia Acnistus, Physalis, etc. 1C + much 2Q (varieties)

further Capsicum Some 1L: steroid alkaloids

3b Thyridia; New genus Cyphomandra 1C, IL, 2R: saponins

4a Napeogenes, Rhodussa, — Solanum subgen. 1L: glycoalkaloids
Oleria Bassovia, Potatoe (sect.

4b Scada, Sais, ieee UM 2R; 1A; possible 1C
Callithom

5a Garsauritis, H PIE Solanum (subgenera Leptoste- IL, 2R, 1A; occasional

yalyris monum and Brevantherum) nicotine, pungent oils

5b Mechanitis, Dircenna,
Pteronymia

6b Ceratinia, Pteronymia, Solanum (sect. Geminata), IL, 2R; 2Z: pungent oils

Godyris, Hypoleria, Cestrum

Pseudoscada, Greta

For example, it could be predicted that the
Solandra-Markea-Juanulloa group of genera,
which support the danaioid larvae of Melinaea
(Fig. 8K, V), might have some protective chem-
icals similar to those of Apocynaceae-Parson-

to be identified (cardenolides are strongly sus-
pected) and should be investigated. Similarly, the
strong-smelling essential oil of Solanum sect.
Geminata should be chemically very similar to
the like-smelling oil of Cestrum; this also seems
to have escaped chemical identification. With-
eringia could have been predicted to show phy-
salins by Ithomia, which also recognized these
chemical components with withanolide-struc-
tures in Physalis and divided its specificity among

c
pound; at least one major substance, possibly
with this type of structure, is present in the neu-
tral fraction (A) of two Capsicum species inves-
tigated (Table 3).

Do the genera Deprea and Athenaea contain
withanolides also? Start by testing some coop-
erative females of Ithomia and Epityches. Are

any other Solanaceae besides Datura/Brugman-
sia especially rich in scopolamine? One cou
begin with some Placidula females or perhaps
even larvae. If the latter were found on a Sola-
num or Cestrum, I would bet on the presence of
tropanes in the leaves. Are there glyco-alkaloids
in Dunalia? I might start by asking Hypothyris,
known only from Solanum, or perhaps even
Pteronymia, which specializes on the high-al-
kaloid (Bradley et al., 1979; Table 3) Geminata
section of Solanum. This analysis could be ex-
tended even to individual constituents once ap-
propriately biis populations of Ithomi-
inae have n identified. It may also give some
false UE fee these will hold even more in-
terest than the expected ones, pointing out new
types of plant components. As a preliminary sur-
vey, it should be strongly suggestive of certain
types of chemicals.

CHEMICAL CONTRIBUTIONS OF THE
SOLANACEAE TO PREDATOR DEFENSE
IN THE ITHOMIINAE
ITHOMIINE CHEMISTRY

Chemical investigation of the passage of So-
lanaceae toxins to adult Ithomiinae started with

382

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

TABLE 5. Summary of PA content and distribution, and bioassay of adult Ithomiinae and Danainae.
No. of spp.
Ana-
Tribe Genus In Genus lyzed Typical Species? Source? Host Plants Analyzed
ITHOMIINAE®
B Tithorea* l harmonia? VB 45 21/16
C Aeria" 3 3 olena: C 43, 44 31/20
elara G 10/15
E tho 7 5 themisto'e C 41, 42 36/27
F Placidula" l l euryanassa* C, J, SL 33 24/17
G ras” 2 l rathisk V 37 4/2
Melinaea” 10 7 ne RO 36 5/4
ludovica* M 36 50/1
menophilus RO 37 15/15
H Thyridia l l psidiï C 20, 21 10/10
Sais 2 l rosalia' G 19 12/12
Scada 6 2 reckia* RO 16 35/13
karschina ES 16 15/4
Mechanitis 5 5 polymnia' C 6-13 44/37
M 4 12/12
lysimnia'9 C 13-15 28/14
I Hyposcada 8 4 egra* M 36 11/1
Oleria 30 13 astrea BJ 12 4/12
RO 12 13/7
J Epityches l l eupom C 25-31 30/33
Rhodussa l l cantobrica RO 8 17/9
Napeogenes 18 8 cyrianassa ES 18 10/8
inachi RO 18 10/11
Garsauritis l l xanthostola M 13-14 48/20
Hypothyris 16 13 ninonia daeta* C 5-14 51/40
euclea’ C 9 22/22
(males attracted) M 10 16/12
V 10 17/14
K Ithomia 21 7 agnosia'* C 30, 31 29/38
L Callithomia" 3 2 lenea xantho* MP 16 18/21
Dircenna 7 4 ero* C 6-13 29/18
M Ceratinia 5 4 neso” BJ, RO 13 24/24
Ceratiscada 3 2 doto BJ 1? 6/7
Prittwitzia 1 l hymenaea" C l 10/12
Episcada 15 5 clausina ES l 17/14
carcinia C 1? 25/29
Hyalenna 5 l pascua“ J 7 1/9
Pteronymiq" 41 10 carlia* € l 31/17
dyris 10 3 zavaleta* RO 38 10/9
Pseudoscada 6 4 ca* C 39 31/28
reta 10 l andromica" V 38 4/4
Mcclungia l 1 salonina* C 39 11/11
H ypoleria 11 10 arzalia G, VB 39 8/4
Heterosais 3 3 nephele VB, RO 40 16/11
giulia V 40 7/4
DANAINAE
Danaus (Anosia) 2 2 gilippus' C Oxypetalum 25/16
Danaus (Danaus)^ 1 (S.A.) l plexippus C Asclepias 22/14
Ttuna* l ilione C I 6/5
Lycorea" 2 2 cleobaea C Ficus 12/11

1987] BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY 383
TABLE 5. Continued.
Individual PA Analysis (5/9)? Odes Fractions:
Average Maximum Bioas.
s Average PA conc. (% av. Ce average dry wi, am Fract. Re-
(mg) % N-Ox.) dry wt.) A (neut.) C (polar) F (fat) Tested sults!
60/75 0.3/0.3 (48/45) 0.9/0.6 8.4 8.4 13.9 Ex +
9/13 2.5/1.6 (55/55) 7.1/4.0 3.9/5.2 14.7/15.8 13.9/16.3 Ex +
9/13 1.3/1.1 (61/57) 1.9/1.9 NT
100/120 0.06/0.1 (41/48) 1.6/0.9 3.2/1.9 7.2/7.0 10.8/26.6 Ex +
54/65 0.2/0.2 (48/48) 1.6/0.9 Ex +*
110/110 1.2/0.3 (54/49) 1.8/0.5 NT
55/60 6.3/1.2 (54/49) 10.5/1.5 NT
55/60 0.6/1.3 (57/59) 2.5/— 5.9/3.7 7.2/9.6 15.7/13.9 Ex +
55/60 3.6/1.3 (50/49) 6.5/4.4 NT
64/70 2.7/2.2 (70/68) 5.1/3.9 3.5/4.3 9.3/7.1 12.3/20.7 Ex *h
25/27 3.6/1.6 (59/58) 6.8/3.6 7.0/7.2 12.7/7.4 20.7/21.6 Ex +
6/7 9.7/3.2 (47/54) 20.6/5.3 NT
8/9 13.1/6.7 (61/65) 20.0/10.4 NT
33/45 1.8/2.4 (60/60) 5.8/4.0 8.9/5.8 6.4/6.7 14.1/23.8 Ex, B, C +
September 5.6/6.1 5.4/3.0 9.7/18.9 A, F _
35/45 4.4/2.9 (58/57) 7.6/4.7 6.2 8.3 10.7 NT
25/35 3.8/1.8 (50/55) 8.8/6.4 Ex +
20/22 0.7/0.2 (51/48) 3.8/— NT
13/15 2.0/1.9 (53/58) 4.5/4.0 NT
13/15 4.0/1.8 (50/43) 8.2/3.4 NT
24/26 6.9/3.2 (56/58) 11.4/4.3 2.3 11.4 11.9 Ex +
18/20 3.1/3.3 (48/52) 8.6/6.1 NT
18/20 7.7/3.5 (55/56) 12.9/5.9 NT
13/14 5.4/3.6 (56/54) 11.7/6.2 NT
20/23 0.9/1.0 (53/53) 4.9/3.2 NT
25/27 3.3/2.8 (51/57) 9.2/6.5 3.5 6.3 21.8 Ex +
22/25 2.2/1.5 (48/44) 9.3/5.4 4.2 10.3 12.2 Ex +
20/23 0.8/1.5 (55/46) 2.9/4.5 5.3 10.8 10.4 NT
20/22 5.1/2.8 (58/60) 9.0/5.7 NT
14/16 5.4/2.0 (56/58) 12.9/7.1 1.9 10.6 23.2 Ex +
24/28 0.5/0.4 (52/53) 2.6/1.5 NT
40/45 1.7/0.9 (65/69) 4.3/2.4 4.0 9.9 12.3 Ex +
15/17 2.0/1.0 (55/56) 5.7/3.3 Ex +
14/15 1.1/0.5 (52/48) 1.9/1.2 NT
14/16 2.8/1.7 (57/60) 5.8/3.5 4.6 7.4 19.5 Ex, B, C +
16/17 5.7/5.3 (53/58) 10.0/7.0 A, F NT
17/18 1.6/2.0 (51/58) 6.0/5.2 Ex +
18/20 1.1/2.0 (47/52) —/3.4 NT
14/17 7.8/3.6 (58/62) 13.3/5.8 Ex +
33/33 1.5/1.4 (43/51) 2.9/3.1 NT
15/17 4.9/2.7 (57/54) 9.7/5.8 4.9 14.0 9.6 Ex F
15/17 7.2/5.0 (56/52) 10.6/6.0 NT
16/17 1.7/1.1 (70/82) 3.3/2.2 5.2 7.5 15.5 Ex +
14/15 1.1/0.5 (56/59) 2.3/0.9 NT
21/26 1.6/0.6 (55/53) 4.4/1 NT
20/24 2.7/0.8 (53/43) 5.3/1.7 4.8/— 9.6/— 8.2/— Ex +
80/75 2.1/1.8 (57/52) 4.5/4.5 NT
180/170 0.1/0.1 (50/51) 0.4/0.2 NT
160/140 1.1/1.2 (54/53) 2.5/2.1 NT
130/110 2.7/1.2 (58/55) 4.8/3.2 NT

384 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74
TABLE 5. Continued.
ANALYSIS OF PARTS:™ VINEIS Appendag
Species & Sex Basal Hairpencil Apical (Legs, esti ) Rest of Head
Mechanitis polymnia 1.13/88 4.15/64 0.14/100 0.26/77 0.60/70
casabranca (males) 1.70/100 1.10/100 1.03/65 0.52/92 1.03/72
1.03/58 1.70/58 0.83/51 0.98/97 1.43/75
2.60/85 4.40/100 2.10/48 4.20/77 3.40/75
Ithomia agnosia 4.50/81 — 1.38/78 7.50/68 12.5/69
agnosia (females) 0.69/42 - 0.63/45 1.60/55 2.00/68
0.70/67 — 0.75/48 0.72/100 1.05/44
“pockets”
Danaus gilippus (male) 3.10/63 1.00/55 0.73/49 2.00/57 1.75/55
(female) 1.45/50 — 0.47/42 1.20/46 1.50/51
Displaying Male
Rest of
Insect Spermatophore
Mechanitis lysimnia 3.09/61 8.50/54
3.32/59 8.85/55
3.06/52 9.75/49
0.78/54 2.40/64
4.20/52 25.3/55
6.40/56 17.4/54
Mechanitis polymnia 2.23/52 8.60/62
0.26/46 1.05/54
0.92/62 3.92/55
Aeria olena 3.00/50 12.3/67
3.60/69 10.0/72

* Authors’ and subspecies names are in Figure 5 or Brown (1985).

> See Table 3, footnote b. Sp din localities: BJ —
various parts of Rondónia. VB = Vila Bela, western

Bujaru, Para. ES = various parts of Espirito Santo. RO =

at ro
Numbers refer to the plants aire in Table 3. Italicized numbers indicate probable food plants or species

near to that number.

* Just-captured butterflies were cut up under 2 ml absolute ese As ml for heavier species) and after at

least one day's standing aliquots (usually 1/20, or 1/4 for componen

were assayed directly for total PA

and N-oxide, following Mattocks (1967, 1968) sien l Bingley (1968); see sol for details.

* See fractionation scheme in Figure 4 for oo
' See footnote g of Table 3 for bioassay co

* No secondary compounds from larval teh plants have been found stored in adult Ithomiinae. Additional

and Hyalyris oulita metella, 13; K Miraleria cymothoe, 31 +

39 and Hypomenitis dercetis, 39 and H. libethris, 39.

Rothschild (1973), who obtained only negative
results. In August 1978, 3,200 dry bodies of Me-
chanitis polymnia (75 g, representing over 300 g
fresh weight of insects) were mailed to Dr. Desiré
Daloze of the Collectif de Chimie Bio-Organique
in Brussels, where ant bioassays were used to
follow repellent activity in the fractionation. No
alkaloids, cardiac glycosides, or other interesting

33: L Velamysta pupilla, 17; N Dygoris dircenna,

active compounds, or even their degradation
products, could be found (D. Daloz
comm.,
sults suggested that the protective compounds o
adult Ithomiinae might be labile, degraded after
death or upon storage, and possibly unrelated to
the larval food plant poisons.

Total MeOH-H.O extracts of fresh ithomiines

1987]

TABLE 5. Continued.

BROWN—SOLANACEAE/ITHOMIINAE CHEMISTRY 385

THORAX
I ABDOMEN
ton Muscles Exoskeleton Fat Reprod. Organs Intestine
0.29/75 0.21/100 0.21/92 0.49/65 0.12/91 0.02/100 NR
1.18/42 0.92/48 1.82/91 0.81/66 2.54/99 0.63/89
1.02/66 0.63/65 0.50/100 0.19/69 0.60/100 0.14/56
4.10/67 4.40/68 6.60/72 3.20/100 4.10/92 4.00/97
Eggs
1.06/79 3.56/50 4.45/65 1.76/65 52.0/45 0.13/91 8.85/78
0.35/45 0.47/20 0.47/38 0.42/54 1.67/40 0.28/14 1.00/30
0.62/48 0.40/33 0.87/100 0.26/18 0.34/100 0.15/60 NR
1.90/57 1.20/50 2.20/56 0.73/65 4.20/58 Hairpencil 3.50/55
1.60/52 1.10/48 1.40/62 3.30/59 0.90/62 ggs 0.90/12
Pairs captured in copula
Male: Rest Spermato- Female: Rest
of Insect ore of Insect Abdomen Ducts
2.12/52 12.8/65 6.26/52 3.50/59
2.23/64 11.9/54 0.40/74 0.35/57
3.71/54 12.7/66 2.26/62 0.83/100
0.44/56 1.31/78 3.00/62 3.30/60
1.29/50 22.5/60 3.30/51 3.51/65
0.25/54 4.70/48 3.58/63 5.10/62
0.87/52 9.10/61 2.52/48 3.26/58
0.38/48 14.6/72 2.06/72 7.20/88
2.96/59 12.6/72 5.06/61 13.6/63
2.26/51 2.32/57 3.90/58 40.5/64
8.90/62 18.7/77 4.10/54 5.40/59

h Larvae of these species may be considered as aposematic in a and behavior.
PAs.

! Eggs of ee subspecies contained up to

i Larvae of these species feeding on the indicated plants did al contain any PAs nor did their extracts protect

9% of dry weight a

Biblis Medus predation by Nephila; possible exceptions are Methona themisto, Aeria olena and Tithorea har-

monia.

* Recently emerged adults of both sexes of these species reared from larvae on the indicated food plants, were

consumed without hesitation by Nephila, which in most cases had just cut out an adult of the same species and

sex gia in the

m

a are giv

field. Notice that almost all S
n as percent of dry weight of part/% of PA a -oxide.
usually papie pila PA present as A ropy bem DB dd confirmed. Especially

a m a an E aga in the food plants.

oteworthy values are printed in boldface type.

(prepared as in Fig. 4), when applied to the edible
Biblis hyperia, were at least as repellent to ur
ila spiders as were the live butterflies. re
pellency was then located in the alkaloid taa
B (Fig. 4) and also in the polar fraction c. Wh
these fractions were compared with the corre-
sponding fractions of the larval food plants, there
appeared to be no compounds in common. Fur-
thermore, all food plant extracts and fractions
tested in the spider bioassay were negative (Table
3). To put the final nail in the coffin, both sexes
of 30 species in 26 genera of Solanaceae-feeding

N-oxide value is high, it

Ithomiinae, reared from the larvae in the labo-
ratory on fresh leaves of natural food plant, upon
emergence from the pupa and introduction into
Nephila webs were promptly and enthusiastically
eaten (Table 5). In most cases, the Nephila had
just as efficiently rejected a field-captured adult
ithomiine of the same species and sex.

In all, 142 species of Ithomiinae in 45 genera
and all 14 tribes were examined in parallel with
48 host plants in 16 genera of Solanaceae, three
genera of Apocynaceae, and one of Gesneriaceae
(see Tables 2, 3, 5), both in chemical analysis

386

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Ithomiinae found in the Campinas region, interior of Sào Paulo state, EE Brazil. Missing are

FIGURE 9.
*Episcada philoclea (Hewitson) and TPseudoscada quadrifasciata, very similar to U.

e to
aa ninonia daeta (8%).
—O. Dirc

thona themisto (s š; on tree-lined city streets and in gardens
ian ,

7596 in some seasons). PD. Mechanitis lysimnia (1796). —
—L. Ithomia agnosia (5.
nna dero (upper D. celtina, lo ower D. rhoeo) (0.8%).—G. Hypothyris euclea (laphria
x nina, mixed Smal aod (0.4% but can be comm i
j i 's common in September). -

B. *Melinaea ludovica paraiya. —

—E. Mechanitis polymnia n (about 50% of
S. Mcclungia salonina
5%).—N. Aeria olena (2. 5%).—P. Prittwitzia

. The
over 20,000 captures, mostly for marking and recapture population studies. For BM order of genera and
species (primitive to advanced), see Tables 2 and 5; for authors’ names, see Figure 5.

and bioassay. In terms of the hypothesis of se-
questration of defensive chemicals by ithomiine
la arvae, these results were thoroughly disappoint-

ing; no important compounds were detected that
were shared by butterflies and their larval food
plants. All butterfly extracts were positive and
all plant total extracts and fractions were nega-
tive in the Nephila tests. However, all butterflies
showed a strong Mayer's test on the acidified
total aqueous extract, suggesting the presence of
some kind of alkaloids.

Since adult ithomiines, like danaines, are
known to be strongly attracted to sources of de-
hydropyrrolizidine alkaloids (Pliske, 1975a,
1975b) and use these at least for pheromone syn-

thesis (Edgar et al., 1976), the extracts were ex-
amined for these compounds, using on TLC
the iodine/Ehrlich test (de

presence of similar moderately polar PAs as the
major components of the alkaloid fraction B; in
all cases, this fraction was greatly augmented and
often became over 9096 of a single compound
after the zinc-reduction loop in Figure 4, indi-
cating that much of the PAs were present as a
single alkaloid and N-oxide in the butterflies, the
latter probably responsible for the activity seen
in fraction c. Pure pyrrolizidine alkaloid frac-
tions accounted for essentially all the activity

387

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1987]

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388

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

E ll. Feeding of dui at PA sources.
Bess ree rs, Amarais, m
flowers, Colorado, Rondóni

seen in the Nephila bioassay, protecting the adults
against predation by this spider.

he most common species in the Ithomiinae
communities in the Campinas area, Mechanitis
polymnia casabranca (Brown & Vasconcellos-
Neto, 1976; Vasconcellos-Neto, 1980, 1986; Fig.
9), was chosen initially for detailed chemical in-
vestigation, using the fractionation scheme shown
in Figure 4. e ethyl acetate extract (F) from
455 butterflies captured in Amarais in August
1982 (19.1 g dry weight, 13 of this as insoluble
tegument) weighed nearly 5 g and readily solid-
ified at 8°C to an off-white crystalline mass, in-
dicating nearly pure saturated triglyceride; its ex-
act composition is presently under investigation,
but it was negative in the Nephila bioassay —
probably quite nutritive, in fact. This extensive
storage of high-energy fat (2696 of dry weight at
the height of the winter dry season) reflects the
long (up to six months) reproductive diapause of
these species (May-October; Vasconcellos-Neto,
1980) and also helps to explain the advantages
of the learned predation behavior of the tanager
(Brown & Vasconcellos-Neto, 1976), which
squeezes this fat out of the abdomen of the but-
terflies. It is interesting that the fat reserves fell
to only half of August levels in butterflies ex-
tracted in late September; the butterflies are
long-lived and have few exogenous resources i

ward (here confirmed) and more diffi-
culty in capture as the days become warmer
Brown & Vasconcellos-Neto, 1976).

The total alkaloid extract B, isolated after the
zinc reduction loop (which increased its weight
from 60 to 260 mg equivalent yield from 19 g
butterflies — 1.496 of dry weight) showed on TLC

—A. Mechanitis polymnia casabranca on Trichogon
s, SP. — B. Mechanitis lysimnia elisa (Guérin) on Eupatorium ape baile hii

a single major spot (90%) that gave a positive I,/
Ehrlich test for PAs. The 60-MHz NMR spec-
trum of this noncrystallizable fraction (Fig. 10)
showed it to be almost exclusively a 60: 40 mix-
ture of lycopsamine and intermedine (Fig. 3E, F)
compared with a spectrum of pure reference ly-
copsamine contributed by J. A. Edgar (Fig. 10C).
Very minor impurities due to other dehydro-
pyrrolizidines could be seen in the NMR spec-
trum. The alkaloid fraction co-chromatographed
with authentic lycopsamine (donate J. A
Edgar) on TLC in four different systems, giving
indications of latent separation of the isomers in
some solvent mixtures.
he NMR spectrum of the total alkaloid frac-

tion showed no signs of the “methylene enve-
lope” or quaternary methyl signals typical of ste-
roid alkaloids of Solanum, nor of the anomeric
protons of glycosides, nor of the N-methyl groups
of solanines, nicotine, and tropane alkaloids (Figs.
, 10). This supports nonsequestration of al-
kaloids from the larval food plant. The spectrum
also indicated that most of the butterflies ob-
tained their alkaloid from a single source; the
structure and the mixture strongly support flow-
ers of Trichogonia gardneri, common in Amarais
and intensively visited by M. polymnia (Fig. 11A;
Table 6).

When the zinc reduction was not performed,
the much diminished alkaloid fraction showe

—

he NMR spectrum of the whole
fraction was perum complex, indicating several
additional components of diverse structures in-
cluding saturated PAs, but still showed no signs
of the presence of other classes of alkaloids.

1987]

In expansion of the work on M. polymnia pre-
liminary fractionation gave PAs representing
various mixtures of the five isomers of a single
structure (Figs. 3, 10) in up to 1396, and in Scada
over 2096 of dry weight in some individuals, in
all 141 additional Ithomiinae species indicated
in Table 5, many captured in the Campinas re-
gion but also sought in more distant places for
comparison and to verify the generality of the
phenomenon. No PAs were seen in the Zn-re-
duced extracts of any of the food plants tested
(summarized in Table 3).

ITHOMIINE DEFENSE

That th the prot ti of adult Ith
predation by Nephila is due to PAs was con-
firmed by feeding 200—400 ug of echinatine
N-oxide (Fig. 3J) in dilute honey solution to new-
ly emerged adults of Mechanitis lysimnia and
Pseudoscada erruca (which normally were eaten
by Nephila); within an hour, the butterflies were
routinely rejected by the spiders (experiments
performed together with J. R. Trigo). Reared but-
terflies kept alive for many days did not biosyn-
thesize any protective chemicals; protection was
lost within a day after death ofa butterfly rejected
by the spider, again indicating the instability of
the PAs. Very fresh field-captured Ithomiinae
often showed no PA and could be consumed by
Nephila, but in general the adults seemed to be
able to accumulate sufficient defensive com-
pound from different sources in their environ-
ment within a very short period after emergence
from the pupa.

The strong dependence of adult ithomiines on
PA sources, including for their courtship pher-
omones, perhaps made it inevitable that they
should also retain the PAs for their defense. The
lack of storage of Solanaceae defensive com-
pounds, already verified in early investigations,
increased the probability of alternate defensive
compounds in the Ithomiinae—labile com-
pounds not detectable in dead insects. The uni-
versal rapid cutting out of Ithomiinae from

ephila webs suggested that such alternate de-
fense substances did not derive from the widely
variable larval foods but from a more homoge-
neous adult food source. In the only other re-
ported case of a lepidopteran being cut out from
spiders’ webs— Utetheisa ornatrix, an aposemat-
ic day-flying arctiid whose larvae feed on Cro-
talaria and pass sequestered PA diesters on to
the adults, where they are also used in phero-
mone synthesis— it has been shown that PAs are

.

BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY

389

baise: for this deterrent activity (Conner et
1981; Eisner, 1982).

The Solanaceae poisons have thus been ab-
solved, at least for the time being, from partic-
ipation in the unpalatability of adult Ithomiinae
to their most dangerous predators. It seems pos-
sible that exceptions will be found to this, es-
pecially as more predators are incorporated into
the bioassays and more butterflies and food plants
into the chemical analysis. However, the PAs
and especially the biologically active (hepato-
toxic, tumor-inhibiting) lycopsamine-group
monoesters and their N-oxides should be re-
garded as the principal, perhaps nearly universal

inae

tiidae moths (Ctenuchinae, Pericopinae, and
Arctiinae), whose adults sometimes inherit them
from the larvae but inevitably seek them out at
the same sources visited avidly by the Ithomi-
inae, and similarly use them for defense and
pheromone synthesis. Indeed, lycopsamine and
its stereoisomers have been found in wild pop-
ulations of essentially every species of Danainae
and Ithomiinae investigated (Edgar, 1982; Table
5), though their presence is erratic in the moths.
These compounds seem to represent a very ef-
fective “ancestral predator defense" that has been
retained in diverging phyletic lines up to the pres-
ent (Edgar, 1975) in spite ofa variety of habitats,
food plants, and behaviors.

Adult Ithomiinae sequester their PAs from a
variety of sources: decomposing borages and
composite flowers (Eupatorieae) especially, but
also orchids, Crotalaria, apocynes, and less tra-
ditional PA-containing materials. These are
inevitably abundant wherever Ithomiinae occur
in numbers.

The proponderance of PAs in adult Ithomiinae
defense could also help to explain the cryptic
coloration and behavior observed in most ith-
omiine larvae (Fig. 8), notable exceptions being
Methona on Brunfelsia (Fig. 8W, X), the feeders
on Apocynaceae (Fig. 8T, U), Melinaea with
similar larvae (Fig. 8K, V) feeding on Markea,
Juanulloa, and related genera, and a few showy
larvae scattered on other plants. Newly emerged
ithomiine larvae eat their eggshells (which con-
tain PA derived from their mother; see below)
and immediately move to the underside of the
same leaves (if not already there), usually ac-
quiring a cryptic coloration. The fact that most
ithomiine immatures are cryptic translucent
green, closely matching their substrate, and feed

[Vor. 74

ANNALS OF THE MISSOURI BOTANICAL GARDEN

390

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BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY

1987]

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392

on the underside of leaves (Fig. 8) may indicate
rapid metabolism of the Solanaceae poisons
(though cryptic and toxic insects are not rare); at
least the poisons are not carried through to the
adults, whom they would not protect in any case,
and do not seem to be stored in the larvae. Apo-
sematic but nontoxic insects are rare (except for
obvious Batesian mimics that diverge from their
taxonomic relatives), the main case being auto-
mimics in the Browerian sense (Brower et al.,
1971; Rothschild, 1979). For this reason the
strongly aposematic larvae in a few ithomiine
genera may incorporate or synthesize some sorts
of unpleasant substances that have yet to be iden-
tified.

In previous work reported without details, a
lycopsamine/intermedine mixture was found in
Hypomenitis dercetis and Oleria makrena from
northern Venezuela (Edgar et al., 1976) and re-
garded as primarily precursorial to pheromones
(only males were analyzed) rather than as defen-
sive. Later work (Edgar, 1982) emphasized the
potential defensive role of these compounds, here
confirmed

SELECTIVE PA STORAGE

When male and female Mechanitis polymnia
were separately extracted in late September 1982,
some interesting differences were observed, in-
dicating that careful work on this chemical in-
teraction should always maintain the sexes apart
in analysis. At least in September, at the begin-
ning of the reproductive season, the females con-
tained much more fat and PA than males (Table
5). That this difference is closely related to the
respective reproductive tasks is supported by the
previous pheromone study (Edgar et al., 1976)
and by analysis of Mechanitis eggs. A raft of 31
M. polymnia eggs, weighing | 1 mg, was extracted
with 2 N H,SO, and directly reduced with zinc,
alcalinized, and extracted with CHCl,-MeOH.
This showed on TLC the presence of abundant
lycopsamine/intermedine mixture, perhaps as
much as 1% of the fresh weight or 0.1 mg. All
other Ithomiinae eggs analyzed also showed the
presence of appreciable PA (Table 5). The bright
white Mechanitis eggs, laid in rafts of 5-100 on
top of the host plant leaves, with which they
contrast vividly (Fig. 8I, N, Q), could be de-
scribed as a true collective display of aposematic
insects, just ie the adult assemblages in dry
season **poc

As ei in PENIS 5, male Ithomiinae gen-
erally accumulate more PAs than the females;

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

indeed, in many species they are the principal or
only sex found on the sources (Table 2; Pliske,
1975a, 1975b). The females of most species seem
to get the majority of their PAs in the spermato-
phores received from males during mating, which
is repetitious (Ehrlich & Ehrlich, 1978); these
small sacs often have 20-50x the PA concen-
tration as the rest of the male (Brown, 1984,
1985).

Both males and females are able to selectively
distribute the collected PAs to different tissues,
including tegument and wings (possibly by re-
gurgitation with fat as a wetting agent) and es-
pecially to reproductive parts (pheromone glands,
spermatophores, ovaries; Table 5), thence in fe-
males to the eggs.

Males attracted to Heliotropium baits placed
in relatively PA-poor areas and females attracted
to displaying males showed lower average and
maximum PA content than random samples of
the same populations, indicating an "appetite ef-
fect" that obviously would be highly adaptive in
these organisms.

More complete information on PA storage, use,
and distribution in Ithomiinae may be found in
Brown (1985).

SOURCES OF DEHYDROPYRROLIZIDINE
LKALOID MONOESTERS

In the same paper in which he predicted that
PAs would be used for defense in the Ithomiinae,
Edgar (1982) predicted that they would be found
in the nectar of Eupatorieae, in view of the heavy
dependence of Ithomiinae on these plants and
their characteristic occurrence in the Boragina-
ceae, also frequently visited by Ithomiinae and
source of precursor for the pheromone (Fig. 3D)
as well as of the most attractant esterifying acids
for ithomiine males (Pliske et al., 1976). Indeed,
many genera and most females are more strongly
attracted to Eupatorieae flowers than to Heli-
otropium (Table 2), the difference being espe-
cially pronounced in Mechanitis and allies (Fig.

Extraction of the flowers of 16 species in four
genera of the Eupatorieae actively visited by Ith-
omiinae in the field led to the isolation and iden-
tification ofa variety of PAs but usually only one
isomer and structure in each species (Table 6;
see Figs. 3, 10). Alkaloids of this structure were
also found in Heliotropium and Tournefortia
flowers (Boraginaceae-Heliotropoideae) often
visited by Ithomiinae, as well as in the leaves of

1987]

Removed by pollinator AN

N

Concentrated
in nectary

Translocated
to flowering
tips & leaves

IVA
P WA

Synthesized
in roots

FIGURE 12.

various Heliotropium species (Table 6), but not
in Mikania (Eupatorieae), Cordia (Boraginace-
ae), or several other flowers sporadically visited
by Ithomiinae. When the plants were analyzed
for PAs by parts (Table 6), the highest concen-
trations always appeared in the nectaries of still
unopened flowers, suggesting that they might be
a reward to specific PA-seeking pollinators; the

BROWN —SOLANACEAE/ITHOMIINAE CHEMISTRY

tegument
Used for pheromone
synthesis

Concentrated in
reproductive organs,
especially in the

spermatophore

Transferred d
to female in
mating

393

Pheromones define communities and
territories, attract females

Spread out

over tegument,
\ concentrated in
UN i

N NN reproductive
NN \ NN. tissues, and
Nec A N placed on

` Nee! eggs.
e

\ KS
VERNY

Flow of dehydropyrrolizidine alkaloid monoesters in natural ecosystems.

concentration was often 2-4% of dry weight of
the whole flower.

In open vegetation or poor soil areas with few
Itl ii t, Eupatori ] yields

of less pure isomers in the flowers and showed
dramatically reduced seed set, even though pol-
lination was effected by other groups that also
depend on PAs (Danainae and Arctiidae). Areas

394

with few E r only very seasonal species
showed scattered and transient Ithomiinae pop-
ulations, whereas in areas with abundant Ith-
omiinae there were always common Eupatorieae
in flower throughout the year, especially Tricho-
gonia (Fig. 11A; see also Pliske, 1975a). It is
evident that this mutualistic relationship, pro-
foundly affecting the reproduction and abun-

the interaction, in a coevolutionary picture prob-
ably much stronger and more stable than that of
the Ithomiinae with the Solanaceae.

FLOW OF DEHYDROPYRROLIZIDINE
ALKALOIDS IN NATURE (FIG.

The analysis of PA monoesters in different parts
of plants and butterflies in various physiological
and reproductive states (Tables 5, 6; Brown, 1984,
1985), made possible by the selectivity and sen-
sitivity of the Mattocks-Bingley assay (Fig. 3),
permits a diagram to be drawn of the synthesis,
Tu, use, and eventual dissipation of these highly
(Fig. 12)
Thus the PAs are probably synthesized in the
roots of Apocynaceae, Boraginaceae, and Com-
positae-Eupatorieae (diesters are also elaborated
by Compositae-Senecioneae and Leguminosae:
Crotalaria), young plants show highest concen-
trations in the roots and even mature plants show
a bimodal concentration distribution between
flower heads and roots. The compounds also may
be concentrated in the leaves when this will give
important protection against herbivores; in a few
cases, these leaves are attacked by specific in-
sects, including larvae of some Danainae, Ith-
omiinae, Ctenuchinae, Pericopinae, and Arctii-
nae among the Lepidoptera, and a variety of
Hemiptera and Coleoptera. Many of these spe-
cific herbivores store and use the compounds
directly for defense or adult reproduction, where-
as others excrete them.

As the plant comes into flower the alkaloids
are translocated to the flowering tips and then
into the nectaries, where they guarantee attrac-

lies, the first three pantropical (with scattered
species in north temperate areas), the fourth neo-
tropical and the last cosmopolitan. The alkaloids
are also retained in the seeds to deter predation.

In the frequent case of selective attraction of

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

male butterflies to the PA sources, these spread
the PAs over their tegument for predator defense
and channel them into pheromone synthesis
glands (Fig. 3), which sometimes are formed only
with PA stimulation (Schneider et al., 1982), and
other reproductive tissues, especially spermato-
phores. The spermatophores are transferred to
females during mating and the PAs are similarly
spread out over the tegument and channeled into
reproductive tissues, eventually being deposited
on the eggs for protection. Newly hatched larvae
consume the eggshells but quickly lose their PAs
unless they find themselves on leaves that con-
tain them.

This chemico-ecological flow scheme (Fig. 12)
opens ample perspectives for the localization,
selection, and cultivation of PA-producing plants.
Indicine N-oxide (Fig. 3K) and at least one iso-
mer (Fig. 3J)
now in advanced clinical testing; some E upato-
rium in the right ecologico-evolutionary setting

ight become important sources for these c
pounds (Table 6). The free bases, however, and
especially the diesters, are very hepatotoxic and
carcinogenic, representing a serious problem in
human and veterinary medicine. Knowledge of
the flow of PAs in nature should be useful for
the control of both plants and PA content in
natural and agricultural systems. It may also help

when used as herbal teas or fortifying salads, to
avoid permanent liver damage to the unsus-
pecting consumers.

CONCLUSIONS, SYNTHESIS, AND
PERSPECTIVES

The integrated ecological, phylogenetic, and
biochemical investigation of the Solanaceae/Ith-
e interface, with the collaboration of
Nephila clavipes, a inre spider that is a major
LUN predator of the butterflies, has shown

m Although there exists a reasonable and
geographically widespread specificity in the usage
of 19 genera of Solanaceae by Ithomiinae larvae,
especially at the level of host plant secondary
chemistry, there is very little evidence for parallel
phylogenetic diversification of these interacting
groups over evolutionary time; in general, ad-
vancing phyletic lines of butterflies use ever more
primitive hosts. The New World Ithomiinae seem
to have colonized the already generically diver-

1987]

sified Solanaceae through sequential preadapta-
tion to, and encounter and tolerati ion of pro-
hei

a.
y

ssively m
foc plants. This has undoubtedly affected the
distribution, population structure, habit, habitat,
phenology, and exceedingly diversified and vari-
able chemistry of these plants, but it should not
be called **coevolution."

(2) The variable and diversified Solanaceae
toxins are not stored by larval Ithomiinae and
do not protect newly emerged adults against the
spider predator, which rejects field-caught indi-
viduals. Adult Ithomiinae depend heavily on de-
hydropyrrolizidine alkaloid monoesters for de-
fense and reproduction. These are sought out and
sequestered from decomposing Boraginaceae and
especially from a constant source, stabilized by
mutualistic interaction: flowers of Compositae-

upatorieae, which place in their nectar a single
chemical structure (usually as only one of five
different stereoisomers) to attract the pollinators
that need these alkaloids. This intimate relation-
ship has undoubtedly determined many aspects
of morphology, physiology, population struc-
ture, distribution, abundance, and the highly
i aliens chemistry of these plants and their
pollinator

Eus additional problems that have aris-
en during this research, presently under active
investigation with similar methodology, include:

(1) The special relationship of the Ithomiinae
Tithorea and Aeria to Apocynaceae-Parsonsieae,
which sometimes contain PAs that may be stored
by the aposematic larvae of these genera (re-
search under way with J. R. Trigo).

(2) The additional aposematic larvae of Ith-
omiinae, which feed mostly on tropane-contain-
ing plants and Brunfelsia (though some Solanum
are also included), and which may be storing
effective predator deterrents from the food plant
and, in the case of Methona (which contains al-
most no PAs in the E possibly passing them
on to the adult butterflie

(3) Reasons for the qM nonstorage (and
perhaps nonstorability) of most Solanaceae tox-
ins by herbivores.

(4) Possible participation of further com-
pounds, volatile or unstable and derived from
PAs or similar precursor, in defense of adult Ith-
omiinae.

(5) Physiological or behavioral mechanisms
for the spreading out of PAs on the tegument of
the butterflies and their use in synthesis of var-
ious pheromones.

BROWN -SOLANACEAE/ITHOMIINAE CHEMISTRY

395

(6) The great diversity and variability of tox-
ins in both the Apocynaceae and the Solanaceae
used by larval Ithomiinae and the importance of
these in relation to evolution, oviposition, larval
feeding, survivorship, and reproduction in their
usual herbivores and in other potential enemies.

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NEW TAXA OF RUBIACEAE FROM VENEZUELA

JULIAN A. STEYERMARK!

ABSTRACT

r new taxa of Rubiaceae are described from Meridie a SOMME apurense, Faramea
Ps sei Psychotria buntingii, and Psychotria subimbrica

Coccocypselum apurense Steyerm., sp. nov. TYPE:
Venezuela. Edo. Apure: Dtto. Pedro Ca-
mejo, Cano El Cabello, 16 airline km NW
of Mata de Guanabana, between the Río
Meta and Río Cinaruco, 6?19'N, 68?19'W,
75 m, 27 Feb. 1979, Gerrit Davidse & Angel
Gonzalez 15825 (holotype, MO; isotype,
VEN)

erba radicans, caulibus 1.5-2.5 mm diam. glabris;

vel subobtusis majoribus 6-7 cm longis 3-4 cm latis,
adultis supra costa adpresso-pilosa atque marginibus
minute adpresso-ciliolatis, pilis 0.1-0.3 mm longis
munitis, ceterum glabris, subtus santas Donius
ervis latera-
libus utroque latere ca. 7 ULLA subtus im-
pressis; inflorescentia ca. 12-flora subcapitata pedun-
culata; pedunculo 4.5 mm longo glabro; calycis lobis
inaequalibus nen lanceolatis 2-2.5 x 7mm;
corollis coeruleis 5 mm longis extus glabris.

Creeping herb with rooting glabrous stems 1.5-

.5 mm diam. Stipules subulate, to 6 mm long,
glabrous. Leaves petiolate, appressed-pilosulous
on upper margins, otherwise glabrous; leaf blades
ovate, shortly acute or subacute at apex, obtuse
or subobtuse at base, the larger ones 6-7 cm long,
3-4 cm wide; adult leaf blades appressed-pilose
adaxially on midrib, minutely appressed-pilo-
sulous on margins with hairs 0.1—0.3 mm long,
elsewhere glabrous, abaxially glabrous; young leaf
blades pilose abaxially on midrib and lateral
nerves; lateral nerves ascending, 6 or 7 each side;
petioles 8-13 mm long. Inflorescence ca. 12-
flowered, subcapitate, the heads 8 mm wide, 5
mm high, pedunculate; peduncle 4.5 mm long,
glabrous. Flowers sessile; bracts subtending flow-
ers linear-ligulate, obtuse, 3 mm long, 0.5 mm
wide, glabrous; hypanthium campanulate, 1.5
mm long, 1.1 mm wide, glabrous; calyx lobes
unequal, linear-lanceolate, acute, the larger pair
2.5 mm long, 0.7 mm wide, the smaller pair 2

mm long, 0.3-0.5 mm wide, sparsely ciliolate to
glabrescent; corolla blue, infundibuliform, 5 mm
long, glabrous without, tube 2 mm long, 1.2 mm
wide, glabrous within, the lobes ovate-oblong,
subacute, 2 mm long, 1.5 mm wide, papillate-
puberulent within. Anthers slightly exserted, lin-
ear, 2 mm long. Style exserted, 6 mm lon

This species resembles the newly described C.
croatii Steyerm. but is readily distinguished from
that taxon by the longer petioles, very short pe-
duncles, less conspicuous, shorter hairs of the
ciliate-margined leaf blades, shorter stipules, in-
florescences with more flowers, and fewer lateral
leaf veins that are less arcuate.

Faramea cazaderensis Steyerm.,
Venezuela. Edo. Tachira: Dtto. Lobatera, La
pude 2,000 m, 22 Jul. 1983, H. van der
Werff & R. Ortiz 5457 (holotype, MO; iso-

type, VEN).

Sp. nov. TYPE:

rutex 4-metralis, ramis juvenilibus quadrangula-
vato-oblongis apice abrupte obtuse
acutis basi subcordatis vel omen 6.5-13 cm longis
3.5-7 cm latis prominente ne vatis, nervis lateralibus

pedicellis 2.5-4 mm longis; calyce hypanthioque 5.5
mm longo, Rd e ein urceolato l mm longo; calyce
tubuloso 4.5 m
dentibus qeloideis acuminatis 0.3-1 mm longis, deinde
uno latere fiss

Shrub 4 m tall, glabrous throughout; branches
quadrangular. Stipular sheath shallowly and
broadly semilunar, 3 mm long, 2 mm wide, end-
ing in an awn 2-3 mm long. Leaves subsessile,
subamplexicaul, oblong or ovate-oblong, abrupt-
ly obtusely acute at apex, subcordate or rounded
at base, 6.5-13 cm long, 3.5-7 cm wide, prom-
inently nerved; lateral nerves 9-12 each side,
nearly horizontai or F widely spreading at an angle
of 5-1 5°, ele imp
anastomosing 3-6 mm before the margin, ter-

adaxially,

' Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

ANN. Missouni Bor. GARD. 74: 398—400. 1987.

STEYERMARK—

VENEZUELAN RUBIACEAE

399

1987]

tiary veins prominently reticulate, marginal
nerves somewhat thickened; petioles 2- m
long. Inflorescence terminal, umbellately cy-
mose, 2-2.5 cm wide, 1.5-1.7 cm high, 12-15-
flowered, pedunculate; peduncle erect, 1.7—4 cm
long, | mm wide; primary axes 4-5, ascending,
5-7 mm long, 0.7-0.9 mm wide, each bearing 3
flowers. Flowers on pedicels 2.5-4 mm long, 0.5—
0.7 mm wide; bracts subtending axes and pedi-
cels deltoid-ovate, acute, 0.5—0.8 mm long. Calyx
and hypanthium 5.5 mm long; hypanthium ur-
ceolate, | mm long; calyx tubular, 4.5 mm long,
shallowly and unequally dentate, teeth broadly
deltoid, acute to acuminate, 0.3-1 mm long,
eventually splitting on one side. Corolla purple
(in bud), abruptly acute, 7 mm long

w
3

Paratypes. VENEZUELA. EDO. TACHIRA: Dtto. Lo-
batera, La Cazadera, 7?55'N, 72°18'W, 1,600 m, 24
Jul. 1983, van - Werff & Ortiz 5587 (MO, VEN);
o, Quebrada Cazadero, 16 km NW of

1981, Liesner & Guariglia 11668 (MO, VEN).

This species is well marked by the subam-
plexicaul leaves and relatively few-flowered um-
bellate inflorescence. It may be distinguished from
F. sessilifolia (Kunth) A. DC. by the umbellately
disposed, smaller inflorescence of 4 or 5 axes,
each of which is only 3-flowered, the obtusely
acute leaves with fewer pairs of lateral nerves,
and by the tubular, longer calyx which becomes
split on one side. Further, the habitat of the new
taxon in the cooler montane forests of the Andes
of Táchira is in contrast with that of F. sessilifolia
in the lowlands of the Orinoco, Rio Negro, and
Madeira river basins.

Psychotria buntingii Steyerm., sp. nov. TYPE:
Venezuela. Edo. Zulia: Dtto. Perijá, alrede-
dores de la Estación Hídrológica Aricuaisá-
Pie de Monte, 9?35'N, 72?53'55"W, en zona
de bosque siempreverde, 100-250 m, 25
Feb.-3 Mar. 1982, G. S. Bunting, G. Pan-
apera & H. Lobo 10876 (holotype, MO; iso-
types, JBM, VEN).

Suffrutex 0. .3- metralis, ramis minute erg ue sti-
5. 5m

n-

gis; foliis elliptico-lanceolatis apice subfalcato- acumi-
M basi acutate angustatis 4. 5- 8.5 cm longis 0. 9-1.9

ute puberulentibus, nervorum lateralium axillis
domatiis barbellatis, ceterum. glabris, subtus punctis

latis cymoso-umbellatis gracillimis sub anthesi 0.4 cm
alto 1.2 cm lato sub fructu 1.2 cm alto 2 cm lato, 12-

17-floris, axibus primariis tribus filiformibus 3-7-flor-

is; pedunculo filiformi 1.6-2 cm longo 0.5 mm lato

dense puberulo; floribus pedicellatis, pedicellis 0. 5-0. 8
is

alyce 1.2
mm lato; corolla (immatura) 3. 5 mm longa.

Slender subshrub; branches minutely tomen-
tellose with numerous minute cystoliths. Ter-
minal stipule and those on next lower node lan-
ceolate, acute, up to 5.5 mm long, 1.5 mm wide,
glabrous or glabrate except for the ciliate mar-
gins, prominently marked with pale cystoliths

near margins, nodes below brown-fimbrillate on
the stipular scar. as nom elliptic-lanceolate,
subfalcately apex, acutely narrowed
to the base, 4.5-8.5 cm long, 0.9- 1.9 cm wide,
4'4—4V5 times longer than broad, glabrous adax-
ially, minutely puberulent abaxially on midrib
and main lateral nerves, otherwise glabrous, the
surfac tely brown-dotted, axils of the nerves
ae barbellate with domatia, margins
sparsely ciliolate; lateral nerves 7-8 each side,
ascending at an angle of 45°, faintly anastomos-
ing 1.5-2 mm from margin; petiole very slender,
4-7 mm long, moderately tomentellose. Inflo-
rescence axillary or terminal, cymosely umbel-
late with 3 main axes, 12-17-flowered, 0.4 cm
high, 1.2 cm wide in anthesis, 1.2 cm high, 2 cm
wide in fruit, pedunculate, the main axes with
two additional shorter central axes 1 mm long;
main axes filiform, the lateral 3-7-flowered. Pe-
duncle 1.6-2 cm long, 0.5 mm wide, densely
puberulent and conspicuously marked with cys-
toliths. Flowers pedicellate, pedicels filiform, 0.5-
0.8 mm long, 0.3-0.4 mm thick; bract subtend-
ing pedicel subulate, 0.2 mm long. Calyx and
hypanthium 1.5 mm long, the hypanthium tur-
binate, 0.5 mm long, 0.5 mm wide, tomentellose;
calyx tube 0.5 mm long, 1 mm wide, lobes gla
brous throughout except for ciliolate margins,
longer than the tube. Corolla white (in bud) cy-
lindric-infundibuliform, 3.5 mm long, glabrous
without, sparsely pilose within the tube, sparsely
marked with cystoliths, tube 1 mm long, lobes
2 mm long. Fruit (immature) elliptic-oblong, 3
mm long, 2 mm wide, sparsely puberulent.

This species resembles the pubescent P. hor-
izontalis var. glaucescens (Kunth) Steyerm., from
which it is readily distinguished by the minute
calyx and hypanthium, shorter, filiform pedun-
cles and pedicels, much smaller inflorescence,
and narrowly elliptic-lanceolate leaf blades with
barbellate domatia in the abaxial nerve axils.

400

It is a pleasure to associate this new taxon with
the name of Dr. George S. Bunting, who has
contributed greatly to our knowledge of the flora
of the state of Zulia in Venezuela.

Psychotria subimbricata Steyerm., sp. nov. TYPE:
Venezuela. Edo. Miranda: Parque Nacional
Guatopo, laderas pendientes pedregosas a lo
largo del Rio Santa Cruz de Rio Grande,
lado occidental de la carretera, al sureste de
Los Alpes, 10?4'45"N, 66?29'30"W, 400 m,
31 Oct. 1981, Julian A. Steyermark & Bruno
Manara 125483 (holotype, VEN).

Planta subherbacea l.5-metralis; foliorum laminis

subtus glabrescentibus, nervio medio iege minute
puberulo; inflorescentiae. peduncu lo | 1-6 c m longo,
1 H 1 hel hv

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

petiole 3-8 x 5 mm. Inflorescence axillary in
upper leafy axils and in leafless middle axils, fas-
ciculate with 1-3 elongated peduncles 1-6 cm
ong, glabrous, trichotomously branched at ma-
turity with the central longer axis up to 13 mm
long and two lateral shorter axes up to 7 mm
long, or the peduncle with two lower axes and
an upper trichotomously branched portion; 2
bracts subtending the base of the principal axes
naviculate, lanceolate to ovate-lanceolate, acute
to acuminate, 7-9 mm long, 4-5 mm broad near
base, glabrous. Flowers in small compact groups
subspicately arranged, each group subtended by
imbricate, ovate, obtuse, glabrous bracts 6.5 x
4 mm; flowers numerous in an inflorescence, 3
or 4 of them ene bs a larger, glabrous,
group. Calyx lobes

ypanthio
glabro.

Stem subherbaceous, simple, hollow, leafy be-
low, 1.5 m tall. Stipule spreading with revolute
margins, broadly ovate-oblong, subacute, 6-10 x
6-8 mm. Leaves broadly obovate or elliptic-ob-
ovate, eens and shortly acute to shortly acu-
minate or obtusely acute at apex, conspicuously
narrowed to the decurrent base, the lamina 21-
25 cm long, 10-15 cm wide; lower midrib mi-
croscopically appressed-puberulent, the middle
and upper portion with two parallel lines of pub-
erulence along the margins of the midrib; abaxial
surface glabrous to glabrescent; adaxial surface
glabrous; lateral nerves 13-16 each side, arcuate-
ascending, arising at angle of 15—20?*, strongly
sulcate adaxially, elevated abaxially; decurrent

5, narrowly ligulate, unequal, subobtuse, cilio-
ate, 3-3.5 x 0.4-0.6 mm; calyx tube and hy-
panthium 2 x 1.5 mm, glabrous; | scale-like
squamella, oblong-triangular or lanceolate, acute,
in the sinus between 2 calyx lobes; corolla tu-
bular, 7 mm long, glabrous except for the barbate
orifice. Anthers exserted, oblong, 1.2 mm long.

tyle 3.5 mm long; style branches filiform, 1.3
mm long, puberulous.

—

The new taxon differs from P. humensis Stey-
erm. and P. decurrens Steyerm. in the glabrous
hypanthium, more branched inflorescence with
longer peduncules, glabrous, subimbricately ar-
ranged bracts and bracteoles, and minutely pu-
berulent lower midrib of a glabrescent abaxial
leaf surface.

SIX NEW SPECIES OF NEOTROPICAL LAURACEAE!
HENK VAN DER WERFF?

ABSTRACT

Among recent collections of Lauraceae and older collections received on loan from various her ba ria
de e

grandiflorum van der Werff,
placements of these new species are discussed.

Lauraceae, comprising about 40 predomi-

small, inconspicuous flowers. Genera are often
hard to recognize and nearly all are in need of
modern treatments.

During identification of the large number of
unidentified Lauraceae in MO and among the
various loans I received, several undescribed
species were found. Of those, the following six
are here described as new

AIOUEA AUBLET

Aiouea, recently monographed by Renner
(1982), comprises about 20 neotropical species
and is best represented in Brazil, Venezuela, and
the Guianas.

Having seen specimens of most species attrib-
uted to Aiouea, it seems quite likely to me that
Aiouea, as circumscribed by Renner (1982) and
accepted here, is polyphyletic. Most of the South

a typical aspect, with large, many-flowered inflo-
rescences, flowers with rather long pedicels, leaves
fal ipia green. ane itwigs with smooth bark. The

ral A p quite different but
w to Aiouea because of their 1 nine 2-celled an-
thers; they do not fit in the other, better defined,
neotropical genera with nine 2-celled anthers.

it is quite well possible that the Central American
Aiouea species are derived from local Ocotea or
Nectandra species which have lost two of their
four anther cells. Variation in the number of an-
ther cells in Aiouea lundelliana Allen, a species
excluded from Aiouea by Renner (1982), has been

We
and P. westphalii van der Werff, are described. Difficulties with the generic

reported earlier (van der Werff, 1984). A study
of the wood anatomy could probably answer the
question whether these Aiouea species share more
characters with the South American Aiouea
species or with Central American Ocotea or Nec-
tandra species.

Aiouea inconspicua van der Werff, sp. nov. TYPE:
Mexico. Vera Cruz: 0-2 km S del campa-
mento Hnos. Cedillo, rumbo a Río Alegre,
por la desviacion al E, Hidalgotitlan, 140 m,
22 Apr. 1974, Brigada Dorantes 2929 (ho-
lotype, MO; isotypes, MO, BM, UC). Fig-
ure 1

Frutex vel arbor parva. Ramuli tenues, teretes, ju-

ores minute tomentelli, vetustiores glabrescentes.
Folia alterna, lanceolata (8-11 x 2-3 cm), c
acuminata es caudata, subtu

ac

tomentellae vel glabrae, ines » raeter nervorum ax-

illas barbellatas glabrae, sicco olivaceae praeter nervos

virides. Inflo rescentiae axillares, minute to mentellae,
Flore

a a 9 mm
longa: 6 exteriora introrsa, 3 interiora extrorsa et basi

tia. Ovarium label stigmate crasso sessilique.
Bacca globos m diametro, basi cupulae planae,
in une ae attenuatae insidens, tota exser-
ta.

Shrub to 7 m tall. Twigs terete, glabrous; young
twigs minutely tomentellous. Terminal buds
slender, greyish pubescent. Leaves alternate, 8—
11 cm long, 2-3 cm wide; lamina chartaceous,
lanceolate, with numerous small gland dots, the
base acute, plane, the apex acute or caudate, with
a slender tip 1 cm long, the margin thickened

! I thank the curators of A, BM, BR, GH, U, UC, US, and VEN for the often large loans that I received. Dr.
J. Dwyer kindly checked the Latin. John Myers made oe eee The designation CORO is used for the

herbarium of the Proyecto Flora Falcón in Coro, Venez

2 Department of Botany, Missouri Botanical Garden, P. (5 “Box 299, St. Louis, Missouri, 63166-0299, U.S.A.

ANN. MissouRi Bor. GARD. 74: 401-412. 1987.

1.5mm

FIGURE |. Aiouea inconspicua.—A. Habit. — B. Detail of twig.—C. Flower. — D. Fruit.—E. Two outer an-
thers. — F. Inner anther with basal glands. — G. Ovary and stigma.

1987]

and cartilaginous, glabrous below with the ex-
ception of small tufts of hair at the bases of the
lowermost lateral veins, puberulous above on the
midveins of young leaves, otherwise glabrous;
lateral veins 4—6 pairs, slightly raised below and
lighter than the leaf tissue, not reaching the mar-
gins but forming conspicuous arches ca. 3 mm
from margin; tertiary venation reticulate, slightly
raised on upper and lower surface. Inflorescences
in axils of deciduous (rarely persistent) leaves,
minutely tomentellous, to 4 cm long, few-flow-

red (4-7 flowers per inflorescence in material
seen). Flowers greenish white, funnel-shaped,
glabrous, with small terminal openings revealing
the anthers, the pedicels 2-3 mm long. Tepals 6,
equal or the outer 3 slightly shorter than the inner
ones, roundish or slightly wider than long (0.9—
1.2 x 1.2-1.4 mm), concave. Fertile stamens 9,
ca. 0.9 mm long; anthers 0.3 mm long, with gland
dots, all with 2 anther cells, these filling the entire

long, as wide as anthers, pubescent, anista of
the inner 3 stamens with two basal glands. Ovary

1 mm long, elliptic, glabrous; style lacking, the
stigma sessile, large, 0.4 mm wide. No stami-
nodia seen. Fruit round, 1.5 cm diam. Cupule a
shallow disc, ca. 8 mm diam., gradually nar-
rowed into the pedicel.

Additional specimens examined. MEXICO. VERA-
CRUZ: Hidalgotitlan, ou Mor" entre Hnos. Cedil-
. 1974 (fl.), Brigada Vazquez

3 Feb. 1981 (fl, fr), Wendt,
L. y» — González ei dedii) (MO).

AT PT. ALTA VERAPAZ: najá, 150-
700 m, 1-2 yen "1942 (sterile), poesi 45 578 (A).

Aiouea inconspicua represents the first Aiouea
species reported trom € It resembles most
closely A. g is (Lundell) Renner, known
only from Guatemala. dte. most striking differ-
ences between the two species are listed in Table
1. In addition, A. guatemalensis has longer, wider
leaves than A. inconspicua, and the leaves dry
dark green with contrasting lighter venation in
A. inconspicua, a feature lacking in A. guate-
malensis.

An unusual feature of A. inconspicua is the
absence of staminodia. In both earlier generic
keys for Lauraceae (Kostermans, 1957; Hutch-
inson, 1964) and by Renner (1982) presence of
staminodia was considered a characteristic fea-
ture of Aiouea, although Renner (1982) men-
tioned two exceptions. Because other lauraceous

I

VAN DER WERFF—NEOTROPICAL LAURACEAE

403

TABLE 1. Selected characters of Aiouea inconspicua
and A. guatemalensis.

inconspicua guatemalensis

Young twigs + puberulous glabrous

terminal bud
Staminodes lacking present (teste

Renner, 1982)

Axillary tufts of present absent

hairs in lower-

most veins
Leaf texture chartaceous chartaceous-cori-

aceous

genera (Ocotea, Nectandra) include species with
and without staminodia, I do not see the absence
of staminodia in A. p.e as an obstacle
for its inclusion in Alou

Two of the aia, dide were distributed by the
Flora Veracruz project and may be present in
several additional herbaria: Brigada Dorantes
2929 was distributed as Nectandra salicifolia
HBK and Brigada Vazquez 1368 as Nectandra
sanguinea Rolander ex Rottboel. The Steyer-
mark collection was distributed as Ocotea effusa
(Meissner) Hemsle

LICARIA AUBLET

The genus Licaria, endemic to the Neotropics,
was recently monographed by Kurz (1983), who
recognized about 40 species ranging from south-
ern Florida to southern Brazil. Among the neo-
tropical Lauraceae, Licaria can be recognized
readily based on its three 2-celled stamens, dou-
ble-rimmed cupule, and alternate (rarely oppo-
site, never clustered) leaves.

Licaria bracteata van der Werff, sp. nov. TYPE:
Guatemala. Alta Verapaz: Sacté, large tree
in dense humid forest, 900-1,050 m (fl), Z.
Kunkel 7 (holotype, BR). Figure 2.

rbor magna. Ramuli obtuse angulati, glabri. Folia
iptica, 25-40 x

magnop

mino odia 9, 6 2 late lanceolata, ad 1 mm longa,
3 interiora lanceolata, ad 0.8 mm longa, staminibus

404 ANNALS OF THE MISSOURI BOTANICAL GARDEN

FIGURE 2. Licaria bracteata. Habit and flower seen from aside and above.

[Vor. 74

1987]

alternantia. Ovarium ED, tubo florale dense pu-
bescente. Fructus igno

Large tree. Twigs roundly angled, glabrous, with
small, light-colored lenticels, the terminal bud
drying black, glabrous. Leaves alternate, char-
taceous, glabrous on both surfaces (only a small
part ofthe lower leaf surface visible on specimens
seen), elliptic, the apex not seen, the base acute,
25-40 x 11-15 cm, the upper surface opaque,
the midvein and lateral veins (11—15 pairs) sunk-
en, the tertiary venation slightly raised, the lower
surface opaque, with midvein, lateral veins, and
tertiary venation raised. Inflorescences in the ax-
ils of bracts, these ultimately deciduous but pres-
ent in young inflorescences, up to 5 cm long, very
ubescent, with 10-20 flowers per in-

of the twigs above the leaves or along short, leaf-
less spurs in the axils of persistent leaves. Bracts
at bases of inflorescences black, glabrous, 1-1.5
cm long. Flowers 3-4 mm long, 2.5-3 mm wide,
sparsely grey-puberulous, urn-shaped, abruptly
widened at the base and gradually narrowed to-
wards the tip. Tepals 6, erect or somewhat in-
curved, unequal, the outer 3 triangulate, ca. 1
mm long, touching each other and largely ob-
scuring the inner 3 tepals, these ca. 0.6 mm long,
1.1 mm wide. Fertile stamens 3, ca. 1.4 mm long,
each with 2 basal glands, 2-celled, the anther cells
small, apical, opening towards the centers of the
flowers. Staminodia 9, the outer 6 broadly lan-
ceolate, ca. 1 mm long, the inner 3 lanceolate,
alternating with the 3 fertile stamens, ca. 0.8 mm
long, occasionally with minute anther cells. Ovary
glabrous, the floral tube densely pubescent in-
side. Fruit unknown.
Additional specimen examined. GUATEMALA. ALT

VERAPAZ: Sacté, large tree in dense humid forest, 900-
1,050 m (fl), r Kunkel 56 (MO).

In Kurz’s treatment (1983) L. bracteata keys
to the subgenus Guianensis (correctly: subg. Li-
caria, because it includes the type species of the
genus) and within this subgenus to a small group
of three Central American and West Indian

have much smaller leaves, smaller flowers, fewer
staminodia, and (with the exception of L. urceo-
lata) pubescence on leaves and/or twigs. Licaria
urceolata, which has glabrous leaves and twigs,
lacks the pubescence of L. bracteata on the in-
florescence. Licaria bracteata is an unusua
species because of its large, rather persistent

VAN DER WERFF—NEOTROPICAL LAURACEAE

405

bracts, large flowers, and, uniquely in subg. Li-
caria, the presence of nine staminodia.

That Mrs. Kunkel-Westphal, who was not
trained as a botanist, collected two interesting
new species of Lauraceae in Guatemala, is both
a compliment to her qualities as a collector and
an indication that many surprises await the col-
lector of large tropical trees.

PHOEBE NEES

A good set of characters to separate the genus
Ocotea from Phoebe has unfortunately not yet
been found. Ocotea is a particularly large and
vaguely defined genus which includes various as-
semblages of species. Phoebe likewise is poorly
understood. In the New World there are two main
centers of Phoebe species: one in northern Cen-
tral America and one in central-southern Brazil.
In the intervening areas Phoebe is very poorly
represented, and it is not clear yet whether the
Brazilian and Central American species form a
monophyletic group. An added difficulty is that
the type of Phoebe is an Asian species and that,
according to Kostermans (1961), the Asian
Phoebe species are not congeneric with the neo-
tropical Phoebe species. Kostermans (1961)
transferred all neotropical Phoebe species to Cin-
namonum, an Asian genus generally not consid-
ered native to the Neotropics. He did not discuss
the difficulties in separating Ocotea from Phoebe,
nor the heterogeneous assemblage of species clas-
sified in neotropical Phoebe.

Earlier authors relied on two characters to sep-
arate Ocotea from Phoebe. In Ocotea the stam-
inodes were said to be inconspicuous, in Phoebe
conspicuous; in Ocotea the tepals do not persist
in the fruiting stage and a cupule is present, while
in Phoebe tepals harden and persist in fruit, but
no cupule is present. The staminodial character
is relative and difficult to interpret. Because flow-
ers of Ocotea and Phoebe are small and the stam-
inodes almost never exceed | mm, one may
rightly ask what makes minute staminodes con-
spicuous in some flowers and inconspicuous in
others. Besides, the presence of staminodes is not

is also a doubtful character. Of the 18
American species attributed to Phoebe for which
I have data, only five have persistent tepals in
fruit. However, fruits are unknown for several
species. Cupule development in Ocotea is very
variable and ranges from a widened pedicel with

406

a minute cupule to well-developed cups. Place-
ment of new taxa in Ocotea or Phoebe has been
subjective and will remain so until Cinnamo-
mum, Ocotea, Phoebe, and their satellite groups
are studied critically.

I use the following characters as indicators for
Phoebe: tepals erect in flower and persistent in
fruit, flowers with relatively long pedicels, stam-
inodes present, and a tendency toward tripli-
veined leaves. Because the new species has four
of the five characters, I place it in Phoebe.

Using these indicator characters for Phoebe,
species such as P. helicterifolia (of which Nec-
tandra corzoana Lundell is a synonym) and its
allies [P. bourgeauviana (of which Ocotea tene-
japensis Lundell is a synonym), P. obtusata, P.
valeriana, Nectandra capituliforma, and N. lon-
gicuspis] would be excluded from Phoebe. These
species should all be placed in the same genus,
probably in the already variable Ocotea, or, if
future study turns up additional characters, in a
new genus.

Phoebe glabra van der Werff, sp. nov. TYPE: Mex-
ico. Oaxaca: Municipio Matias Romero, tree,
13 m, Wendt et al. 4813 (holotype, MO).
Figure 3.

Arbor, ad 13 m. Ramuli glabri, teretes. Gemma ter-
minalis glabra. Folia glabra, alterna, chartacea, basi
acuta vel rotundata

vata, axillis vorum basalium domatiis cavitatibus

m ng
Fr uctus ellipsoideu oo `cupula plana mar-
gine integra sore incrass

Tree, 8-13 m. Twigs terete, slender, glabrous.
Terminal bud glabrous. Leaves chartaceous, al-
ternate, the apex acuminate, the base acute or
somewhat rounded, elliptic, 12-16 x 4-6 cm;
lateral veins 3-5 pairs, the lowest pair most
strongly developed and with slitlike domatia in
the axils, these visible as small lumps on the
upper surface; veins and reticulation promi-
nently raised on both surfaces; petioles to 1.5 cm
long, glabrous. Inflorescences paniculate, gla-
brous, 5-10 cm long, in the axils of persistent
leaves or deciduous bracts, often appearing ter-
minal. Pedicels glabrous, 2-3 mm long. Flowers
glabrous, 2.5-3 mm long, funnel-shaped. Tepals
6, equal, ca. 1 mm long, ovate. Stamens 9, all
4-celled, the filaments narrower than the anthers,
1.3-1.4 mm long, glabrous; outer 6 anthers with

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

cells introrse, inner 3 with cells extrorse. Stam-
inodia 3, ca. 0.9 mm long. Ovary globose, gla-
brous, ca. 1.8 mm long; style short, ca. m
long. Fruit an ellipsoid berry, 2 cm long, cupule
platelike, the margin entire, 1.4 cm wide, pe-
duncle gradually widened towards the fruit.
Additional specimens examined. MEXICO. VE-
RACRUZ: Municipio Minatitlan, Wendt et al. 3217 (MO);
Municipio Hidalgotitlan, Wendt et al. 3141 (MO).
OAXACA: Municipio Matias Romero, Wendt et al. 3064
(MO).

Phoebe glabra is closely related to Ocotea eu-
venosa Lundell. Both species have twigs, leaves,
and flowers glabrous, raised reticulation, and slit-
like domatia (these were not mentioned by Lun-
dell (1965), but the GH isotype of O. euvenosa
shows the domatia clearly). Ocotea euvenosa dif-
fers in its larger leaves, laxer reticulation (even
a small leaf of O. euvenosa, of comparable size
to a regular leaf of Phoebe glabra, has a laxer
reticulation), and in its pinnately veined leaves,
not subtripliveined as in P. glabra. The type of
O. euvenosa is in young fruiting stage, which
makes comparison difficult. In the fruiting stage
the pedicels become much larger, to 2.5 cm long
and I therefore attach little taxonomic value to
the longer pedicels reported by Lundell (1965)
for O. euvenosa. Three of the four collections of
P. glabra come from limestone-derived soils.

PLEUROTHYRIUM NEES

The genus Pleurothyrium is separated from the
other neotropical lauraceous genera by a set of
characteristics associated with strong enlarge-
the staminal glands. Each of the fila-

ments of the inner three anthers carries two
glands, which, in other genera, are roundish and
rather small. There is no doubt these glands pro-
duce nectar. In the genus Pleurothyrium these
glands become strongly enlarged and grow out-
ward between the outer six stamens. In the re-
lated genera Ocotea and Nectandra the outer sta-
mens usually form a tight ring, with the anthers

cells are not possible because the anthers form
such a tight ring. However, in Pleurothyrium the
stamens are separated from each other by the

rather isolated. There are no spatial constraints
against lateral cells on the anthers, and indeed
one characteristic of Pleurothyrium is that the
lower pair of cells on the six outer anthers is
laterally positioned.

1987]

FiGure 3. Phoebe glabra. — A. Habit

VAN DER WERFF—NEOTROPICAL LAURACEAE

407

— B. Leaf base showing bero —C. Flower. — D. Fruit. — E. Two outer

stamens. — F. Basally united inner anther, glands, and two stamino

A second characteristic of Pleurothyrium is that
in many species the staminal glands become fused
into a ring at the margin of the flower or even
form a cushionlike glandular mass with only the
anthers and the pistil protruding. This glandular

ass has been called a disc by some authors
(Hutchinson, 1964). Sometimes one can still dis-
cern that the g nass is the result of fusion
of six glands, and Ears that in Pleurothyrium

nine stamens have two glands each are erro-
neous, as Rohwer & Kubitzki (1985) pointed out.

Additional characteristics for Pleurothyrium
are the relatively large, warty cupules, distinctly
larger than in most Ocotea and Nectandra species
and quite like the cupules in Aniba. The leaves
have many lateral veins for their size, a character
that, in addition to the large cupules, makes iden-
tification of fruiting material possible. Several

408

Pleurothyrium species also have hollow twigs,
which are frequently inhabited by ants

I noticed in several species that the margins
of the tepals are thinner than the centers and that
in old flowers the margins of the tepals curl
downward and become revolute. I have not seen
this in other neotropical Lauraceae.

Pleurothyrium costanense van der Werff, sp. nov.
TYPE: Venezuela. Edo. Falcon: Sierra de San
Luis, above La Chapa, 1,200 m, 10 Aug.
1979, van der Werff 3654 (holotype, U; iso-
type, CORO). Figure 4.

Arbor, 15-20 m. Ramuli eg dense ferrugineo-
pubescentes. Folia alterna, chartacea, elliptica vel late
Md apice rotundata vel dece acuta, basi obtusa
vel acuta, 20-35 x 8-17 cm. Venatio super immersa,
Mr elevata reticulatione conspicua. Inflorescentiae
axillares, pubescentia ferruginea, 10-15 cm longae. Te-
pala 6, aequalia, patentia, elliptica. Stamina 9, 4-lo-
cellata, 3 interiora locellis extrorsis, 6 exteriora locellis
inferioribus lateralibus, locellis superioribus introrsis.

tylum crassum dense pubescens. Glandulae stami-
nium in muro humile connatae, staminibus gynoeci-
oque in centro exposito. Cupula magna, verrucata.

ee, 15-20 m. Twigs more or less terete,
densely bisce, pubescent, glabrescent with
age. Terminal bud densely ferruginous pubes-
cent. Leaves alternate; laminae chartaceous, el-
liptic or broadly elliptic, the apex rounded or
shortly acute, the base obtuse or acute, 20-35 x
8-17 cm; with upper surface opaque, the vena-
tion sunken, glabrous except for some pubes-
cence on midvein and main lateral veins; lower
surface with raised midvein, lateral veins, and
reticulate venation, the midvein and lateral veins
with brown pubescence, otherwise with spread-
ing hairs in flowering stage, but glabrous when
fruiting; lateral veins 10-15 pairs, the veins arch-

nected with the superior vein; petioles 2-3 cm
long, ferruginous pubescent. Inflorescences in ax-
ils of deciduous or persisting leaves, 10-15 cm
long, ferruginous-pubescent, rather laxly flow-
ere owers white or buff, ca. 1 cm diam. Te-
pals 6, equal, spreading at anthesis, the outer 3
ferruginous-pubescent outside, the inner 3 pu-
berulous outside except for a narrow, median,
pubescent strip, puberulous inside, elliptic, ca. 4
mm long. Stamens 9, all 4-celled, the inner 3
with extrorse cells, the outer 6 with the lower
cells lateral, the upper ones introrse. Staminal
glands fused into a low wall surrounding the sta-
mens and gynoecium. Style short, stout, densely
grey pubescent; stigma flat, about as wide as the

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

style, glabrous; ovary glabrous. Flowering in Au-
gust. Cupule large, ca. 2 cm high when pressed,
conspicuously warty.

Additional € examined. |. VENEZUELA. SUCRE:
Peninsula de a, Cerro Espejo (fl.), Stevermark &
Rabe 96072 (US. VEN) MONAGAS: Y ucucual, E of Ca-
ripe (fr.), Lao 10 (MO).

Pleurothyrium costanense is known from three
collections in the Cordillera de La Costa and the
Sierra de San Luis, in northern Venezuela. Di-
agnostic characters are the reticulate venation of
the leaves, the broad leaves, and the ferruginous
pubescence. The only other Pleurothyrium species
in northern Venezuela are P. reflexum Lasser and
P. zulianense Lasser (these two names represent
the same species, for which I have used the name
P. zulianense Lasser). Pleurothyrium zulianense
differs in the lack of reticulate venation, in having
narrower leaves and smaller flowers, and in the
absence of dense ferruginous pubescence.

The epithet costanense refers to the Cordillera
de La Costa, where this species is found.

Pleurothyrium grandiflorum van der Werff, sp.
nov. TYPE: Colombia. Choco: Rio Mecana,
5-10 m. Tree, 15 m, along river. Flowers
yellow, with pleasant but strange fragrance
(as in some euglossine bee-pollinated flow-
ers), A. Juncosa 1675 (holotype, MO; iso-
types to be distributed). Figure

or, 15 m. Ramuli teretes dense breviter pubes-
centes. Folia elliptica, 15-25 x 4-7 cm, membranacea,
glabra, apice acuta, basi argute acuta vel decurrente.
Inflorescentiae ad 10 cm longae, pauciflorae, e foliorum

interiora angustiora, d
ina 9, 4-locellata, 3 interiora locellis extrorsis, 6 ext
iora 2 locellis inferioribus a alibus, 2 = ellis PANE
loribus laterali-introrsis. Glandulae stam nium in muro
connatae, staminibus styloque in nne expositis.
Fructus ignoti

Tree, 15 m. Twigs more or less terete, slender,
with dense, short, brown pubescence, this vel-
vety-shiny on the terminal bud. Leaves elliptic,
15-25 x 4-7 cm; laminae membranaceous, gla-
brous on both surfaces, but midvein with some
pubescence on lower surface, the apex acute, the
base sharply acute or decurrent on the petiole;
lateral veins 15-20, departing from the midvein
at angles of almost 90°, scarcely or not at all
decurrent along the midvein, the venation sunk-
en above, slightly raised on lower surface; peti-
oles 1-1.5 cm long, canaliculate, with same pu-

1987]

T JH F

VAN DER WERFF—NEOTROPICAL LAURACEAE

409

FIGURE 4. Pleurothyrium costanense. Habit, flower, and cupule.

bescence as twigs. Inflorescences to 10 cm long,
inserted in axils of deciduous leaves below the
leafy apices of the twigs, with same pubescence
as twigs, few-flowered (only 2 or 3 flowers present
on each inflorescence, but with scars indicating
former presence of more buds); peduncles ca. 5
mm long, with same pubescence as flowers.
Flowers very large for the family, 1.5-1.7 cm
diam., yellow, fragrant, densely short brown pu-

bescent, the tube 2-3 mm long. Tepals 6, spread-
ing, 6—7 mm long, subequal; the outer 3 broadly
elliptic, the inner 3 narrower, especially near the
base, all with dense, short pubescence on the
inner surface. Fertile stamens 9, all 4-celled, the
inner 3 extrorse, the outer 6 with the lower pair
of anther cells lateral, the upper pair lateral-in-
trorse; anthers glabrous. Staminal glands form-
ing a large wall surrounding the stamens or seem-

410

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FIGURE 5. Pleurothyrium grandiflorum. Habit and flower.

ingly covering the stamens in very young flowers.

Stigma platelike, ca. 0.3 mm diam., of almost

same color and texture as the anthers. Fruit un-
known.

Pleurothyrium grandiflorum is known only
from the type collection made in the Chocó area,
Colombia, an area rich in endemics. The species
is very distinct in its large flowers and few-flow-
ered inflorescences. In contrast to most Laura-
ceae, the lateral veins are hardly or not at all
decurrent along the midrib.

d westphalii van der Werff, sp. nov.
TYP uatemala. Alta Verapaz: Sacté. Tree
in Psp humid forest, 20 Apr. 1976, J.

Kunkel 9 (holotype, BR). Figure 6.

Arbor, 20 m. Ramwuli glabri, juvenales hinnulei pu-
bescentes. Folia alterna, elliptica, apice acuta, basi acu-
a, basi acuta vel decurrenti, membranacea, glabra. In-
lorcscentiae ad 8 cm longae, parviflorae, pubescentia

a,e foliorum
m Mem ij Mig patentia sub anthesim, pu-
scent tamina 9, 4-locellata, 3 interiora
locellis ae 6 exteriora 2 locellis inferioribus la-

£

1987]

VAN DER WERFF—NEOTROPICAL LAURACEAE

Ficure 6. Pleurothyrium westphalii. Habit and flower with bud.

mah É eek superioribus introrsis. Glandulae
sta in muro connatae staminibus styloque in
odi eR En ignoti

Tree, to 20 m. Older twigs glabrous, young
twigs with brown, dense, short pubescence.
Leaves elliptic, the apex acute, the base sharply
acute or somewhat decurrent along petiole,
membranaceous, 15-20 x 4-7 cm, glabrous or
nearly so above, with some appressed pubes-

cence, especially near base, but glabrescent with
age below; lateral veins 5-8, fading out near the
margin, not looping upward and not connected
with other lateral veins; upper leaf surface dark,
dull, venation sunken; midvein, secondary, and
tertiary venation slightly elevated; petioles to 1
cm long, pubescent as the young twigs. Inflores-
cences ca. 8 cm long, densely grey-brown pu-
bescent, few-flowered (fewer than 10 flowers per

412

inflorescence), in the axils of deciduous leaves or
bracts on the young twigs but attached below the
leaves, narrowly pyramidate. Flowers with 6 equal
tepals, these spreading or slightly reflexed at an-
thesis, densely short tomentose outside, slightly
less so inside, the tepals ovate, ca. 4 mm long.
Fertile stamens 9, all 4-celled, the inner 3 ex-
trorse, the outer 6 with the lower two e lateral,
the upper two cells introrse. Anthers glabrous,
staminal glands not seg nce! visible, but fused
into circular mass ca. 3 mm in diam. with the
anthers and pistil iS in e middle. Pistil
platelike, gray. Fruit unknown.

Additional specimens examined. GUATEMALA. ALT
VERAPAZ: Sacté, Kunkel 17 (fl.) (MO), 298 (sterile) (BR).

Pleurothyrium westphalii is the northernmost
species of its genus and is known only from one
locality in Guatemala. Two other Pleurothyrium
species, as yet undescribed and under study by
W. Burger, are known from Central America.
The species represented by Allen 5885 (GH, US)
from Costa Rica differs in having leaves with
acuminate apices, persistent bracts in the inflo-
rescence, leaf bases acute but not decurrent, and
the inner surfaces of the tepals papillose-puber-
ulous, not pubescent. The leaves of Allen 5885
also have more lateral veins which arch upward
and become connected, forming almost a mar-
ginal vein, as is often seen in Pleurothyrium

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

species. The second species, represented by Bunt-
ing & Licht 872 (NY) from the Rio San Juan
area, Nicaragua, differs in having a many-flow-
ered inflorescence (20-30 flowers per inflores-
cence), darker pubescence, and the staminal
glands not fused into a large wall as in P. west-
phalii

This species is named after the collector, Irene
Kunkel, née Westphal, who collected two new
and very interesting species of Lauraceae in Gua-
temala.

LITERATURE CITED

HUTCHINSON, J. 1964. The Genera of Flowering
Plants. Clarendon Press, Oxford.
KOSTERMANS, A.C. J. H. 1957. Lauraceae. Commun.
Forest Res. Inst. 57: 1—64.
. The New World species of Cinna-
momum Trew. (Lauraceae). Reinwardtia 6: 17-
24.

Kunz, H. 1983. due dui cu inr eodd Gat-
tungen neotropischer Lauraceen und Revision der
Gattung d Ph.D. D na a Peu
of Hambur

LUNDELL, C. L. 1965. Additions to the Lauraceae of
Guatemala. Phytologia 12: 243-

RENNER, S. 1982. Aiouea. Fl. Neotrop. “Monogr. 31:
85-116.

ROHWER, J. & K. Kusirzki. 1985. Entwicklungsli-
nien im Ocotea-Komplex (Lauraceae). Bot. Jahrb.
Syst. 107: 129-

WERFF, H. VAN DER. E Notes on neotropical Lau-
raceae. Ann. Missouri Bot. Gard. 71: 1180-1183.

A NEW SPECIES OF OCOTEA (LAURACEAE) FROM
SOUTHEASTERN MEXICO!

ToM WENDT? AND HENK VAN DER WERFF?

ABSTRACT

The new species Ocotea een is a common riparian tree of the Uxpanapa area of extreme

southern Veracruz, Mexico. It appea

s to be most closely related to O. eucuneata of Gua

temala and

Belize. It is yet another species M endemic to the rain forests of the Uxpanapa area.

Ocotea uxpanapana Wendt & van der Werff, sp.
nov. TYPE: Mexico. Veracruz: Municipio
Minatitlán, Zona de Uxpanapa, terracería
La Laguna-Uxpanapa, orilla Oeste del Río
Oaxaca, un poco al Oeste de Uxpanapa,
17?13'N, 94?13'15"O, 160 m, 14 Feb. 1981
(fü, Wendt, Villalobos & Olmstead 2869
(holotype, MEXU; isotypes, BM, CAS,
CHAPA, ENCB, MO, NY, and others).

Arbor ad 30 m. Ramuli hornotini sericei cito gla-

longiores quam latiores, apice acumin
vel cuneata, supra glabratae, ate sibi: oes
axillis barbatis.

3 mm longa; stamina externa stipitata. Fructu
ellipticus, ad 2.2 cm longus; cupula valde 6-lobata.

Tree, to 30 m, to 1 m d.b.h. or often with
several trunks from near the base; buttresses small
to medium-sized or lacking; bark medium gray-
brown to dark chocolate-brown, finely to prom-
inently warty, soft; slash of bark aromatic, light
brown or yellowish-brown, oxidizing in ca. 1 mi-
nute to darker orange- RIO a, sapro pale
cream-brown. Shoot portions of
stems and new leaves) and axillary buds densely
and finely sericeous; twigs soon glabrate, green
(drying dark or black). Leaves alternate; blade
obovate to elliptic, usually narrowly so, 8-25 cm
long, 2.2-8 cm wide, some leaves smaller, mostly
3-4.3 times as long as wide, firmly membrana-
ceous, slightly and irregularly conduplicate, me-
dium-dark green, finely and densely glandular-
punctate, distally acute or rounded to a short- or
long-acuminate (to 2.5 cm) apex with a minutely

rounded tip, the base acute to usually cuneate;
venation slightly raised above, prominently so
below, laterals 7-14, diverging at 40—65?, the
lowest 1—2 pairs usually more strongly ascending
(20-35°), basically eucamptodromous but often
more or less brochidodromous distally, the ter-
tiary venation more or less transverse between
laterals, the fine venation reticulate; adaxial sur-
face glabrous, abaxially finely strigose on surface
and sides of midvein, at length sometimes sub-
glabrate, lowest 1—several pairs of lateral veins
prominently barbate in axils; petioles 1-2.3 cm
long, canaliculate adaxially, at first finely stri-
gose, soon glabrate. Inflorescence complex formed
by a group of paniculate cymes, each of these
arising from the axil ofa quickly deciduous brac-
teate leaf (or rarely a foliage leaf), the apex of
each complex a small, temporarily dormant veg-
etative shoot, each inflorescence arising either in
the axil of a mature leaf or along the apical por-
tion of the main stem, several inflorescences oc-
curring together to form a large pseudoterminal
inflorescence complex, the vegetative apices usu-
ally elongating after flowering and producing ma-
ture leaves, the fruiting panicles thus more ob-
viously lateral; individual inflorescences 5-20 cm
long, 2-13 cm broad, cylindrical to pyramidal,
with a more or less straight central axis bearing
strongly diverging laterals, the latter sometimes
similarly rebranched, bearing dichasial groups of
flowers, the axes flattened, green to pinkish, light-
ly canescent to glabrate in basal parts, to densely
canescent distally; flower bracts lance-ovate to
deltate, 2-2.5 mm long, densely and finely ca-
nescent, very quickly deciduous; pedicels stout,
1.5-3.5 mm long, densely and finely canescent,
green. Flowers (4—)4.5-6 mm diam. at anthesis,

! We thank Hans Georg Richter (Hamburg) for bark and wood analysis, and Eduardo Merino, of the Centro

de Estudios del Desarrollo Rural of the Colegio de Postgraduados, for the drawing. Fieldw
author was jy que by the Centro de Botánica of the Colegio de Postgraduados and th
.; thanks are due to all who have participated in that fieldwork, a Ache "Villalobos

ork 2 b senior
n del Pa-

ete
? Centro de Botánica, Colegio de Postgraduados, 56230 Chapingo, Edo. de México, Mexico.
A.

? Missouri Botanical Garden, P.O. Box

ANN. Missounmi Bor. GARD. 74: 413-415. 1987.

299, St. Louis, Missouri 63166-0299, U.S.

414 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FIGURE |. Ocotea uxpanapana. — A. Branchlet € iota idee inset, lower surface of leaf. — B. Portion

of inflorescence, pre- -anthesis. — C. Flower. — D. Parts e flower: above, outer stamen, adaxial view; center,
inner stamen, abaxial view; . staminode (frequent absent) right, gynoecium.—E. Fruits with unique
lobed calyx. Bar alee A, E = 1 cm; B, C, D = m. Vouchers: A-D, Wendt et al. 2869 (staminode in D,

2865); E, Wendt et al. 2777.

1987]

tepals at anthesis ascending 25-45? or more,
broadly ovate, 1.923 mm long, 1.5-2.4 mm wide,
thick-textured, cream to creamy-green, abaxially
canescent with short, thick, gray hairs mostly
0.05-0.15 mm long, adaxially less densely so, the
apex broadly acute to acute; hypanthium 0.6-1
mm long. Outer 6 stamens 1.1-1.6 mm long;
filaments 0.5—0.9 mm long, pubescent; anthers
quadrate-ovate, with truncate to emarginate api-
ces, and with the upper pair of thecae more or
less above the lower pair; inner 3 stamens 1.5-
2 mm long; filaments 0.7-1.2 mm long, pubes-
cent, the basal pair of glands short-stipitate to
subsessile; staminodes absent or, when present,
less than 1 mm long, linear, pubescent. Ovary
1-1.5 mm long, glabrous; style (0.5—)1-1.4 mm
long, shorter than to slightly longer than ovary.
Fruit (nearly mature) broadly ellipsoid, to 2.2
cm long, to 1.9 cm thick, green, seated in a very
shallow strongly 6-lobed red, fleshy cupule, the
lobes 5-7 mm long, 3-5 mm high, 2-3 mm thick;
flesh of fruit ca. 1 mm thick, pale green. Testa
dark brown; flesh of cotyledons pink.

Paratypes. MEXICO. VERACRUZ: Zona de Uxpana-
m Mpio. Hidalgotitlán: Campamento La Laguna, along
o Las Cuevas, 6 Mar. 1984 (fl.), Taylor 387 (F, MO,
n. Campamento Hnos. Cedillo, 18 Jun. 1974 (fr),
P. E. Valdivia Q. 808 (MO, XAL); Propiamente en la
Escuadra, 7 Aug. 1974 (fr), P. E. Valdivia Q. 1312
(MO, XAL); A 600 m del Campamento Hnos. Cedillo
hacia Paso Moral, 8 Aug. 1974 (fr), P. E. Valdivia Q.

Brigada. Vázquez 1029 (MEXU, MO); Brecha Hnos.
Cedillo-Agustin Melgar, 5 Mar. 1974 (fl), Brigada
Vázquez 535 (MO, XAL); La Escuadra-Hnos. Cedillo,
por el Río Soloxuchil, 3 May 1974 (fr), Husad Váz-
A 729 (MO, XAL); Campamento La Laguna, Rio
Las Cuevas, 17?16'30"N, 94*30'W, 130 m, 17 Mar
1982 (fl), Wendt, Villalobos & Navarrete 3725 (CHA-
PA, MEXU, MO, and exu
Uxpanapa, desde el pobla

17°12'N, 94*10'W, 130 m, y pe 1. 1980 MEUM 2i
Wendt, Villalobos & Lara 2564 (CHAPA,
and oth

a
W de La Garganta, 17?13'30"N, 94?15' 30"W.
18 ES Feb. 1981 (fl), Wendt, Villalobos & Olm-
sy rp 2865 (CHAPA, MEXU, MO, and others); type
locality and date, Wendt, Villalobos & Olmstead 2869A
(CHAPA, MEXU, MO, and others).

Common names. Aguacatillo, laurel.

Ocotea uxpanapana is a common riverside tree
of the Uxpanapa area of southern Veracruz, a
region of lowland rain forests with high ende-
mism (see Wendt et al., 1985, and references
there cited); it is interesting that this riparian tree

WENDT & VAN DER WERFF—OCOTEA

415

has never been collected farther downstream in
the Río Coatzacoalcos basin. In the Uxpanapa
area, the new species and Ficus insipida Willd.
are the most common large riparian trees. The
root system of O. uxpanapana is apparently deep
and well adapted to resist flooding; during the
major flooding of the Río Oaxaca in October of
1980, all trees (including Ficus insipida) except
for many individuals of O. uxpanapana were
toppled and carried I Ocotea uxpanapana
also occurs scattered sites on deep
limestone-derived S It flowers mostly in Feb-
ruary and March, fruiting in September and Oc-
tober.

Ocotea is a large, mostly neotropical genus with
a few hundred species. The last comprehensive
treatment was by Mez (1889), and since that time
many additional species have been described.
The Central American species were treated by
Allen (1945), but the genus has remained difficult
and almost inaccessible to the nonspecialist.
Under these circumstances, we were initially hes-
itant to describe yet another Ocotea species, but
the striking, deeply six-lobed cupule is such a
unique feature in the genus, even in the family,
that we decided to publish a new species. It ap-
pears to be most closely related to O. eucuneata
Lundell (type from Belize). In addition to its less
strongly lobed cupule (fruiting material collected
in the mountains of Guatemala, identified by C.
Allen), O. eucuneata differs in its inflorescences
borne in the axils of mature leaves (not forming
a pseudoterminal inflorescence as in O. uxpan-
apana), its smaller flowers, and in having the
reticulation on the upper leaf surface immersed
(not raised as in O. uxpanapana).

Specimens collected by the Brigada Vázquez
and P. E. Valdivia Q. are distributed under sev-
eral names aaa s loeseneri Mez, N. salici-
folia Kunth, Persea americana Miller, and Phoebe
gentlei mand Standley & Steyerm.) and are
likely filed as such in several additional herbaria.

Wood and bark samples of the new species
(Wendt et al. 3725) were examined by H
Richter. He noted (pers. comm.) that the SM
and bark anatomy are typical of Ocotea in gen-
eral and are nondescript within the genus.

LITERATURE CITED

ALLEN, C. K. 1945. Studies in Lauraceae, VI. J. Ar-
nold Arbor. 26: 280—434.
m i 1889. Sende Se; Americanae. Jahrb. Kónigl.
t. Gart. Berlin 5: 1-556.
E T. S. A. RI & G. T. PRANCE. 1985. Esch-
weilera mexicana (Lecythidaceae): a new family
for the flora of Mexico. Brittonia 37: 347-351.

FOUR NEW SPECIES OF AXONOPUS
(POACEAE: PANICEAE) FROM TROPICAL AMERICA!

GERRIT DAVIDSE?

ABSTRACT

The following species of 4 xonopus sect. jai ipsis are described: A. rupestris, from Goiás, Brazil,

A. casiquiarensis from Guainía, Colombia, and Amaz

Venezuela, and A. jeanyae from Coclé, Panama.

Axonopus is a Aper i pond
ican grass genus with a few spec di to
the American subtropics i pon Old World
tropics. Black (1963) monographed the genus and
recognized 109 species. With continued explo-
ration it has become apparent that new species
remain to be described. On the other hand, it is
also probable that some of the species recognized
by Black must eventually be synonymized as in-
termediate populations become known.

Axonopus belongs to the tribe Paniceae and is
distinguished by the absence of a lower glume,
lower palea, and lower flower, and by its solitary,
dorsally compressed spikelets borne inversely
(i.e., with the back of the upper glume facing
away from the rachis) in two rows on one or
usually several to many racemes.

This paper reports on recent fieldwork in trop-
ical America that has brought to light four un-
described species of Axonopus sect. Axonopus.
These are published now so that the names will
be available for two forthcoming floras, Flora
Mesoamericana and Flora of the Venezuelan
Guayana.

Axonopus rupestris Davidse, sp. nov. TYPE: Bra-
zil. Goiás: Mun. Presidente Kennedy, road
from highway BR-153 to Itaporá, 12 km W
of village of Presidente Kennedy, Fazenda
Primavera along Ribeirão Feinho, ca. 3?25'S,
40°37'W, 400—500 m, | Feb. 1980, T. Plow-
man, G. Davidse, N. A. Rosa, C. S. Rosário
& M. R. dos Santos 8216 (holotype, MG;
isotypes, F, MO, NY). Figure 1.

xonopus rupestris Davidse, sp. nov. A. triglochi-
noides (Mez) Dedecca affinis sed spiculis i ici

MARN

and NSF in dept It

dos Santos,

for the stations, Per O. Huber,
manuscri

nas, Venezuela, A. chimantensis from Bolívar,

racemis paucispiculis, glumis minus valde nervis, et
anthoeciis glabris differt.

Caespitose perennial; culms 10-35 cm tall, the
internodes 1-2 mm wide, hollow, glabrous, the
nodes l-2, glabrous, the uppermost geniculate.
Leaves erect; sheaths to 7.5 cm long, strongly
distichous, conduplicate and keeled, apically
winged, mostly glabrous, sparsely hirsute near
the juncture with the blade; flag leaf sheath to 10
cm long; collar not differentiated, the sheath with
a slight marginal constriction apically but oth-
erwise merging imperceptibly with the blade; lig-
ule a ciliate membrane 0.5-0.6 mm long, the
membrane 0.1—0.2 mm long, the cilia 0.4—0.5
mm long; blades erect, mostly 5-10 cm long,
folded, mostly 1-2 mm wide as folded, sparsely
papillose-hirsute abaxially and adaxially along
the midrib to glabrescent, the apex acute. Inflo-
rescences 4.5-7 cm long; peduncles 1 or 2, the
uppermost to 19 cm long; racemes usually 2,
conjugate, sometimes 3 with the lowermost borne
5-10 mm below the upper pair, 4.5-6.5 cm long,
widely spreading at maturity, the axil puberulent;
rachis ca. 0.6 mm wide, glabrous, 14—16 spikelets
per 25 mm; pedicels 0—0.1 mm long, glabrous;
spikelets 2.9-3.5 mm long, 0.8-1 mm wide, el-
liptic-oblong, obtuse, the upper glume 0.2-0.3
mm longer than the lower lemma, 5-nerved, the
lateral nerves marginal, the midnerve weakly de-
veloped, prominently pubescent marginally and
between the nerves with hairs 0.5-1 mm long,
the hairs longest apically, the upper '4 glabrous;

idne

ally and between the nerves; anthoecium 0.5-0.9
mm shorter than the upper glume, stramineous,

' Fieldwork was conducted with the support of the Projecto Flora Amazónica in Brazil; NSF, CONICIT,
R, NGS, Fundación para el Desarrollo de las Ciencias Físicas, Matemáticas y Naturales in Venezuela;

. Huber, S. S. Tillett, B. Nelson, T. Plowman, N. A. Rosa, C. S. Rosário, M. R
milton, J. s. Miller, and C. Brewer-Carías for help with my personal fieldwork, J. K. Myers
J. D. Dwyer, and E. Kellogg for useful suggestions for improving Nie

2 pepa Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

ANN. Missounmi Bor. GARD. 74: 416—423. 1987.

1987]

FiGUREl. Isotype of Axonopus rupestris. — A. Habit

DAVIDSE—AXONOPUS

417

2.0cm

—B. Abax upper glume. —

C. Adaxial view of spikelet showing the lower lemma. [After POM et al. 8216 (MO). 1

nearly glabrous at the tip, obtuse; lodicules 2,
fleshy, truncate, glabrous; stamens 3, the anthers
1-1.2 mm long, dark yellow; styles 2, separate;
stigmas plumose, white; caryopsis not seen.

This species is known only from the type col-
lection, which was encountered in cracks and
shallow, gravelly soil pockets of a black, granitic
rock outcrop (Fig. 2). The epithet alludes to this
rocky habitat.

This species, by virtue of its glabrous rachis,
stramineous anthoecia, perennial habit, few ra-
cemes, and weakly developed midnerve of the
glumes, belongs to sect. Axonopus ser. Axonopu.
of Black (1963). However, it does not seem closely
related to any of the species grouped by Black in
this series. Rather, A. rupestris seems most closely
related to A. triglochinoides, a species of the Rio
Atabapo and upper Rio Negro drainages, which

[97]

Black placed in sect. Axonopus ser. Barbigeri
subser. Barbigeri because the midnerve of the
glume is developed, although not as strongly as
in typical members of this series.

Axonopus rupestris differs from A. triglochi-
noides in the smaller, much more conspicuously
pubescent spikelets (2.9-3.5 vs. 3.9-4.8 mm),
greater spikelet density on the inflorescence
branches (14-16 vs. 6-7 spikelets per 25 mm),
less prominently nerved glumes, and lack of a
prominent tuft of hairs at the tip of the anthoe-
cium. The two species are similar in the general
facies of their inflorescences and leaves, espe-
cially in the prominently distichous, densely tuft-
ed sheaths. Both species are relatively uncom-
mon, although local populations may consist of
hundreds of individuals. Axonopus triglochi-
noides also grows in cracks of granite outcrops
and shallow soil pockets.

E 2. Axonopus rupestris habit and habitat,
type loal. Goiás, Brazil.

Axonopus chimantensis Davidse, sp. nov. TYPE:
Venezuela. Bolívar: Distr. Piar, Macizo del
Chimantá, sector centro-noreste del Chi-
mantá-tepui, cabeceras orientales del Caño
Chimanta, vegetación litófita y riberena al-
rededor del comienzo E del Cañón recto del
Río Chimantá superior, ca. 5?18'N, 62°09'W,
ca. 2,000 m, 26-29 Jan. 1983, O. Huber &
J. A. Steyermark 6931 (holotype, MO; iso-
types, K, NY, US, VEN). Figure 3.

4xonopus chimantensis Davidse, sp. nov. A. villosus

Swallen affinis sed foliis culmis ligulis et aie mi-

noribus differt

Caespitose perennial; culms 25—60 cm tall, the
internodes 1-2 mm wide, hollow, somewhat flat-
tened, glabrous or apically with two lines of pu-
bescence along the margins, unbranched, the
nodes 1, rarely 2, densely pilose; sheaths 3-9 cm
long, very strongly distichous, conduplicate and
keeled, apically winged, glabrous except for the
upper '4 of the margin and keel where ciliate, the
margin membranous; flag leaf sheath 10-21 cm
long; collar prominent, densely pilose; ligule a

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

ciliate membrane 0.3-0.5 mm long, the mem-
brane 0.1—0.2 mm long, the cilia 0.2-0.4 mm
i blades folded, 6-18 cm long, 3.5-5 mm
e as folded, divergent, usually prominently
Hem on the margins, the keel sparsely ciliate to
glabrescent, the tip obtuse, scabrous, adaxially
very sparsely pilose to glabrous, abaxially gla-
ous. Inflorescence 3-10.5 cm long; axis 5-18
mm long; peduncles 1—2, usually well-exserted;
racemes 2-4, 2.5-10 cm long, spreading at acute
angles, the axils pubescent, the rachis 0.5-0.8
mm wide, scaberulous or pubescent marginally,
9-13 spikelets per 25 mm; pedicels 0.2-1.1 mm
long. Spikelets 3-3.4 mm long, 0.9-1.1 mm wide,
oblong-elliptic, obtuse; upper glume as long as
the lower lemma or to 0.4 mm longer, 5-nerved,
the midnerve developed, inconspicuously and
sparsely short-pubescent between the nerves;
ower lemma similar to the upper glume; an-
thoecium 0.3-0.7 mm shorter than the upper
glume, stramineous, apically with a distinct tuft
of hairs, obtuse; lodicules 2, fleshy, minutely
erose, glabrous; stamens 3, the anthers 1.2-1.5
mm long, purple; styles 2, separate; stigmas plu-
mose, purple; caryopsis not seen.

Paratypes. "VENEZUELA. BOLIVAR: Distr. Piar, Ma-
cizo del Chimanta, sector centro- noreste del Chiman-
ta-tepui, cabeceras orientales de

] mantá, altiplanicie en la base me-
riodional de los farrallones superiores del Apacará-
tepui, sector norte del Macizo, ca. 5?20'N, 62°12’W,
ca. 2,200 m, 30 Jan.-1 Feb. 1983, Steyermark et al.
128405 (K, MO, VEN), Huber & Steyermark 7043
(MO, VEN), 7032 (MO, VEN); Chimantá Massif, cen-
tral section, island in Río Tirica above Middle Falls
e

ma

central section, swampy savanna along tributary valley

> branch of headwaters of Rio Tirica, 2,120 m, 13
Feb. 1955. Steyermark & Wurdack 836 (US, VEN).

Axonopus chimantensis belongs to Black's
(1963) sect. Axonopus ser. Barbigeri subser. An-
cipites because of the perennial habit, non-pa-
pillose-pilose rachis and spikelets, stramineous
anthoecia, developed midnerve of the glume, and
broad, folded blades with obtuse apices. This
group of species is best developed in northern
South America, especially in the Guayana Re-
gion.

Axonopus chimantensis seems most closely re-
lated to A. villosus and A. steyermarkii Swallen.
Both of these are much larger than A. chiman-
tensis. In addition A. villosus has slightly larger

1987]

DAVIDSE—AXONOPUS

2.0cm

FIGURE 3. Holotype of Axonopus chimantensis. — A. Habit.
— C. Adaxial view of spikelet showing the lower lemma. [After Huber & Steyermark 6931 (MO

glume.

and broader spikelets with more prominent
nerves, larger ligules, relatively longer anthoecia,
and a tendency toward densely pubescent blades
and sheaths. Axonopus steyermarkii differs fur-
ther in its nearly glabrous foliage, blades notice-
ably narrowed at their bases, and usually longer
pedicels.

Many of the cited specimens of 4. chimanten-
Sis have smut-infected spikelets.

The rachis of the racemes in A. chimantensis
varies from pubescent to scabrous, a character-
istic also attributed to A. villosus by Black (1963).
Black cited Steyermark & Wurdack 463 and 863

—B. Abaxial view of spikelet showing the upper

as A. villosus, but I refer them to A. chimantensis.
As far as presently known, A. chimantensis is
endemic to the large Chimanta-tepui imn in
Bolivar and derives its epithet from t

Axonopus steyermarkii is nd to Cake
Marahuaca and nearby Cerro Duida in Ama-
zonas, Venezuela. Axonopus villosus occurs in
the same area as A. steyermarkii and may reach
as far north as the Rio Manapiare.

S —— eg sp. NOV. TYPE.
mazonas: Dpt. Atabapo, Cu-
Mine de ptem. ç S rds ofthe middle part

420

FIGURE 4. Holot type of Axonopus casiquiarensis. — A. Habit
glume. — C. Adaxial view of spikelets showing the lower lemma. [After Davidse et al. 16907A (MO).]

of Caño Caname, 67?22'W, 3?40'N, ca. 100
m, 30 Apr.-1 May 1979, G. Davidse, O.
Huber & S. S. Tillett 16907A (holotype, MO,
mounted as 2 sheets; isotypes, K, US, VEN).
Figure 4.

Axonopus casiquiarensis Davidse, sp.
mans (Trin.) Kuhlm. et A. camargoanus Hu affini
sed stolonibus, basibus culmi gracilioribus et foliis la-
tioribus differt

Stoloniferous perennial; stolons infrequent,
when present well-developed, to 70 cm long, leafy,
frequently rebranching and producing tufts of
leaves at the nodes, the internodes to 20 cm long;
culms 24—90 cm tall, the internodes ca. 1 mm
wide, hollow, slightly flattened, glabrous, the
nodes 1-2, usually appressed-pubescent, rarely

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

—B. Abaxial view of spikelet showing the upper

glabrous. Sheaths mostly 3-12 cm long, rounded
on the back, not strongly distichous, not winged,
usually glabrous, sometimes pilose at the base,
auricular hairs 1-3 mm long usually present; flag
leaf sheath to 29 cm long; collar not differen-
tiated; ligule a ciliolate membrane 0.1-0.2 mm
long, the membrane nearly obsolete to 0.1 mm
long, the cilia ca. 0.1 mm long; blades mostly 7—
60 cm long, 1-3 mm wide, flat to convolute,
commonly ciliate in the lower !^, otherwise gla-
brous, the apex acute. Inflorescence 4-1 5 cm long;
axis 2-30 mm long; peduncles 1 or 2, usually
well-exserted; racemes 2-3(-5), 4-14.5 cm long,
nearly appressed to spreading at acute angles, the
axils puberulent; rachis 0.3-0.5 mm wide, sca-
berulous, 10-15 spikelets per 25 mm; pedicels
0.1-0.7 mm long. Spikelets (2.5—)2.7—3.7(-4.1)

1987]

mm long, 0.8-1.3 mm wide, oblong-elliptic to
lanceolate-elliptic, obtuse; lower glume as long
as to 0.3 mm shorter than the lower lemma, 5-
7-nerved, the midnerve developed, slightly sul-
cate between the nerves, usually sparsely pubes-
cent between the nerves, sometimes nearly gla-
brous or strongly pubescent; lower lemma similar
to the upper glume; anthoecium 0.1-0.7 mm
shorter than the upper glume, stramineous, api-
cally with a distinct tuft of hairs, obtuse; lodicules
2, fleshy, inconspicuously 3-lobed, glabrous, vas-
culated; stamens 3, the anthers 1.5-2.2 mm long,
purple; styles 2, separate; stigmas plumose, pur-
ple; caryopsis ca. 1.5 mm long and 0.7 mm wide,
elliptic-obovate, the hilum elliptic-punctate, the
embryo ca. '2 the length of the caryopsis.

Paratypes. COLOMBIA. GUIANÍA: Rio Atabapo, ca.
7 km S of San Fernando de Atabapo, 67°43’W, 3*55'N,
28 Apr. 1979, Davidse 16825 (MO, VEN). VENEZUELA.
MAZONAS: DPT. ATABAPO: Cano Caname, 67?22'W,
3*40'N, 29 Apr.-4 May 1979, Huber et al. 3643 (MO,
VEN), 30 Apr. 1979, Huber et al. 3651 (MO, VEN),
2 May 1979, Davidse et z 17049 (COL, K, MO, TFAV,
VEN), 67°13'W, 3°40'N, 3 May 1979, Davidse et al.
17126 (MO, VEN), 5 W, 3°41'N, 2 May 1979,
Davidse et al. 17084 p VEN); Cano Yagua, 66?34'W,
3*36'N, Davidse et al. 17421 (MO, VEN), 66*41'W,
3°29' N. 7 May 1979, Davidse et al. 17314 (MO. VEN):
5 km al S de ^a Laguna Yagua, 66?38'W, 3?43'N, 22
Jul. 1980, Huber & Tillett 5476 (MO, VEN); Rio Ata-
bapo, 20 km S of San Fernando de Atabapo, 67?39'W,
3°50'N, 29 Apr. 1979, Davidse et al. 16856 (INPA,
MG, MO, VEN); Cano Cotuá, cerca Cerro Yapacana,
66°50'O, 3?40'N, 14-28 Feb. 1978, Huber 1605, 1679
(MO, VEN), 66°52'W, 3?*38'N, 6 May 1979, Davidse
et al. 17262 (MO, NY, VEN), 17229, 17239 (MO,
DPT. CASIQUIARE: medio Río Temi, 67?29'O,
2°57'N, 24 Feb. 1979, Huber 3410 (K, MO, TFAV,
) DPT. RIO NEGRO: lower Río Baria, 66°32'—
pag har 1°27'-1°10'N, 22-23 Jul. 1984, pu ie
F, XU, MO, NY, TFAV, VEN); Rio 43
Ed Eus W. 1?53'21?27'N, 23-25 Jul. 1984, D
vidse 27757 (MO, NY, VEN), 66°32'W, 1?38'N, 9 Feb.
1981, Huber & Medina 5888 (MO, VEN), 66?33'W,
1°35'N, 8 Feb. 1981, Huber & Medina 5858 (VEN);
io Siapa, 66?25'W, 2?05'N, 7 Feb. 1981, Huber &
Medina 5807 (VEN).

By virtue of its few, scaberulous racemes, stra-
mineous anthoecia, perennial habit, well-devel-
oped midnerve of the glume, and narrow leaves
A. casiquiarensis belongs to sect. Axonopus ser.
Barbigeri subser. Barbigeri of Black (1963).

It is related to A. comans and A. camargoanus
of central to southern Brazil both of which lack
stolons, and have consistently convolute blades
and thickened, usually deeply buried culm bases.

Both 4. comans and A. camargoanus inhabit
wet savannas, whereas A. casiquiarensis grows in

DAVIDSE—AXONOPUS

421

white-sand savannas and river banks, including
sandy pockets on granite rock outcrops.

This species is distributed in Colombia and
Venezuela in the lowland area (elev. 80-125 m)
centering around the Departamento Casiquiare
from whence it derives its epithet. So far it has
been collected from the Río Baria and the Río
Siapa in the south to Cerro Yapacana and the
Río Atabapo in the north.

Approximately one-fifth of the cited collec-
tions have some specimens with conspicuous,
long stolons. Apparently in this species, as in
many of its congeners capable of producing sto-
lons, stolon production is negatively correlated
with density of plants and in general is highly
dependent upon local growing conditions. Those
plants (e.g., Davidse et al. 17421, Huber & Medi-
na 5858) from completely open exposures in her-
baceous, white-sand savannas have very short,
erect, stiff leaves, whereas those from more fa-
vorable sites, such as sabanetas (e.g., Davidse et
al. 16856) and the margins of savannas (e.g.,
Davidse et al. 16907A), have a more luxurious,
lax growth habit.

Axonopus jeanyae Davidse, sp. nov. TYPE: Pan-
ama. Coclé: area between Cano Blanco del
eni Caño Sucio and Chorro del Rio Tife,
80°36'30"—80°38'W, 8?42'19"—8?43'06"N,
spray basin of waterfall, 3 Feb. 1983. G.
Davidse & C. W. Hamilton 23570 (holotype,
MO: isotypes, ISC, PMA, US). Figure 5.

Axonopus jeanyae Davidse, sp. nov. A. ciliatifolius
Swallen affinis sed nodis et spiculis p et
gluma et lemmate anthoecio longiore

Densely iue a perennial; culms 50-
75 cm tall, the internodes 1-2 mm wide, solid
or nearly so, aa: flattened, especially to-
ward the base, the nodes mostly 2, blackish, often
noticeably elongated, appressed pubescent with
hairs to 1.5 mm long. Sheaths mostly less than
10 cm long, keeled, not strongly USA gla-
brous, the ciliate at least when
young; flag leaf sheath to 16 cm ww 2s not
differentiated; ligule a ciliolate membrane 0.5 mm
ong, the membrane 0.2 mm long, the cilia 0.3
mm long; blades to 25 cm long, 2-3 mm wide,
folded at the base, flat to folded above, papillose-
ciliate towards the base, otherwise glabrous, the
apex obtuse, ps Inflorescence 6-12 cm
i m long; peduncles pore 2,
well-exserted; racemes 2-5, 4-11 cm long,
spreading at acute angles, the axils bend to

422

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

ee 5.

Holotype of Axonopus jeanyae. —
glu — C. Adaxial view of spikelet showing the lower lemma. [After Davidse & Hamilton 23570 (MO).]

pubescent, the rachis 0.4—0.6 mm wide, d
10-13 spikelets per 25 mm; pedicels 0.2-0.5 m

long, glabrous. Spikelets 1.9-2.5 mm long, 0. E
0.8 mm wide, elliptic-oblong, obtuse; upper glume
about as long as the lower lemma, 4.5-nerved,
the midnerve not or only weakly developed, with
appressed lines of pubescence between the nerves;
lower lemma similar to the upper glume; an-
thoecium 0.1—0.4 mm shorter than the upper
glume, brown, apically minutely papillate, ob-
tuse; lodicules 2, minutely 3-lobed, fleshy, vas-
culated, glabrous; stamens 3, the anthers 1.0-1.1

—A. Habit.

—B. Abaxial view of spikelet showing the upper

mm long, purple; styles 2, separate; stigmas plu-
mose, purple; caryopsis 1.3 mm long, 0.7 mm
wide, oblong, broadly obtuse, the hilum elliptic-
punctate, the embryo ca. 2 the length of the cary-
opsis.

The perennial habit, glabrous racemes, and
brown anthoecia place A. jeanyae in sect. Ax-
onopus ser. Suffulti of Black (1963). This new
species appears to be most closely related to A.
ciliatifolius Swallen, a species so far known only
from Belize. Axonopus ciliatifolius differs in its

1987] DAVIDSE—AXONOPUS 423

glabrous nodes and more fragile, glabrous, less generated by Missouri Botanical Garden collec-
strongly nerved glume and lower lemma, both tors in Mesoamerica.
of which are as long as the anthoecium.

The epithet recognizes the contributions of my LITERATURE CITED
wife, Jeany Vander Neut Davidse, to the Flora Biack, G. A. 1963. Grasses of the genus Axonopus.
Mesoamericana project. She is largely respon- Adv. Front. Pl. Sci. 5: 1-186.

sible for processing the thousands of collections

TWO NEW MESOAMERICAN SPECIES OF CHUSQUEA
(POACEAE: BAMBUSOIDEAE)!

LYNN G. CLARK?

ABSTRACT

ew Mesoamerican species of Chusquea are described and ee both from complete

Two
one ss of vegetative and flowering material.

Chusquea grandiflora, a cloud-forest species from

nama and northwestern Colombia, has infravaginal branching, long, relatively narrow foliage leaves,

open panicles, and rather large spikelets. It is allie

e smaller, wider, yellowish-green, and strongly eae

WV This species is probably most n. related to C. nelsonii Scribn. & Smith, and C. mueller
Munro

Duri tion ofa treatment of Chusquea
Kunth for Flora Mesoamericana, examination
of herbarium material revealed the existence of
two more undescribed species of this diverse
bamboo genus. Both are represented by complete
collections of vegetative and flowering material.
For explanation of terminology relating to the
vegetative parts, especially buds and branching,
see Clark (1985). In the specimen citations, a
flowering specimen is indicated by the insertion
of (fl) after the collection date.

Chusquea grandiflora L. G. Clark, sp. nov. TYPE:
Panama. Panamá: along road past Cerro Jefe
toward La Eneida, 6 Jan. 1971 (fl), Croat
13070 (holotype, ISC; isotypes, COL, CR,
K, MEXU, MO as two sheets, PMA, US).
Figure 1A-D.

Culmi scandentes, usque ad 18 m longi. Folia cul-
morum 16-50 cm longa; vaginae 12- 33 cm longae, 2-
3-plo longiores quam laminae, abaxiales scabrae su-
perne; laminae 4-17 cm longae, Abas aias scabrae; cin-

glabrae, non tessellatae, ap-
icibus brevi-setosis, basibus attenuatis. Inflorescentia
aperta, paniculata, 6-11 cm longa; rhachis ron
ngulati pubescent

"ai 9.7-12.6 mm longae, scaberulae. ae 2
s E .4 mm longa, l-nervis; gluma II 2.2-4. 8

uen a, 1-3-nervis. Lemmata sterilia 2; lemma ste-
rile I DES , 5.2-6.7 mm longum, 5-7-nerve; lem-
ma sterile II subulatum, 7-9.5 mm longum, 7-nerve.
emma fertile subulatum, 8.4-10.9 mm longum, 7-9-
nerve. Palea bicarinata, 6.9-10.2 mm longa, 6-8-ner-
vis.

Culms clambering, viny, to 18 m long. Inter-
nodes terete, smooth to scabrous-hirsute just be-
low the nodes. Culm leaves 16-50 cm long, the
juncture of sheath and blade abaxially indistinct;
sheaths 12-33 cm long, 2-3 times as long as the

lade, abaxially smooth at the base, scabrous
toward the apex; blades erect, usually persistent,
4-17 cm long, abaxially scabrous; girdle well-
developed, 2-3 mm wide, glabrous; corky ridge
present at the juncture of the sheath and girdle.
Nodes with one triangular central bud subtended
by numerous (up to 50) smaller, subequal sub-
sidiary branches; sheath scar dipping markedly
below the bud/branch complement. Branching
infravaginal; the central bud developing tardily
or sometimes not at all; leafy subsidiary branches
not rebranching. Foliage leaves with sheaths gla-
brous, the margins ciliate near apex; blades 8-
20 cm long, (0.5—)0.7-1.4 cm wide, L: W = 11-
27, the adaxial surface glabrous, the abaxial sur-
face glabrous, not tessellate, the apex short se-
tose, the base attenuate; pseudopetiole more or
less distinct, 2-4 mm long; outer ligule evident,
o 1 mm long; inner ligule truncate, asymmet-
rical, 1-1.5(-2) mm long. Inflorescence an open
panicle 6-11 cm long, usually not fully exserted
from the subtending sheath; rachis triquetrous,
pubescent; branches strongly spreading, angular,
pubescent, the lower ones short, up to 3 cm long;
pedicels 1-2 mm long, angular, pubescent. Spike-
lets 9.7-12.6 mm long, scaberulous. Glumes 2;
glume I ca. % the spikelet length, 2-2.4 mm long,
apically acute, abaxially scaberulous, 1-nerved;
glume II ca. '^ the spikelet length, 2.2-4.8 mm
long, apically pointed, abaxially scabrous, 1—3-

thank Dr. Richard W. Pohl for reviewing this manuscript, and Dr. Gerrit Davidse for editorial and

agrostological assistance

? Department of Botany, Iowa State University, Ames, Iowa 50011, U.S.A.

ANN. MissounRi Bor. GARD. 74: 424-427. 1987.

1987]

FIGURE 1.

B. Inflorescence, scale = 1 cm.— C.

scale = 1 cm. w^ D based on E 201 14, B, ir
cm.—F. Foliage leaf blade, adaxial view, scale =

with emerging branches, scale

CLARK— CHUSQUEA

kelet, scale - 1 mm
based on Croat 13070.) E-H. "i per e af

425

bero nin grandiflora and C. aperta. ard C. grandiflora. — A. Culm leaf with emerging branches,

m. — D. Foliage leaf beni abaxial view,
Culm le

orescence,
)

e = | cm.—H. Spikelet, anis — | mm. (E based on Soderstrom 2237, F-H ES on pa iae 2239.

nerved. Sterile lemmas 2; sterile lemma I ca. 2
the spikelet length, 5.2-6.7 mm long, apically
subulate, abaxially scaberulous, 5-7-nerved;
sterile lemma II ca. % the spikelet length, 7—9.5
mm long, apically subulate, abaxially scaberu-
lous, 7-nerved. Fertile lemma 8.4-10.9 mm long,
apically subulate, sence scaberulous, 7—9-
nerved. Palea 2-keele te except toward the

base, 6.9-10.2 mm long, apiculate, abaxially sca-
berulous, 6-8-nerved, the sulcus pubescent. Lod-
icules 3; 3—3.5 mm long. Stamens 3; anthers 4.3—
5.5 mm long. Fruit unknown.

Additional specimens examined. COLOMBIA.
tween Alto del Tigre and
1944 (fl), Core 817
(US). CHOCO: abis. La Sirena, 4 km W of La Mansa

426

at top of Cord. Occidental, Gentry & Renteria 24203
US). PANAMA. CHIRIQUÍ: Vicinity of Gualaca ca. 7.8
mi. from Planes de Hornito, La Fortuna on road to
dam site, Antonio 5182 (MO, US); camino hacia la
finca Landau, NE del oo e Fortuna (sitio
de presa), i Sep. 1976 (fl), Correa et al 2185 (F, MO,
NY). coctÉ: El Valle, Soderstrom 2014 (US). PANAMA:

17 Oct. 1977 (fl), Folsom & Page 5944 (MO, PMA,

, VEN); La Eneida, 12 km N of Goofy Lake, Sod-
erstrom 2006 (US). vERAGUAS: Cerro Tute, trail past
agricultural school near Santa Fe, Antonio 1878 (MO).

Chusquea grandiflora is characterized by in-
fravaginal branching, culm leaves abaxially sca-
brous (except basally), open panicles with short,
spreading primary branches, pubescent rachis,
and scaberulous spikelet with glume II usually
twice as long as glume I. This species is known
from montane forests at altitudes of 700 to 1,700
m in Panama and northwestern Colombia. Gen-
try & Renteria 24203, a vegetative specimen re-
ferred to this species, was collected at 2,300-
2,400 m

Chusquea grandiflora lacks the leafless, fibril-
lar branchlets so characteristic of C. scabra, but
otherwise the two species are strikingly similar
vegetatively. Chusquea scabra usually exhibits
much more rounded leaf bases and consistently
scabrous internodes. Chusquea grandiflora is also
closely related to C. foliosa L. G. Clark, C. lon-
gifolia Swallen, and C. patens L. G. Clark, shar-
ing infravaginal branching and long, relatively
narrow foliage leaves with these three species.

e open panicles with strongly spreading
branches of C. grandiflora and C. patens are sim-
ilar in some specimens, but C. grandiflora is dis-
tinguished by its pubescent rachis, shorter inflo-
rescence branches, larger spikelets, and wider
foliage leaf blades.

Chusquea aperta L. G. Clark, sp. nov. TYPE: Mex-

ico. Oaxaca: 107 km SW of Tuxtepec, 2,650

m, 4 Oct. 1977 (fl), Soderstrom 2239 (ho-
lotype, US). Figure 1E-H.

Culmi usque ad | cm diam., usque ad 1-2 m alti.

Folia culmorum: vaginae persistentes, 6.8-8.5 cm lon-

alis. Laminae foliorum rigidae, 7-12 cm longae,
1.5 cm e, L: W = 7-11, glabrae, valde a
inferne, apicibus brevi-setosis, basibus rotundatis =

apic
Doe aiden aperta, paniculata, 7-11 c
dern triquetra, glabra; rami eius
us ue a ongi, angulati, glabri. Spiculae £z:
mm longae, Pues adpressae. Glumae 2, squam

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

formes; gluma I 0.3-0.5 mm longa, enervis; gluma II
.4-0.8 mm longa, enervis. Lemmata steri
sterile I apiculatum, 4-4.6 mm longum, 1-3-nerve;
lemma sterile II subulatum, 4.5-5.1 mm longum, sna
nerve. Lemma fertile subulatum, 5.7-7.1 mm, S nerv
Palea bicarinata, 5.5-6.8 mm longa, 4—6-nerv

Culms to 1 cm diam., to 1-2 m tall. /nternodes
terete, solid, smooth. Culm leaves with the sheaths
persistent, the overlapping margin fused to the
sheath to 5 mm above the base, 6.8-8.5 cm long,
abaxially smooth; blades evidently not persis-
tent, in the one example seen, blade triangular,

cm long, abaxially smooth; girdle asymmetri-
cally developed, prominent only near the bud
complement, to 5 mm wide; inner ligule 2-3 mm
long. Nodes swollen, the central bud subtended
by up to 25-30 subsidiary branches; supranodal
ridge prominent. Branching infravaginal, the
central bud often developing; leafy subsidiary
branches occasionally rebranching. Foliage pun
>= the blades stiff, 7-12 cm long, 0.8-1.5 c

e, L:W - 7-11, yellowish-green, the RE
si smooth and not tessellate, the abaxial
surface smooth and strongly tessellate, the apex
short-setose, the base rounded to attenuate;
pseudopetiole usually distinct, 2-3 mm long;
outer ligule a stiffrim to 0.5 mm long; inner ligule
elongate, 2-9 mm long, rounded to acute at the
tip. Inflorescence an open panicle 7-11 cm long,
completely exserted from the subtending sheath
on a peduncle up to 12 cm long; rachis trique-
trous, glabrous; branches spreading, angular, gla-
brous, the lower branches to 4—6 cm long; ped-
icels variable, to 14 mm long, angular, glabrous.
Spikelets 6-8 mm long, glabrous, appressed to
branches. G/umes 2, scalelike, less than Y,, the
spikelet length; glume I 0.3-0.5 mm long, nerve-
less; glume II 0.4—0.8 mm long, nerveless. Sterile
lemmas 2, ^—h the spikelet length; sterile lemma
I 4—4.6 mm long, apiculate, glabrous, 1-3-nerved;
sterile lemma II 4.5-5.1 mm long, subulate, gla-
brous, 1-5-nerved. Fertile lemma 5.7-7.1 mm
long, subulate, glabrous, 7-nerved. Palea
2-keeled, the upper half sulcate, 5.5-6.8 mm long,
acute, glabrous, 4—6-nerved. Lodicules 3. Sta-
mens 3; anthers 4-5 mm long. Fruit unknown.

oe specimens examined. MEXICO. OAXACA:
a. 0.5 N i. SW

ue
75 to Tuxtepec, 16 Aug. 1975 (fl), Davidse
& Davidse 9733 (MO, US); on road between Oaxaca
and Tuxtepec, 11 Apr. 1975 (fl), Fisher & Engleman
75-41 (MO); Distrito de Ixtlan, Sierra de Juarez, ruta

1987]

175 Tuxtepec a Oaxaca, ca. 0.5 km al S de Cerro Pelon,
13 Apr. 1982 (fl), Lorence et al. 4004 (US), 108 km
SW of Tuxtepec on road to Oaxaca, Soderstrom 2237
(US).

According to label data, Chusquea aperta oc-
curs in pine-oak cloud forests at elevations from
1,670 to 2,750 m. At present, this species is known
only from a relatively restricted area of Oaxaca,
Mexico, on the road between Oaxaca and Tux-
tepec. Chusquea aperta is characterized by in-
fravaginal branching, deciduous culm leaf blades,
stiff, yellowish-green foliage leaves with blades
strongly tessellate abaxially, and long-exserted,
open panicles with spikelets appressed to the
spreading branches. Vegetatively, C. aperta re-
sembles C. nelsonii Scribn. & Smith, but can be
distinguished by its glabrous, abaxially tessellate
foliage leaf blades, in contrast to the narrower,
abaxially pubescent, non-tessellate leaf blades of
the latter species. The inflorescences and spike-
lets of C. aperta are similar to those of C. muelleri

unro (7 C. carinata Fournier and C. mexicana
Hackel), but the inflorescences of C. muelleri are
smaller, less open, and not long-exserted. With
regard to the spikelets, the glumes of C. aperta

CLARK—CHUSQUEA

427

are both small and scalelike, while the glumes
of C. muelleri are more developed, with the sec-
ond glume usually about paoe as long as the zt
The ty tly older, spen
T Sod (but still with some spikelets), Fw
a few younger, fresh inflorescences derived from
regrowth of subsidiary branches. These newer
spikelets are just at anthesis, and are slightly larg-
er and fuller than the old spikelets produced dur-
ing the primary flowering period. The newer
spikelets are unusual because they all have a one-
or two-keeled palea-like bract in the axil of the
second sterile lemma. I observed individual
spikelets with two fertile florets subtended by two
sterile lemmas in some specimens of a few Chus-
quea species, but this is the first indication ofa
rudimentary floret in the position of a sterile
lemma. No other flowering specimens of C. aper-
ta show this condition. The abnormal spikelets
were left out of the description.

LITERATURE CITED
CLARK, L. G. 1985. Three new species of Chusquea
(Gramineae: Bambusoideae). Ann. Missouri Bot.
Gard. 72: 864-873.

NOTES

NEW COMBINATIONS IN CHUSQUEA
(POACEAE: BAMBUSOIDEAE)

The genus Swallenochloa McClure was seg-
regated from Chusquea Kunth solely on the basis
of differences in certain vegetative features
(McClure, 1973). To my knowledge, McClure
never collected Swallenochloa, and only limited,
incomplete herbarium material of this group of
species was available to him. Soderstrom & C.
Calderón (1978) applied the terms extravaginal
and intravaginal respectively to the branching
patterns in Chusquea and Swallenochloa, and
discussed other vegetative differences in addition
to those listed by McClure. As the result of ex-
tensive field and herbarium studies, I determined
that the species of Swallenochloa constitute a
coherent, natural group that falls within the ge-
neric limits of Chusquea. Details of the reduction
of Swallenochloa to a section of Chusquea will
be presented in a later publication (Clark, in
prep.). For the present, this transfer necessitates
new combinations for three species.

Chusquea angustifolia (Soderstrom & C. Cal-
derón) L. G. Clark, comb. nov. Swallenoch-
loa angustifolia Soderstrom & C. Calderón,
Brittonia 30: 303. 1978. TYPE: Venezuela.
Tachira: subparamo y bosque enano, faldas
inmediatamente debajo del Paramo de
Tama, cerca de la frontera Colombo-Ve-
nezolana, Steyermark, Dunsterville & Duns-
terville 98615 (holotype, US; isotype, VEN).

ANN. MISSOURI Bor. GARD. 74: 428. 1987.

Chusquea longiligulata (Soderstrom & C. Cal-

erón) L. G. Clark, comb. nov. Swallenoch-

loa longiligulata Soderstrom & C. Calderón,

Brittonia 30: 305. 1978. TYPE: Costa Rica.

San José: along the Carretera Interameri-

cana, ca. 5 km SE of Empalme, near Tres

de Junio, Poh! & Selva 12842 (holotype, US;
isotypes, F, ISC, MO).

Chusquea vulcanalis (Soderstrom & C. Calde-
rón) L. G. Clark, comb. nov. Swallenochloa
vulcanalis Soderstrom & C. Calderón, Brit-
tonia 30: 309. 1978. TYPE: Costa Rica. Car-
tago: abrupte dominante La Playita, Volcán
Irazu, Pittier 14126 (holotype, US; isotypes,
ISC, US)

LITERATURE CITED

MCcCLUuRE, F. > - Genera of bamboos native to
the New rld (Gramineae: Bambusoideae).
— pens Bot. 9: ix-xii, 1-148. [Edited

y T. R. Soderstrom.]
SODERSTROM, T.R. & ALDERÓN. 1978. Chus-
qu ea and Swallenochloa (Poaceae: Bambuso -
species. Brit-

tonia 30: 297-312.

—Lynn G. Clark, Department of Botany, Iowa
State University, Ames, Iowa 50011, U.S.A.

NOTES ON CORNUS (CORNACEAE) IN SOUTH AMERICA

The family Cornaceae is represented in South
America by the genera Cornus L. and Griselinia
Forster f. The latter, often segregated as Grise-
liniaceae, occurs in Brazil and Chile, and in New
Zealand, while the former has been known pre-
viously from Bolivia north to Colombia in the

ndes. We report here, for the first time, Cornus
in Venezuela, ~ viai the first record of the
family in Venezu

Macbride pene was first to record the genus
from South America. Based on fragmentary
fruiting material collected from Peru and Boliv-
ia, he described two species, C. peruviana and C
boliviana, the former based on the collection
Macbride 3439 from Cani, Depto. Huánuco,
Peru, the latter based on Bang 1799 from Bolivia
without exact locality indicated. His decision to
separate the two collections as distinct taxa was
based on his observation. of the "equally two-
armed" trichomes on the abaxial leaf surface of
the Peruvian plant as contrasted with the “un-
equally branched or pronged" hairs with “one of
their two ‘arms’ being much longer than the oth-
er" in the Bolivian material. A year later, how-
ever, Macbride (1930) changed his mind and
concluded that Cornus peruviana was identical
with an unpublished species of Viburnum from
Peru collected by Ruiz & Pavon which he ex-
amined at B, and, accordingly, he transferred it
to Viburnum peruvianum (J. Macbr.) J. F.
Macbr. At the same time, he decided that the
Bolivian material did not belong to Cornus, but,
being uncertain as to its proper family position,
did not assign it to any genus.

Subsequently, Standley (1935) re-examined the
specimens described by Macbride and concluded
that they did indeed belong to Cornus, and that,
moreover, they were conspecific. Standley chose
the epithet peruviana for them. In the Flora of
Peru Macbride (1959) accepted Standley's con-
clusions and placed C. boliviana J. F. Macbr. in
synonymy.

During an expedition to the Andes of Estado
Táchira, Venezuela, near the Colombian border,
the authors found a fallen branch from a tree that
proved, upon later examination, to belong to the
genus Cornus. In determining that this collection
pertained to Cornus, and not to Viburnum, the
criterion of leaf pubescence, as indicated by So-
lereder (1899) and Metcalfe & Chalk (1950),

ANN. Missouri Bor. GARD. 74: 429-430. 1987.

proved critical. A comparison of the Steyermark
& Liesner collection from Venezuela with other
South American specimens initiated the present
study of the genus.

In general, we have observed, after examina-
tion of available herbarium material, that the
specimens from Venezuela and Colombia show
a more appressed type of indument on the calyx,
hypanthium, and lower surface of the leaf in con-
trast with collections from Ecuador, Peru, and
Bolivia, which have a more loosely crisp, spread-
ing, and often denser type of pubescence. How-
ever, the total number of specimens examined
from South America remains inadequate for con-
clusions as to the variability present. The ap-
parent differences noted between the northern
and southern populations may disappear when
more abundant material becomes available. For
the present, therefore, we do not recognize any
segregation of taxa, and conclude that the plants
from Venezuela are conspecific (sensu lato) with
those from the other parts of South America. In
this connection, we are employing the name C.
peruviana J. F. Macbr. as selected by Standley
(1935) when he concluded that the Peruvian and
Bolivian populations were conspecific.

ecimens examined (listed ee ssa
uth). VEN

shale substrata, south of El Reposo, 14 km SE
licias, 7°31’N, 72?24'W, 2,150-2,300 m, 22-23 Jul.
rmark & Liesner 118420 (MO, V

1935, pes 271 (F).
HUAN co: ped NE of Mito, 2,575 m
= 1923, Macbride 3439 (type en puaa F);
. CAJAMARCA: Prov. Ce sor canyon of the Río
Marañon above Balsas, 5 km bel mit of the road
to oo 24 deed 1964, Fehon: "i Wright 5336
(US); D AZONAS: Leimebambo, 2,100 m, 23 Dec.
1962, Woytowski 7796 (MO, US); Ambay, 3,100 m,
Tus. 1938, Univ. Cuzco s.n. O). BOLIVIA. DEPT.
HABAMBA: Ayopaya, Sailapata, 2,800 m, Nov. 1935,
Cardenas 3355 (US); Bang que (type of C. boliviana.

,

The following key indicates the main charac-
ters distinguishing Cornus from Viburnum:

1. Abaxial surface of leaves with 2-branched,
unicellular, nodose trichomes incrusted with
lime carbonate; flowers tetramerous; petals
free, separate; l Cornus

l. Abaxial surface of leaves not as above, the

430

pubescence, when present, of simple, stellate,
peltate, tufted, or glandular hairs; flowers pen-
tamerous; corolla HE ovary uniloc-

| Nn SOS OS MOSA UPS ks Viburnum

LITERATURE CITED

CRONQUIST, A. 1981. Pp. 668-670, 1006-1008 in An
Integrated System of Classification of Flowering
Plants. Columbia Univ. Press, New York.

MACBRIDE, J. F. 1929. To a genus new to South

merica. Trop. Woods 19: 5.
19 outh NA viburnums incor-
rectly desorbed as new species of Cornus. Trop.

Woods 24: 29.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

. 1959. Cornaceae. Fl. Peru, Field Mus. Nat.

Hist., Bot. Ser. 13(5), no. 1: 44.

METCALFE, C. R. & L. CHALK. Pp. 735-737,
752-754 in Anatomy ofthe Dicotyledons, Volume
2. Clarendon Press, Oxf

SOLEREDER, H. 1899. Pp. 487 -495 in Systematische
Anatomie der vio ace Verlag von Ferdi-
nand Enke, Stuttgar

STANDLEY, P. C. 1935. "The genus Cornus in South
America. Trop. Woods 43: 16-17.

—Julian A. Steyermark and Ronald Liesner,
Missouri Botanical Garden, P.O. Box 299, St.
Louis, Missouri 63166, U.S.A.

A NEW COMBINATION IN LUCILIA
(COMPOSITAE—INULEAE)

Lucilia pusilla (Kunth) Hieron. (1900) is the

generally accepted name for a small, caespitose
herb from the high Andes of Venezuela, Colom-
bia, Ecuador, and Peru. While working on a no-
menclator of Conyza (Compositae—Astereae), I
found that its basionym Conyza pusilla Kunth
(1818) is illegitimate, being a later homonym of
C. pusilla Houtt. (1779) from South Africa. De
Candolle changed the name of Conyza pusilla
Kunth to C. kunthiana DC. (1836). Since under
article 55.1 of the 1983 International Code of
Botanical Nomenclature the specific epithet of
the former correct name must be retained on
generic transferences, the correct name for this
species in Lucilia is L. kunthiana (DC.) Zardini,
comb. nov.

iiu

1 yva Y"
hu mm

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Lucilia kunthiana (DC.) Zardini, comb. nov.
Based on Conyza kunthiana DC., Prodr. 5:
379. 1836 (new name for C. pusilla Kunth).
Gnaphalium kunthianum (DC.) Kuntze,
Revis. Gen. Pl. 3(2): 152. 1898. Conyza pu-
silla Kunth, Nov. Gen. Sp. 4: 69. 1818, non
C. pusilla Houtt. (1779). Lucilia pusilla

TYPE: Ecuador: in Andean region, Humboldt
& Bonpland s.n. (P holotype; P isotype, B
isotype photograph F 15128).

I thank Dr. Zijlstra (U) for confirming the ne-
cessity of this new combination.

— Elsa Zardini, Missouri Botanical Garden, P.O.
Box 299, St. Louis, Missouri 63166, U.S.A.

FiGURE 1. Holotype of Conyza pusilla Kunth (P, photograph Zardini n. 1036, MO).

ANN. Missour!I Bor. GARD. 74: 431. 1987.

CHROMOSOME COUNTS OF MISSOURI ASTERACEAE
AND POACEAE

Few native Missouri plants are chromosomal-

ly known from Missouri populations. The list of

chromosome counts for composites and grasses
in Table 1 is a small contribution in this area.
The counts were made from standard anther
squashes stained in propionic-carmine and dis-
sected from buds fixed in Carnoy's or Newcom-
er's fixative. Except for Paspalum laeve, meiosis
was normal in all collections, and the observed
numbers agree with previously reported num-
bers as summarized in Federov (1969), Moore
(1973, 1974, 1975), and Goldblatt (1981, 1984,
1985). Vouchers are deposited in MO and KUH.

The chromosome number for Paspalum laeve
has previously been reported as 2n = 40 (Brown,
1948), 2n — 60 (Burton, 1942, as P. longipilum
Nash), and n = 40 (Banks, 1964). Our count of
n — 29 apparently represents an aneuploid re-
duction from the hexaploid level of n = 30. Mei-
osis and pairing were normal. The second pop-
ulation sampled had the heptaploid number, n =
35, and the chromosomes were almost complete-
ly asynaptic in early meiosis. No more than five
bivalents were ever observed at diakinesis or
metaphase I (Fig. 1). When chromosomes were

paired, the association was very loose. Structures
that we believe may be micronucleoli were pres-
ent in variable numbers at diakinesis (Fig. 1A).
We suspect, based on a similar pattern of asyn-
apsis in the tetraploid apomictic cytotype of P.
conjugatum Berg. (Fang & Li, 1966; Mehra,
1982), that the plant may have been apomictic.
Since only one plant was examined, we do not
know whether this condition was isolated or
widespread in the population. The species is
morphologically variable and further cytotax-

y be help-
ful in relating some of this variation to ploidy
levels

Supported by NSF grant INT-8510317.

LITERATURE CITED

BANKS, D. J. 64. Chromosome counts for Paspa-
lum. Rhodora 66: 368-370.

BRowN, W. V. 1948. Acytological study in the Gra-
mineae. Amer. J. Bot. 35: 382-395.

BuRTON, G. W. 1942. A cytological study of some
species in the tribe Paniceae. Amer. J. Bot. 29:
355-359.

FANG, J. S. & H. W. LI. 1966. Cytological studies in

TABLE l. Chromosome numbers of Missouri Asteraceae and Poaceae.
Taxon n Voucher?
Asteraceae

Erigeron strigosus Muhlenb. 18 Vahidy & Davidse 21

Eupatorium coelestinum L. f. 10 Vahidy & Davidse 20

Lactuca floridana (L.) Gaertner 17 Vahidy & Davidse 12

Rudbeckia laciniata L. 27 Davidse & Vahidy 30845

Rudbeckia missouriensis Engelm. 19 Davidse & Vahidy 30838

Solidago nemoralis Aiton 27 Vahidy & Davidse 15

Solidago ulmifolia Muhlenb. 9 Davidse & Vahidy 30835

Poaceae

Leersia virginica Willd. ca. 24 Davidse & Vahidy 30840

Panicum capillare L. 9 Vahidy & Davidse 17

Paspalum fluitans (Elliott) Kunth 10 Davidse & Vahidy 30854

Paspalum laeve Michaux var. /aeve 35 Vahidy & Davidse 1

Paspalum laeve Michaux var. pilosum Scribner 29 St. Louis, Tower Grove Park, Da-
vidse & Vahidy 30846

Paspalum pubiflorum Rupr. var. glabrum Vasey 30 Vahidy & Davidse 24

Paspalum setaceum Michaux var. muhlenbergii 10 Vahidy & Davidse 14

(Nash) D. Banks
Sorghastrum nutans (L.) Nash 20 Vahidy & Davidse 16

“Collected at Shaw Arboretum, Franklin County, Missouri, unless indicated otherwise.

ANN. Missouni Bor. GARD. 74: 432-433. 1987.

1987]

URE 1. Diakinesis in Paspalum laeve var. laeve, n =

NOTES

b
e? vh.
< e ?
è A []
; ~ .
y ° ° E b
o3 * se e
4 e e °
; , ° £ r^
dil" . 4 4 °
< e $
e
e
B *
LJ * °

35.—A. Photomicrograph showing nearly complete

FIGURE 1.
asynapsis.—B. Camera lucida drawing of A with 5, (marked) + 60,

pd a conjugatum Berg. Bot. Bull. Acad. Sin.

Feoesov - A. (editor). 1969. Chromosome Num
ers of PUE Plants. V. L. Komarov Botanical
stitute, Lenin
M secre P. (editor). 1981. Index to plant chro-
e numbers 1975- xn Monogr. Syst. Bot
Missouri Bot. Gard. 5: 3.
(editor). ce to plant chromosome
numbers 1979-1981. Monogr. Syst. Bot. Missouri
Bot. Gard. 8: 1-427.
(editor). 1985. Index to plant chromosome
numbers 1982- MAS Monogr. Syst. Bot. Missouri
Bot. Gard. 13: 1-224.
MEHRA, P. N. vus Cytology of East Indian Grasses.
P. P. Kapur, New Delhi.

Moore, R. J. (editor). 1973. Index to plant chro-
mosome numbers 1967-1971. Regnum Veg. 90:

(e di tor. 1974. Index to plant chromosome

numbers for 1972. Regnum Veg. 91: 1-108.
(editor). 1975. Index to plant chromosome

numbers for 1973/74. Regnum Veg. 96: 1-257.

hsan A. Vahidy, Department of Genetics,
c

Box 299, St. Louis, Missouri 63166, U.S.
Youji Shigenobu, Department of Natural uis
Naruto University of Teacher Education, Taka-
shima, Naruto-shi 772, Japan.

A NEW SPECIES OF THRASYA (POACEAE: PANICOIDEAE)
FROM THE MOSQUITIA OF NICARAGUA AND HONDURAS

Recent work for the account of the Poaceae
for the Flora Mesoamericana has brought to light
a previously undescribed species of Thrasya.

ups —— Davidse & Burman, sp.
nov. TYPE: Nicaragua. Zelaya: Along banks
oa pee gallery forest of Rio Likas
near Silima Lila, ca. 14?30'N, 83°50’W, 50
m, 5 Mar. 1979, John J. Pipoly 4107 (ho-
lotype, MO; isotypes, HNMN, SP, US). Fig-
ure |

Gramen perenne; culmi 65-110 cm longi; ligula
membranacea, 1.5-2.4 mm longa; laminae lineares,
-32

papillosum minute; flos superus perfectus.

Perennial herb; culms 65-110 cm long, erect,
sometimes rooting at the lower nodes, the inter-
nodes slightly compressed, hollow, mostly gla-
brous, the upper portans ang the nodes ap-

uber ulent
toward the apex, keeled, the collar appressed pu-
bescent; ligule a membrane 1.5- m long;
blades 16-32 cm long, 9-16 mm de linear,
flat, glabrous or puberulent toward the base be-
low, acuminate, the base slightly rounded to
gradually narrowed; leaf subtending the inflores-
cence distinctly smaller. Racemes terminal and
axillary, solitary, 10—16 cm long, arcuate, usually
well-exserted from the sheath; peduncles puber-
ulent near the tip or entirely glabrous; rachis 2.0—
2.8 mm wide, winged, dorsally and ventrally gla-
brous, the margin minutely scabrous, the base
with an inconspicuous tuft of hairs. Spikelets 3.4—
4.0 mm long, 1.4-1.8 mm wide, glabrous, ellip-
tic, broadly acute, arrayed in one row, oriented
back-to-back, paired, the pairs borne on alter-
nate sides of the rachis; pedicels puberulent, un-
equal, the upper 1.8-2.0 mm long, its lower half
adnate to the rachis, the lower 0.2-0.4 mm long;
glumes with the bases clasping and slightly swol-
len; lower glume dimorphic between the short-
and long-pedicellate spikelets, variable in the
short-pedicellate spikelets, 0.6-2.5 mm long, tri-
angular, membranous to herbaceous, 0-1-nerved,

ANN. Missouri Bor. GARD. 74: 434-436. 1987.

obtuse to acute or aristate, uniform in the long-
pedicellate spikelets, 0.7-1.2 mm long, triangu-
lar, membranous, nerveless, acute; upper glume
2.8-3.4 mm long, ca. 3⁄4 as long as the spikelet,
herbaceous, 5(-7)-nerved; lower lemma as long
as the spikelet, subindurate, deeply sulcate on
the back, sometimes splitting at maturity, 6(—7)-
nerved, the midnerve often absent, the inner pair
of nerves well-developed, slightly keeled and
crested toward the tip; lower palea as long as the
lower lemma, 2-nerved and 2-keeled; lower flow-
er staminate: lodicules 2, the stamens 3, the an-
thers 1.3-1.9 mm long, purple; upper lemma 3.0-
3.6 mm long, 1.4-1.6 mm wide, indurate, ellip-
tic, minutely papillose, the tip bearing a tuft of
minute hairs; upper palea of the same texture as
the upper lemma; upper flower perfect: lodicules
2, the stamens 3, the anthers 1.4-1.8 mm long,
purple, the styles 2, separate, the stigmas plu-
mose, purple; caryopsis (immature) ca. 1.7 mm
long, 1.1 mm wide, the embryo nearly '^ as long
as the caryopsis, the hilum narrowly elliptic, ca.

!^ as long as the caryopsis.

Paratypes. NICARAGUA. ZELAYA: Comarco del Cabo,
Kornuk Creek, Puente Pozo Azul, 10 Jul. 1972, Sey.
mour 5802A (MO), 5803 (MO). HONDURAS. GRA
A DIOS: 5 km de Puerto Lempira, 28 Jan. 1984, oe
& Cruz 8621 (TEFH).

The generic placement of this new species must
be considered since the limits of the genus Thra-
sya have always been difficult to circumscribe

posed of species that have the following features:
1) winged, one-sided, solitary racemes; 2) solitary
spikelets arrayed in a single row; 3) back-to-back
orientation of the spikelets, i.e., the backs of the
lower glume and lower lemma of the short-ped-
icellate spikelet face the backs of these same
structures of the long-pedicellate spikelet. The

problem of ihe outlying Species that show vary-
ing i between the core
group of Thrasya and the group of
Paspalum (sensu Chase, 1929) has been dis-

spikelets is superficially lost through the fusion
of both the long and short pedicels to the rachis

1987]

so that the free portion of all the pedicels is ba-
"x the same length.

. mosquitiensis the spikelets are arranged
ina gibt row in a back-to-back orientation along
a winged, one-sided, solitary raceme. Although
the spikelets are arranged in a single row, the
spikelets are clearly paired and the longer pedicel
is only partially fused to the rachis (Fig. 1). The
spikelet

along the rachis. With respect to | the distinctly

in T. mosquitiensis the basal half of the upper
pedicel is adnate to the rachis, in contrast to the
Decumbentes group where both pedicels of a
spikelet pair remain completely free. In addition,
TE : aay ea ý

in p p g
in several rows and the regular back-to-back po-
sition of the spikelets is never attained. There-
fore, although T. mosquitiensis is intermediate
between the core group of Thrasya and the De-
cumbentes group of Paspalum, similarities with
Thrasya are greater, and this is the reason for its
inclusion in that genus.

There is a tendency for the spikelets to be di-
morphic in the development of the lower glume
within single inflorescences. In the short-pedi-
cellate spikelets the lower glume may vary from
nerveless, membranous, and obtuse to 1-nerved,
herbaceous, and aristate. In the long-pedicellate
spikelets the lower glume is uniformly nerveless,
membranous, and acute. Also, the length of the
upper glume relative to the upper lemma is great-
er in the short-pedicellate spikelet than in the
long-pedicellate spikelet. This is most easily ob-
served as the length of the upper lemma that is
not covered by the upper glume: (0.1—)0.4—0.5
mm in the short-pedicellate spikelets, 0.5—0.7
mm in the long-pedicellate spikelets. Although
such dimorphism is characteristic of Paspalum
species of the Decumbentes group, it also occurs

in two Thrasya species: T. campylostachya
(Hackel) Chase and 7. petrosa (Trin.) Chase.

Thrasya mosquitiensis seems to be most closely
related to T. campylostachya, the only other Me-
soamerican species with glabrous spikelets.
Thrasya mosquitiensis is distinguished from the
latter by the larger size of all its parts, most no-
ticeably the spikelets (3.4-4.0 mm vs. 2.6-3.0
mm long and 1.4-1.8 mm vs. 0.8-1.5 mm wide),
racemes (10-16 cm vs. 4-10 cm long), and blades
(9-16 mm vs. 3-10 mm wide and 16-32 cm vs.
6—20 cm long). Moreover, the upper glume of
the long-pedicellate spikelet is proportionally

NOTES

435

= —— a 1

pe teen ee
He LLILLVA

.
`... Tes... mom RR RTI Tm
—— LB T xa e L n xx dero on ate

RE

FIGURE 1.

Spikelet m. of Thrasya mosquitiensis
Davidse & Burman showing the back-to-back orien-
tation ofthe PUN ud rachis (partially removed

n one side), and partially oyy maa of the long-
pedicellate sakaki. Scale line =

longer (34 vs. Y} the length of the spikelet),

roader, more clasping, and less papery in tex-
ture. Finally, the upper pedicel in 7. mosqui-
tiensis tends to be less adnate to the rachis than

436

in 7. campylostachya, although the latter species
is somewhat variable in this regard.

The specific epithet refers to the area of Nic-
aragua and Honduras, dominated by pine sa-
vannas, that is commonly referred to as the Mos-
quitia or sometimes as the Costa de Miskitos.

We thank ken Myers for the illustration, and

Peter Lowry an
ful comments on oe manuscript.

CI 101 USC-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

LITERATURE CITED
BURMAN, A, G. 1982. Three iid md of Thrasya
(Gramineae) from Brazil. Bri a 34: 458-462.
— (In pre Xd Revision of um zo Thrasya.
Acta Bot. Vene
CHASE, A. 1929. The North American species of Pas-
palum. Contr. U.S. Natl. Herb. 28: 1-310.

—Gerrit Davidse, Missouri Botanical Garden,
P.O. Box 299, St. Louis, Missouri 63166, U.S.A.;
and Alasdair G. Burman, Instituto de Botanica,
Caixa Postal 4005, 01000 Sao Paulo, Brasil.

A NEW SPECIES OF BULBOSTYLIS (CYPERACEAE)
FROM TROPICAL AMERICA

Bulbostylis, in the sense employed here, in-
cludes ca. 100 species centering in the tropics
and subtropics of both hemispheres, with best
representation in Africa and South bidon It
is usually placed in the tribe Scirpeae and is
taxonomically (some state Sd ane he related
both to Abildgaardia and Fimbristylis. It is dis-
tinguishable from both by the sheaths which bear
a sparse to dense beard of long, firm, often bristly
hairs at the orifice and by the achenes which are
surmounted by a persistent tubercle formed from
the style base. Many species have weedy tenden-
cies, are heliophytes, and thus grow abundantly
in open areas that are artificially or naturally
disturbed, in soils that are typically acid, sandy,
azonal, and seasonally moist or wet. Thus they
are much a part of vegetation of savanna, acid
rock outcrops, natural or artificial clearings, and
fluctuating, sandy-silty shores. Since such habi-
tats are common in many areas of Amazonia,
Bulbostylis species are an important component
of the herbaceous vegetation. One species, fre-
quently collected in the last decade along banks
and bars of major streams in Amazonian Brazil,
Colombia, and Venezuela, represents a new one
that we describe here.

Bulbostylis fluviatilis Kral & Davidse, sp. nov.
TYPE: Venezuela. Amazonas: Dpto. Río Ne-
gro, south side of hamlet of Santa Lucia,
sandy recently cleared area for heliport, by
Río Negro, common, 25 Nov. 1984, R. Kral
71973 (holotype, VEN; isotypes, F, FSU,
GH, ISC, MICH, MO, NY, P, U, US, VEN,
VDB). Figure 1.

Bulbostylis tenuifolia (Rudge) Macbride affine a quo
glumis mucronibus excurrentibus et antheris 2, 0.2-
0.3 mm longis differt

Annual, densely cespitose, 5-30 cm high. Roots
capillary-fibrous. Leaves few per culm, polysti-
chous, sub-basal, the lowest ns bladeless or
with blades shorter than the sheaths; principal
leaves usually '^—'^ as long as the scapes, some-
times as long, green; sheaths medially convex or
keeled, multicostate, the costas strongly hirtel-
lous, the broad, scarious, tan borders hirtellous,
narrowing acutely to a long ciliate apex; blades
5-15 cm long, capillary, erect or spreading,
proximally involute, 0.2-0.3 mm broad, penta-

ANN. Missouni Bor. GARD. 74: 437-439. 1987.

costate, distally triquetrous with costas scabrid-
ulous, the tips narrow but blunt. Scapes 0.2-0.3
mm thick, capillary, erect, sharply 5—7-costate,
smooth or proximally sparsely scabridulous. In-
florescence a broad anthella 2-4 cm high, sub-
umbellate, diffuse; involucral bracts 3-5, leaflike,
subtending prophyllate Tays, ins longest bract
equalling or slightly o he anthella, the
central spikelet sessile, 8-4: mm "aa its lowest
bracts also subtending primary rays; primary rays
0.5—3 cm long, erect to spreading, capillary and
costate, 1—3-spicate; secondary rays 5-15 mm
long. Spikelets mostly 3-6 mm long, lance-ovoid
to lance-cylindric, pedicellate except for the cen-
tral spikelet, acute; bracts 1.0-2.0 mm long,
loosely spirally imbricate, lance-ovate, navicu-
lar, glabrous or hispidulous, 1-nerved, medially
carinate, the midnerve green, the apex acute, vil-
losulous, the margins sparsely ciliolate, the cusps
and mucros 0.1-0.3 mm long, excurvate, sca-
bridulous, the broad sides scarious, tan or red-
brown, glabrous or hirtellous; lowest bract 2-3
mm long, usually sterile, strongly cuspidate, the
cusp to 2 mm long; stamens mostly 2, the fila-
ments flat, the anthers 0.2—0.3 mm long, basi-

xed, bilocular, laterally dehiscent, ellipsoidal,
apiculate; style glabrous, branched medially to 3
slender, papillate stigmas. Achene 0.5-0.7 mm
long, broadly obovoid-trigonous, the faces nearly
plane, the adaxial face broadest, the surfaces fine-
ly papillose-lined longitudinally, transversely ru-
gulose, whitish gray to brown; tubercle ca. 0.1
mm long, subglobose to oblate, apiculate, dark
brown.

Paratypes. COLOMBIA. GUAINÍA: near Coitara, ca. 7
km S of San Fernando, 67?43'W, 3°55'N, Davidse 16827
(COL, MO, VDB, VEN); Finca Buena Vista on the Río
Negro, ca. 15 km S of San Felipe and San Carlos de
Río Negro, 67°00'W, 1°42'N, Liesner 8780 (COL, MO,

Betun along the Río C ;
Davidse & González 13121 (MO, VEN); 27 km WSW
of Paso de Cinaruco along the Río Cinaruco, 67°45'W,
6°31'N, Davidse & González 12574 (MO, VEN); bank
of Río Orinoco on Isla Poyatón, 67°05'W, 7°02'N,

vidse & González 122164 (MO, VEN); 9 km W of Paso

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

438

SOD Osco o

O6o0cG6o00o090 ad
0005500 009.29 2203 099
O

aoa

- v.
ay `
Se

iw
ES

1987]

de Cinaruco along the Río Cinaruco, 67?35'W, 6?35'N,
Davidse & González 12495 (MO, VEN). BOLIVAR: Au-
yan- tepui, coche caps n. (VEN). BRAZIL. MATO GROSSO:
Rio Aripua Humboldt Campus, 59?21'N,
10°12’S, d et al. 18319 (MO, NY).

Among the tropical American species of Bul-
bostylis, B. fluviatilis seems most closely related
to B. tenuifolia (Rudge) Macbride, and some of
the cited paratypes have been distributed under
that name. Bulbostylis fluviatilis is similar to B.
tenuifolia in its general facies and annual life cycle
but differs in having one-nerved glumes with an
excurrent mucro and two stamens with anthers
0.2-0.3 mm long. Bulbostylis tenuifolia has three-
nerved glumes with the midrib slightly or not at
all excurrent and one stamen with anthers 0.4—
0.5 mm long.

In superficial appearance B. fluviatilis also re-
sembles some species of the Trichelostylis group
of Fimbristylis. However, the prominent hairs
along the mouth of the sheath and the distinctly
tuberculate style base clearly indicate that this
species belongs in Bulbostylis.

Bulbostylis fluviatilis grows along large rivers
(hence the epithet) at elevations of 65-120 m. It
grows primarily on the sand bars that become
prominently exposed in these rivers during the
dry season and on accumulations of sand on ex-
posed granite outcrops along the rivers.

NOTES

439

We believe that the cited collection of Vareschi
from Auyan-tepui is not a reliable record and
that it is probable that the wrong label was
mounted with the plant. It is perhaps possible
that the plant originated along one of the rivers
at the base of Auyan-tepui, but it is extremely
unlikely that it originated from the highly dif-
ferent habitats atop Auyan-tepui. The Brazilian
collection differs from the others in having the
leaves about as long as the inflorescences.

The fieldwork that resulted in some ofthe cited
collections was supported by the National Sci-
ence Foundation, National Geographic Society,
Consejo Nacional de Investigaciones Científicas
y Tecnológicas (Venezuela), and Fundación para
el Desarrollo de las Ciencias Físicas, Matemá-
ticas y Naturales (Caracas). We thank Otto Hu-
ber and Charles Brewer-Carías for providing op-
portunities for fieldwork in southern Venezuela,
and the Herbario Nacional (VEN), INP-
ARQUES, for use of their facilities.

— Robert Kral, Department of General Biology,
Vanderbilt University, Nashville, Tennessee
37203, U.S.A.; and Gerrit Davidse, Missouri Bo-
tanical Garden, P.O. Box 299, St. Louis, Mis-
souri 63166, U.S.A.

—

FIGURE 1.
sketch. — b. Leaf blade apex.—c.
Outer view of the leaf base. — f. Anthella base, showin

Bulbostylis fluviatilis Kral & Davidse, drawn from the type Epsum. Kral 71973.—a. Habit
id-sector of the leaf blade, dorsal view. —d. I

view of the leaf base.— e.

t a al and bases of primary
ruit

rays. —g. Sector of the scape.—h. Fertile bract.—i. Two sorts of spikelets. —j. Floret.—k.

A NEW CAREX SECT. OLIGOCARPAE (CYPERACEAE)
FROM WESTERN ARKANSAS AND EASTERN OKLAHOMA

In late April of 1977, during fieldwork along
the summit of Rich Mountain, the highest range
of the Ouachita trend in western Arkansas, the
senior author discovered on the dry, oak-pine
slopes of the southwestern-facing side numerous
clones of a Carex similar to C. hitchcockiana
Dewey but distinct in habit, indument, and foliar
character. On being informed of this plant, Ar-

ductive searches for it, not only in the original
locality but elsewhere on Rich Mountain (in-
cluding Oklahoma) and southward. We are aware
now of an abundance of it, often in association
with C. hitchcockiana and C. oligocarpa Schk.
of sect. Oligocarpae over a substantial range of
arenaceous oak-hickory-pine uplands in Arkan-
sas (Howard, Polk, Scott counties) as well as in
LeFlore County, Oklahoma. This habitat system
is so well represented in the Interior Highlands
physiographic province, particularly the Ouach-
itas, that a sedge thought to be rare and local may
indeed be widespread and locally abundant with-
in that physiography, as has now been shown in
the case of C. /atebracteata Waterf. (Kral, 1983).
The new Carex is named after the mountain
complex where it first was found.

Carex ouachitana Kral, Manhart & Bryson, sp.
nov. TYPE: United States. Arkansas: Polk Co.:
sandy rocky woods at summit of Rich
Mountain, just W of Queen Wilhelmina Park
Headquarters, 26 Apr. 1977, R. Kral 59803
(holotype, MO; isotypes, BM, CTB, GA, GH.
MICH, MO, US, VDB). Figure 1.

Spec. nova e sectione Oligocarpae. Planta perennis,
30-80 cm a glabra, laxe cespitosa, valde squam ate
rhizomat s vel
paulo brevioribus, Rhizoma repens vel ascendens, plus

-5 c nga
acuta vel acuminata scopu rpurea, carinata. Culmi
dimorphi, lateralibus Stenilibus foliosissimis. Folia

principalia 30-70 cm longa. Laminae compressae, 3-

5 mm latae, scabridae, apicem versus attenuatae, sca-
bromarginatae; pagina ad medi

ae, ae eeu ascendentibus vel erectis, 2-6 mm lon-
sis subtendentes, infimae laminis 10-13 cm longis et

m erecto rostro ca. 4.5-5.0 mm longa duds
concava, pallide brunneola, minute papillosa

Perennial 30-80 cm long, smooth, loosely ces-
pitose, strongly scaly-rhizomatous, the principal
leaves slightly longer or slightly shorter than the
culms. Rhizome creeping to ascending, + lig-
neous, 2-4 mm thick, branching, the scales ovate,
acute, spirally imbricate, multicostate, brown,
gradually passing into cataphylls. Cataphylls ob-
long, 1-5 cm long, acute or acuminate, red-brown-
purple, multicostate (costas often white), cari-
nate. Culms dimorphic, spreading, slender,
acutely trigonous, scabrid, the lateral ones sterile,
more leafy. Principal foliage leaves longer toward
culm base and approximate, 30-70 cm long. Leaf

sheaths carinate, smooth; ligule erect, a narrow,
horseshoe-shaped or acute scale. Blades spread-
ing or recurved, flattened, linear, 3-5 mm wide,
scabridulous, apically attenuate, triquetrous, the
edges harsh, the surfaces at mid-blade above sca-
brellous, impressed-nerved, abaxially elevated-
nerved, with midcosta sirongly raised, scabrid.
Spikes 2-4, narrow, the uppermost all male, nar-

GURE l. j. Carex ouachitana. —a. Hab

c. quies rare side. — d. Axial (left) and abaxial (right) sides of midsector of leaf blade. —

— b. Ventral (left) and lateral-oblique view of leaf sheath apex

ssa ~ staminafe

flower.—f. Anther and upper part of filam —g. Perigynium and accompanying scale, spike . Peri-
gynium and accompanying scale, spike apex, pomi stigmatic branches and m of included fruit. — i. Detail,
further anc of perigynium.—j. Fruit. From Kral 59803.

ANN. Missouni Bor. GARD. 74: 440-442. 1987.

441

NOTES

1987]

“scu ti ESE

z5

“ease ass nee

442

rowly densely ellipsoid-cylindric, 2-4 cm long,
ca. 3 mm thick, 2-3 times shorter than the pe-
duncle. Spikes beneath all female, or androgy-
nous with 1-7 male florets at the tip, narrowly
linear, few-to-many-flowered, 3-6 cm long, ba-
sally interrupted, at apex with the flowers more
approximate. Male spike bract without sheath,
lanceolate, 10-15 mm long, narrowly acuminate
or cuspidate, carinate; bracts of lateral spikes fo-
liaceous, erect, subtending erect to ascending pe-
duncles 2-6 cm long, the lower with blades 10-
15 cm long and with closed sheaths to 4 cm long.
Male scales scarious, oblong-lanceolate, ca. 2.5-
3.0 mm long, acute, carinate, the sides pale brown,
the costal area green. Female scales ovate to lan-
ceolate, navicular, 5-15 mm long including the
cusp, acute to narrowly truncate, the midzone
green, unicostate, the sides stramineous to brown,
the tips of the lower scales strongly cuspidate.
Perigynia obovoid, 4-6 mm long, obscurely tri-
gonous, the faces slightly concave to level, im-
pressed-nerved, the beak short, excurved. Fruit
tightly included, stipitate-obovoid, trigonous,
4.5-5.0 mm long including the short erect beak,
the faces concave, pale brown, minutely papil-
ate.

Additional specimens examined. UNITED STATES.
ARKANSAS: Howard Co.: shaley wooded river bluffs,
9.6 mi. E of Wickes, 9 May 1979, Kral 63507; NW of
AR Hwy. 4 and Cossatot River on slopes above river
under mixed hardwoods, 10 May 1986, Bryson 4332.
Polk Co.: Rich Mtn., along Ouachita Trail near Hwy.
88 N 0.3 mi. W of Rich Mtn. tower, 27 Apr. 1981,
Rettig & Davis 239; Queen Wilhelmina State Park,
Sect. 11, R32W, TIS, below and to N of visitor center
and AR Hwy. 88 along trail on N-facing slope under
mixed hardwood forest, rich rocky soil, 10 May 1986,
Bryson 4289; NW of Rich Mtn. Lookout Tower, N of
AR Hwy. 88, SW '4 Sec. 8, R31W, TIS, along Ouachita
Trail on N-facing slope under hardwood forest with
dense herbaceous undergrowth, 10 May 1986, Bryson
4290; NE of AR Hwy. 246 and Cossatot River crossing
NE ‘4 Sec. te R30W, T4S peal pos hardwood forest

with few pines, on W-facing slope cky soil with thick
leaf E "10 May 1986, Do: 4326. Scott Co.: N
of Men

slopes of Black Fork
a

Oklahoma-Arkansas state line near a

#1, 30 Aug
1979, Taylor 28122; just W of Sta Line Historical
nds on roc

hickory forest with few eg toward top, thick brushy
and herbaceous unders rich rocky soil in associ-
ation with Carex prede € : posses C. hitchcock-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

iana, C. albursina, 10 May 1986, Bryson 4310. Du-
plicates are to be distributed later, thus acronyms for
recipients are not cited.

Staminate terminal spikes, female spike scales
with harsh cusps and costae, perigynia with many
impressed nerves, and short toothless beaks place
this sedge squarely in sect. Oligocarpae, a section
first designated by Carey in Gray (1848), and
subsequently revised by Kukenthal (1909), Mac-
kenzie (1931-1935) and Fernald (1950). In most
modern classifications this complex comprises
just C. oligocarpa and C. hitchcockiana. Both of
these are densely cespitose and erhizomatous;
whereas C. ouachitana is loosely cespitose,
sometimes forming colonies up to 0.5 m across
in open areas and up to 1—2 m across in shaded
areas, and produces strong, ligneous, imbricate-
scaly rhizomes. The staminate spike is larger,
with a longer peduncle, thicker, and bears more
florets. The lateral spikes are frequently androg-
ynous, while in C. oligocarpa and C. hitchcock-
iana they are solely carpellate. The perigynia are
much like those of C. hitchcockiana in size and
shape but differ in that the beak is shorter, broad-
er, and more bent outward. On the other hand,
the fruit beak resembles that of C. oligocarpa by
being erect or suberect. However the members
of this triad are related morphologically, it is
significant that, even though they have been ob-
served to share habitat in the same localities, no
apparent naturally occurring hybrids have thus
far been seen.

LITERATURE CITED

iae M. L.

1950. Gray's eium i Botany, 8th
tion. American Book Com

, New York,

n
A Manual of the Botany ofthe North-
ern United States. James Munroe & Co., Cam
ndon.

. Carex eda eii lied Pp. 108-
112 in A report o threatened, or en-
dangered forest- ated ae sae of the south.
Volume I. USDA Forest Service Technical Publ.
R8-TP 2. Atlanta, Georgia.

KUKENTHAL, G. 1909. Cyperaceae—Caricoideae.

1931-1935. Cariceae. InN. Amer.
Fl. 18(7): 1-478.

— Robert Kral, Department of Biology, Vander-
bilt University, Nashville, Tennessee 37235,
U.S.A.; James Manhart, Division of Biological
Sciences, The University of Michigan, Ann Arbor,
Michigan 48109, U.S.A.; and Charles T. Bryson,
Southern Weed Science Laboratory, P.O. Box
350, Stoneville, Mississippi 38776, U.S.A

NOTES ON CHROMOSOME CYTOLOGY OF
RUTACEAE— DIOSMEAE

The tribe Diosmeae R. Br. emend. Benth. &
Hook. f. (1862) of the Rutaceae is strictly African
in distribution and consists of the widespread
tree Calodendrum capense (L.f.) Thunb. (south-
ern Cape to north Kenya) and some 280 species
of small to medium-sized shrubs, mostly of er-
icoid habit. The tribe is, with the exception of
Calodendrum, restricted to southern Africa and
largely to the Cape Region of the south and west
coast of South Africa. Taxonomic revisions of
the tribe have recently been published for all but
the large genus Agathosma (Strid, 1972; Wil-
liams, 1975, 1978, 1981a, 1981b, 1982) but
chromosome cytology of the group is poorly
known.

Apart from several counts by Strid (1972) for
Adenandra, there are only scattered reports for
a few genera. The monotypic Calodendrum is a
high paleopolyploid with 2n = 54 (Honsell, 1954;
Smith-White, 1954). In Agathosma, A. crenulata
(L.) Pill. has been reported as having 2n = 45
(as Barosma) (Riley & Hoff, 1961), while Guerra
(1984a) found 2n = 26 in A. apiculata E. Meyer
(also as Barosma) and A. lanceolata (L.) Engl.

There are two counts for Coleonema pulchel-
lum I. J. Williams (as C. pulchrum Hook.), Smith-
White (1954) reporting 27 — 36 and Guerra
(1984a) 2n = 34; Guerra also reported 27n = 34
in the closely related C. album (Thunb.) Bartling
& Wendl. Numbers reported in Adenandra are
2n — 28 in two species, 38 in one more, 42 or
ca. 42 in two species including A. fragrans, and
2n — 48-50 in another three. In conjunction with
continuing systematic studies of Diosmeae a pre-
liminary investigation of the chromosomes of
some genera was undertaken and this report is
the result of the study.

All counts were made from mitotic metaphase
in root tip squashes of germinating seeds, fol-
lowing a method described fully elsewhere
(Goldblatt, 1979, 1980). Chromosome number
and voucher information are as follows:

Diosma subulata Wendl. 2n — 30. South Af-
rica, Cape, ex hort. Williams, Williams s.n.
(NBG); Wortelgat, Caledon Div., Williams 1721
(NBG).

Diosma aristata I. J. Williams 2n = 30. South
Africa, Cape, ex hort. Williams, Williams s.n.

(NBG)
ANN. Missouni Bor. GARD. 74: 443-444. 1987.

Diosma oppositifolia L. 2n = 30. South Africa,
Cape, Vogelgat, Caledon Div., Williams 2406
(NBG).

Euchaetes avisylvana I. J. Williams 2n = 28.
South Africa, Cape, Grootvadersbos, Heidelberg
Div., Williams 2377 (NBG).

Adenandra frag (Sims) Roemer & Schultes
2n = 42. South Africa, Cape, Grootvadersbos,
Heidelberg Div., Williams 2378 (NBG).

DISCUSSION

The count of 2n = 42 for Adenandra fragrans
confirms the earlier report by Strid (1972) for
this species. Reports here for Euchaetes and
Diosma are the first records for these genera. On
the basis of four counts for three species of Dios-
ma, all 2n = 30, we suggest that x = 15 is basic
for the genus. In Euchaetes the single count sug-
gests a generic base of x = 14; x = 7 seems un-
likely since all other counts in Diosmeae are at
paleotetraploid or paleohexaploid levels.

Basic chromosome number for Rutaceae is
most likely x = 9 as suggested by Smith-White
(1954). This hypothesis is founded on the pre-
dominance of n = 18 and 36 in the unspecialized
Xanthoxyleae and Flindersieae, and Ehrendorfer
(1981, 1987) has also pointed out the occurrence
of only n = 9 and 18 in Aurantieae. Data for
Diosmeae throw no direct light on the possible
base number for the family, as they are relatively
specialized. However, the number in Ca/oden-
drum, n = 27,is most likely paleohexaploid based
on x = 9, and the same base number is by ex-
tension probable for all Diosmeae, given the bas-
al position of Calodendrum in the tribe. Other
genera of Diosmeae that have been counted ap-
pear fundamentally paleotetraploid, a view en-
dorsed by Guerra (1984b) for Coleonema based
on a comparison of genome size in Rutaceae.
Thus Coleonema, either n = 18 or 17, is probably
hypotetraploid based on x = 9. The base of x =
15 in Diosma may represent either further aneu-
ploid decrease from an ancestral x — 18, or less
likely, allopolyploidy from hypothetical ances-
tors with x = 7 and 8. Euchaetes, closely related
to Diosma, with n = 14 fits with the apparent
trend for decrease from an ancestral tetraploid
base. Adenandra appears basically paleotetra-
ploid (? x = 14) but some species may be octo-

444

ploid and derived from the secondary hypotet-
raploid base. This may best explain the range
from n = 25, 24, 21, and 19, as well as n = 14
in two species.

Further counts in Diosmeae are needed to re-
solve the several questions that this initial study
has posed. The variation in chromosome num-
ber in the tribe is unexpected and is likely to be
of considerable value in future studies of generic
limits and of the relationships of genera and
species, and ultimately of phylogeny.

LITERATURE CITED

BENTHAM, G. & J. D. HOOKER. 1862. Rutaceae. In
enera denies 1: 278-306.
EHRENDORFER, Speciation Mpeg in woody
angiosperms roi l origin. Pp. 479—509 in C.
Barigozzi (editor), Mechanisms of Speciation. Alan
Liss, New York.
87. Affinities of the African dendroflora:
suggestions from karyo- and chemosystematics.
onogr. Syst. Bot. Missouri Bot. Gard. (in press).
GOLDBLATT, P. 1979. Chromosome cytology and
hien Pd in Galaxia (Iridaceae). Pl. Syst.
Ev 133: 61—69.

Redefinition of Homeria and Moraea
ue in the light of pb: data, with
me nov. Bot. N 5.
GUERRA, M. pos S 1984a. New chromosome num-
ers in Rutaceae. Pl. Syst. Evol. 146: 13-30.

. Cytogenetics of Rutaceae II. Nuclear
DNA content. Caryologia 37: 219-226.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

HoNSELL, E. 1954. Osservationi sulla struttura
dell'ovulo e sulla cariologio di Calodendron ca-
pense Thunb. e Pilocarpus pennatifolius. Ann. Bot.
(Rome) 24: 438-447.

RirEv, H. P. & V. J. Horr. 1961. Chromosome stud-
ies in some South African dicotyledons. Canad. J.
Genet. Cytol. 3: 260-271.

SMITH- WHITE, S. 1954. Chromosome numbers in the

luni mid development of the tribe in rem Austra:
ustral. J. Bot. 2: 287-303.
. Revision of the genus Adenandra
(Ru ta ceae). Opera Bot. 32: 1-112.
WILLIAMS, ION. 1975. Studies in the genera of Dios-
meae (Rutaceae): 5. J. S. African Bot. 41: 180-
i

978. Studies in the genera of Diosmeae (Ru-
23

981a. Studies in the genera of Diosmeae
(Rutaceae): 9. A revision of the genus Coleonema.
J. S. African Bot. 47: 63-102.

1981b. Studies in the genera of Diosm
(Rutaceae): 10. A review of the genus E riii
J. S. African Bot. 47: 157-193.

1982. Studies in the genera of Diosmeae (Ru-
tacea e): 14. A review of the genus Diosma. J. S.
African Bot. 48: 329-407.

— Peter Goldblatt, B. A. Krukoff Curator of Af-
rican Botany, Missouri Botanical Garden, P.O.
Box 299, St. Louis, Missouri 63166, U.S.A.; and
Ion Williams, 29 Tenth St., Voelklip, Hermanus
7203, South Africa.

THE FRUITS OF DECARYDENDRON (MONIMIACEAE)

Decarydendron Danguy is a poorly known ge-
nus of Monimiaceae (subfamily Mollinedioi-
deae, tribe Hedycareae) comprising three species
endemic to Madagascar (Danguy, 1928; Cavaco,
1958a, 1958b; Lorence, 1985). Fruiting material
of the genus, hitherto unknown, is described here
for the first time based on a specimen of D. per-
rieri Cavaco recently collected by L. J. Dorr and
L. Barnett in Madagascar. The following descrip-
tion is based on FAA-fixed fruits of Dorr & Bar-
nett 3203 (Madagascar. Province Tamatave: en-
virons d'Andasibe-Périnet, 18?56'S, 48?25'E, 2-
4 Nov. 1984). Voucher specimens have been de-
posited at MO and TAN, and the fixed collection
is at PTBG.

Fruiting receptacle attached at or below ground
level, terminal on a finely velutinous peduncle
2.5-3 mm diam., swollen, fleshy, obconical-tur-
binate, initially 3 cm diam., during development
curling outwards and splitting irregularly into 3
partly everted segments, externally and inter-

carpels 12, numerous
aborted carpels. Mature carpels sessile to sub-
sessile, obpyriform, 2-2.5 cm long including the
constricted, curved, beak-like apex 5-7 mm long,
1.3-1.6 cm diam., externally yellowish brown,
corky, bearing circular to elliptic corky lenticels

S. 1
1cm Ld
T
faber

with attached carpels.—

ANN. MissouRi Bor. GARD. 74: 445-446. 1987.

over most of the surface; exocarp fleshy, to ] mm
thick (thicker at apex), the surface corky, par-
enchymatous, the tissue replete with densely ag-
gregated brachysclereids, underlain by a vascu-
larized zone, the mesocarp parenchymatous with
scattered + cuboidal idioblasts; endocarp hard,
white, + smooth, discontinuous apically at the
micropyle, 0.8-1 mm thick, co

densely packed, radially oriented layer of narrow,
thick-walled, fusiform columnar sclereids; testa
0.1-0.2 mm thick, brown, composed of 2 layers
of tangentially elongated cells with slightly thick-
ened walls, underlain by an endotesta many lay-
ers thick of short, rounded tracheids with sca-
ariform thickenings; tegumen 3-4 layers thick,
composed of elongate thin-walled cells. Endo-
sperm about 1 cm diam., white, copious, oily,
interspersed with crystals. Embryo situated api-
cally in the endosperm below the micropyle, cy-
lindrical-clavate, 2-3 mm long, | mm diam. dis-
tally, the cotyledons erect, + appressed, about
half the total embryo length. Figure la, b.

DISCUSSION

In a recent monographic treatment of the

DANIEL nein

Fruiting receptacle and carpels of Decarydendron perrieri. —a.
b. Carpel, longitudinal section. Based on Dorr & Barnett 3203 (PTB

Mature, irregularly split receptacle
G).

446

any of the Malagasy genera: both androecious
and gynoecious flowers with shallowly concave
receptacles bearing 1—2 whorls of large, imbricate
tepals; flowers that unfold gradually at anthesis
without splitting into segments as in the other
three genera (i.e., Ephippiandra Decne., Monim-
ia Thouars, and TambourissaS
flowers with numerous (ca. 300—1,000) free, sub-
sessile, clavate carpels. As Decarydendron closely
resembles Hedycarya Forster & Forster f. from
Oceania and various Pacific islands in terms of
gynoecious floral morphology, it was postulated
that mature fruits of the former genus would most
likely resemble those of the latter one (Lorence,
1985

) gynoecious

The findings reported here confirm that ma-
ture fruits of Decarydendron perrieri do strongly
resemble those of other genera in the tribe He-
dycareae (Philipson, unpublished) and support
its placement there. Members of this tribe are
characterized by gynoecious flowers with a non-
calyptrate floral cup that encloses few to many
free carpels, and fleshy discoidal or cupuliform
fruiting receptacles that gradually split or become
everted as the carpels mature. Among the other
Malagasy genera belonging to the Hedycareae,
the free, sessile to subsessile carpels of Decary-
dendron are less specialized than those of Ephip-
piandra, which are also free but partly immersed
in cupules formed by the receptacle. Fruits of
both these genera are in turn less specialized than
the inferior, syncarpous carpels of Tambourissa,
which are completely immersed in and united
with the ovary wall (Lorence, 1985). Thus, fruit
morphology also supports the suggestion that the

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

three Malagasy genera of Mollinedioideae form
part of an increasingly specialized evolutionary
series.

Among the extra-Malagasy members of the
tribe Hedycareae of the subfamily Mollinedioi-
deae, Decarydendron is most closely allied to
Levieria Becc. from Malesia and Australia, and
particularly to Hedycarya, in terms of floral and
fruit morphology. Decarydendron differs from
both these genera in being monoecious, in having
sexually mixed cauliflorous inflorescences, and
in having gynoecious flowers with more numer-
ous clavate carpels having the stylar canal situ-
ated in the basal half of the carpel.

I thank L. Dorr of the Missouri Botanical Gar-
den for providing fixed material of Decaryden-
dron perrieri and Danial Cobian of the Instituto
de Biología for the illustration.

LITERATURE CITED

CAVACO, A. Sur le genre Decarydendron
(Monimiacées). ips Soc. Bot. France 105: 38-39
195 espéce nouvelle de Decaryden-

ontribution à l'étude des Moni-
cées de Madagascar. Bull. Mus. Hist. Nat.
(Paris) ae 278-280.
Lor 5. A monograph of the Monim-
iaceae a in the Malagasy Region segera
st Indian Occan). Ann. Missouri Bot. Gard. 7
1-16
PHILIPSON, w. R. A m of the Monimiaceae
(unpublished manuscript).

— David H. Lorence, Pacific Tropical Botanical
Garden, P.O. Box 340, Lawai, Kauai, Hawaii
96765, U.S.A

RECONSIDERATION OF THE GENERIC PLACEMENT
OF PALICOUREA DOMINGENSIS
(RUBIACEAE: PSYCHOTRIEAE)

Although the Caribbean species sometimes
known as “Palicourea domingensis” has com-
monly been treated in Palicourea, it lacks the
distinguishing features of this genus and is better
placed in Psychotria.

Palicourea Aublet (Rubiaceae: Psychotrieae) is
distinguished within its tribe by comparatively
long (tubes 5—40 mm long), tubular corollas that
are -— us elation! gwalen at the base with
aringoftri
abgve the basal swelling. Characteristically, the

ally and bright

ly colored, usually red, yellow, blue, or purple.
The thyrsiform inflorescences are typically open,
with well-developed branches, bracts, and ped-
icels. This genus has been loosely circumscribed
by many authors, who have used variously one
ora few of these features or characters to separate
it from Psychotria

The species of Psychotrieae now usually called
“Palicourea domingensis” (Jacq.) DC. is a shrub
or small tree of moist forest on soils derived from
limestone. It is distributed through most of the
Caribbean islands (Cuba, Jamaica, Hispaniola,
Puerto Rico, St. Thomas, St. Croix, Tortola,
Antigua, St. Kitts, St. Eustatius, Guadeloupe,

Nicaragua, although it has not been previously
reported from these areas (Standley & Williams,
1975).

This species is characterized by salverform co-
rollas with tubes 5-15 mm long. The corollas are
predominantly white, although often tinged with
rose or purple. Their tubes are glabrous exter-
nally and internally, and are not at all swollen
or asymmetrical at the base, although the tube
may be curved somewhat through its length. The

inflorescence bracts are poorly developed or
lacking.

Thus, “Palicourea domingensis” lacks the dis-
tinctive floral features and the common inflo-
rescence characters of Palicourea. It has been
placed in Palicourea by most authors apparently
based on its relatively long, tubular, often rosy
corollas, combined with an inconsistent circum-

ANN. MissouRi Bor. GARD. 74: 447—448. 1987.

scription of Palicourea and a respect for previous
practice.

This species is better placed in Psychotria subg.
Heteropsychotria Steyerm., which is closely re-
lated to Palicourea. The floral features of Psy-
chotria domingensis are similar to those of other
species of Psychotria, and the inflorescence char-
acters described above that are rare or lacking in
Palicourea are common conditions in this sub-
genus. This relationship was recognized previ-
ously by Standley when he described this same
species from Nicaragua under the synonym Psy-
chotria mombachensis Standley. Although the
corollas of Psychotria are described in most treat-
ments as comparatively short, and typically do
have tubes 5 mm long or shorter, long corollas
occur in the genus as well, as in Psychotria chia-
pensis Standley which has corollas with tubes 30-
50 mm long. In fact, within subg. Heteropsy-
chotria, Psychotria domingensis 1s similar in sev-
eral characters to Psychotria chiapensis and to
Psychotria gardenioides (Scheidw.) Standley:
these species share glomerulate, subsessile flow-

or rosy corollas with internally glabrous tubes 20
mm long or longer. (The last two species differ
from Psychotria domingensis in their longer co-
rolla tubes and well-developed inflorescence
bracts.)

efore, “Palicourea domingensis” seems
better classified in Psychotria. In this case, the
correct name of this species is Psychotria dom-
ingensis Jacq. A complete list of synonymy and
citations of specimens from Central America are
presented below.

NOMENCLATURE

Psychotria domingensis Jacq., Enum. Pl. Carib.

60. Palicourea domingensis (Jacq.)

DC., Prodr. 4: 529. 1830. TYPE: Santo Do-
mingo.

Hk pavetta d) Prodr. 45. 1788. Pavetta pen-
andra Sw. . Occ. 1: 233. 1797. Palicourea
And (Sw) DC. Prodr. 4: 525. 1830. Palicourea
pentandra (Sw.) K. Schum., Nat. Pflanzenfam.
Bis 115. 1891, nom. illeg.. Art. 63. TYPE: Ja-
otographs A, N

aica ( NY).
m ones tabernaefolia Poir., Encycl. 5: 704. 1804.

448

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FIGURE 1.

Palicourea ig sib Lene Ad ) DC., Prodr. 4: 525.
0. TYPE: Santo ingo.
Psychotria angustifolia Poir. ps 5: 703. 1804. TYPE:
anto Domingo.

—— mombachensis Standl.,
Mus., Bot. Ser. 8: 188. 1930. TYPE: Nicara-
ie coffee plantation, Mombacho Volcano, 600-
750 m, Maxon, Harvey & Valentine 7818 (holo-

type, F)

Specimens examined from Mexico and Central
America. co. TABASCO: La Palma, Balancan,
Matuda 3236 (NY).

GUATEMALA. PETEN: Tikal, Contreras RG 1413
(TEFH); Dolores, Contreras 2503 O); Macanche,
Contreras 5847 (DS); Chincila, pede ease 6360 (MO);
Isla del Armadillo, Laguna Macanche, Contreras 7 1 A
(MO); Sayaxche Road, km 59, Contreras 7247 (M
TEFH); Guayacuan, La Pita, Contreras 7477 MOS
Tikal, Contreras 7707 (MO); between Paso Caballo and
Carmelita, Cox 2835 (EAP); Tikal, Molina 15750

)

Publ. Field Colom-

BELIZE. CAYO: vicinity of La Flor at Río de La Flor,
6 mi. S of San Grano de Oro, 1,700-2,700 ft. [520-
820 m], Croat 23739 (MO), Dwyer 10876 (MO); in
advanced forest, Lundell 6199 (NY); W of Humming-
bird Hwy. 7 mi. S of junction with Western Hwy., N

Distribution of Psychotria domingensis Jacq. in Mexico, Central America, and the West Indies.

boundary of Roaring River Estate, 80 m, Spellman &

1915 (MO); 1,700 yd. W of Hummingbird Hwy., 7 mi.
of junction with Western Hwy., 80 m, Spellman &
Newey 2010 (MO); TOLEDO: Dwyer 9863 (MO
I thank the curators of the following herbaria,
who very kindly made specimens available for
study: A, CAS, DUKE, EAP, F, GH, M, MO,
NY, TEFH, and US; I thank in particular Dr.
James Ackerman of UPRRP, who provided
working space. I also thank Orlando Matos Con-
cepción for help in preparing the distribution
ata

LITERATURE CITED
STANDLEY, P. C. & L. O. WiLLIAMs. 1975. Rubiaceae.
In Flora of Guatemala, Part XI. Fieldiana Bot.
24(11): 1-274
— Charlotte M. Taylor, Department of Biology,
Universidad de Puerto Rico— Colegio Universi-
tario de Cayey, Cayey, Puerto Rico 00633

CLARIFICATION OF THE TAXONOMY OF
LEUCAENA SALVADORENSIS
STANDLEY EX BRITTON & ROSE

The nomen nudum Leucaena salvadorensis
Standley was first published by him in Standley
& Calderón (1925) in a list of El Salvador plants,
without a complete description or designation of
type. In their revision of the genus, Britton &
Rose (1928) supplied a description and desig-
nated a type, giving the authority as Standley.
Later, Standley & pue uem (1946) reduced the
species, cited as L. salvadorensis Standley ex
Britton & Rose, to synonymy under L. shannonii
J. D. Smith. Prewbaker et al. (1972) and Brew-
baker in a mimeographed leaflet (1978) placed
L. salvadorensis under the synonymy of L. leu-
cocephala (Lam.) de Wit, and more recently
Brewbaker & Ito (1980) treated it as a subspecies
of L. leucocephala (also see Brewbaker’s mim-
eographed revision, 1984). Zarate (1982, 1985)
concluded that it was the same as the small-leaf-
let form of L. shannonii and returned it to syn-
onymy with that species.

Recently, Colin Hughes, Brian Styles, and An-
gela Laguna have collected material in Honduras
and Nicaragua (C. Hughes 332, 334, 446 (FHO,
MEXU); C. Hughes & B. Styles 37, 102, 129
(FHO, MEXU); A. Laguna 427 (HNMN,
MEXU)) referable to this taxon. From study of
these specimens it became apparent that it de-
serves recognition as a subspecies of L. shan-
nonii.

The dome-shaped or truncate-conic, oblong
petiolar glands and the corolla-to-calyx length
ratio of ca. 0.7 in L. shannonii and L. salvador-
ensis clearly distinguish these taxa from L. /eu-
cocephala with suborbicular or elliptic or ob-
ovate thick-bordered and furrowed to concave
glands and a corolla-to-calyx length ratio of ca.
0.5. The more numerous pairs of pinnae and
leaflets, 6-11 and 23-35, respectively; consis-
tently narrow oblong leaflets, ca. 1.5 cm long;
florets 4 mm long; and pods strongly stiped, 1 1-
21 cm long and ca. 2.5 cm wide, separate it from
the typical element of L. shannonii with (2—)4—

ANN. MISSOURI Bor. GARD. 74: 449. 1987.

6(-8) pairs of pinnae, 8—22 leaflet pairs, very
variable leaflets (0.8—)1—2 cm long, florets 3-3.5
mm long, and slender-stiped pods ca. 15 cm long
and 1.4-2.2 cm wide. Therefore, the following
combination is made.

Leucaena shannonii J. D. Smith subsp. salva-
dorensis (Standley ex Britton & Rose) S. Zá-
rate, comb. et stat. nov. Basionym: L. sal-
vadorensis Standley ex Britton & Rose, Fl.
N. Amer. 23(2): 125. 1928. TYPE: El Sal-

vador. Morazán: Jocoro, Calderón 2031
esum NY).

LITERATURE CITED

BREWBAKER, J. L. 1978. Guide to the systematics of
the genus Leucaena (Mimosaceae). C.I.A.T. Cali.
dp h.

. 1984. Revision ofthe genus Leucaena (Mim-

osoideae: Leguminosae). Honolulu. Mimeograph.

TO. 1980. Taxonomic studies of the

. 1972. Va-
rietal variation and yield trials of Leucaena leu-
cocephala (Koa pik in Hawaii. Haw. Agric. Exp.
Sta. Res. Bull. 166:

BRITTON, N. L. & J. N. Rost.

N. Amer. 23(2): 12

PE si as PC & S. abu 1925. Lista Preli-
minar de Plantas de El Salvador. San Salvador.

. STEYERMARK. 1946. Flora of Guatemala.

denar Bot. 24(5): 47-48.

TE, P. S. 1982. Las especies de Leucaena Benth.
de Oaxaca con notas sobre la sistemática del gé-
nero para México. Thesis. Facultad de Ciencias,

, México.

1984 [1985]. Taxonomic revision of the ge-

nus Leucaena Benth. from México. Bull. IGSM

12: 24-34

1928. Mimosaceae. Fl.

—Sergio Zárate Pedroche, Instituto de Biología,
Herbario Nacional de México (MEXU),
U.N.A.M., México. Presently graduate student
supported by CONACYT (Reg. No. 49993), De-
partment of Biology, Indiana University, Bloom-
ington, Indiana 47401, U.S.A.

AN UNUSUAL NEW SPECIES OF HELMIOPSIS H. PERRIER
(STERCULIACEAE) FROM MADAGASCAR

Eight species of He/miopsis H. Perrier (Ster-
culiaceae) currently are recognized in the Flore
de Madagascar et des Comores (Arénes, 1959).
All are characterized by monodelphous androe-
cia with staminodes opposite the petals, sessile
ovaries, caducous petals, capsular fruits with api-
cally winged seeds, and overall vestitures of pel-
tate or fimbriate (i.e., peltate with fimbriate mar-
gins) scales. All species described to date are
restricted to western Madagascar. A new species
of Helmiopsis is described here from the north-
ern mountains of Madagascar, also part of the
phytogeographical Domaine de l'Ouest of the is-
land (Humbert, 1965). This species combines
several features that are unusual for the genus.
For example, its spheroid, rather than conical,
capsule morphology is unique and provides the
basis for the name He/miopsis sphaerocarpa L.
Barnett. The seed wings of H. sphaerocarpa are
atypical for the genus in being reduced to only a
thin, ventral keel; nnn E angs of Hel-
e, to 10 mm
in length. The inflorescence i IS ssa in struc-
ture, being multibranched and many-flowered,
as opposed to the few-flowered cymose inflores-
cences more commonly found in the genus. The
terminal nature of the inflorescence is also un-
usual in this species, although terminal inflores-
cences also have been observed in isolated col-
lections of H. pseudopopulus (Baillon) Capuron
(e.g., Capuron 24661, P). Finally, this new species
is unique in bearing glandular tissue both on the
calyx lobes and on the petals; in all other Hel-
miopsis species with glandular tissue, the tissue
is restricted to either the calyx or the corolla, but
is never found in both perianth whorls.

Helmiopsis sphaerocarpa L. Barnett, sp. nov.
TYPE: Madagascar: Centre (Nord) jusqu’aux
confins de l'Ouest (Nord), Massif de la Mon-
tagne d'Ambre, créte entre les bassins de la
riviére des Makis et de la riviére d'Anka-
zobe, entre 800 et 600 m d'alt., 26-27 May
1970 (fl), Capuron 29194 SF (holotype, P;
isotype, TEF). Figure 1

rbuscula vel arbor, squamarum lepidotarum de-

nales, 15-30 flor
bis intus Mn duce Petala 5, E glandulosa. Tubus

ANN. Missouni Bor. GARD. 74: 450-452. 1987.

staminalis | mm altus; staminibus 10, staminodiis 5.
Ovarium 5-loculare, loculis 2-ovulatis; stylo glabrato,
stigmate leviter 5-lobata. Capsula didi loculici-
da. Semen carina ventrale membranace

Large shrub or tree to 8-12 m tall, branches
terete, finely striate, new growth sparsely peltate-
scaled, older twigs glabrate. Leaves alternate, de-
ciduous, blades broadly ovate to narrowly ob-
ovate, 6.5-12 cm long, 4—9 cm wide, apex acute
to acuminate, base shallowly cordate to cordate,
palmately 5(-7) nerved, midvein and secondary
veins conspicuous, raised below and slightly de-
pressed above, discolorous, margins irregularly
crenulate, lower surface with scattered peltate and
fimbriate scales, becoming glabrate; petioles 1-
6 cm long, with scattered peltate and fimbriate
scales. Inflorescences terminal and in the axils of
uppermost 3-5 leaves, determinate, branching
3-5 times, bearing 15-30 flowers. Floral buds
ovoid, 4-5 mm long, 4-4.5 mm wide. Calyx
5-lobed, lobes connate only at the bases, lanceo-
late, ca. 7 mm long, 3 mm wide, outer surfaces
densely covered with peltate scales, inner sur-
faces glabrate and with an arc of glandular tissue
at the base. Petals 5, white (according to label
data of Capuron 29194 SF), obovate, asymmet-
rical, 7-8 mm long and 7 mm wide, inner surface
gland-dotted toward the base. Androecium co-
roniform, staminal column ca. 1 mm tall; sta-
mens 10, ca. 3 mm long from the base of the
column, each of the 5 antisepalous pairs alter-
nating with a staminode; filaments 1-1.5 mm
long; anthers ca. 2 mm long; staminodes 5, ob-
lanceolate, 4-5 mm long, ca. | mm wide. Ovary
5-locular, densely peltate-scaled; ovules 2 per
locule, collateral; style 5-6 mm tall, glabrate;
stigma obscurely 5-lobed. Fruit capsular, woody,
spheroid, dorsally loculicidal, 6-8 mm long, 8-
10 mm wide, subtended by a woody, persistent
calyx. Seeds 2 per carpel, asymmetrically ovate
and laterally flattened, 5-6 mm long, 3-4 mm
wide, each with a narrow, membranous, ventral

eel.

Additional specimens examined. MADAGASCAR.
NORTHERN SECTOR. DISTRICT DIÉGO-SUAREZ: Versant Est
du massif de I’ at ati 17 Dec. 1966 (fr), Capuron
27349 SF (P, TE ontagne des Français, 11 e Fla
1952 (fr), Service Forestier 5673 SF (P, TEF);
lafa mbodipo-Antsahalalina, 7 June 1956 (fl), po
vice Forestier 15962 SF (P, TEF).

1987] NOTES 451

Ficure 1. A-I. Helmiopsis sphaerocarpa. — A. — B. Schematic diagram of an inflorescence axis. — C.
p . —D. Inner surface of calyx lobe showing ndun ce gii Inner surface of petal showing glandular
—F. Detail of the Werben ies —G. Gynoecium. —H. Fruit. —I. Seed. (A, B. Capuron 27349 SF; C-G.

EL... 29194 SF; H, I. Service Forestier 5673 SF.

452

Helmiopsis sphaerocarpa occurs on mountain
slopes between 50 and 800 m altitude. It has been
reported on black volcanic soils, but its presence
on the Montagne des Francais suggests that it
also may occur on limestone (Lemoine, 1906).

The 5-carpellate gynoecium and reduced stig-
mata of Helmiopsis sphaerocarpa place this
species in Helmiopsis subg. Helmiopsis. Its
broadly ovate leaves, glandular petals, and ten
stamens ally it with sect. Glandulipetalae Arénes
in subg. Helmiopsis. Helmiopsis pseudopopulus
(Baillon) Capuron, a member of the same sec-
tion, may be the most closely related species; it
also has a many-flowered inflorescence, occa-
sionally with a terminal axis. Glandular tissue at
the base ofthe calyx, however, has been observed
in only one other He/miopsis species, H. inversa
H. Perrier (sect. Helmiopsis).

This research was supported by NSF Doctoral
Dissertation Improvement Grant BSR-8414032.
thank the curator of P for the loan of specimens,
and of TEF for permission to examine material.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

The illustration was prepared by P. Parker. L.
Dorr and B. Stein provided helpful comments
on the manuscript. Work in Madagascar was
made possible by collaboration between the Mis-
souri Botanical Garden and the Parc de Tsim-
bazaza, Antananarivo, Madagascar.

LITERATURE CITED

ARÈNES, J. 1959. 131e Famille—Sterculiacées. Jn H.
Humbert (editor), Flore de Ma adagascar et des Co-
es. Mus ist. Nat., Ph

Notice de la carte, Madagascar. S
fique et Technique de l'Institut Francais de Pon-
dichéry, Pondichéry, India

LEMoINE, P. Etudes géologiques dans le nord
de Madagascar. Librairie Scientifique A. Her-
mann

—Lisa C. Barnett, Department of Botany, Uni-

versity of Texas at Austin, Austin, Texas 787 13-
7640, U.S.A

WITHERINGIA FOLLICULOIDES (SOLANACEAE):
A NEW SPECIES FROM COSTA RICA

The genus Witheringia L'Hér. embraces about
a dozen species. One species, W. solanacea
L'Hér., is weedy in nature and ranges from north-
ern Mexico to Bolivia. The other species tend to
be more localized, and the genus seems to be best
developed in mature wet forests of Costa Rica,
Panama, and northern South America. Wither-
ingia is a member of subfamily Solanoideae. The
species of Witheringia are mostly shrubs or small
trees with few- to many-flowered inflorescences
of yellow or green flowers held under the leaves,
and berries held erect above the foliage. The ca-
lyces range from minute and nonaccrescent to
arge and, as in the present species, completely
enveloping the fruit.

Witheringia folliculoides D'Arcy & J. L. Gentry,
sp. nov. TYPE: Costa Rica. Puntarenas: moist
forest in valley bottom, tropical wet forest
with open understory on steep slopes and
ridges and areas of secondary vegetation
north and west of the air field 5
Rincón de Osa, 50-200 m, J. L. Gentry &
Burger 2844 (holotype, MO-2825976; iso-
type, F)

°
amc

Frutex 3-5 m altus, ramis hornotinis dun d
dads Folia integra, cupias lanceolatave, saepe o
qua, 9.5-23 cm longa lata, venis n: D lanis
14

tiolis gracilibus. nET pauci, 3-4 mm longi. Flores
5-meri, calyce parvo, 2.5-4 mm longo, glabrato, cam-
panulato, apice tru Bae in statu fructificanti mag-

2.5-3 mm
ee ae obtusis, 3
escentibus, antheris oblongis, haud apiculatis,
longitudinaliter dehiscentibus. Acinus subglobosus, 12
olliculo tectus, calyce in folliculum
e wA um maturescenti.

Small tree 3-5 m tall; young branches slender,
glabrate, drying smooth. Leaves entire, drying
concolorous, mostly elliptic or lanceolate, some-
times oblanceolate, often oblique, 9.5-23 cm long,
2.3-7 cm wide, apically short acuminate, basally
mostly cuneate, the veins plane or slightly im-
pressed above, sometimes drying somewhat ru-
gose, the costs and major lateral veins elevated
beneath, the lateral veins arching, 6-14 on each
side; minor leaves lanceolate to broadly elliptic
or rotund, 2.3-7.8 cm long, 1.4-4.2 cm wide,
acute to acuminate or sometimes rounded api-

ANN. Missouni Bor. GARD. 74: 453-454. 1987.

cally, cuneate to rounded basally; petioles slen-
der, 4-9 mm long, those of the minor leaves 2-
3 mm long. Inflorescence with the peduncle
mostly subobsolete but sometimes 2-4 mm long
and the flowers appearing fasciculate but only
one or two appearing at a time; pedicels few, 1—
7 (evidenced by scars), 3-4 mm long, becoming
4—7 mm long, perhaps all but one flower abort-
ing. Flowers 5-merous; calyx campanulate, api-
cally truncate, 2.5-4 mm Hong, accrescent in fruit,
drying dark, minutely y pubescent with
short curved simple hairs anoetly on the apical
portion, glabrescent, inconspicuously ribbed; co-
rolla yellowish, apparently unmarked, glabrous
outside, tubular campanulate, the tube 2.5-3 mm
long, the throat pubescent with reduced hairs,
the limb lobed less than halfway down, the lobes
ovate, 3-4 mm long, marginally puberulent; sta-
mens with the filaments pubescent at the base,
apically glabrous, 1.5 mm long, the anthers yel-
low, oblong, 2 mm long, not apiculate; ovary
glabrous, the style equalling the stamens, gla-
brous, the stigma minute. Fruit a subglobose
?juicy berry 10-12 mm across, loosely and com-
pletely enveloped by the strongly accrescent ca-
lyx; fruiting calyx rotund, much enlarged and
inflated around the berry but open at the apex,
apparently unribbed, glabrous, 15-20 mm across,
approximately twice the length of the fruit; seeds
numerous (?ca. 40), dark brown, flattened, re-
niform, 2-3 mm across, the testa wavy retizulate
on the face, scalariform on the rim; embryo
broadly horseshoe-shaped, terete throughout
(lacking any notching along its length).

Paratypes. CosTA RICA. PUNTARENAS: Osa Penin-
sula, tropical wet forest along dry stream bed parallel-

d cón de Osa, 50-200 m, Gentry & Burger 2842 Adi
X.S tto airport

Rincón de Osa, 20-300 m, . Liesner 1887 (MO).

This species is distinct from most other species
of Witheringia in its bladdery fruiting calyx, whic
loosely and completely envelops the fruit. It oc-
curs in southern Costa Rica where a number
of other species of the genus occur. It is perhaps
most similar to W. exiguiflora D’Arcy and W.
morii D'Arcy, which also occur in western Pan-

454

ama and southern Costa Rica, and which both
have accrescent calyces, but their calyces do not
form the large bladderlike enclosures of W. fol-
liculoides. Witheringia folliculoides differs fur-
ther from W. exiguiflora in its tubular campan-
ulate corolla, in its smaller and much less
coriaceous leaves, and in its larger fruiting calyx.
From W. morii it differs in its broadly open co-
rolla and its few-flowered inflorescences. In hab-
itat W. folliculoides also differs from these two
species: it occurs near sea level in the seasonally
dry Osa Peninsula of Costa Rica, whereas W.
exiguiflora and W. morii occur in Am
rainy forests. Both W. exiguiflora and W. mo

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

occur in the uplands of 1,100-1,800 m in Costa
Rica and Panama, but W. exiguiflora also ranges
down to near sea level on the Atlantic slopes of
Panama.

This work was supported by National Science
Foundation grant no. BSR-8305425 (William G.
D’Arcy, principal investigator).

— William G. D'Arcy, Missouri Botanical Gar-
den, P.O. Box 299, St. Louis, Missouri 63166;
and Johnnie L. Gentry, Jr., The University Mu-
seum, University d Arkansas, Fayetteville, Ar-
kansas 72701, U.S

ESCHWEILERA COSTARICENSIS (LECY THIDACEAE):
A NEW SPECIES FOR THE FLORAS OF
COSTA RICA AND NICARAGUA

Eschweilera C. Martius ex DC. is the largest
(ca. 90 species) and most poorly known genus of
neotropical Lecythidaceae. The genus is char-
acterized by zygomorphic flowers with coiled an-
droecial hoods, absence of anthers and produc-
tion of nectar in the androecial hoods, bilocular
ovaries with basal placentation, seeds often with
lateral arils, and lack of differentiated cotyledons
(for illustrations of these features see figs. 9, 10G,
13, and 18 in Prance & Mori, 1979). The Central
American species of Eschweilera provide ex-
amples of how poorly collected the species of
Eschweilera are. Woodson (1958) included only
four species of Eschweilera from Panama, where-
as today we know ofat least 14 from that country.
Further north, species diversity of Eschweilera
is markedly reduced. There are only three species
in Costa Rica, two or possibly three in Nicaragua,
one in Honduras, and one in Mexico. Most of
these are known only from one or very few col-
lections. The recently described E. mexicana
(Wendt et al., 1985) is the first species of Lecythi-
daceae recorded from Mexico. It escaped collec-
tion until 1983 despite being common locally.

Although it might seem more appropriate to
describe E. costaricensis in our forthcoming
monograph of the zygomorphic-flowered genera
of Lecythidaceae (Mori & Prance, manuscript),
the need to have a name available for a treatment
of the family for the Flora of Nicaragua (Prance
& Mori, in press) mandates separate publication.

Eschweilera costaricensis Mori, sp. nov. TYPE:
Costa Rica. Heredia: Tropical wet forest
along Guacimo ridge trail, La Selva Protec-
tion Zone, 275 m, 18 Jan. 1983 (fl), Hart-
shorn 2555 (holotype, NY; isotypes, BM,
CR, F, K, MO, PMA,

Ab E. pittieri venis impressis in pagina adaxiali fo-

liorum et sine lobis calycis amplificatis in fructibus
differt.

Understory tree, 5-10 m tall. Leaf-bearing
branches 3-5 mm thick. Leaf blades elliptic to
widely elliptic, 15-26 x 7-15 cm, glabrous, char-
taceous, with 10-11 pairs of adaxially impressed
lateral veins; apex long acuminate; base obtuse
to rounded; margins entire; petiole 7-8 mm long.

ANN. MissounRi Bor. GARD. 74: 455-456. 1987.

Inflorescences simple racemes or weakly once-
branched, terminal, in axils of uppermost leaves,
or from branches, the principal rachis 1.5-3 cm
long, pubescent, the pedicels ca. 7 mm long, pu-
bescen nt. Flowers 3-4 cm in diam., calyx with 6,
d Pini 6-

imbricate for '2 length; petals 6, widely ellibtie
to orbiculate, cream colored or light yellow. Hood
ofandroecium 17 x 17 mm, forming double coil
with slight beginning of triple coil; staminal ring
with 169—181 stamens; filaments ca. 1 mm long,
expanded towards apex; anthers 0.5 mm long.
Hypanthium sulcate, pubescent; ovary 2-locular,
with 6—9 ovules in each locule, ovules attached
to a hemispherically shaped placenta arising from
floor of locule; style obconical, oblique, 2 mm
long, arising from umbonate ovary summit 1
mm high. Fruits cup-shaped, 2.5 x 4 cm (ex-
cluding operculum), the calycine ring inserted
near apex of fruit base, supracalycine zone 0.5
cm wide, bearing ie ern calyx lobes and ped-
icel when mature, pericarp 2-3 mm thick, with
rough, lenticellate koi surface; operculum
umbonate, 2 cm high, o 5 mm high. Seeds
triangular in cross or. ca. 20 x 15 mm, lat-
erally arillate.

Distribution. Known only from wet forests
in the Caribbean foothills of Costa Rica and Nic-
aragua where it is an occasional (0.1—1.0/ha)
understory tree (Hartshorn, pers. comm.). Flow-
ers have been collected in January and August
and mature fruits have been gathered in January.

uidi specimens examined. NICARAGUA. RIO
o Indio, Caño Negro, 4 Dec. 1982 (im-

mature fr), Araquistain baie (M
Co dui CA. : La Selva Protection Zone,
cimo o Ridge Trail 300 m, 18 Jan. 1983 (fr,

ahue eren (NY), 25 Aug. 1979 (fl), Davidson & Don-
ahue NY).

Eschweilera costaricensis is most closely re-
lated to E. pittieri R. Knuth, which is relatively
common and distributed from the Pacific coast
of western Panama to the Magdalena Valley in
the east and to the coastal forest of northern
Ecuador in the west. Eschweilera pittieri does not

456

co-occur in the Caribbean forests of Costa Rica
and Nicaragua with E. costaricensis.

Eschweilera costaricensis differs from E. pit-
tieri by having impressed lateral veins on the
adaxial leaf surface and fruits without knobby
calycine protuberances.

I am grateful to Gary Hartshorn who first
brought this new species to my attention and to
G. T. Prance and G. Hartshorn for reviewing the

script. My research on Eschweilera, done
in collaboration with G. T. Prance, has been
sponsored by National Science Foundation grant
DEB-8020920.

LITERATURE CITED
Moni, S. A. & G. T. Prance. Eschweilera. In S. A.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Mori & G. T. Prance, Lecythidaceae — Part II.
Mani act to be submitted for publication in FI.

Neotrop.
PRANCE, G. T. & S. A. ‘Mori. 1979, Lecythidaceae—
Part I. Fl. Neotrop. Monogr. 21: 1-270

n press.) Lecythidaceae. Jn Flora

RI & G. T. PRANCE. 1985. Esch-
ida aceae): k new family
: 347-351.

; ether eld In Flora of
Panama. Ann. Missouri Bot. Gard. 45: 115-136.

— Scott A. Mori, The New York Botanical Gar-
den, Bronx, New York, 10458-5126, U.S.A.

A NEW VARIETY OF HEDYOSMUM (CHLORANTHACEAE)
FROM NICARAGUA

Hedyosmum Swartz is a genus of ca. 40 species
of primarily montane neotropical trees and shrubs
with one herbaceous species in the Far East. In
the New World species are found from southern
Mexico to Panama, Venezuela, Colombia, Ec-
uador, Peru, Bolivia, Brazil, and Paraguay, as
well as in the West Indies. The greatest density
of species occurs in the northern wet Andes. In
the field the genus is distinguished by its opposite
leaves with connate sheathing petiole bases,
stipular appendages along the distal margin of
the sheath, staminate inflorescences composed
of many ebracteate flowers of single stamens, and
pistillate inflorescences composed of solitary or
clustered, bracteate flowers.

Until a recent monograph (Todzia, 1986), the
genus has been in a state of taxonomic confusion,
in part because in the last hundred years Hed-
yosmum had never been studied over its entire
geographic range. Several species proposed from
Central American material had, in fact, already
been described from South America. Such is the
case of Hedyosmum goudotianum Solms, a
a originally described from Colombia but

ow recognized to range from Nicaragua to Peru
» bd 1986). This species is distinguished from
all other Hedyosmum species by its elongate ra-
cemose or paniculate pistillate inflorescences with
many-flowered cymules on short peduncles, no-
tably coriaceous leaves with impressed second-
ary venation and sharp, closely spaced teeth, long
leaf sheaths with striate triangular patches of ap-
pressed hairs below the stipules, and trichomes
(when present) restricted to the abaxial sides of
the primary, secondary, and sometimes tertiary

eins.
Although populations throughout its range are
variable with respect to trichome density, leaf
size, and inflorescence length, the northernmost

Rica (Burger, 1973, 1977). They are unusual in
being glabrous and in having shorter internodes
with often overlapping leaf bases, shorter leaf
sheaths that lack the striate patch beneath the

ANN. MissouRi Bor. GARD. 74: 457—459. 1987.

stipular processes, and leaves with more widely
spaced teeth

A key to the varieties is provided below with
a full description for Hedyosmum goudotianum
var. mombachanum.

Hedyosmum goudotianum Solms in A. P. de
Candolle, Prodr. 16: 482. 1869. TYPE: Co-
lombia: “‘Quindiu, El Inciendial, La pal-
milla." Nov.-May 1844 (pist.), Goudot s.n.
[lectotype (here designated), P.]. This is the
only specimen of Goudot seen from this lo-
cality. It 1s clearly designated as type ma-
terial in Solms's handwriting.

KEY TO VARIETIES OF
HEDYOSMUM GOUDOTIANUM

1. Internodes (3-)4-9 cm long; leaves with teeth

sheaths well-spaced and not overlapping, (1—)
1.8-3 cm long, with a triangular patch of ap-
pressed hairs below stipular processes; Costa
Rica, . Venezuela, Colombia, Ecua-
dor, a ia

oudotianum var. se d
Internodes l- Perd cm long; leaves with tee
—4 mm distant and with secondary vein

flush with surface, these glabrous beneath; leaf
sheaths often overlapping, (0.7—)1—1.6 cm long
without a striate patch below stipular pro-
cesses; Nicaragua .

. H. goudotianum var. mombachum

—

Hedyosmum goudotianum Solms var. goudotian-
um

ae nada (Solms) O. Kuntze, Revis. Gen.
: 566. 189

a monta ion Burger, Phytologia 26: 133.
1973. TYPE: Costa Rica. Heredia: Río Vueltas (up-

Nov. 1969 (pist.), Burger & Liesner 6336 aiam
type, F; isotypes, BM, COL, CR, GH, MO).

Distribution. Costa Rica, Panama, Venezue-
la, Colombia, Ecuador, and Peru in montane
cloud forest. In Peru this variety has been re-
corded in association with Chusquea, while in
Costa Rica it is known to occur with Podocarpus,
Weinmannia, and Clethra. In Central America
flowering is most common May through August,

458

while in South America flowering occurs spo-
radically throughout the year.

Hedyosmum goudotianum Solms var. momba-
chanum Todzia, var. nov. TYPE: Nicaragua.
Granada: “En las últimas antenas del Vol-
can Mombacho, 1,200-1,220 m, 23 Feb.
1981 (pist.), Moreno & López 7134 (holo-
type, MO; isotypes, F, HNMN not seen).

Differt a var. goudotianum internodiis 1-4(-5) cm
longis, laminis foliorum subtus glabris dentibus 2.5-4
mm distantibus, venis lateralibus subtus non elevatis,
vaginae glabrae superpositae

Dioecious, aromatic shrubs or small trees, 2—
4 m tall, with prop roots; bark whitish-gray to
gray, smooth; young stems quadrate, brittle, usu-
ally rugose, sometimes glabrous; large stems te-
rete, with tubular leaf bases persisting and be-
coming fibrous with age; internodes 1-4(-5) cm
long, the nodes slightly swollen. Leaf blades nar-
rowly elliptic, elliptic, ovate to obovate, 3.3-13.6
cm long, (1-)2.7-5.3(-7) cm broad, with acu-
minate tips 0.2-0.7 cm long, cuneate to obliquely
cuneate at base, at margins sharply serrulate with
teeth. 2.5-4 mm distant continuing to apex,
sometimes revolute, smooth, dull, light green
above and beneath when fresh, drying charta-
ceous to subcoriaceous, slightly scabrous, gray to
brown above and beneath; midveins impressed
above, raised beneath, glabrous; lateral veins 6—
8, 7-13 mm distant, arcuate, flush with surface
and glabrous beneath; intersecondary veins ex-
tending '4 to 2 distance to margin; free portions
of petioles smooth to asperous, 0.5-0.8 cm long,
narrowly winged; petiolar sheaths smooth to as-
perous, glabrous, (0.7—)1-1.6 cm long, 0.6-0.8
cm broad at apex, slightly inflated, terete or
quadrangular, with or without 2 raised longitu-
dinal ciliate lines extending down length of sheath
from stipular appendages, overlapping, persis-
tent, becoming gray and fibrous with age, not
extending beyond free portions of petioles, at
distal margin with 2 caducous, linear to slightly
fimbriate stipular appendages ca. 1 mm long. Sta-
minate inflorescences terminal or axillary, 1.5—
4 cm long, composed of 1-2 opposing pairs of
spikes on a short rachis terminated by a single
spike; bracts subtending terminal inflorescence
linear to obovate, 0.7-2 cm long,
broad; mature spikes 1.5-4.5 cm long, 0.3-0.6
cm broad, with ca. 100 stamens, sessile or borne
on short peduncles 1-2 mm long, each subtended
by a small, dentate or entire, spatulate to obovate

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

bract 0.2-1 cm long, 1-2 mm broad; stamens
congested on axis but becoming 0.5-1.5 mm dis-
tant; rachis 1-2.5 mm thick with a thick, irreg-
ularly margined, basal annulus; anthers yellow-
ish-green, 1.3-1.8 mm long, 0.6-1 mm thick;
connectives extended ca. 0.2 mm beyond thecae,
acute. Pistillate inflorescences axillary or termi-
nal, racemes or panicles (1.3-)2.5—4.4 cm long
bearing 8-13 cymules; subtending inflorescence
bracts 15-20 mm long, 3-5 mm broad; cymules
with (2—)3-4 clustered flowers, 3-4 mm long and
broad, borne on short peduncles 1-3(-5) mm
long, alternate or opposite on inflorescence axis,
2-6 mm distant; subtending floral bracts green,
connate in lower 4 to 34, 2-3 mm long including
acuminate tips 1-2 mm long, 2-4 mm broad,
ciliate or entire along free margins, enclosing '^
to *⁄4 of flower. Pistillate flowers trigonous, 2-3
mm long, 1.5-2 mm thick with a minute to large
pore on each face of the ovary; perianth lobes
deltate, acute, 0.2-0.5 mm long, basally connate;
stigmas white, 2-3 mm long, irregular in shape,
linear to irregularly lobulate, 2- or 3-angled, with
long papillae. Fruiting cymules white, irregularly
globose, 5-8 mm diam.; seeds ca. 3 mm long,
brown, trigonous, minutely papillate.

Distribution. Elfin and cloud forest on Vol-
cán Mombacho and Volcán Maderas in Nica-
ragua in association with Cavendishia, Clusia,
and Freziera at elevations of 740-1,220 m.
Flowering and fruiting occur sporadically
throughout the year.

Additional "a is examined. NICARAGUA. GRA-

ADA: Volcán mbacho, Atwood 7771 (MO, US);
upper slopes of Volcán Mombacho along W shore of
Lake Nicaragua, ca. 15 km S of Granada, 1,100-1,200
m, Croat 39092 (F, MO); Volcán Mombacho, Plan de
Flores, 740 m, Grijalva 2506 (HNMN, TEX); N slope
of Volcán Mombacho, above Finca El Progreso, Neill

somewhat above Plan del Flores, 950—1,150 m, Stevens
4332 (F, HNMN, MO, TEX). Rivas: near summit and
upper slopes of Volcán Maderas above Balgüe, Isla
ae - E 200 m, Nee & Robleto Téllez 28086
(HNM X).

I gratefully acknowledge the support ofthe Na-
tional Science Foundation (doctoral dissertation
improvement grant BSR-8400923) and The
University of Texas at Austin (graduate con-
tinuing fellowship). I thank B. B. Simpson, J.
Henrickson, and B. A. Stein for comments on
various drafts of this paper, and M. C. Johnston
for help with the Latin diagnosis. The curators

1987]

and staff of the following herbaria generously
provided loans of specimens: BM, COL, CR, F,
GH, MO, MSC, P, TEX, WIS, US.

LITERATURE CITED
BuRGER, W. C. Notes on the flora of Costa
Rica, 2. ages of the Chloranthaceae. Phy-
tologia 26: 131-

NOTES

459

1977. Chloranthaceae. Jn W. C. Burger (ed-
or), Flora Costaricensis. Fieldiana 40: 1-10
Systematics and evolution of
the genus Hedyosmum (Chloranthaceae). Unpub-
lished Ph.D. Dissertation. The University of Tex-
as at Austin, Austin, Texas.

— Carol A. Todzia, Missouri Botanical Garden,
P.O. Box 299, St. Louis, Missouri 61366, U.S.A.

BOOK REVIEW

Tomlinson, P. B. 1986. The Botany of Man-
groves. Cambridge University Press. ISBN
0-521-25567-8. Price: $69.50. (Initiating a
new series: the Cambridge Tropical Biology
Series.)

Almost any botanist who has visited the trop-
ics or subtropics will have encountered man-
groves, and this refreshingly written book is the
perfect introduction to both their taxonomy and
biology. Overall, the book is a pleasing potpourri
of selected data from many different fields about
many different aspects of mangroves. The bulk
of the book (211 pages) is a family-by-family
summary of mangrove taxa, including original
and very useful keys to the different species and
genera of most of the mangrove families. Also
included are discussions of various aspects of
morphology, architecture, reproductive biology,
and biogeography, as well as miscellaneous taxo-
nomic notes and elegant illustrations of impor-
tant species and genera. Although a key to "strict
mangrove" genera is provided in the floristic sec-
tion, no attempt is made to key out the different
back-mangrove families. Thus the reader must
first know the genus or family of the plant in
which he is interested in order to take full ad-
vantage of Tomlinson's taxonomic section.
Nevertheless, from the viewpoint of the system-
atic botanist, the gathering together in one place
of the available nomenclatural and taxonomic
data for mangrove species and genera is a high
point of the book, as are the author's personal
Observations on identification and biology of the
individual species.

Throughout, but especially in the photo cap-
tions, the author's puckish sense of humor often
comes to the fore. A Nypa stem is aptly described
as "resembling nothing so much as a series of
overlapping cowplats." The fruits of Avicennia
alba are said “to resemble a gorged leech.” An
“obliging butterfly visitor" in a photo of Lum-
nitzera racemosa is contrasted with lack of an
obliging bird visitor in L. /ittorea.

Another strong point of the book, as might be
expected considering the author's research pre-
dilection, is the series of chapters devoted to the
fascinating morphological and anatomical spe-
cializations of mangroves. No fewer than four
chapters and 71 pages treat shoot systems, root
systems, water relations and salt balance, an
seedlings and seeds. In contrast, the five chapters

ANN. MissounRi Bor. GARD. 74: 460—462. 1987.

devoted to ecology, floristics, biogeography,
flowering, and utilization and exploitation
amount to only 68 pages

I found the treatment of mangrove ecolog
disappointing. The author acknowledges this de-
ficiency and justifies it by noting that ecology is
generally outside his area of expertise. However,

tremendous amount of effort has been devoted
to this field, and a few extra pages devoted to
mangrove zonation and summarizing some of
the abundant and often conflicting literature that
implicates different toleration of salt concentra-
tions vs. soil texture and edaphic conditions vs.
rainfall would have been a welcome addition,
especially if accompanied by the author's own
trenchant evaluations.

A minor but bothersome ecological problem
is Dr. Tomlinson's adoption of the word **man-
gal" to refer to the mangrove community while

"mangrove' is reserved for the t plants
I find this to serve no useful puis and to be
distinctly cumbersome in the same way as the
plethora of Braun-Blanquet lene end-
ings. Luckily, after emphasizing “mangal” in the
first chapter, the author himself largely abandons
it in later chapters where traditional terms like
"mangroves, mangrove associates," and
"mangrove communities" are often used in-
stead. In this review I will use exclusively the
traditional terms in hopes that “mangal” (as well
as Rhizophoretum, etc.) will soon pass into well-
deserved oblivion.

Another, more significant, ecological problem
is the author's attempt to separate mangrove
species into "strict mangroves" (subdivided as
to major and minor components) and **man-
grove associates." While this should be ue a
natural way to divide mangrove taxa, in
opinion many taxa are wrongly placed in is
tables on pages 27-30, and others, even those
treated elsewhere in the book, are outright omit-
ted. This is especially obvious for the Pacific-
American mangrove taxa, and inclusion of some
of the omitted full-mangrove species would give
rise to very different conclusions. For example,
the taxonomic distinctiveness of strict mangrove
species is far more often at the level of species
than Tomlinson implies. Glaring outright omis-
sions from these lists include Tabebuia palustris
and Phryganocydia phellosperma, which are strict
mangroves, not even **back-mangroves"'; indeed

”°” 66

1987]

in the case of Bignoniaceae the back mangroves
are correctly listed but the full mangroves are
completely ignored! Crenea patentinervis, men-
tioned as a potential herbaceous mangrove in the
text, is not included in any of the tables of man-
groves or mangrove associates on the grounds
that as an herb it cannot be a true mangrove.
Actually Crenea is a subshrub and is at least as
woody as such included genera as Acrostichum,
Tuberostylis, Acanthus, or Batis. Even were it an
herb, that would be no reason to exclude it, es-
pecially as it is a full-mangrove species, not a
back-mangrove. Both species of T'uberostylis are
mangroves and they are mostly epiphytic on the
roots and lower trunks of full-mangroves rather
than on back-mangroves. Some major mangrove
components are relegated to the “minor” list,
e.g., Pelliciera which forms pure mangrove for-
ests many km? in extent in the delta of the Rio
San Juan in Colombia. Gymnosperms are spe-
cifically mentioned as playing no role in man-
groves even though Zamia roezli is typically
found in mangroves and is restricted to them and
to the adjacent coastal fringe. Muellera (p. 263)
is stated to be most commonly recorded well
away from the sea, but I have seen it only in the
back-mangrove regions with a distinct tidal in-
fluence and strongly suspect that the entire genus
is restricted to this habitat.

There are also serious problems with the num-
bers of species given for mangrove genera, es-
pecially since the data in these tables are used as
evidence of the taxonomic distinctiveness of
mangrove plants. Thus the single b

ofthis 18- -species genus, and Hippomane has four
noncoastal species as well as the well-known
coastal one. Mangrove epiphytes are implied to
be basically plants from nearby terrestrial com-
munities that transgress into the mangroves. Yet
many epiphytes seem unique to mangroves (such
as the entire genus Tuberostylis) and others are
certainly more characteristic of mangroves than
other habitats. Similarly, Tomlinson states that
there are no climbers in mangroves, presumably
because climbers have wide vessels subject to
extreme water tension, yet there is at least one
clear exception. Phryganocydia phellosperma is
not a species that roots behind the mangroves
and scrambles into them (although its unlisted
confamilial Cydista aequinoctialis is and should
probably be added to the list of mangrove as-
sociates); it is a strict mangrove and roots in (and
only in) the mangroves themselves.

BOOK REVIEW

461

The biogeographic discussion focuses so much
on the dichotomy between the relatively depau-
perate western mangroves vs. the more diverse
eastern ones that the almost equally striking dif-
ference between the richer mangrove flora of the

acific Coast of South America as compared
with the Atlantic one (possibly related to the rel-
atively recent opening of the Atlantic Ocean?) is
overlooked. Only eight true mangroves are said
to occur in the western hemisphere, ANS a
local concentration of “incipient mangroves”
western Colombia is acknowledged. While dese
species are said to be mere mangrove associates
that “lack complete fidelity to mangal” (p. 55),
several of them are true and obligate mangroves
in the strictest sense (although there are also en-
demic back-mangrove species in this region). It
may well be that the western Colombian man-
groves are fundamentally different from other
mangroves in their greater habit diversity and in
being mostly individual mangrove species of
otherwise nonmangrove genera, but they are not
less true mangroves for that. Knowledge of man-
groves would be better served by focusing on the
unusual aspects of these species rather than by
trying to sweep them under the rug.

There are a number of inconsistencies, espe-
cially in the biogeographic discussions. Rhizoph-
ora racemosa is on both Atlantic and Pacific sides
of tropical America as reported on p. 334, but
in the key (p. 329) it is characterized as being
only on the Atlantic coast of South America.
There are five Pacific coast collections in the MO
herbarium, as well as collections from Honduras,
Costa Rica, and Panama, all outside the Vene-
zuela and Guianas to West Africa range indicated
by Tomlinson (p. 335). Rhizophora harrisonii is
rather precisely mapped as having a disjunct
population in the middle of the Peruvian coastal
desert outside the range of any mangrove, but
the numerous records from coastal Ecuador and
Colombia were apparently overlooked. On the
other hand, the range of R. samoensis is hypo-
thetically extended to include the Pacific coast
of Sout merica, where it may occur but has
not yet been documented; that species does reach
the Galapagos, according to R. Horna (pers.
comm.).

Inevitably a few insignificant errors in spelling
of Latin binomials, taxonomic authorities, etc.
are unavoidable in a book of this scope. Exam
du include *Anaemopegma" (p. 32), " Mouri-

` and **Pachyra" (p. 56) and the authors of
AUR species of Phryganocydia (p. 214). More

462

problematic is Lysianthus (presumably = Lys-
iana?) supposed to be a mangrove mistletoe (p.
33). Dalbergia amerimnion Benth. (p. 261) has
long been regarded as a synonym of D. brownei.
Phryganocydia phellosperma is not distinguished
from P. corymbosa by a simple tendril, a trait
shared with the entire genus. Tabebuia palustris
is not deciduous as stated (p. 214), which would
have been quite remarkable in a mangrove, but
oi like all other mangroves known to me.
milar minor errors in biogeographic distri-
butions include Tuberostylis rhizophorae, whose
aimed “wider distribution in Central America”
-consists of a single collection from southernmost
Darién, and Pavonia rhizophorae, supposed to
be recorded only for Colombia but reported in
the Flora of Panama to cross the Panama border
in the same part of southern Darién.
Obviously, it is difficult to eliminate such mis-
cellaneous errors. Whether they are as frequent

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

in other taxa and regions as among the man-
groves I happen to know personally (i.e., mostly
Bignoniaceae and Pacific American), I do not
know. But it seems likely that it would have been
useful for the author to run a draft of his manu-
script past a few taxonomic specialists or field
botanists as well as checking a few more herbaria
for distributional data. Such minor imperfec-
tions detract very little from the message of the
book, except that inasmuch as this is the closest
thing to a monograph of many mangrove taxa
that we are likely to have in the foreseeable fu-
ture, it is a bit of a shame that some of the work
that would have gone into an actual monograph
was neglected. Overall The Botany of Mangroves
succeeds admirably in its purpose of providing
a concise and highly readable introduction to the
world’s mangroves.— 4/wyn Gentry, Missouri
Botanical Garden, P.O. Box 299, St. Louis, Mis-
souri 63166, U.S.A.

Volume 74, No. 1, pp. 1-182 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN was published on 15 June
987

A 7 ri F
P $ |
FE j
i DUNS. š . š ç
| F tí S" r m A ri
ETE " d t £ ;

I x t. Oligocarpae |
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I

Charlotte M. eH 2223

CONTENTS

> Bada

S * New Species of Thrasya TL Panicoidaea) bo diy Mow of aise and 52
g avidse & A

Cerri t se Adni ir G. Burman dus 63A 43M

um Contents c continued on nec

Vascular Epiphytism: Taxonomic Participation and kes Diversity David H. Ben-
zing 183
Diets and Ships of Neotropical Vascular Epiphytes Alwyn H. Gentry & C.
H. Dod. 205
Nus Fixation by Epiphylls in a Tropical Rainforest Barbara L. Bentley -aana 234
Adaptive Radiation of Salamanders in Middle American Cloud Forests David B. Wake 242
Revision of Eremanthus (Compositae: Vernonieae) Nanda F. F. Macebeish 2 265
Biosystematics of Tetraploid Eucharis (Amaryllidaceae) Alan W. Meerow 2 s 291
The Shrubby Gentian Genus Macrocarpaea in Panama Kenneth J. Systm „= 310
. Menoecy and Sex Changes in Freycinetia (Pandaceae) Hans-Helmut Poppendieck .. 314
A Guide to Collecting Lecythidaceae Scott A. Mori & Ghillean T. Prance .cc:2cccco 321
Chromosome Cytology of Aq (Asteraceae- Mutisieae) Peter Goldblatt ......... 331
` Notes on Cipura (Irid hand Central America, and a New Species from Venezuela
Peter Goldblatt & Da E. Henrich . 333
n ‘tenis (Lepidoptera: Nymphalidae): Summary of Known Larval Food Plants Bone
A. Peu IE Keith S Brown dro V ens 341
. Chemistry at the Solanaceae/Ithomiinae Interface Keith S. Brown, Jr. 399
: 5 New Taxa of Rubiaceae from Venezuela - Julian A. Steyermark PR euer cic . 398
E New Species of Neot pical | Lauraceae Henk van der Werff... ipii, VOR
* A New Species of Ocotea shee from Southeastern Mexico Tom We nš må Henk
van der Verf... 2 Mic EL gs Pe E OSEE
Four New: Species o of Axonopus (Poterie: Panicese from Tropical America Gerrit ;
! se T
| Two Mean ERS : : quea (Poaceae: Bambusoideae) Lynn G. Clark 424
New Caitlin in TRE ea blue Lynn G. Clark ——— M8
Me Ne Cor nus cad in South America Julian A. Steyermark & Ronald
ilia | FRESI sian Else Zardini .— u aN Em ;
Chromosome Counts of Missouri Asteraceae and Poaceae — Ahsan A. Vahidy, ! Gerrit LO TS
— Davidse & Youji . Shige

— EE d e W

Volume 74 \ Z
N 7A
N u mber 3 ! Ac N ae i P rd

Volume 74, Number 3
Fall 1987

Annals of the
Missouri Botanical Garden

The Annals, published quarterly, contains papers, primarily in systematic botany, con-
tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the
Garden will also be accepted. Authors should write the Editor for information concerning
arrangements for publishing in the ANNALS. Instructions to Authors are printed on the
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Missouri Botanical Garden

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Missouri Botanical Garden &
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- Missouri Botanical Garden — s

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Volume 74
Number 3
1987

Annals
of the
Missouri
Botanical

Garden

NZ

A REVISION OF PANICUM SUBGENUS PANICUM
SECTION RUDGEANA (POACEAE: PANICEAE)!

FERNANDO O. ZULOAGA?

ABSTRACT

Panicum subg. Panicum sect. Rudgeana, herein revised, includes six species: P. cayennense, P.
campestre, P. cervicatum, P. ligulare, P. rudgei and P. vinaceum. It is characterized mainly ideis n

upper anthecium. The stipe consists of two portions: a membranous portion towards the ve
rtion towards its dorsal face. The position of the section within

of the spikelet and an indurate po

ntral fac

subg.
Panicum is discussed, as is the relationship of Rudgeana with other sections containing species Rin

a stipitate upper anthecium.

Hitchcock & Chase (1910) included P. rudgei
Roemer & Schultes and P. rotundum A. Hitchc.
& Chase within the ungrouped species of the
genus and noted their close relationship. In 1915
they repeated this treatment, indicating also that
the species they had described was the same as
P. campestre Nees ex Trinius. The same year,
Hitchcock also placed P. rudgei in an informal
group he named Rudgeana. Chase, in unpub-
lished manuscripts, later placed both species in
the Rudgeana group, which she characterized as
“Rather stout much branched perennials with
usually harshly pilose or papillose sheaths.
Spikelets abruptly pointed, the first glume point-
ed, more than half the length of the spikelet, the
midnerve scabrous.”

The two species mentioned above plus P. cay-
ennense Lam., P. ligulare Nees ex Trin., P. vi-
naceum Sw., and P. cervicatum Chase share char-
acters that allow them to be included in sect.
Rudgeana (A. Hitchc.) Zuloaga.

Section Rudgeana falls within subg. Panicum,
having the following characters in common with
the rest of the sections in the subgenus [which
are sects. Panicum, Repentia Stapf, Urvilleana
(A. Hitchc. & Chase) Pilger, and Dichotomiflora
(A. Hitchc. & Chase) Honda].

Species of subg. Panicum are characterized by
the presence of the C, photosynthetic pathway
of the NAD-me subtype (Brown, 1977) and are
distinguished anatomically by having a double
sheath around the vascular bundles. The inner

! I wish to thank the Consejo Nacional de Investigaciones Científicas y Técnicas de la Repüblica Argentina

(CONICET) for a grant that allowed me to spend 1982
doctoral fellow. I owe appreciation to Cecilia Ezcurra and Em
of the manuscript. The line drawings were made by Vladimiro ‘Duda as,

and 1983 at the Smithsonian Institution as a post-
met J apt sva for help during the preparation
am always grateful

2 Instituto de Botanica Darwinion, Casilla de Correo 22, San Isidro, peciit

ANN. Missouni Bor. GARD. 74: 463-478. 1987.

464

one is a mestome sheath with thick-walled cells.
It is surrounded by a Kranz outer sheath con-
taining specialized chloroplasts that are usually
disposed centripetally. Between each vascular
bundle there are two or three tabular cells ar-
ranged radially. The number of secondary vas-
cular bundles present between each primary bun-
dle varies from two to six.

In sect. Rudgeana, as in most sections of subg.
Panicum, the plants are cespitose and short rhi-
zomatous with erect, few- to many-noded culms.
The ligule is membranous at the base and short-
to long-ciliate at the upper portion. The leaf blades
are lanceolate to linear-lanceolate, with or with-
out involute borders. The species are usually
found in dry and open places, but some species
in sect. Dichotomiflora and in sect. Repentia grow
in wet places and have decumbent culms that
root at the lower nodes.

The inflorescences are pyramidal, lax and dif-
fuse, and have ellipsoid to lanceolate spikelets
dispersed on the branches.

The nervation of the glumes and lemmas and
ornamentation of the upper anthecium are dis-
tinctive characters that hold together the sections
of the subgenus. The upper glume and lower lem-
ma are 7- to 9-nerved (1 1- to 15-nerved in species
of sects. Rudgeana, Panicum and Urvilleana),
with a few exceptions in species of sects. Dichoto-
miflora and Repentia, in which these bracts are
5-nerved. The upper anthecium is smooth and
shiny over the entire surface, and compound or
both compound and simple papillae are present
near the apex of the upper palea.

Panicum sect. Rudgeana differs from sect. Di-
chotomiflora by the length of the lower glume ('⁄4
to 3 the length of the spikelet in sect. Dichotomi-
flora) and by the absence of papillae on both
surfaces of the leaf epidermis; also, as noted be-
fore, species of sect. Dichotomiflora grow in hu-
mid places with the culms decumbent and root-
ing at the lower nodes. Section Rudgeana is
separated from sect. Repentia by the absence of
stout rootstocks at the base of the plant. Section
Urvilleana is distinguished from sect. Rudgeana
by having long macrohairs at the base of the
upper lemma and numerous, whitish oa cov-
ering both glumes and the lower lem

Section Rudgeana can be aoaaa clearly
from sect. Panicum and the sections mentioned
above by the occurrence of a well-developed stipe
at the base of the upper anthecium. Two seg-
ments of the stipe can be distinguished: a) a por-
tion of membranous tissue towards the palea of

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

the upper anthecium (Figs. la, e, 2d, e, 3b) and

) a portion of indurate, smooth, and shiny tissue
towards the lemma of the upper anthecium (Figs.
I b, f, 2a, 3c-e). The texture of the indurate por-
tion is similar to that of the main portion of the
upper anthecium. The membranous portion of
the stipe is reduced in P. cayennense and P. cam-
pestre (Figs. le, 4g, h) or is larger and more ex-
panded in P. rudgei (Fig. 3b) or remarkably no-
ticeable in the rest of the species of the section.
In P. ligulare and P. vinaceum the membranous
portion of the stipe is prominent and may be
prolonged into one (Fig. 5h, i) or two wings (Fig.
2e) that cover the base of the upper anthecium.
In all cases the membranous portion appears tur-
gid when the spikelet is rehydrated, and it is free
from the base of the upper anthecium.

The indurate portion of the stipe is found be-
low the upper anthecium and is appressed to the
membranous portion, at least when the spikelet
is immature (Figs. la, b, 2a, 3d, e). At maturity,
it extends behind the upper anthecium as a mu-
cro (Fig. 1f). Size and length of this mucro vary
among species of the section, but it usually re-
mains on the rachilla when the upper anthecium
falls (Fig. 3c).

In Australia there exists a group of Panicum
species with a structure similar to the stipe found
in sect. Rudgeana. These species were trans-
ferred from Ichnanthus Beauv. to Panicum by
Lazarides (1959), who noted that the appendages
found at the base of the upper anthecium are not
adnate to the upper lemma (as in Ichnanthus)
but rather originate from the apex of a noticeable
stipe. Shaw & Webster (1983) supported this
concept, emphasizing distinctness of the ap-
pendages in /chnanthus from Australian species
of Panicum.

More recently, Lazarides & Webster (1984)
removed these “ichnanthoid” species from Pan-
icum, erecting for them the new genus Yakirra.
Included in it were four species previously treat-
ed in Panicum: Y. muelleri (Hughes) Lazarides
& Webster, Y. majuscula (F. Muell. ex Benth.)
Lazarides & Webster, Y. australiensis (Domin)
Lazarides & Webster, and Y. pauciflora (R. Br.)
Lazarides & Webster; also included was a new
species, Y. nulla.

They provided a table of features separating
Yakirra from Ichnanthus and Panicum and stat-
ed that there were no conclusive characters to
differentiate Yakirra from Panicum besides the
presence of a stipe at the base of the upper an-
thecium. I regard this as correct, since the other

ZULOAGA— PANICUM SUBG. PANICUM SECT. RUDGEANA

GURE 1. Scanning electron mt of the upper anthecium of Panicum species. a-d. P. ligulare.—a.
Lateral view of the base showing the stipe.—b. Dorsal view of the base showing the indurate portion of the
stipe.—c. Apex of the upper anthecium i diit: papillae at the tip of the palea.—d. Detail of the papillae. e, f.
P. campestre. —e. Ventral view of the base of the upper anthecium showing the membranous portion of the
stipe.—f. Dorsal view showing indurate portion of stipe. a-d, based on Irwin 1 4904; e, f, based on Chase 8645.
Scale bars: a—c, f, x 100; d, x 500; e, x150.

ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FIGURE 2. Scanning electron micrographs of the upper anthecium of Panicum species. a—c. P. cervicatum. —
mbranous and indurate portion of the stipe. — b. Apex showing compound papillae
i f. P. v

pound papillae at the tip of the palea. a—c, based on C ase 7 0737; d-f, based on Steyermark 59173. Scale bars:
a, d, x 50; b, x 100; c, x 500; e, x 70; f, x 300.

468 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

FIGURE 4. Panicum campestre. —a. Habit.—b. Spikelet, lateral view.—c. Spikelet, ventral view.—d. Spikelet,
dorsal view. —e. Lower palea, dorsal view. —f. Lower palea, ventral view.—g. Upper anthecium, dorsal view. —
h. Upper anthecium, ventral view.—i. Caryopsis, embryo side.—j. Caryopsis, hilum side. Based on Sendulsky
637.

1987] ZULOAGA — PANICUM SUBG. PANICUM SECT. RUDGEANA 469

FicunE 5. Panicum ligulare.—a. Habit. —b. Spikelet, lateral view.—c. Spikelet, ventral view.— d. Spikelet,
dorsal view.—e. Lower palea, ventral view.—f. Lower palea, dorsal view.—g. Upper anthecium, dorsal view.—
h. Upper anthecium, ventral view.—i. Upper anthecium, lateral view. Based on Irwin 14904.

470 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE 1.

Comparison of sections of Panicum with stipitate upper anthecia.

[VoL. 74

Subg. Phanopyrum

Sect. Sect. Sect.
Characters Stolonifera Parvifolia Phanopyrum
Stipe a Homogeneous Homogeneous ' Homogeneous
Stipe pre In all species In a few species In the only species
Ended: pathway š C,
Compound papillae at tip of palea Absent Absent sent
Inflorescence type Racemose Paniculate Racemose branches
branches branches

Upper glume and lower lemma

3- to 5-nerved

3- to 5-nerved

3- to 5-nerved

nervation

Panicle dimorphism Absent

Absent Absent

characters listed for Yakirra (habit, life form,

ret, and photosynthetic path-
way) are common features of different sections
of subg. Panicum.

Lazarides & Webster (1984) noted the pres-
ence ofa stipitate flower in the American P. gym-
nocarpon Ell. On this basis they accepted this
species as correctly placed by some authors in
the monotypic genus Phanopyrum (Raf.) Nash
and asserted that “The acceptance of Phanopy-
rum as a valid genus makes Yakirra morpho-
logically distinct from Panicum, based on the
presence or absence of a stipitate flower.”

However, stipitate upper florets are also pres-
ent in species of sects. Lorea Zuloaga, Agrostoi-
dea (A. Hitchc. & Chase) Hsu, Stolonifera (A.
Hitchc. & Chase) Pilger, Dichanthelium, and
Parvifolia (A. Hitchc. & Chase) Pilger. wean
ters distinguishing these taxa are summarize
Table 1

Section Rudgeana is similar to Yakirra in de-
tails of habit leaf blades, ligules, inflorescences,
spikelet compression and length, form and ner-
vation of glumes and lower lemma. It differs from
the Australian genus mainly by having a heter-
ogeneous stipe below the upper anthecium, a
lower palea (almost absent in Yakirra) well-de-
veloped, and the upper anthecium with com-
pound papillae at the tip of the palea only. In
species of Yakirra the anthecium has simple pa-
pillae in longitudinal rows all over the lemma

Therefore, th fal
in sect. Rudgeana is a good character for its de-
limitation within subg. Panicum, but I judge it

to be an insufficient one for removing species
from Panicum.

The elongation of the rachilla could help in
opening the spikelet and posterior dispersal of
the caryopsis. Davidse (in press) has pointed out
that the stipe below the upper anthecium in P.
cervicatum is an elaiosome involved in ant dis-
persal of the diaspore and noted that a similar
elaiosome might be present in P. vinaceum and
P. trinii. Berg (1985) reported a similar elaio-
some in the stipe of Panicum australiense Do-
min.

METHODS AND MATERIALS

Classical taxonomic studies have been carried
out in this paper, utilizing a Wild M5 dissecting
microscope and a Wild M20 microscope. For

LE, M, MO, NY, P, R, RB, S, SI, SP, and US.

SYSTEMATIC TREATMENT

Panicum subg. Panicum sect. Rudgeana (Hitch-
up R n

1915. TYPE: Panicum rudgei Roemer &
Schultes.

Cespitose perennials or occasionally annuals,
with erect, more or less branched culms and usu-
ally pilose leaves. Ligule membranous, short- to
long-ciliate. Leaf blades lanceolate to linear-lan-

1987] ZULOAGA — PANICUM SUBG. PANICUM SECT. RUDGEANA
TaBLE l. Continued.
Subg. Subg. Subg.
Phanopyrum Dichanthelium Agrostoides Subg. Panicum Genus Yakirra
Lorea Dichanthelium Agrostoidea Rudgeana

Homogeneous Homogeneous Homogeneous Heterogeneous Homogeneous

In a few species In a few species In a few species In all species In all species

° C, C4, NADP-me 4 NAD-me C, NAD-me
Absent Absent Present Present
Paniculate Paniculate Paniculate Paniculate branches

Absent

Paniculate
branche branche

3- to 5-nerved 7- to 9-nerved

Absent

Absent Present

3- to 5-nerved

7- to 11-nerved 7- to 9-nerved

Absent Absent

ceolate, flat. Inflorescence a single, terminal and
lax panicle or a terminal and several axillary pan-
T ple an elongated, compound arrange-
pedicels long, flexuous. Spikelets obovoid
to van falling from the pedicels, pilose with
long, rigid hairs to glabrous, pale to nearly pur-
plish; glumes and lower lemma with 5-9(-11)
prominent nerves, gaping and exposing the fertile
floret at maturity. G/umes unequal, the lower
glume '^ to 3⁄4 as long as the spikelet; upper glume
and lower lemma a little longer than the anthe-
cium, pointed at the apex. Lower palea conspic-
uous, membranous, with or without a male flow-
er. Upper anthecium stipitate, ovoid, glabrous,
smooth and shiny, indurate; stipe membranous
ventrally, indurate dorsally; palea with co
pound papillae at the apex (papillae occasionally
absent in P. cayennense). Stamens 3; stigmas 2,
plumose and purple; /odicules 2, membranous,
glabrous. Caryopsis with the hilum punctiform.
Embryo less than half the length of the caryopsis.

Species of sect. Rudgeana grow in open and
sunny places, and are common in savannas of
Central and South America and in the cerrado
of Brazil; they are frequently found in sandy soils
from sea level to ca. 1,500 m

KEY TO THE SPECIES OF SECTION RUDGEANA

. Panicles terminal, lax; axillary m usu-

ally absent; spikelets ` ate i ong, 1.2-2.5

mm wide; stipe prominen mm or longer.

2a. Stipe glabrous pam or sheaths pa-
pillose-pilose, with glassy hairs; E

8-3.2 mm long; spikelets 4. w

long Plu

2b. Stip pil t ll J? l fst th e

—
£

to grins but without es pe pini
m long; spikelets m lon
3a. Spikelets 7-9 mm long, 2. 122. 5 nn
m upper anthecium 4-4.5 m
P. Bos P
3b. Spikelets 5.9-6.7 mm long, 1.5-
m m upper anthecium 3-3.5
eee e 6. P. vinaceum
lb. Panicles terminal and axillary, Pormang x

3.5 mm long, 1-1.3 mm wide; stipe 0.5 mm

or shorter.

4a. Viri 3-3.5 mm long; leaf sheaths
ually with glassy hairs 5. P. rudgei

4b. Spikelets 2.1-2.8 mm long; leaf sheaths

hairs.

5a. Plants perennial; spikelets ig
sparsely pilose, 2.6-2.8 m g uo

us

5b. Plants annual; spikelets d
glabrous, 2.1-2.6 mm lon

. P. cayennense

]. Panicum campestre Nees ex Trin., Gram. Pan.
197. 1826. TYPE: “V. sp. Brasil (N. ab
Esenb.)." Not seen.

P. ise A. Hitchc. & Chase, Contr. U.S. Natl.

. 15: 139. 1910. TYPE: Brazil. Minas Gerais:

TH Widgren s.n. (holotype, US; fragment of
holotype, BAA).

Perennial, 30-74 cm tall, usually with thick
adventitious roots. Cu/ms erect or geniculate at
the base, rooting or not at the lower nodes, many-
branched; internodes cylindrical or compressed,
3-10 cm long, hirsute with appressed, rigid hairs
to glabrescent; nodes densely pilose, with long
and whitish hairs. Leaf sheaths 3—7 cm long, usu-
ally shorter than the internodes, the lower ones
longer, hirsute with long, tuberculate hairs; mar-

472

gins ciliate. Ligule 1.5-2.5 mm long, with hairs
on the back towards the base of the blade; collar
pilose, stramineous to brownish. Leaf blades lin-
ear-lanceolate, 7-25 cm long, 0.5-0.7 cm wide,
flat or with involute borders, acuminate apically,
rounded or subcordate basally, densely hirsute
on both surfaces, with scabrous and ciliate mar-
gins, the midnerve manifest. Panicles terminal
and axillary, forming an oblong, compound in-
florescence, sometimes the terminal panicle dis-
tant from the other ones, lax and diffuse; axis
sparsely hirsute, at least in the lower portion,
longitudinally ridged, flexuous, scabrous, the
branches alternate, divaricate, flexuous and sca-
brous, the axils of the branches pilose; pedicels
long, scabrous. Spikelets ovoid, 2.6-2.8 mm long,

2 wide, sparsely pilose, pale to purple
toward the apex to completely purplish. Lower
glume 1.8-2.2 mm long, '^—À as long as the
spikelet, acuminate to subulate, with rigid hairs
toward the apex, 5-9-nerved, the midnerve sca-
brous. Upper glume and lower lemma subequal,
acuminate, 2.6-2.7 mm long, 7—-9-nerved,
sparsely pilose on the inner surface and with or
without long and sparse hairs on the outer sur-
face. Lower palea elliptic, 1.8-2 mm long, pres-
ent or absent. Upper anthecium broadly ellip-
soid, 1.8-2 mm long, 1 mm wide, pale; lemma
7-nerved; palea with compound papillae at the
apex; stipe with the membranous and indurate
portion ca. 0.2 mm long. Caryopsis 1.2-1.3 mm
long, 1 mm wide. In flower December-May. Fig-
ure 4

Distribution. Brazil, from Para and Bahia to
Paraná; 0—1,500 m; growing in sandy or red clay
soils in campos or cerrados.

Selected specimens examined. BRAZIL. BAHIA: Ser-
ra Geral de Caitité, 9.5 km S of Caitité on road to
Brejinhos da Ametistas, Harley 21319 (CEPEC).
DISTRITO FEDERAL: Universidad de Brasilia, Clayton
4795 (NY, US), 4842 (MO, NY, US); E of Lagoa Par-
anoa, Irwin et al. 11181 (F, GH, NY, US); 15 km S of
Brasilia, Irwin & Soderstrom 5700 (US); Sobradinho,
Clayton 4875 (NY). Goias: 6-7 km E of Alto Paraíso,
Anderson 6515 (MO, NY); 26 km NE of Catalào, Irwin
et al. 25210 (F, MO, NY, US); 75 km N of Corumbá
de Goiás, Irwin et al. 19000 (F, GH, MO, NY, US);
14 km S of le Irwin et al. 34386, 34387 (F,
NY, US); 16 km N of São Joao da Aliança, Dawson
14442 (US); between Viannápolis and Ponta Funda,
Chase 11315 (US); Serra do Rio Preto, 14 km E of
Cabeceiras, Irwin et al. 10354 (US); Corumbá, Macedo
4482 (BAA, US). MATO GROSSO: | km NE of Garapé,
1 Oct. “es "Flee be ee s.n. (US-2642542).

MATO G rande, Chase 10790
(GH, RB. "US. poi 9598 (US). MINAS GERAIS: Bar-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

bacena, Serra Mantiqueira, Chase 8645 (F, GH, NY,
RB, US); Corinto, Fazenda Diamante, Mexia 5567 (F,
: 5 11 km N of Gouveia, Anderson et
ine Faria, Serra da Bocaina, Chase
m W of Barao de oo Irwin et
(F, MO, S US); 7 km W of Campanha,
Davidse et al. 10651 (MO, NY); 33 km NE of Francisco
Sá, Irwin et al. 23071 (F, MO, NY, SP); el km SW of
Gouveia, Anderson et al. 35148 (F, MO, NY); 9 km
NE of Estiva, Davidse et al. 10544 (MO); Ouro Preto,
Chase 9354 (F, NY, US); 2 km S of Itacolumy, Irwin
et al. 29364 (F, MO); Pocos de Caldas, Chase 10637
(US); Piloes, Macedo 4876 (NY, US); lower slopes of

Chase 8856 bus pire Chase 8815 (US), Maia 18
(RB); Jardim, Widgren 908 (US); Lagoa Santa, Chase
$995 (US); Hermilo Alves, Duarte 6352 (US). PARA:
Marajó ate Natal, Schwacke 62 (R); Fazenda d
js Rio Aurá, Black 54-16109 (R). PARANA: Jagua
iahyva, Dusén 16393 (F, GH, NY, US), 10074 (US),
Swallen 8678 (US); 2 km W of Rio Itararé and road
PR-11, Davidseet al. 11375 (MO, NY). RIO DEJANEIRO:
Monte nin Serra da poss = hase 8358 (F, GH,
MO, NY, US); Resende, Hoe
US). SAO PAULO: ps N of recon C

NY, SP, US); 3 km from Cajurü, Sendulsky 126 (SP,
US); 16 km NNE of Padua Sales, Eiten 1669 (NY).

Although it was not possible to examine the
type specimen of this species, P. campestre is
clearly differentiated by the diagnosis given by
Trinius and by the illustration of this same au-
thor (1829).

Nees (1829) published tł
same name, the type specimen being completely
different from the species described by Trinius.
Panicum campestre Nees of 1829 was validly
renamed as P. peladoense by Henrard (1940).

pecies with the

2. Panicum cayennense Lamarck, Tabl. Encycl.
1: 173. 1791. TYPE: “Cayenne, D. Stoupy”
(holotype, P, not seen; fragment of holotype,
BAA, US-81397)

P. sessilicaule Desv., Opusc. 95. 1831. TYPE: “Habitat
in Carolina” (holotype, P; fragment of the holo-
A

type, BAA).
P. floribundum Rich. ex Lam., Encycl. 4: 742. 1798,
pr . P. ca nse.
P. pedunculare Willd. ex Steudel, Syn. P1. Glum. 1: 77.
YPE: “P. cayennense ie Agr. Bras. 195.
Brasil” (fragment of the syntype: “America me-
ridionale, from Humboldt,” US. 2907507).
P. g var. curtatum Doell, in C. Martius, Fl.
s. 2(2): 220. 1877. TvPE: "extra fines in via
imer Cayenne et Baduel (Yelski, inter plantas a
tafinski D mecum communicatas)"
ues US-80517

1987]

Annual, to 110 cm tall. Culms erect or spread-
ing, usually branching at the lower and middle
nodes, often zigzag, few-noded; internodes hol-
low, compressed, hispid to glabrous; nodes dark,
covered with whitish hairs. Leaf sheaths 2-8 cm
long, shorter or longer than the internodes, pi-
lose, with thick, tuberculate hairs; margins cil-
iate. Ligule 0.8—1.6 mm long; collar pilose, pale.
Leaf blades linear-lanceolate, 5-28 cm long, 0.4—
| cm wide, flat, acuminate apically, rounded at
the narrowed base, hispid on both surfaces to
glabrescent, the margins scabrous and ciliate, the
midnerve manifest. Panicles several, terminal and
from the upper leaf axils, forming an elongated
compound inflorescence 5-32 cm long, 3-12 cm
wide, reaching 73 to almost the entire height of
the plant, each panicle included at the base; axis
longitudinally ridged, flexuous, scabrous and his-
pid towards the base, the branches divaricate,
alternate to opposite, sometimes pseudoverticil-
late, scabrous and flexuous, the axils of the
branches pilose to glabrous; pedicels long, flex-
uous and scabrous. Spikelets obovoid, 2.1-2.6
mm long, 1.1-1.3 mm wide, glabrous, ¿lobos
pale to purplish. Lower glume 1.2-1.8 mm long,
more than half the length of the spikelet, acu-
minate apically, pilose on the inner surface,
5-nerved, the midnerve scabrous toward the apex.
Upper glume 2.2-2.5 mm long, acute apically,
7-nerved, pilose to glabrous on the inner surface.
Lower lemma 2-2.4 mm long, 7-nerved. Lower
palea elliptic, 1.6-1.9 mm long, 0.6-1.1 mm wide,
membranous, glabrous; male flower absent. Up-
per anthecium broadly ovoid, 1.5-1.8 mm long,
0.9-1.2 mm wide, pale; stipe less than 0.3 mm
long, the indurate portion prolonged beyond the
upper anthecium as a mucro. Caryopsis broadly
ovoid, 0.9 mm long, 0.7 mm wide, pale. In flower
all year.

Distribution. Mesoamerica, West Indies
(Cuba, Jamaica and Dominican Republic), and
South America, from Venezuela to Bolivia; 0—
1,500 m; occurring in savannas, in sandy or clay
soils.

Chromosome number. n =
Pohl, 1974).

27 (Davidse &

Selected Vae aee examined. MEXICO. CHIAPAS:

IZABAL: S of Río Dulce, at Shell Station, LeDoux et at

ZULOAGA — PANICUM SUBG.

PANICUM SECT. RUDGEANA 473

6 (NY). PETEN: Santa Rita, 20 km al S de Santa Elena,

5 (F,

ALAJUELA:
601 (F, US), 605 (F). PANAMA. CHIRIQ
Boquete, McDaniel 6807 (MO); vicinity of David,
Hitchcock 8372 (F, MO, NY, US). P A: Near Ar-
raijan, Woodson, Jr. 1402 (MO, NY. US) pia ISLA
DE PINOS: Near Nueva Gerona, Curtiss 267 (MO, US);
vicinity of San Pedro, Britton et al. 14455 (MO, US);
Isla de Pinos, Taylor 34 (MO, US). oRIENTE: Cayo del
Rey, Ekman 10028 (US). PINAR DEL RIO: Herradura,
Tracy 9073 (US), 9093 (MO), Britton et al. 6520 (US);
Sierra de Cabra on Guane road, Britton et al. 7275
(US); Laguna Jovero and vicinity, Shafer 10510 (US).
JAMAICA. Halliss Savanna, Upper Clarendon, Harris
12226 (MO). DOMINICAN REPUBLIC. DISTRITO NA-
CIONAL: Sierra Prieta, Villa Mella, Liogier 17408 (US).
LA VEGA: Vicinity of Pedra Blanca, Allard 16060, 16067
(US); Cordillera Central, Sabana de la Mar, El Valle,
Ekman 15694 (US

Simba, Fernández 11 (MO). Without department and
locality, Mutis 5359, 5378, 5498, 6110 (US).
VENEZUELA. AMAZONAS: Puerto Ayacucho, Williams
13085 (F, US); near Capuana, Davidse & Huber 16811
(MO); 23 km NE of Puerto Ayacucho, Davidse & Huber

Davidse et al. 4548 (MO). FRENCH GUIANA. Cayenne,
Pu hie (MO- ES US- epa 54 ES route
de R mbeau, Hoock s. . SURIN n distr
Pará, yon 1495 (MO). Cmn . CRUZ: Buena
Vista, Steinbach 6935 (BAA, F, GH, LIL, MO, NY
US). BRAZIL. AMAPA: Rio Pedra Fróes & Black 17 322
(US); Macapá, Fazendinha, Black & Lobato 50-9659
uu: gente s: 2 km S of HO Prance et al. 8177
(F,G , NY); km 27 of road Humaitá-Porto Vel-
p: Prance al. 3517 (MO). BAHIA: Col. Valença, Pinto

US). coras: 2 km SW of Araguiana, Eiten 10154

US MATO GROSSO: 20 km S of Garapü, Irwin & Soder-
strom 6485 (US). MATO GROSSO DO SUL: 100 km W of
Coxim, Bommer 54 (NY, US); Paiaguás, Fazenda Al-
vorada, Allem & m 1001 (MO); Xavantina-Cach-
imbo road, W of k

3674 te dicia depu a

2- 4 km Eof Mutumparaná, Prance et al. 8831 (F, MO).

Panicu m differs from P. campestre
mainly by its smaller, glabrous, and obovoi
spikelets. It also differs in its annual habit; in P.
cayennense the culms are generally short,
branched, and bear numerous panicles nearly

474

from the base, the axillary ones aggregating with
the apical ones. Nevertheless, there are some
specimens with elongated culms in which the
terminal panicles are somewhat separated from
the axillary ones.

This species was included by A. Hitchcock &
Chase (1915) in the Capillaria group, along with
P. miliaceum, P. capillare, and others, but the
presence ofthe characteristic stipe of sect. Rudge-
ana clearly separates it from these species.

3. Panicum cervicatum Chase, J. Wash. Acad.
Sci. 32: 164, f. 10, 1942. TYPE: Brazil. Mato
Grosso do Sul: Tres Lagoas, 4 Feb. 1930, A.
Chase 10737 (holotype, US-1500814; iso-
types, RB, US-1816795)

Perennial, 40-100 cm tall. Culms erect, simple
or occasionally branched; internodes 7-23 cm
long, terete, glabrous to sparsely pubescent just
below the nodes; nodes densely pilose to gla-
brous. Leaf sheaths 7-13 cm long, the lower ones
overlapping, pale, densely hirsute to glabrous;

margins ciliate. Ligu/e 1.5-2 mm long; collar dark
brown, short- to long-pilose. Leaf blades lanceo-
late, sf, 16-36 cm long, 0. 8-1. 6 cm wide, long-
y, flat or the
margins involute i in n drying, hispid or strigose to
glabrous on both surfaces, the margins scabrous
and largely ciliate with papillose-pilose hairs
(these hairs caducous), the midnerve prominent.
Panicles lax, diffuse, many-flowered, 25-60 cm
long, 12-35 cm wide, the spikelets in pairs; axis
longitudinally ridged and scabrous, the branches
alternate or opposite, scabrous, the axils of the
branches pilose and pale; axillary panicles usu-
ally absent, when present similar in shape and
smaller than the terminal one; pedicels scabrous,
2-20 mm long, the spikelets set obliquely on the
pedicels. Spikelets ellipsoid, 7-9 mm long, 2.1—
2.5 mm wide, glabrous, pale to purplish. Lower
glume 3.5-3.8 mm long, acuminate, 7-1 1 -nerved,

m long, sparsely pilose to gla-
brous, long pilose at the base, the inner surface
pilose towards the apex, 7-1 1-nerved, the mid-
nerve scabrous. Lower lemma glumiform, 6.2—
7.3 mm long, long pilose at the base, the inner
surface pilose, purplish, 7—9-nerved. Lower palea
elliptic to obovate, 4-5.8 mm long, 1.3-2.2 mm
wide, membranous, the borders pilose; male
flower absent. Upper anthecium ovoid to ellip-
soid, 4—4.5 mm long, 1.8-2.1 mm wide, at ma-
turity 2.5 mm wide and dark brown; stipe ca. 1

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

mm long, somewhat fleshy with an expanded
summit and a thick, indurate process on the back,
prolonged beyond the base of upper anthecium
at maturity; rachilla pilose below the stipe. Cary-
opsis 2.8-3.2 mm long, 1.5-2.2 mm wide. In
flower December-September.

Distribution. Bolivia and Brazil; sandy or
sandy-clay savannas, campos or open cerrados;
400-1,300 m

Additional specimens examined. BOLIVIA. SANTA
CRUZ: Santiago de Chiquitos, San pae Cárdenas

4506 (US); Men esp cerca de El Carmen, Cá d as
4503 (US). BRAZIL. BAHIA: Ro De 305 k SW
of Barreiras, Irwin B al. 14657 (MO, NY, SP, US):

Espigão Mestre, 100 km WSW of Barreiras, Anderson
et al. 36654 (F, MO, NY, US); Rio Roda Velha and
highway BR-020, Davidse et al. 12084 (MO, NY).
DISTRITO FEDERAL: Universidade de Brasilia, Clayton
4809 (SP), 4839 (NY, SP); 20 km E of Brasilia, Jrwin
et al. 9213 (F, MO, NY, US); Brasilia, Belém 1970
(CEPEC); 15 km E of Brasilia, Irwin &

121 93 (NY); between Jatahy and Rio Araguaya, Chase
11736 (US); vicinity of Annapolis, Chase 11519 (US);

38 km N of Sao Jose da Alianga, Dawson 14354 (US);
40 km W of Rio Verde, Chase 11713 (US); W of Santa
Rita do Araguaya, Chase 11863 (US); between Vian-
napolis and Ponta Funda, Chase 11281 (US). MATO
Grosso: Rodovia Cuiaba-Santarem, Lemes 4125 (RB);
Rondonópolis, Rio Paguba, Rondon 2566 (RB, US);
Diamantina, Weddell 3081 (US). MATO GROSSO DO SUL:
Xavantina-Cachimbo road, 85 km from Xavantina,
Hunt & Ferreira 5739 (NY, SP, US); NW of Sao Lou-
m Chase 11959 (US). MINAS GERAIS: Lagoa Santa,
14 Feb. 1864, Warming s.n. (US); 26 km NE of Pa-
ire dk et al. 25582 (F, NY, SP); Serra do Cipó,
110k of Belo Horizonte, Chase 9138 (F, GH,

MO, m ead between Sucupira and Omega, S of
Uberlandia, Chase 11167 (US); 3-4 km de Prata, Sen-
dulsky 18 (SP), 37 (SP, US); Frutal, Valls 649 (US);

Caldas, Linc TI 1369 (US); Pratinha, Dorsett 189b

S). R

Pickel 5887 (US); = Sant à n-
dulsky 148 (US); 4 raguacu Paulista, Clay-
ton 4596 (SP, US); sp E agai Black 51-11072
(B MARANHAO: orda to Grajahü,
Swallen 3648 (RB, SP, US).

When publishing this species, Chase described
and illustrated the frag ment of rachilla below the
sai anthecium, show the two constituent

rts. Although she indice that she had not
due this character in any other species of

=e

1987]

the genus, she related P. cervicatum to P. oly-
roides Kunth, and in unpublished manuscripts
placed both species in the **Olyroides" group;
recently, Renvoize (1984) also related P. cervi-
catum and P. ligulare to P. olyroides.

However, in P. olyroides the characteristic stipe
of sect. Rudgeana is absent, and there are long,
acintate hairs at the base of the upper anthecium
on its ventral face (Fig. 4d).

The spikelet is frequently obliquely disposed
on its pedicel in P. cervicatum, a character pres-
ent also in other species of Panicum (e.g.,
hirtum).

Iconsider one ofthe paratypes, Williams 13221
of Venezuela, to belong to P. vinaceum Swallen.
Consequently P. cervicatum remains known only

rom Brazil and northern Bolivia.

4. Panicum ligulare Nees ex Trinius, Gram. Pan.
206. 1826. TvPE: *V. sp. imperfectum Brasil
(N. ab Esenb.)" (lectotype here designated:
floriferous part, LE), non Nees, Agrost. Bras.:
196. 1829. TYPE: “Hab. in campis prope Al-
meirim provinciae Paraensis" (lectotype here
designated: floriferous part of number 3800,
M)

Perennial, 1.30-2 m tall, with thick adventi-
tious roots and lanose cataphylls. Cu/ms erect,
many-noded; internodes 8-24 cm long, solid or
hollow, pilose to glabrous; nodes dark, pilose to
glabrous. Leaf sheaths 8-23 cm long, greenish to
purplish, papillose-pilose, the hairs urticant and
caducous; margins ciliate. Ligule 1.8-3.2 mm
long, with long hairs on the back towards the
base of the blade; collar pale, densely villous.
Leaf ioa Tac kapuni 8 30-55 cm long, 0.9—
1.9 cm wide, flat, acuminate apically, subcordate
basally, panes pilose on both surfaces to gla-
brescen

lax, diffuse, many-flowered, 47-65 cm
30 cm wide, the branches spreading; axis lon-
gitudinally ridged, pilose towards its base, oth-
erwise scabrous, the branches alternate or op-
posite, sometimes verticillate at the base of the
panicle, scabrous, the axils of the branches pi-
lose, pale to brown; axillary panicles usually ab-
sent, when present similar to the upper one but
smaller; pedicels claviform, 2-20 mm long, sca-
brous. Spikelets ellipsoid, 4.4-5.7 mm long, 1.2-

mm wide, glabrous, greenish to purplish.
Lower glume 2.9-3.8 mm long, ⁄2—⁄4 the length
of the spikelet, subulate apically, shortly pilose

ZULOAGA — PANICUM SUBG.

PANICUM SECT. RUDGEANA 475

towards the apex on the inner surface, 7—9-nerved,
the midnerve scabrous. Lower lemma glumi-
form, 4.1-4.9 mm long, acuminate apically, pi-
lose towards the apex in the inner surface, 5-7-
nerved. Lower palea elliptic, 3-3.3 mm long, 0.9-
1.5 mm wide, glabrous, whitish, membranous,
the margins with or without short hairs; male
flower absent. Upper anthecium ovoid, 2.5-3.2
mm long, 1.1-1.5 mm wide, pale; palea with
compound papillae at the apex; stipe conspicu-
ous, glabrous, with 1 or 2 wings nearly 0.8-1.1
mm long, the indurate portion 0.4—0.7 mm long.
Caryopsis 2.4 mm long, 1.3 mm wide. In flower
March-October. Figure 5.

Brazil, from Maranhào and Ba-
rosso; cerrado; 500-1,100 m.
Capim elefante.

Distribution.
hia to Mato
Common name.

Additional specimens examined. BRAZIL. BAHIA: 150
km SW of Barreiras, Irwin 14904 (F, MO, US). DISTRITO
FEDERAL: Chapada de Contagem, ca. 20 km NE of Bra-
silia, Irwin & Soderstrom 5166 (US), Irwin et al. 9653
s: 20 km N of Cristalina, Serra

3700
et al. 21525 (F, US): Serra Dourada, Glaziou 22525
(US); vicinity of Goiás, Chase 11460 (F, GH, NY);
26 Ey By S of Goiania, Davidse et al. 12278 (MO).
MA AO: Carolina to San Antonio de Balsas, Swal-
len 4094 (US); Serra do Penitente, Miranda 128 (RB).

Xava ntina, Irwin et al. 16122
Azul, 77 km from Barra do Gargas, Hunt 6075 (NY,
US); Serra do Roncador, 86 km N of Xavantina, /rwin
F, NY, US); Xavantina-Cachimbo road,

215 km from Xavantina, Hunt & Ferreira Ramos 5606

Y,
atter et al. 2090 (NY, RB); Campos Novos, Kuhl-
mann 1745 (RB).

Trinius (1826), in attributing P. /igulare to
Nees, described the species as possessing a lan-
ceolate, membranous ligule 6-10 mm long, and
used the epithet /igu/are in reference to this char-
acter. After examining abundant material of P.
ligulare and studying the type of P. /igulare in
Leningrad, I discovered that the type sheet con-
tains a mixture of material. The panicle of this
specimen does correspond to what I consider P.
ligulare (which agrees with the description given
by Trinius for the floriferous part), but the veg-
etative portion (which is separated from the flo-
riferous part) is markedly different from the veg-
etative parts of the species. The leaf sheaths and
leaf blades are completely glabrous, and the
membranous ligule is exceptional because of its

476

size. This type of ligule has never been found in
any species of Panicum up to now.

In 1829, Nees described P. /igulare as Trinius
did, mentioning that the type of ligule he ob-
served was unique in Panicum. In his descrip-
tion, Nees reported the type locality as “Hab. in
campis prope Almeirim provinciae Paraensis."
On studying the type material in Munich, I found
two specimens collected by Martius in that lo-
cality, one with the number 3798 (attached to
the plant) and the other identified as 3800. In
3800 there is a mixture of material similar to the
specimen from Leningrad. Specimen 3800 is un-
doubtedly the one Nees used in his diagnosis. In
specimen 3798 there is no mixture, and it fits
perfectly with what I have described as Panicum
ligulare. In this specimen there is a note on which
Trinius stated that this material is different in its
vegetative parts to the one examined at Lenin-

grad.

Trinius (1835) and Steudel (1855) treated the
species in the same way as Nees and Trinius did
before

Doell (1877), in Flora Brasiliensis, noted the
difference between specimens 3798 and 3800 of
Munich. He considered 3800 to be P. ligulare
“in sensu strictiore," but erroneously judged 37 98
to be P. virgatum (a completely different North
American species

I select the floriferous portion ofthe Leningrad
material as the lectotype of Panicum ligulare Nees
ex Trin., and the floriferous portion of the Mu-
nich specimen 3800 as a lectotype of P. ligulare
Nees.

5. Panicum rudgei Roemer & Schultes, Syst. Veg.
2: 444. 1817. Based on P. scoparium Rudge,
Pl. Guian. 1: 21, pl. 29. 1805, non Lam.,
1798. TYPE: “Panicum scoparium Rudge, ex
herb. Rudge" (fragment, US-2830540).

P. ior e Roth ex BERS ERER Syst. Veg.
4 817. TYPE: “Roth nov. plant Spec. Ms. ...

In Essequebo, Mertens” ea at US- 2830939).

P. asl var. brasiliense Raddi, , Agrost. Bras. 48. 1823.

nse ;
P. riiophtlum Mns Syn. Pl. Glum. 1: 1855.
s Salzm. Hrbr. Bahia" sss,

US-4 w
P. cayennense var. divaricatum Doell, in C. Martius,
Fl. Bras. 2(2): 220. 1877. TYPE: same as the species.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Perennial, 30-130 cm tall, with short rhizomes
and pilose, scaly cataphylls. Cu/ms decumbent
or geniculate to erect, often zigzag, rigid, hollow,
branching from the lower and upper nodes, many-

densely to sparsely pilose; nodes covered wit

whitish, appressed hairs to completely glabrous.
Leaf sheaths 4-13 cm long, densely papillose with
thick, glassy hairs; margins ciliate. Ligule 1.5-2
mm long, with long hairs on the back towards
the base of the blade; collar pale, densely to
sparsely pilose. Leaf blades eee 20-
40 cm long, 0.6-1.1 cm wide, acuminate apically,
narrowed Tai flat or with involute borders,
densely hispid to sericeous on both surfaces to
glabrescent, the margins scabrous, ciliate or not
in the lower portion, the midnerve manifest.

rescence !^ or more the length of the plant, 25-
50 cm long, 10-20 cm wide; axis and branches
longitudinally ridged, scabrous to pilose, the ax-
ils of the branches long-pilose to glabrous,
brownish to pale, the branches alternate and di-
varicate, somewhat flexuous; pedicels d
long-pilose, flexuous. iii ovoid, acum
nate, 3-3.5 mm long, I-1. m wide, pale i
nearly purplish, sparsely ous with stiff hairs
irregularly distributed. Lower glume 2-2.7 mm
long, 7^ as long as the spikelet, acuminate api-
cally, with stiff, whitish hairs on the upper part
to completely pilose, the inner surface pilose, 3-
5-nerved, the midnerve scabrous. Upper glume
2.7-3 mm long, acuminate apically, pilose on
the inner surface, 7—9-nerved, the midnerve sca-
brous. Lower lemma 2.5-2.9 mm long, acute api-
cally, long-pilose to glabrous, 7—9-nerved, the
midnerve scabrous. Lower palea elliptic, 1.8-2.3
mm long, 0.5-1.1 mm wide, membranous, the
margins shortly pilose; male flower present, the
anthers purplish; rachilla with or without whitish
hairs. Upper anthecium ellipsoid, 1.8-2.2 mm
long, 0.8-1.1 mm wide; palea with compound
papillae towards the apex; stipe with the mem-
branous portion ca. 0.4 mm long, the indurate
portion 0.5 mm long, prolonged beyond the back
of the lemma as a mucro. Caryopsis pale, 1.5
mm long, 1 mm wide. In flower all year.

Distribution. Mesoamerica, West Indies (Ja-
maica, Trinidad) and South America from Co-
lombia to Bolivia and Brazil; 0-1,000 m; in open
savannas, campos or cerrados, usually in sandy

ils

1987]

Common names.
carricillo (Venezuela).

Chromosome number. n= 9 (Davidse & Pohl,
1974, 1978); 2n = 18 (Pohl & Davidse, 1971)

makuna-ta (Colombia);

Selected specimens examined. MEXICO. TABASCO:
Achotal, Matuda 3087 (F, GH, US). GUATEMALA. IZA-

39673 (F). BELIZE. Cabbage Hall, Dwyer et al. 454 (F.
MO); Swasey Branch, Monkey River, Gentle 3862 (F,
GH, MO, NY, US); Machaca, Gentle 6893, 6923 (F,
NY, US). Costa RICA. ALAJUELA: Buenos Aires, León
1184 (US), Tonduz 3679, 4875 (US); Los Palmares,
Pittier 10588 (US). PUNTARENAS: Buenos Aires, Molina
2739

73 , MO, US); east of CIA, road to Buenos Aires,
Pohl 13116 (F, MO). SAN JOSE: Vicinity of
General, Skutch 3065 (GH, MO, Pittier 12064

). NICARAGUA. ZELAYA: Entr iu a y Limbaikán
Seymour 497 : n Mea Jaboga, Killip 4163
s Canal Zone, ne rt Randolph, Standley 285 98
(MO, US); Perlas eee San Jos and, John-

324 (GH, HL sami Clar-
endon, Harriss 12845 (NY, US); Halliss Savanna, Up-
don, Harriss 12235 (MO, NY, US); Bunkers

S. ; Mason River

O, NY, US); St. Joseph, Hitchcock
abadie, Britton 688

Hitchcock 10083 (M
10181 (US); Piarco d So

Pan de Azúcar, Orozco et al. 7

Guainía, Puerto Colombia a et al. 17936 (US)
META: 73 km W of Las Gaviotas, Davidse 5390 (MO);
43 km NE of Puerto , Davidse

Wilches y au “a km 16 Killip & Smith
T (F, GH, MO, NY, US). ToLIMA: El Con
W of San Lorenzo, PD 3509 GH, MO. ‘NY.
US). VAUPES: hans I Circasia, Eb 7201 (US).
VICHADA: 25 k of Cumaribo, Davidse 5325 (MO);
10 km W of Las “sasa Davidse 5367 (COL, MO,
NAS: Cerro Duida, Maguire
29424, 29060 (NY); Serrania Parü, Cowan 31486 (NY,
US); 20 km S of Puerto Ayacucho, Davidse 2641 (MO);
5 km NE of San Carlos de Río Negro, Liesner 3703
(MO); 25 km S of Samariapo, Gentry & Berry 14600
(MO); Yavita, Williams 13879 (F, US); pie del Cerro
Huachacamari, Huber 4990 (MO); El Manguito, 1 km
N of Cafio Caname, Davidse et al. 17482 (MO); alrede-
dores de Canaripo, Huber 1981 (MO); 8 km S de Puerto
Ayacucho, Davidse & Huber 14916 (MO). ANZOATEGUI:
Vicinity of Santo Tomé, Chase 12841 (GH, US). APURE:
end of the — de Cinaruco, Davidse & González
14667 (MO); nea Fundo El Algarrobo,
Davidse & ar den 14217 MC. BARINAS: 16 km SW
of the Merida rsa dg just outside of Barinas, Da-
vidse 3182 (M

raima, Steyermark 59429 (F, US); 0.5 km NE of

ZULOAGA — PANICUM SUBG.

PANICUM SECT. RUDGEANA 477

ides Steyermark & Wurdack 22 (F, NY, US).
ONAGAS: E de Maturin, ca. caserio La Pica, Ariste-
iun 4048 (F, MO, NY); Laguna Most, 12 km N de
Capirito, Trujillo 14194 (F). ZULIA: 60 km NW of Santa
Bárbara-San Carlos del Zulia, near Campamento El
Rosario, de Bruijn 1473 (MO, NY, US). GUYANA. Tu-
matumari, See os 40 (GH); Waini River, de la Cruz
(F, Waramuri Mission, Horuka

e Tu
Hills Estate, Hitchcock 17191 (F, MO, NY, yea Kaie-

St. Laurent

ERU. LORETO: , Rio anon, Gentry et al. 29965
(MO); via Nauta- jn E Díaz & Jaramillo 1270 (MO).
BOLIVIA. BENI: 15 km de erin, camino a
Riberalta, Krapovickas & Bator 35068 (U 9). LA PAZ:
San Carlos, Buchtien : polo,

Williams 1020 (NY); San Antonio, Buchtien 1 15 9 (US)

(CEPEC, MO, NY); Marau, Belém &
(CEPEC), Zuloaga et w deg (RB

418, 16 km del cruce A- 01.
(CEPEC, NY). MATO GROSSO: Serra Azu
Xavantina, Irwin et al. 17302 (F, MO, NY, US); 270
km N of Xavantina, Ratter 2069 (NY
s dos pn iion: antina, Irwin et al. 15961 (F,

US). M I DO SUL: Tres Lagoas,
Pen por (US) P PARA: Santarém, fisv 3721 (US);
Soure, Ilha do Marajó E a Acara,
Thomé Assu, Mexia 592 io

US); 73 km NE se eid FUE "E 17939 (MO.
NY); 17 km SE of Vigía, along road Pa-140, Davidse
et al. 17610 (MO, NY). PERNAMBUCO: Vicinity of Re-
cife, Chase 7675 (F, US), Poazeves, Pickel 3137 (US).
DE JANEIRO: Silvestre, Holway et al. 1116 (US);
Merity, 20 km N of Rio de Janeiro, Chase 8465 (US).
RONDONIA: Porto Velho, Black & Cordeiro 52- 15348

mann 4110 (US); 7 km de São José dos Campos, Eiten
& Mimura 3351 (MO, US).

6. Panicum vinaceum Swallen, Fieldiana. Bot.
28(1): 27. 1951. TYPE: Venezuela. Bolívar:
Gran Sabana, between Kun and waterfall at
Rue-Meru, south of Mount Roraima, elev.
1,065 m, 2 Oct. 1944, J. A. Steyermark 59173
(holotype, US-1911661; isotype, F).

Perennial, 40-100 cm tall. Culms erect, few-
noded; internodes 6-11 cm long, pilose; nodes

478

pilose. Leaf sheaths 4-14 cm long, covered by
long dense hairs or glabrescent; margins ciliate
to glabrous. Ligule 0.6-2 mm long; collar pale,
pilose. Leaf blades linear-lanceolate, 15-42 cm
long, 0.5-1.2 cm wide, flat, acuminate apically,
subcordate basally, with appressed hairs on both

long, 6-20 cm wide, the branches spreading; axis
longitudinally ridged, scabrous, the branches al-
ternate to opposite, scabrous, the axils of the
branches pilose, pale; axillary panicles usually
absent, when present similar to the terminal one
but smaller; pedicels scabrous. Spikelets ellip-
soid, 5.9-6.7 mm long, 1.5-2 mm wide, globose
and glabrous, pale to purplish, the inner surface
ofthe glumes and lower lemma densely to sparse-
ly pilose. Lower glume 2.9-3.8 mm long, subu-
late apically, 5-9-nerved, the midnerve scabrous.
Upper glume 5.3-6.4 mm long, acuminate api-
cally, 7-1 1-nerved, glabrous. Lower lemma 5.1—
5.5 mm long, acute apically, 7-9-nerved. Lower
palea obovate, 3.1-4 mm long, 1-1.5 mm wide,
whitish, membranous, the margins pilose; male
flower absent. Upper anthecium ovoid, 3-3.5 mm
long, 1.4—1.8 mm wide, pale; stipe with the mem-
branous portion 0.7-1.2 mm long, with or with-
out wings, the indurate portion ca. 0.9 mm long,
obtuse; rachilla pilose below the upper anthe-
cium. Caryopsis 2.4 mm long, 1.5 mm wide. In
flower September-A pril.

Distribution. Brazil and Venezuela; 100-

,000 m; savannas.

Additional specimens examined. BRAZIL. GOIAS: Rio
da Prata, 6 km Posse, Irwin et al. 14509 (US).
PARA: Serra do Cachimbo, BR-163 Cuiabá-Santarém,

km 823, Prance et al. 24993 (MO, NY); Serra do Ca-

Y, US); Estación Bo-
livar, en sabanas de Santa P Tamayo 2964 (MO,
NY, US).

This species is closely related to P. cervicatum,
from which it can be separated only by the sizes
of the spikelet, lower palea, and upper anthe-
cium. The rest ofthe differential characters noted
by Swallen (size and pilosity of the plants and
size of the panicles) have no value in separating
the two species. Irwin 14509 is exceptionally large
and differs from Swallen's description. Never-
theless, spikelet size (a constant character in the

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

scant material available for this species), shows

that Irwin 14509 must be included in P vina-
ceum, thereby partially di s con-
cept of this species.

LITERATURE CITED

BERG, R. Y. 1
traliense (Poaceae): taxonom
plications. Austral. J. Bot.

Brown, W. V. 1977. The Kranz syndrome and its
subtypes in grass ee Mem. Torrey Bot.
Club 23(3): (ii + 7.

DavipsE, G. Fruit pee and seedling iones
ment in the Poaceae. Proceedings of the
tional Symposium on Grass Systematics ind Evo.
lution. Smithsonian Contributions to Botany (in
press).

985. Spikelet structure in Panicum aus-
c and ecological im-
3: 79-583.

W. POHL. 1974. Chromosome numbers,
meiotic behavior, and notes on tropical American
ine i a Canad. J. Bot. 52: 317-328.
. Chromosome numbers of
a "nd grasses (Gramineae): 5. Ann.
Missouri Bot. Gard. 65: 637-649.
DoELL, J. C. 1877. Tribe 3, Paniceae. In C. Martius
aa Flora Brasiliensis 2(2): 33-342, pls. 12-

eee J. TH. 1940. Notes on the nomenclature
of some grasses. II. Blumea 4: 503-507.

HirTCHCOCK, A. S. 1915. Panicum. North American
Flora 17: 198-289.

& A. CHASE. 1910. The North American

species of Panicum. Contr. U.S. Natl. Herb. 15:
6

& — 1915. Tropical North American
species of Panicum. Contr. U.S. Natl. Herb. 17:
35.

LAZARIDES, M. 1959. The Australian species referred
to Ichnanthus Beauv. (Gramineae). Austral. J. Bot.
7: 328-346.

& R. D. WEBSTER. 1984. Yakirra (Paniceae,
Poaceae), a new genus for Australia. Brunonia 7:
289-296.

NEES VON ESENBECK, C. G. D. 1829. Agrostologia
Brasilien A .... Stuttgart & Tübingen. (This is

.l, Gramineae, of Flora Brasiliensis seu
oer tara .., edited by C. F. T von Martius.)

G. Davipse. 1971. Chromosome

numbers of Costa Rican grasses. E 23: 293-

324.
RENVOIZE, S. A. 1984. The Grasses of Bahia. Royal
Botanical Gardens, Kew, Englan
SHAW, R. . D. WEBSTER. 1983. Characteristics
ofthe (Poaceae: Pan-
iceae). "Bot. jm iced 144: 363-370.
e UDEL, E. nopsis plantarum Gram
earum. In Synopsis Banta Glumacearum "
-47

usns C. B. —-1836. Species Graminum ico-

nibus et DUM DER UA 3 volumes, 360

plates. Leningrad, U.S.S.R
1826.

De Graminibus Paniceis: Dissertatio
botanica altera. 291 pages. Impensis Academiae
Imperialis Scientiarum. St Petersburg, U.S.S.R.

ARUNDOCLAYTONIA, A NEW GENUS OF THE
STEYERMARKOCHLOEAE (POACEAE: ARUNDINOIDEAE)
FROM BRAZIL!

GERRIT DAVIDSE? AND R. P. ELLIS?

ABSTRACT

i seni ae i: NE iaa Davidse & Ellis, gen. et sp. nov. is described from Amazonian campinas
i a, Bra

e second genus

| quien asiy lt odis aban it is bor ie b a caespitose growth habit, proliferation. and

lignification of the numerous basal culm

unisexual iran ag iin inflorescences Haas into a false panicle, 3—9-flow
er, 3-flowered female spikelets with only the middle floret ertile and its palea

l
fusiform caryopsis with an elliptic- -punctate hilum. Anatomically this species is charact
isodiametric chlorenchyma, and 2 bundle het we is

ternodes, normally developed leaves - spiral phyllotaxy,

red male spikelets

by the absence of stomata and Edu reduction jajn silica bodies, thick epidermis, and hypodermal

ows. Its classification in Steyermarkochloeae

is based primarily on the morphology of the aa

During 1974 an unusual grass was collected
by William R. Anderson and associates in Pará,
Brazil. Although it was recognized as an unde-
T taxon, the inflorescences were too im-
mature to show the exact morphology of the
doe In 1979 Cleofé E. Calderón and co-

workers collected abundant mature material of
a same taxon in Amazonas, Brazil. Our study
of both these collections indicates that they rep-
resent the second genus ila lan — Š again

dse & Ellis, 1984).
We are naming the zenux in pid of = W. D.
Clayton, eminent agrostologist at the Royal Bo-
tanic Gardens, Kew, who has made and contin-
ues to make outstanding contributions to agros-
tology. The compound generic name at the same
time refers to the arundinoid affinity ofthe genus.
The specific epithet alludes to the strongly dis-
similar male and female inflorescences and
spikelets.

DESCRIPTION
x asi ivan salen aig eae & Ellis, gen.
et sp. nov. TYPE: Brazil. Amazonas: Trans-
amazon Highway. ca. "53 km W of the Ari-

puafia River, abundant dominant plant of
the vegetation. Growing in a white sand soil
“campina.” This plant grows in large, open
areas mixed with shrubs and alternating with
narrow strips of islands of low tree forest.
Most of the population reduced to burnt
bases. These trunks look like big candelabra,
some ca. 70 cm or less. From them come
up solid stems with thickened bases formed
by aerial roots. In many cases from the top
of burnt trunks, bunches of leaves start com-
ing again. Few plants still blooming. Plants
ca. 2-3 m tall when flowering. 28 June 1979,

alderón, O. P. Monteiro & J. Guedes

R
SP. US (mounted as 11 sheets)]. Figures

Gramen duced culmi internodiis numerosis in-
ferioribus he
is; phyllot

sam; spicula
glumis disarticulates; glumae 2; spiculae masculinae 3-

! We extend our gratitude to the late Dr. Thomas R. Soderstrom, Smithsonian Institution, who made the

vii uid Calderón et al. enia and photographs available to us and
seful review comments by Dr.

uch appreciate the ve
still dise with him o

do Cuchimbo:
ri Botanical Garden

who encouraged us in our studies. We
Steve Renvoize, but, pending additional m
for drawing

eid tribal classification of Steyermarkochloeae. We thank Jo
r. William R. Anderson, University of Michigan, for information about his collecting itinerary

, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

3 Ry Research Institute, Private Bag X101, Pretoria 0001, South Africa.

ANN. Missouni Bor. GARD. 74: 479—490. 1987.

480

9-florae palea 2-carinata; stamina 2; spiculae femineae

stigmate 2; caryopsis fusiformis-teres hilo punctato.

Perennial 2-3 m tall, erect. Vegetative culms
usually densely covered for 2-70 cm to a thick-
ness of 1.5-6 cm by aerial roots tightly appressed
to the culm and by remnants of leaf sheath bases;
internodes numerous, 2-15 mm long, 1-1.5 cm
diam., solid, lignified; nodes bearing one prom-

ta

intravaginal near t asal cluster o
Flowering culms to 1 cm diam., consisting of
many, often elongated internodes; internodes 1-
16 cm long, glabrous, densely waxy when young,
green in the exposed portions when older, hol-
low, gradually becoming solid toward the base
ofthe plant; nodes glabrous; branching primarily
intravaginal, profuse in the upper !^ of the culm
to form a false inflorescence. Leaves primarily
clustered toward the base, those of the flowering
culms fewer and gradually reduced in size toward
the tip of the culm. Basal leaves with the sheaths
densely overlapping, much longer than the in-
ternodes, stramineous, long persistent, turning
brown and eventually reduced to fibers in age,
rounded and glabrous abaxially, without a dif-
ferentiated midrib, the margins glabrous, free to
the base, the base pilose at the point of insertion
and between the veins or glabrescent, the apex
ciliate with hairs 2-4 mm long, wider than the
base of the blade, rounded, a collar not clearly
differentiated; ligule a ciliate membrane 0.9-2.1
mm long, the membrane 0.3-0.9 mm long, the
cilia 0.5-1.2 mm long; blades 45-80 cm long, 8-
16 mm wide, flat with involute margins or en-
tirely involute, the upper portion always involute
and the apex pungent, the abaxial surface green,
glabrous and smooth, the adaxial surface grayish
green, densely and minutely scabrous, grooved
between the veins, the veins approximately the
same size, a midrib not differentiated, the mar-
gins ciliate with hairs 2-3 mm long in the lower
l^. scaberulous in the upper 7^. Cauline leaves
similar to the basal leaves but smaller, the up-
permost much reduced with the blade shorter
than the sheath and entirely involute. /nflores-
cences numerous, borne on axillary, exserted pe-
duncles, aggregated into a false panicle, unisex-
ual, consisting of hemispherical clusters of 7-20
l or 2 series

florescences produced before the female, 9-13

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

mm wide, 6-11 mm high; female inflorescences
20-36 mm wide, 15-23 mm high; peduncles ge-
niculate and pilose at the base with hairs 0.5-1.5
mm long, sometimes with a line of pubescence
or with a scarious bract 1-3 mm below the cluster
of spikelets, always subtended by a sheath, the
sheath terminating in a sharp point; peduncles
of the male inflorescences usually longer than
those of the female inflorescences. Female spike-
lets 7-19 mm long at anthesis, sessile or short-
pedicellate with pedicels to 0.5 mm long, lan-
ceoloid, rounded on the back, disarticulating be-
low the glumes, falling as a unit, slightly curved,
3-flowered; glumes 2, unequal, herbaceous,
shorter than the lemmas, ovate, broadly acute,
pilose at the base, otherwise scaberulous, the
nerves free or connected by cross-veinlets, the

wer 1.5-2.0 mm long, 1-3-nerved, the upper
2.2-3.5 mm long, 3-5-nerved; lower floret with-
out a flower, the lemma 3.0-5.1 mm long, 7-9-
nerved, ovate, broadly acute, pilose at the base,
otherwise scaberulous, cross-veinlets few, the pa-
lea absent or rudimentary, 0.7-1.5 mm lon
hyaline when present; middle floret unisexual,

ma 5 rved

o

conspicuous cross-veinlets, ovate, acute, pilose
at the base and between the nerves just above
the base with hairs 1-1.5 mm long, otherwise
scaberulous, the palea conspicuously longer than
the lemma, 7.5-17 mm long, 9-13-nerved,
slightly curved in the upper half, convolute, shal-
lowly grooved on the back, spongy-thickened,
smooth and shiny in the lower '2—*4, herbaceous
and scaberulous in the upper '4—'/, ciliate on the
overlapping margin at the base with hairs 1-1.5
mm long; upper floret rudimentary or consisting
ofa single 3-nerved bract, 0.1-4 mm long, ciliate
at the base, borne on a prominent rachilla 3.5—
10.5 mm long, the floret and rachilla fitting into
the palea groove of the middle floret; lodicules
absent; staminodia absent or present as an an-
terior pair of rudiments to 0.2 mm long; gynoe-
cium cylindrical, the ovary wall free from the
ovule, the style one, dividing into 2 inconspic-
uously plumose stigmas slightly below the tip of
the middle palea, the stigmas 2.5-4 mm long,
terminally exserted through an apical, iain
orifice formed by the convolute palea; caryopsis
fusiform-terete, narrowing apically, 6-7 mm long,
0.8-1.2 mm diam., glabrous, the embryo J49-^,,
as long as the caryopsis, the hilum elliptic-punc-
tate. Male spikelets 3.5—7.5 mm long, sessile or
short-pedicellate with pedicels to 0.5 mm long,
rounded on the back, disarticulating below the

1987]

glumes, 3—9-flowered, the florets (except the up-
permost) bearing flowers, the middle florets
slightly larger than those above or below, the
uppermost usually rudimentary; glumes 2, un-
equal, shorter than the lemmas, membranous,
ovate, erose, truncate or obtuse, pilose at the
base, usually with cross-veinlets, the lower 1.3—
2.1 mm long, 1—3-nerved, the upper 1.8-2.5 mm
long, 3-nerved; lemmas similar to the glumes in
pubescence, shape and texture, 2.6-4.1 mm long,
shorter than the paleas, 3—9-nerved; paleas 3.2-
5.8 mm long, broadly obtriangular, truncate,
sometimes erose, 2-keeled (each keel with a
nerve), the base with hairs 1-1.5 mm long, the
back sulcate, the keels ciliolate, the margins over-
Wine lodicules absent; stamens 2, one situated

side of the sulcate palea, terminally
Heer through an opening formed by the over-
lapping palea margins, the filaments separate, ba-
sifixed, the anthers 2.2-2.9 mm long.

Paratypes. BRAZIL. PARA: Alto Tapajós, Rio Cu-
ruru, northwest edge of Serra do Cachimbo, 25 km by

Santos & R. Souza 10950 (MO, NY, UB).

MORPHOLOGICAL OBSERVATIONS

Arundoclaytonia dissimilis when fully mature
and undisturbed by fire has an unusual appear-
ance caused by the thick accumulation of leaf
sheath bases and adventitious aerial roots (Figs.
1, 3). Such plants in the aspect of their basal parts
are more reminiscent of certain species of Vel-
lozia. This unusual appearance is accentuated
after the plants have been moderately or severely
burned (Figs. 2, 4, 5).

Although the plant is fundamentally a tussock
plant, the dense cluster of leaves, which is nor-
mally basal in a tussock grass, is raised up to 70
cm above ground level in older plants of Arun-
doclaytonia (Figs. 2-5). These small “trunks” re-
sult from the proliferation of numerous, short
internodes in the basal portion of the culms. Short
basal internodes are typical of grass culms. What
is unusual in Arundoclaytonia is their large num-
ber, thickness, woodiness, and perennial dura-
tion. Annual culms characterize most grasses.

In typical caespitose perennial grasses the short
basal internodes perennate and bear the buds
from which new tillers are produced for the new
growing season. It is this region of the culm that

DAVIDSE & ELLIS— ARUNDOCLAYTONIA

481

is much elongated by the proliferation of inter-
nodes and gives Arundoclaytonia its trunklike

es, ei
Arundoclaytonia. In contrast, the elongated in-
ternodes of the flowering culm produced above
the cluster of basal leaves gradually become hol-
low, as is common among grasses.

When the sheath bases and mass of aerial roots
have been removed from the lower portion of
the culm (as may happen after severe burning
and the subsequent wearing off of the root and
sheath remains), it becomes apparent from the
position of the axillary buds that the leaf ar-
rangement is not distichous. Every sixth node
bears a solitary, prophyllate, dormant bud (Fig.
7A) that occurs in the same relative position as
the buds five nodes above and below it. Since
two complete turns around the culms must be
made to attain the same position, phyllotaxy is
2/5. The arrangement of the spikelet bracts ap-
pears to be nearly distichous; however, the rel-
ative position ofthe bracts is much more difficult
to observe because the very short internodes of
the spikelet and the broad bases of the glumes
and lemmas obscure the exact point of insertion
of these spikelet parts. Distichous phyllotaxy is
characteristic of the Poaceae (Arber, 1934: 282;
Barnard, 1964: 47). Only one other exception has
been reported: spiral phyllotaxis in Micraira sub-
ulifolia F. Muell., a mosslike plant from Queens-
land (Watson & Dallwitz, 1980: 89)

Branching occurs near the base of the plant to
orm the main culms that constitute the bulk of
the tussock (Figs. 4, 5). These branches originate
from the buds illustrated in Figure 7A. Branching
is very infrequent in the middle portion of the
culms but profuse in the upper portion. At each
upper node, a smaller axillary branch is produced
which itself is rebranched several times (Fig. 8)
into branchlets terminating in inflorescences. The
branching pattern is the relatively simple one

_
5

era. A prophyllum is the first foliar organ pro-
duced at the lowest node of each branch (Fig. 8).
Each prophyllum is many-nerved and promi-
nently two-keeled with narrow wings on the keels.
Buds at subsequent nodes on the branch are sub-
ended by leaves with blades reduced and gen-

Just below the inflorescences they are reduced to
scarious bracts, presumably representing re-
duced sheaths only

482 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FIGURES 1-6. Habitat and habit photographs of Arundoclaytonia dissimilis; E W of the Aripuana River,

Nerio] Brazil.—1. Unburned campina.— 2. Burned campin t rned flowering plant.— 4.
Contrast between a severely burned, killed plant and a moderately burned regenerating plant.—5. Detail of the
base of a burned, regenerating plant.—6. Detail of a section of the com inflorescence showing the small,

long-peduncled male inflorescences below and above and the uel laa seine short-peduncled female
inflorescences in the center. Photographs by Dr. Cleofé E. Calderó

IE
Ii Mi ;

Ill TI |
WU

R

FiGuRE7. Morphology of Arundoclaytonia dissimilis.—A. Basal portion of culm with the sheath bases worn
off and showing the approximately identic m Line of the buds every sixth node.—B. Portion of a leaf n
ligule. — C. Sq inflorescence. — D. Male spikelet. E-I. Components of ipn spikelet.— —E. Lower glume
Upper glume.—G. Lemma.—H. Inner P i palea with the two stamens composing the male flower. E
Stamen. — J. Female inflorescence. — K. Female spikelets. L-R. Components of female Pudet —L. Lower iin me.—

. Upper glume.—N. Lower lemma.— O. Middle lemma, ventral view.— P. Middle lemma, dorsa
Middle palea and rachilla extension with the rudimentary upper floret. — R. T proba bly pollinated
and slightly expanded. S-U. Caryopsis in three different views. — —S. Lateral view.— T. Hilum view. es mbryo
view. Scales: A, B = 1 cm; C, D, I-K, U = 1 mm; magnification for E-H, K-R, and S-U the sa
48

484

key: v =
7

ANNALS OF THE MISSOURI BOTANICAL GARDEN

ff

[Vor. 74

foliar bract

female inflorescence

male inflorescence

prophyllum
leaf

B = meristematic tissue, bud

------- = continuation

d illustration of a portion of the inflorescence of Arundoclaytonia dissimilis showing

FIGURE 8.
the Ein of the internodes,
been altered for clarity nre are shown i in two dimensio

At any node along the flowering portion of a
main culm, male inflorescences are produced be-
fore the female inflorescences, and the male in-
florescences are borne on longer branches than
those of the female inflorescences. Each branch
complement along the main culm ultimately ter-
minates in a male inflorescence. The proportion
of male to female inflorescences varies from 2
male: 1 female in the lowest portion, gradually
changing to 1 male:3 female in the uppermost

portion. wever, since the male spikelets are
functionally 2-8-flowered compared with the
functionally one-flowered female spikelets, the
total number of male flowers is greater than fe-
male flowers. Although the plant is fundamen-
tally protandrous, the large number of inflores-

leaves, prophylla, nie EEA: The proportions of the structures have

cences produced by any mature plant ensures a
significant overlap between the flowering of male
and female spikelets. No information is available
about self-incompatibility or frequency of flow-
ering.

The male and female spikelets are strongly di-
morphic and, besides the difference in flowers
and number of anthoecia, differ significantly in
the anthoecial morphology. Both the lemma and
especially the palea of the functional floret of the
female spikelets are convolute and thicker in tex-
ture, and the palea is greatly elongated and 9-
13-nerved (Fig. 7K, O-Q). In contrast, the lem-
mas and paleas of the male spikelets are mem-
branous, the lemmas are 3-9-nerved and round-
ed on the back, and the paleas are 2-keeled and

1987]

2-nerved (Fig. 7D, G, H). The relative lengths
of lemmas and paleas in the two kinds of spike-
lets are similar, the paleas being longer in both
kinds, although those of the female spikelets tend
to be somewhat longer than those of the male
spikelets.

In both kinds of inflorescences the outer whorl
of spikelets is surrounded by a ring of small ster-
ile bracts that we interpret to represent rudi-
mentary spikelets (Fig. 7C, J). In some cases these
rudimentary spikelets may reach 4 mm in length
in the female inflorescence and consist of four or
five bracts in the same positions as the normal
female spikelet parts. From such rudimentary
spikelets there is a gradual diminution and sim-
plification to small solitary bracts. Occasionally
one of the normally sized female spikelets on the
outside of the inflorescence has an extra bract.
However, those on the inside of the inflorescence
uniformly have the two glumes and three florets.
In the several cases where a small extra bract was
observed in the inner part of the inflorescence,
it clearly originated below the very short pedicel
and presumably also represented a rudimentary
spikelet.

LEAF BLADE ANATOMY
ANATOMICAL TECHNIQUES

Anatomical studies were carried out on leaves
from herbarium specimens and those fixed in the
field in FAA. Preparation of the sections fol-
lowed the methods outlined by Ellis (1984). The
very fibrous nature of the leaf blades frequently
caused the sections to tear as they were cut, mak-
ing it difficult to obtain completely undamaged
sections.

LEAF IN TRANSVERSE SECTION

Outline: open, expanded with the margins
slightly recurved (Fig. 9). Leaf thickness 30 um
laterally to 40 um centrally. Ribs and furrows:
prominent, flat-topped adaxial ribs with straight,
vertical sides present over all the vascular bun-
dles (Fig. 10); ribs associated with first-order and
third-order vascular bundles of identical size and
shape; furrows narrow, cleftlike, penetrating at
least half the leaf thickness. Abaxial ribs or fur-
rows absent. Median vascular bundle: no midrib
or keel developed; median vascular bundle struc-
turally indistinguishable from lateral first-order
bundles. Vascular
25 first-order bundles with metaxylem vessels

1O1€ than

DAVIDSE & ELLIS—ARUNDOCLAYTONIA

485

per leaf section; one third-order bundle without
metaxylem vessels between consecutive first-or-
der bundles, this alternating pattern occurring
across the full width of the blade (Figs. 9, 10).
All vascular bundles located slightly closer to the
abaxial surface. Vascular bundle structure: first-
order bundles round to elliptical in outline (Figs.
9-12); phloem tissue adjoining the inner bundle
sheath; protoxylem lacunae present; metaxylem
vessel elements wide (+5 um) with a diameter
double that of the parenchyma sheath cells, thin-
walled and slightly angular (Fig. 12). Third-order
bundles elliptical with xylem and phloem tissue
distinguishable. Vascular bundle sheaths: first-
and third-order bundles completely surrounded
by an inner bundle sheath (Fig. 11); mestome
sheath cells relatively large, of the same diameter
as the parenchyma sheath cells; secondary walls
heavily but uniformly thickened, almost exclud-
ing the lumen (Figs. 11, 12). Outer bundle sheath
round, adaxially and abaxially interrupted by
sclerenchyma girders (Figs. 11, 12); bundle sheath
extensions absent; cells elliptic, variable in size,
thin-walled and lacking chloroplasts. Scleren-
chyma: adaxial girders inversely anchor-shaped,
following the shape of the adaxial ribs (Figs. 11,
12); abaxial sclerenchyma forming a continuous
hypodermal band with projections toward the
vascular bundles as well as the bulliform cell
groups (Fig. 11). Fibers very thick-walled with
lumens almost completely filled; lignified, except
those projecting toward the bulliform cells which
may have cellulose secondary walls (Fig. 10). Me-
sophyll: chlorenchyma not radiately arranged;
cells small, isodiametric and tightly packed with-
out visible intercellular air spaces (Fig. 11); oc-
cupying the sides of the ribs but divided abaxially
by the bulliform cells, colorless cells, and abaxial
hypodermal sclerenchyma; arm or fusoid cells
absent. Colorless, inflated, thin-walled paren-
chyma cells linking the bulliform cells to the
hypodermal sclerenchyma. Adaxial epidermal
cells: bulliform cells at the base of all furrows
and occurring in restricted, fan-shaped groups
with an inflated central cell. Epidermal cells with
a very thick cuticle, even on the sides of the
furrows; papillae or macrohairs absent; inter-
locking prickles common on the sides of the fur-
rows (Figs. 11, 12). Abaxial epidermal cells: bul-
liform cells absent; epidermal cells small, with
an extremely thick, continuous cuticle equal in
thickness to the diameter of the epidermal cells;
hairs, papillae, and stomata absent; costal and
intercostal zones not differentiated.

486 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FIGURES 9-12. Leaf blade anatomy of Arundoclaytonia dissimilis in transverse section. —9. Outline showing

the absence of a keel.— 10. Alternating first- and third-order vascular bundles and buen nent adaxial ribs and
cleftlike furrows.—11. Anatomical detail showing double bundle sheath, compact mesophyll, Dance anchor-
shaped adaxial sclerenchyma girder, abaxial hypodermal band and reel celis E associated colorless
cells. — 12. Scanning electron e adaxial furrows and structure
of the metaxylem vessel elements. Scales: 9 — 50 u ; 10 = 20 um; 11, 12 = iD um. " Based on Anderson et al.
10950 (Figs. 9, 10) and Calderón et al. 2706 (Figs. 1 1, 12).

ABAXIAL EPIDERMIS rophytic adaptations. This is demonstrated most

clearly by the well-developed abaxial hypoder-
mal sclerenchyma, extremely thickened abaxial
epidermis, and strongly ribbed and furrowed
adaxial surface. These xerophytic modifications
appear to have led to the consequent loss or re-
duction of many epidermal features commonly
well developed in most other grasses. The most
important reductions are the lack of distinction
between the abaxial costal and intercostal zones,
the absence of abaxial stomata and microhairs,
and the reduction of silica bodies. These xero-
phytic features undoubtedly allow a rapid in-
rolling of the leaves and may be responsible for
the involute margins or completely involute
leaves observed on the herbarium specimens.

Zonation: costal and intercostal zones indis-
tinguishable; entire epidermis composed of uni-
form long and short cells (Figs. 13-15). Long
cells: elongate rectangular, length 3 x the width,
anticlinal ben parallel, end walls vertical (Fig.

orizontal and vertical anticlinal walls
kaviy Ghee. pitted and deeply sinuous.
Long cells usually adjoining one anther but in-
frequently separated by cork-silica cell pairs. Sto-
mata: lacking on the abaxial surface (Fig. 13).
Short cells: tall, with irregular ao associated
with silica cell of similar shape (Fig. 14); occur-
rence irregular. Papillae: absent. MS ab-
sent. Silica bodies: tall and narrow, irregular in
outline; scattered throughout the epidermis.

+
WT

CLASSIFICATION AND DISCUSSION
ADAXIAL EPIDERMIS

As mentioned in the introduction, we consider
Arundoclaytonia to belong in the now bigeneric
Steyermarkochloeae despite the fact that Arun-
doclaytonia differs markedly from Steyermarko-

chloa in growth habit, leaf morphology, and in-

Sides and tops of the ribs covered with prickle

rickles obscuring all other epidermal details of
this surface (Fig. 16).

ANATOMICAL OBSERVATIONS
markable rese bibis ance between the genera be-

The anatomy of the leaf blade of Arundoclay- comes evident. Ignoring for the moment a low
tonia is highly modified and exhibits many xe- percentage of bisexual spikelets in Steyermarko-

heavily thickened, sinuous- walled lon

photomicrograph of the adaxial epider

DAVIDSE & ELLIS— ARUNDOCLAYTONIA

BRRRG G

i Aru ann,

COUT
oye

P NMU Y V VENT
^ LUMA CL

i TRAIL
DOR i m

w r^

p

A VU"

ev
t ert try

xr
a `

um
5 = 10 um; magnification for 14-16 = - 10 um. Based on Anderson et al. 10950 (Figs. 13. 14) and Calderón et

al. 2706 (Figs. 15, 16).

chloa, the fundamental structure of the unisexual
spikelets is identical in the two genera. The male
spikelets have two stamens, are cioe
and lack lodicules in both genera. The majo
difference in the male spikelets is that those of
Arundoclaytonia have more florets. This kind of
variation is analogous to that between species of
Eragrostis, Bromus, and Bambusa, to name just
three of the many genera in which this kind of
ariation is well known. The female spikelets are

in the size and eau aes is SEES These
striking and fi et

almost certainly not due to: Ameena evolution
but indicate a fundamental phylogenetic rela-
tionship which is reflected in our classification
of Arundoclaytonia in the Steyermarkochloeae
The generic status of Arundoclaytonia is jus-
tified by the following major differences from
Steyermarkochloa: monomorphic vs. dimorphic

culms; typical vegetative leaves with open sheath,
many-ribbed blade, and ligule vs. highly modi-
fied vegetative leaves with stemlike, solid sheath,
two-ribbed blade, and ligule absent; many small,
hemispherical male and female inflorescences
aggregated into a false panicle vs. single, large,
terminal, spicate inflorescences bearing male, fe-
male, and bisexual spikelets; and in leaf anato-
my—vascular bundles at one level vs. different
levels; absence vs. presence of lacunae; absence
vs. presence of abaxial stomata; and adaxial fur-
rows and ribs associated with all vascular bun-
dles vs. associated only with the median vascular
bundle.

Our decision to place Arundoclaytonia in the
Steyermarkochloeae necessitates a modification
of the description of the tribe (Davidse & Ellis,

1984). Because the leaves of Steyermarkochloa
are unique in the family, leaf characters were
believed very important i
and in differentiating it from others. This i is now

gthe tribe

488

shown to be true only when Steyermarkochloa
was known. In fact, at the macroscopic level, the
tribe now encompasses both “normal” and high-
ly modified leaves. In this light the unusual fea-
tures of the leaves of Steyermarkochloa, both at
the macro- and microscopic levels, must be seen
as adaptations to its seasonally inundated habi-
tat, just as the strongly xerophytic features of
Arundoclaytonia are presumably adaptations that
allow it to cope with the nutrient deficiencies,
frequent moisture stress, and intense solar ra-
diation of the white-sand soils of its campina
habitat (Ab'Sáber, 1982; Anderson, 1981). Such
white-sand soils are considered to be the most
nutrient-deficient soils in South America (Eiten,
1978). Although the campinas are located in high
rainfall regions, they dry out rapidly near the
surface during periods of low rainfall and never
experience the long-sustained inundation of the
sabaneta or morichal habitats of Steyermarko-
e (Ab'Sáber, 1982; Eiten, 1978; Anderson,

ier formal, emended tribal description is the
following:

Steyermarkochloeae Davidse & Ellis, Ann. Mis-
souri Bot. Gard. 71: 994. 1985.

Perennial grasses with mono- or dimorphic
culms and leaves; leaves solitary or numerous
per culm, consisting of a flattened sheath, blade,
and ligule, or a solid, cylindrical sheath and flat-
tened blade without a ligule, or reduced to blade-
less flattened sheaths. Inflorescence spicate, elon-
gate and cylindrical or a hemispherical cluster of
spikelets, bearing male or female spikelets only,
or bearing female spikelets above male and bi-
sexual spikelets. Spikelets solitary, usually uni-
sexual, dorsally compressed, disarticulation be-

m style 1; caryopsis fusiform; male
spikelets 2-9-flowered, the paleas 2-keeled; fe-
male spikelets 3-flowered, the lowest floret ster-
ile, the middle floret fertile, the upper floret ru-
dimentary and borne on a prominent rachilla
segment; palea of the functional female floret
spongy, curved, (5-)7-13-nerved, longer than the
lemma

When describing the Steyermarkochloeae and
including it in

primarily on anatomical characters. At the same

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

time, they recognized its uniqueness in the
subfamily on the basis of the gross morphology
of the leaves, inflorescence, spikelets, and flow-
ers. Much of that detailed discussion is appli-
cable to Arundoclaytonia as we

The discovery of “normal” leaves in the Stey-
ermarkochloeae lessens the importance of the
unique leaves of Steyermarkochloa vis-a-vis the
other genera of the Arundinoideae. In leaf mor-
phology we now consider such leaves to be basic
in the Steyermarkochloeae and thus well within
the norm of the Arundinoideae, and we consider
the leaves of Steyermarkochloa to be a later spe-
cialization that evolved in its own lineage.

n contrast, the 1 flowers of
Arundoclaytonia are deri ore special-
ized than those of mU Bisexual
flowers are universally considered more primi-
tive than unisexual flowers among grasses. Their
Occurrence, along with the more specialized uni-
sexual flowers, in Steyermarkochloa and their
absence in A loclaytonia indicates that the lat-
ter is more specialized in this respect. Similarly,
we consider the occurrence of bisexual and uni-
sexual spikelets in one inflorescence in Steyer-
markochloa to be less specialized than the seg-
regation of the spikelets into strictly male and
female inflorescences in Arundoclaytonia. This
strict separation of male and female flowers may
be considered the ultimate step in the Charles-

orth & Charlesworth (1978) model of the evo-
lution of monoecism through a gynomonoecious
pathway discussed by Davidse & Ellis (1984) for
Steyermarkochloa.

Only in one respect of spikelet morphology
might Arundoclaytonia be considered less spe-
cialized than Steyermarkochloa, and this is in
the number of florets of the male spikelets. The
occurrence of two sterile florets in the female
spikelets of both genera suggests a reduction in
number of fertile florets. This interpretation is
supported by the occurrence of bisexual spikelets
in Steyermarkochloa with two bisexual florets
and others with a male floret below the bisexual
floret. Against this background, the larger num-
ber of florets in the male spikelets of Arundo-
claytonia might be less specialized. This inter-
pretation must be tempered with the fact that
floret number may easily change up or down, as
is evident in the Arundinoideae, Pooideae,
Chloridoideae, and Bambusoideae in general, al-
though reduction seems to be prevalent in the

—

S

ily.
The number of florets in the Arundinoideae

1987]

varies from one to many, but the predominant
trend and probably the primitive condition in

this trend well in the male spikelets. In the female
spikelets both apical and basal reduction are ev-
ident. In reduction of the lowest floret to a sterile
lemma, Steyermarkochloa and Arundoclaytonia
resemble the Panicoideae. However, the termi-
nal rudimentary floret plus the numerous florets
of the male spikelets of Arundoclaytonia suggest
that this similarity is convergent. The rather
rounded female spikelets of Arundoclaytonia and
Steyermarkochloa are presumably due to reduc-

morphology of these two genera is the reduction
of the number of spikelets per inflorescence in
Arundoclaytonia, but, at the same time, the ag-
gregation of the many small unisexual inflores-
cences into a large false panicle. This exactly par-
allels a trend in other tribes in the family, for
example, Saccharum vs. Hyparrhenia in the An-
dropogoneae (Clayton, 1969) and Panicum lig-
ulare Nees vs. P. rudgei Roem. & Schult. in the
Paniceae

Anatomically, the absence of abaxial stomata
and microhairs and the reduction of silica bodies
in Arundoclaytonia complicates the phylogenetic
interpretation of the anatomical structure of the
leaf blade as many of these features are generally

claytonia, therefore, does not exhibit the com
plete set

to assign grasses to a given subfamily, thus lim-
iting our ability to use these characters for de-
termining the affinities of this unusual grass.
However, by a process of elimination certain
possibilities can be discarded.

Arundoclaytonia does not possess arm or fu-
soid cells and, therefore, cannot be accommo-
T in the Bambusoideae. The Chloridoideae
is entirely Kranz with only one possible excep-
tion ee 1984), and the Panicoideae is pre-
dominantly C,. Non-Kranz members of the pan-
icoid group all have a semiradiate type of
mesophyll and do not have the compact, isodi-
ametric emloreschyma a Irunqoerd tona
hows
no anatomical resemblance with ps panicoid
grasses, and phylogenetic relationships with this
subfamily appear most unlikely. The type of
chlorenchyma found in Arundoclaytonia does,

DAVIDSE & ELLIS— ARUNDOCLAYTONIA

489

however, occur in many arundinoid grasses. Some
members of the Pooideae also have this type of
chlorenchyma, but the pooid grasses typically do
not have sinuous long cells or tall, vertical silica
bodies as does Arundoclaytonia. The leaf ana-
tomical evidence, although somewhat limited,
does suggest arundinoid affinities for Arundo-
claytonia, and its systematic position does ap-
pear to lie with the Arundinoideae.

The tribes ofthe Arundinoideae cannot be sep-
arated on anatomical criteria and the decision to
classify Arundoclaytonia in the Steyermarko-
chloeae is based on morphological s. 1.
mainly that of the nius This decision is nei-
ther confirmed nor refuted by the anatomical
evidence. The leaf anatomy of Steyermarkochloa
and Arundoclaytonia differs substantially and
both appear to have highly advanced and derived
leaf anatomy. Because of these great anatomical
differences, they undoubtedly cannot be accom-
modated in the same genus

Clayton & Renvoize (1986) considered the
Steyermarkochloeae to be a tribe in the Pani-
coideae, noting “an obvious resemblance to Hy-
menachne" in features that are not unique. They
furthermore believe Steyermarkochloa to fun-
damentally differ from C, panicoids only in the
lack of microhairs (Renvoize, in litt.). We believe
that the additional evidence presented by us for
Arundoclaytonia gives further support for our
classification of the tribe, although we certainl
recognize the isolated position of the Steyer-
markochloeae in the Arundinoideae, and rec-
ognize that we are adding one more relatively
discordant element to the traditional “dumping
ground" of the family. Except for the lacunae
and stellate cells in the leaves and the gross form
of the inflorescence, but not its branching pat-
tern, we find it difficult to observe any obvious
resemblances between Steyermarkochloa and
Hymenachne.

We do agree with Clayton & Renvoize (1986)
that embryoc would
new information for clarifying the duxi cul
sition of the tribe. Chromosome information
could also be potentially useful. Unfortunately
we were unable t
obtained from an herbarium specimen af Arun-
doclaytonia. In the case of Steyermarkochloa all
our meiotic cytological samples were too young.
Unlike the typical situation in grasses, inflores-
cences of Steyermarkochloa must apparently be
well exserted from the sheath before meiosis takes
place.

crarvnncec

490

The distribution of Steyermarkochloa in the
northern Amazon basin and Arundoclaytonia in
southcentral Amazonia suggests that other taxa

d, since

large areas of XEM Bir ie em botanicallv very
poorly known. Obviously, the area between the
ranges of these two genera would seem to be the
most promising in this respect. Any relatively
open vegetation on white-sand soils (campinas
or Amazonian caatingas) might harbor further
taxa ofthis tribe. These vegetation types are found
in greatest abundance in the drainages of the Rio
Negro and the Rio Branco (Eiten, 1978; Ander-
son, 1981).

LITERATURE CITED

AB'SÁBER, A. N. 1982. oo and paleo-
eco logy of Brazilian Amaro a. Pp. Al -39 i n G.
cal D

Tropics. Columbia 1. pde New York.
ANDERSON, A. 81. White-sand — of
Brazilian een Biotropica 13: 199-
ARBER, A. 1934. The Gramineae. The Du Press,
Cambridge

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

BARNARD, C. 1964. Form and structure. Pp. 47-72
in C. Barnard (aeon, Grasses and Grasslands.
Macmillan & Co., Ltd., New York.

CHARLESWORTH, B. & D. CHARLESWORTH. 1978.

ulation genetics of partial male sterility and ike
evolution of monoecy and dioecy. Heredity 41:

37-
CLAYTON, w. 969. A revision of the genus Hy-
parrhenia. yo Bull. Add. Ser. 2: 1-1

& S. A. RENVOIZE. 1986. Genera Graminum.

DAVIDSE, G. R. LLIS. 1984 [1985]. Steyer-
markochloa unifolia, a a new genus from Venezuela
and Colombia (Poaceae: Arundinoideae: Steyer-
markochloeae). Ann. Miescün Bot. Gard. 71: 994—

1012.
EITEN, G. 1978. Delimitation of the cerrado concept.
Vegetatio 36: 169-1
ErLis, R. P. 1984. Eragrostis walteri—a first record
of non-Kranz leaf anatomy in the sub-family
an :

cation of the Gra

Watson, L. & M. J. ee
Grass em The renee National Univ.,
Canberr:

SIPHOCAMPYLUS OSCITANS
(CAMPANULACEAE: LOBELIOIDEAE), A NEW NAME FOR
BURMEISTERA WEBERBAUERI FROM PERU!

BRUCE A. STEIN?

ABSTRACT

Burmeistera weberbaueri is transferred to the genus Siphocampylus under the new name Siphocam-
pylus oscitans. The species is noteworthy in being one of only three species of Siphocampylus known

to have a dilated anther-tube orifice.

As part of a reassessment of the Peruvian
members of Burmeistera (Stein, 1987), it has be-
come apparent that B. weberbaueri Zahlbr. ac-
tually belongs to the large Andean genus Sipho-
campylus. This paper makes the appropriate
generic transfer and proposes a new name to ac-
commodate this unusual species.

Generic delimitations in Campanulaceae
subfamily Lobelioideae have long been recog-
nized as problematic (Gleason, 1925; McVaugh,
1940). Traditional systems, such as those of Presl
(1836) and Wimmer (1943), rely heavily on fruit
morphology for classification at the tribal level.
In particular, capsular versus baccate fruit is a
fundamental character used to define and align
genera. Strict reliance on this dichotomy in the
classification of the subfamily has separated close
relatives, for example, capsular-fruited Sipho-
campylus and baccate-fruited Centropogon. Al-
though fruit type in conjunction with other fea-
tures can be reliable for clustering related ids
of species, probable convergence in fruit c
acters suggests caution in its application.

Among neotropical Lobelioideae the emphasis
on fruit type, and to a lesser degree on the pres-
ence or extent of a dorsal slit in the corolla
(another seemingly labile character), has yielded
genera of convenience. One of the most natural
groupings, however, appears to be the genus Bur-
meistera, which is characterized by baccate fruits,
oblong or linear seeds, mostly nonbracteolate
pedicels, entire corolla tubes, and distally open
and oblique anther tubes often with little or no
apical pubescence.

The distally open anther tube of Burmeistera
is one of its most distinctive features. In most
genera of Lobelioideae the three dorsal anthers
are longer than the two ventral ones and curve
downward at the apex, effectively closing the
mouth of the anther tube. This allows the inter-
nally released pollen to build up pressure as the
style and stigma elongate, pushing pistonlike
through the anther tube. The characteristic tuft
of stiff hairs at the tip of the ventral anthers func-
tions as a lever that opens the orifice slightly and
allows the pressurized pollen. to discharge. Pre-

traea by Brantjes (1983) and has been observed
in the neotropical genera Centropogon and Sipho-
campylus (Stein, in prep.). Since the dorsal an-
thers in Burmeistera do not curve downward
closing the anther tube, this type of regulated
pollen discharge does not occur. This difference
in pollen presentation probably explains the cor-
related feature of glabrous or only sparsely pilose
anther apices in Burmeistera sect. Imberbes F
Wimmer: apical hairs there have no function as
trip mechanisms. Whether the densely villous
tuft at the tip of the ventral anthers in Burmeis-
tera sect. Barbatae F. Wimmer has a functional
role in pollen discharge is not clear

Burmeistera weberbaueri was indludeti in this
genus by Zahlbruckner (1906) on the basis of
floral features alone, as he had no mature fruit.
This decision was probably based upon his ob-
servation of a naked and dilated anther-tube ori-
fice, the absence of bracteoles, and the somewhat

orter P. Low

! I am grateful to Carlos Reynal for locating the type specimen at MOL and to B. E. — for searching
B. Dan

for type material at B H. Nicolson kindly pro

II and Peter Goldblatt made useful suggestions on an earlier draft. Fieldwork i in Peru was densi by National
Science Foundation "in Dissertation Improvement Grant BSR84-13912 and by a Garden Club of America

Award in Tropical Bot

? Missouri Botanical peA and Washington University. P.O. Box 299, St. Louis, Missouri 63166-0299,
U.S.A

ANN. Missouri Bor. GARD. 74: 491—493. 1987.

492

*burmeisteroid" corolla morphology (a short,
straight tube with an abruptly ampliate throat
and falcate lobes). The strongly turbinate hypan-
thium visible in the Field Museum type photo-
graph suggests the mature fruit of this species to
be capsular rather than baccate. Furthermore,
the coriaceous and rugose texture of the leaves
characterizes many members of Siphocampylus
but is unknown in Burmeistera. An open anther-
tube orifice of the kind described by Zahlbruck-
ner is, however, extremely rare in Siphocampy-
lus.

OBSERVATIONS

In order to investigate further the generic
placement of this species, I visited the type lo-
cality in January 1987 to collect fresh material
and to ascertain the fruit type and anther-tube
morphology. This represents the first collection
since the type was collected by August Weber-
bauer 85 years before. The fruits turned out to
be capsular with half-superior ovaries. Careful
examination of fresh pedicels occasionally re-
vealed bracteoles, a feature often difficult to ob-
serve in dried material. The presence or absence
of these bracteoles was consistent within indi-
vidual inflorescences. The most interesting con-
firmation is of the dilated anther-tube orifice.
Zahlbruckner's description of this feature is true
for the species and not based on an artifact of
preservation as I had previously assumed.

TAXONOMIC TREATMENT

Siphocampylus oscitans B. A. Stein, nom. nov.
Burmeistera iier A. Zahlbr., Bot.
y 451. 1906. Non Siphocam-
pylus ee A. Zahlbr., Bot. Jahrb.
Syst. 37: 456. 1906. TYPE: Peru. Junin: Prov.
Tarma, mountains east of Huacapistana,
3,200 m, Jan. 1902, Weberbauer 2203 (lec-
totype here designated, W; isolectotype, G).
SYNTYPE: Peru. Junin: Prov. Tarma, moun-
tains east of Palca, 3,200-3,600 m, Feb.
1902, Weberbauer 2473 (MOL, photograph
of lost B syntype, F neg. 9074).

Erect many-branched shrubs to 1.5 m tall, gla-

clustered toward branch tips, alternate and spi-
rally arranged, rarely appearing subopposite or
pseudoverticillate, sessile; blades narrowly ovate

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

to lanceolate, (3—)5—9 cm long, (0.7-)1.5-4 cm
wide, apex attenuate, base rounded to obtuse;
margins densely and sharply callose-serrulate, 9—
13 teeth per cm; when fresh the lamina fleshy
and coriaceous with the tips recurving, the adax-
ial surface smooth and dark green, the abaxial
surface pale or occasionally tinged purple, drying
coriaceous with the adaxial surface nitid and ru-
gose, prominently impressed by the 6-8 acutely
ascending, almost straight secondary veins. In-
florescence subcorymbose and auxotelic, the
flowers solitary in congested upper leaf axils, the
rachis later elongating and producing normal fo-
liage leaves beyond the maturing flowers and
fruits; pedicels ascending to spreading, 4-8 cm
long, generally exceeding length of subtending
leaves, occasionally with sub-basal, subulate
bracteoles to 2 mm long. Flowers 35-40 mm
panthium widely obconic, the sides

erect or slightly spreading, triangular, 8-11 mm
long, 3—4 mm wide at base, about twice the length
of hypanthium, the margins entire or minutely
denticulate; corolla green and maroon, yellowish
within, corolla tube straight, 15-18 mm long dor-
sally and 10-15 mm long ventrally, ca. 6-8 mm
wide at base, narrowing and re-expanding slight-
ly above, the lobes lance-oblong with acuminate
slightly revolute tips, mostly abruptly decurved,
the dorsal lobes falcate, 14-17 mm long, ca. 4
mm wide at base, the lateral and ventral lobes
11-14 mm long; filament tube 27-30 mm long,
adnate to corolla ca. 4 mm above base, glabrous
and greenish, well exserted from corolla; anthers
connate, the upper 7-8.5 mm long, the lower 6.5-
7 mm long, glabrous except for sparse, wispy

g
capsule dehiscing by two terminal valves, 15 mm
long (including valves), 10 mm wide; seeds el-
lipsoid, minutely foveate, ca. 0.6 mm long.

Distribution. Known only from shrubland at
ca. 3,200 m on the eastern slope of the Andes
near Palca and Huacapistana in Junín Depart-
ment of central Peru.

Additional specimens examined. PERU. JUNIN: Vi-
toc, 3,600 m, Feb. 1984, Pearce s.n. (BM); Prov. Tar-
ma, mountains E of Palca, rd. to Illic, 10-12 km above
Culebreo, 11?19'S, 75?33'W, 3,250 m, 13 Jan. 1987,
Stein, Kallunki & Diaz 3831 (AAU, B, CAS, F, G, K,
MO, NY, P, US, USM

1987]

DISCUSSION

The presence of capsular fruit in this species
excludes it from Burmeistera as currently cir-
cumscribed. Among the capsular-fruited genera
it is best placed in Siphocampylus based on the
entire corolla tube. Since the name Siphocam-
pylus weberbaueri has already been used by Zahl-
bruckner (1906), I propose the new name Sipho-
campylus oscitans for this species. The epithet
oscitans derives from the Latin for yawning, in
reference to the dilated anther-tube orifice, an
unusual feature in Siphocampylus. Siphocam-
pylus oscitans is easily recognized among Peru-
vian members of this genus by its completely
glabrous, stiff, coriaceous, lanceolate leaves, rel-
atively short and stout corolla with strongly de-
flexed lobes, and erect to spreading sepals that
are about twice as long as the strongly obconic
hypanthium.

open anther tube of the kind found in Sipho-
campylus oscitans is known in only two other
members of the genus: S. sceptrum Decne. of
Venezuela and S. rusbyanus Britton of Bolivia
and southern Peru. While S. oscitans occurs
nearer geographically to S. Past partas, erana
logically it more phases
of the highly variable S. eena. In addition
to the open and thinly pubescent anther-tube ori-
fice, certain collections of S. sceptrum share the
following features with S. oscitans: a half-supe-
rior ovary and shallow hypanthium; thick, ses-
sile, and often lanceolate or narrowly ovate leaves

VIUSVI Y

pedicels; adnation of the filaments to the corolla
tube near its base; and a stout, straight corolla
tube. The two species differ most obviously in
the length of the corolla and in the extent of
fusion of the four upper lobes (the feature that
best characterizes S. sceptrum). On the basis of
these similarities, S. oscitans seems best placed
near S. sceptrum in sect. Brachysiphon F. Wim-

er.

Zahlbruckner (1906) cited two Weberbauer
collections (2203 and 2473) in the protologue of
Burmeistera weberbaueri. Gleason (1925) des-
ignated Weberbauer 2203 as the “type,” but it is
unclear whether this constitutes valid lectotypi-
fication. He gave no indication that the type spec-
imen was actually examined and may well have

STEIN — SIPHOCAMPYLUS OSCITANS

493

been following the then common practice of des-
ignating the first collection listed in the original
description. The type specimens studied by Zahl-
bruckner at Berlin were both apparently de-
stroyed during World War II (B. E. Leuenberger,
pers. comm.), and my search of that herbarium
confirms their absence. Because Zahlbruckner
worked at Vienna and annotated the W sheet of
Weberbauer 2203, I have designated that spec-
imen as the lectotype.

Note added in proof: Recent examination of
the type of Burmeistera splendens at BM shows
this to be a fourth species of Siphocampylus with
a dilated anther tube. This Colombian species
appears closely related to S. oscitans based on
anther tube, hypanthium, sepal, and foliage fea-
tures, and provides support for the above dis-
cussion linking S. oscitans of Peru with S. scep-
trum of Venezuela. I here make the transfer of
B. splendens to Siphocampylus, as has already
been suggested by Jeppesen's annotation on the
type specimen.

Siphocampylus splendens (F. Wimmer) J eppesen
ex B. A

Andes del Norte, July 1857, Tiana 3059/
23 (holotype, BM).

LITERATURE CITED
BRANTJES, N. B. MD 83. Regulated VR issue in
Isotoma, Campanulaceae, and evo n of sec-
onary pollen uit meer Acta Bot. eee 32:

1925. d gem on the flora of northern
s Burmeistera. Bull.

GusAsoN lg A.
h America IV:
Torrey Bot. es 52: 93- 104.
McVAuGH, R. 40. A revision of “Laurentia” and
allied du fe in North America. Bull. Torrey Bot.
Club 67: 778-798.
. Studies in South American Lobelioi-
deae (Campanulaceae) with special reference to
Colombian species. Brittonia 6: 450-493.
. Prodromus Monographiae Lobeli-

. 1987. Synopsis of the genus Burmeistera
in Peru. Ann. Missouri Bot. Gard. 74: 494—496.

WIMMER, F. . Campanulaceae-Lobelioideae.
Pflanzenreich IV. 276b: 1-260 (Heft 106)

ZAHLBRUCKNER, A. 06. Campanulaceae andinae.
Bot. Jahrb. Syst. 37: 451-463.

SYNOPSIS OF THE GENUS BURMEISTERA
(CAMPANULACEAE: LOBELIOIDEAE) IN PERU'

BRUCE A. STEIN?

ABSTRACT

The southern distributional limit of peA has long been obscured by the inclusion in that

genus of several disharmonious elements from cen

southernmost stations known for these two species, and thus the genus as a whole, are at approximately

5°50'S in San Martín Department of northern Peru

Burmeistera Triana is a well-delimited neo-
tropical genus of Lobelioideae distinguished from
other baccate-fruited genera by the combination
of oblong or linear seeds, oblique and distally
open anther tubes, and mostly nonbracteolate
pedicels. The typically distorted, greenish to ma-
roon corollas make the genus easily recognizable
in the field. However, complex morphological
patterns, in particular extreme local differentia-
tion, render it taxonomically challenging at the
species level.

Burmeistera is found primarily in montane wet
forests from Chiapas, Mexico south to Venezuela
and Peru, with its center of diversity in the Andes
of Colombia and Ecuador. The southern distri-
butional limits of the genus have been problem-
atic, owing to the questionable inclusion of sev-
eral central and southern Peruvian taxa. This
paper clarifies the status of Burmeistera in Peru
and establishes its currently known southern lim-

t

Six Peruvian species of Burmeistera were in-
cluded in the Flora of Peru (Wimmer, 1937) and
in Wimmer’s (1943) revision of the genus, the
most recent treatment available. These are: Bur-
meistera asteriscus F. Wimmer, B. macrocarpa
(A. Zahlbr.) F. Wimmer, B. peruviana F. Wim-
mer, B. ramosa F. Wimmer, B. tricolorata F.
Wimmer, and B. weberbaueri A. Zahlbr. Of these,
only B. ramosa is retained in Burmeistera as the
genus is presently circumscribed. McVaugh
(1949) correctly noted the position of B. macro-
carpa, B. peruviana, and B. asteriscus in Centro-
pogon and made the necessary transfers for the
last two. The two remaining species clearly be-
long in Siphocampylus and are discussed below.

An Pici species, Burmeistera micro-
phylla J. D. Smith, was recently collected in
northern ud bringing the number of Peruvian
species to two. This collection at approximately
5*50'S in San Martín Department, along with a
recent collection of B. ramosa from the same
general vicinity, represents the southernmost sta-
tions known for the genus

The erroneous inclusion of several southern
and central Peruvian taxa in Burmeistera has
obscured an i
The Huancabamba deflection of northern Peru
marks the distributional limit for a number of
plant and animal groups and appears to have
been a significant barrier to north-south migra-
tion (Simpson, 1975, 1979; Berry, 1982; Vuil-
leumier, 1984). The geographical range of Bur-
meistera as defined here provides another
example of the Huancabamba deflection com-
prising the boundary for a group. Although ap-
proximately 30 species are present in Ecuador
(Jeppesen, 1981), diversity in the genus falls
abruptly in northern Peru, where only two species
are known to occur. In Peru, these species have
been found only in the area along or near the Río
Maranon gap, the lowest elevation depression
along the eastern slope of the Central Andes and
a major component of the Huancabamba de-
flection.

Why Burmeistera is absent from apparently
suitable habitats further south along the eastern
slope of the Peruvian Andes remains a mystery.
The relatively low elevation of the Rio Maranon
gap (ca. 500 m) does not alone explain the pat-
tern, since B. ramosa and other species occur at
such elevations. Furthermore, members of the

! I thank James S. densa m Kallunki, and Porter P. Lowry II for helpful comments. Fieldwork in Peru

e Foun

Xuan n in Tropica

dation p iin Dissertation Improvement Grant BSR84-13912 and

ub any.
? Missouri Botanical Garden and Washington iM P.O. Box 299, St. Louis, Missouri 63166-0299,
A.

ANN. Missounmi Bor. GARD. 74: 494—496. 1987.

1987]

genus are capable of dispersal past such barriers,
as evidenced by their successful establishment in
areas across far more formidable low-elevation
gaps, in one instance reaching the isolated Cerro
de la Neblina massif of the Guayana Highland.
The complex patterns of local differentiation and
endemism in Burmeistera suggest that dispersal
may often be quite limited, even though the var-
iously spongy, juicy, or inflated fruits often ap-
pear well suited for bird-dispersal. In this regard
it may be significant that B. microphylla is one
of the widest-ranging members of the genus, ex-
tending from Costa Rica to northern Peru. An
additional factor may be our insufficient knowl-
edge of the middle and upper-elevation forests
along the eastern Andean slopes of northern Peru.
These inaccessible areas are among the most
poorly collected in the Andes.

KEY TO THE SPECIES OF BURMEISTERA IN PERU

la. Leaves thick and coriaceous when dry; sepals
5-8 mm long, ipd fruit we less
thon 10 mm in diamet icrophylla
i sepals 2-5 mm long, erect or spreading;
fruit inflated, to 25 mm in diameter .. B. ramosa

1. Burmeistera microphylla J. D. Smith, Bot.
Gaz. (London) 25: 146. 1898.

The single Peruvian collection, as well as col-
lections from southern Ecuador treated by Jep-
pesen (1981) under the synonym B. aurobarbata
F. Wimmer, differ from typical Central Ameri-
can B. microphylla in their more uniformly lan-
ceolate to narrowly lanceolate leaves and in lack-
ing golden external anther trichomes. Further
investigation of geographically intermediate
populations of this wide-ranging and variable
di complex (including B. crassifolia F. Wim-

and B. maculata F. Wimmer) may indicate
E these southernmost populations warrant
treatment as a distinct taxon.

Specimen lacius PERU. SAN MARTIN: Prov.
W of Rioja on road to Pedro Ruíz, 2
with Amazonas
Dept.), 2, na m, 16 Feb. 1985, Stein y Todzia 2198
(MO).

2. Burmeistera ramosa F. Wimmer, Repert. Spec.
Nov. Regni Veg. 30: 16. 1932
This species is now known from two collec-
tions in Peru. The type, Tessmann 4725, was
collected at a relatively low elevation at the mouth

STEIN—PERUVIAN BURMEISTERA

495

of the Rio Santiago along the Rio Marañon, not
“near Iquitos” as indicated by Wimmer (1937).
The second, a fruiting collection, was made in
1983 in the Venceremos region from mid-ele-
vation cloud forest. Burmeistera ramosa is ap-
parently more common in Ecuador (Jeppesen,
1981) and shows a similar wide range in eleva-
tional preference. Since the early botanical ex-
plorers Ruiz and Pavón never ventured into
northeastern Peru (Steele, 1964) where this
species would be expected, one of their collec-
tions, formerly thought to originate from Peru
(Wimmer, 1943), was probably among the Ecua-
dorean collections made by their apprentice J.
Tafalla.

Specimens examined. PERU. AMAZONAS: Rio Ma-
rañon, Pongo de Manseriche, 160 m, Tessmann 4725
(NY; photographs, MO, NY). SAN MARTIN: Rioja Prov.,
Pedro Ruíz-Moyobamba rd., km 390, Venceremos,
1,770-2,150 m, 5-7 Aug. 1983, Smith & Vasquez 4608
(MO).

EXCLUDED SPECIES

1. Burmeistera asteriscus F. Wimmer, Repert.
Spec. Nov. Regni Veg. 38: 5. 1935. = Cen-
tropogon peruvianus (F. Wimmer) Mc-
Vaugh, Brittonia 6: 462. 1949. See discus-
sion under Burmeistera peruviana.

2. Burmeistera macrocarpa (A. Zahlbr.) F.
Wimmer, Repert. Spec. Nov. Regni Veg. 30:
41. 1935. = Centropogon macrocarpus A.
Zahlbr., Bot. Jahrb. Syst. 37: 452. 1906.

3. Burmeistera peruviana F. Wimmer, Repert.
Spec. Nov. Regni Veg. 38: 5. 1935. = Cen-
tropogon peruvianus (F. Wimmer) Mc-
Vaugh, Brittonia 6: 462. 1949

Field studies conducted at the type locality of
Centropogon peruvianus and C. asteriscus aroun
Pillahuata, Cuzco Department, show that the dif-
ferences noted by Wimmer (1935) in describing
these two species, primarily leaf width and shape
and sepal length, are variable within populations.
The type specimens of these two species were
collected in the same general vicinity and merely
represent upper and lower elevation collections
(3,000-3,300 m and 2,200-2,400 m) of a single
species. The names were published simulta-
neously; however, since McVaugh (1949) select-
ed C. peruvianus as the type species for his Cen-
tropogon sect. Peruviani, C. peruvianus is the
preferred name.

496

4. Burmeistera tricolorata F. Wimmer, Repert.
Spec. Nov. Regni Veg. 30: 22. 1932. = Si-
phocampylus rusbyanus Britton, Bull. Tor-
rey Bot. Club 19: 372. 1892

A fruiting collection of this species (Stein 2505)
made recently at the type locality of B. tricolorata

The Peruvian
collections closely match Siphocampylus rusby-
anus, a species recognized previously only from
northern Bolivia.

5. Burmeistera weberbaueri A. Zahlbr., Bot.
Jahrb. Syst. 37: 451. 1906. c CS
oscitans B. A. Stein.

A new collection of this species (Stein 3831)
has shown the fruit to be capsular, excluding it
from Burmeistera. For a discussion of this species,
its transfer to Siphocampylus, and the necessary
proposal of a new name (the epithet weberbaueri
has been used previously in Siphocampylus), see
the accompanying paper ii. 1987
Bb weber-

. trico mprise Wimmer's
(1943) bsc aequilatae of Burmeistera sect.
imberbes F. Wimmer. Since both of these species
are here excluded from Burmeistera, that strictly
Peruvian subsection is no longer recognized.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

LITERATURE CITED

BERRY, P. 1982. The systematics and evolution of
dern sect. eter iii Ann. Missouri
t. Gard. 69:
pee S. um
11-170.
McVAUuGH, R. 1949. Studies in South American Lo-
belioidae (Campanulaceae) with special reference
4

rei EM In Fl. Ecuador 14:

s in the flora
of the high tropical Andes. EOS 1: 273-
294.

9. Quaternary biogeography of the high
s. regions of South America. Pp. 157-1

Siphocampylus oscitans (Cam-
panulaceae: Lobelioideae), a new name for Bur-
meistera prx rn from Peru. Ann. Missouri
Bot. Gard. 7 93.

VUILLEUMIER, E pn Zoogeography of Andean
birds: two major barriers; and speciation and tax-
onomy of the Diglossa caibonaria superspecies.
Natl. Geogr. Soc. Res. Rep. 16: 713-731.

WIMMER, F. Burmeistera: eine e
Pflanzengattung und ihre Arten. Repert. Spe
Regni Veg. 30: 1-52.

. 1937. Campanulaceae. /n Flora of Peru. Field

Mus. Nat. Hist., Bot. Ser. 13: 383-489.

; ; Campanu laceae-Lobelioideae. Pflan-

zenreich IV. 276b: 1-260 (Heft 106).

NEW SPECIES OF PASSIFLORA SUBGENUS PASSIFLORA
ROM ECUADOR

L. B. HoLM-NIELSEN? AND J. E. LAWESSON?

ABSTRACT

new species of Passiflora from Ecuador are described, viz. P. montana, P. palenquensis, P.

. and P. pergrandis.

A revision of Ecuadorean Passiflora subg. Pas-
siflora (subg. Granadilla sensu Killip) for the Flora
of Ecuador (Holm-Nielsen et al., in press) has
revealed four new species: P. montana, P. palen-
quensis, P. deltoifolia, and P. pergrandis, be-
longing to “series” Lobatae, Tiliaefoliae, Men-
ispermifoliae, and Laurifoliae, respectively. The
total number of Ecuadorean subg. Passiflora
species is 19. Killip (1938) divided the subgenus
into 15 series; although these are not validly pub-
lished, we are following Killip for convenience.
The “series” occurring in Ecuador are Digitatae,
Incarnatae, Kermesianae, a Qua-

rangulares, and those mentioned above

Passiflora palenquensis Holm-Nielsen & Lawes-
son, sp. nov. TYPE: Ecuador. Los Rios:
Quevedo—Sto. Domingo road, km 56, Rio
Palenque Science Center, 150-200 m, 6 Oct.
1979, C. H. Dodson, A. Gentry & G. Shupp
8854 (holotype, MO). Figure 1.

Liana ubique glabra. Caule striato et pins Stipulis

oblongo-lanceolatis, acutis, 1 x 0.4 cm, interdum de-
p Nem angulo- alatis, 6. 5-9 em ^ longis cum
3-4n ib r apice

extremo, pi glandes infra 2- 3 cm; Pedum basi-
fixis, late ovatis, ovatis, 14-19 x
cordatis, manifeste pinnatinervibus
aceis, margine integra. Inflorescentia saltem
pedicellis teretis a ess 3-4 cm

nfra flore

o 3 cm, ad b
xe ce lobi calycis oblongi, acuti, 2.5 x 1.3 cm, de-
isi et in Vb n mucronem; petalis oblongis, ob-
3 0.6-1 cm. Corona simplice, filamentis
dx NE in 2 seriebus ordinatis quae ad apicem ex-
tremam tubi calycis locatae sunt; series externa minute
liguliformis, apex filiformis, 4 mm longa; serie interna

crassisima, 4-angulata, 2 cm longa, apex filiformis, fas-
urea et alba; tubus calycis glaber infra co-

rme, 0.5 cm lon Jil
gynophori limini circumcinctae. Ovario gla

Liana, glabrous throughout. Stem terete, striate.
Stipules oblong-lanceolate, acute, 1 x Í
sometimes deciduous. Petioles angular-winged,
6.5-9 cm long, with 3-4 black, sessile urceolate
glands, 1 mm long, one pair at extreme apex,
other glands 2-3 cm below; blades basifixed,
broad-ovate to ovate, 14-19 x 11—20 cm, deeply
cordate, prominently pinnate-veined, lustrous,
coriaceous, the margin entire. Inflorescence ses-
sile, with at least 2 flowers collateral with the
tendril in the axil of the leaf, 5-6 cm diam.; ped-
icels terete, striate, 3—4 cm long; bracts verticil-
late, united halfway, inserted about 5 mm below
ower, oblong-ovate, mucronate, 5 x 2 cm. Hy-
panthium tube funnel-shaped, 3 cm long, 1 cm
wide at base, 2 cm at apex, outside lavender;
calyx lobes oblong, acute, 2.5 x 1.3 cm, ending
in a minute mucro; petals oblong, obtuse, 3 x
0.6-1 cm. Corona composed of 2 series situated
at the edge of the hypanthium; outer series of
short ligulate filaments, 4 mm long, with filiform
apices; inner series of very stout, 4-angled, 2 cm
long filaments with filiform apices, the inner fil-
aments with shifting purple and white, 4 mm
broad stripes; hypanthium smooth below coro-
na. Operculum situated 1 cm above hypanthium
floor, membranaceous, 0.5 cm long, split into
filiform segments. Trochlea present at the an-
rogynophore opposite the apex of the opercu-
lum. Limen surrounding lower 5 mm of andro-
ynophore. Gynophore 1 cm longer than the
androgynophore present. Ovary glabrous. Fruit

! We thank R. R. Haynes and H. Balslev for iwi the manuscript, K. Tind for making the drawings, and

N.-H. Andreasen, who supplied the Latin diagn

Fieldwork was su

the Danish Natural Science

pporte ed by
Research kao grant numbers 11-3038, 11- 3689. and 81-4024. This is deris ore No. 53 from the AAU

Ecuador Pro

? Botanical PM University of Aarhus, 68 Nordlandsvej, DK-8240 Risskov, Denmark.

? Charles Darwin Rese

ANN. Missouni Bor. GARD. 74: 497—504. 1987.

arch Station Galapagos, Casilla 5839, Guayaquil, Ecuador

498 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

A \ (€ B 2cm
FIGURE 1. Passiflora palenquensis Holm-Nielsen & Lawesson.— A. Flowering stem. — B. Longitudinal section
of flower.

1987]

ovoid, 7 x 4 cm, green. Seeds ovate, 5 x 3 mm,
finely reticulate.

Passiflora palenquensis is a member of *'se-
ries" Tiliaefoliae, which is characterized by united
bracts and entire leaves. Passiflora palenquensis
has a distinct gynophore 1 cm longer than the
androgynophore and differs from all other mem-
bers of Tiliaefoliae by the corona having only
two series of filaments. Other species of Tiliae-
foliae have at least three series. It is closely re-
lated to P. seemannii Griseb. which superficially
appears to have two series. However, the two
series of filaments are 1-1.2 cm and 2-2.5 cm
long and have several rows of tubercles inside.
ii iUi palenquensis, on the other hand, has

nly two series, these being 4 mm and 2 cm,
a Rebus The tubercles are absent. The new
species is also related to P. tiliaefolia L., which
has more numerous series of filaments and is
confined to high altitudes (2,000-3,000 m). Pas-
siflora palenquensis is restricted to low eleva-
tions.

Additional specimens examined. Coro
NARINO: railroad Tumaco-Diviso km 86, 27 n: 1952,
Castaneda 3326 (AAU). ECUADOR. LOS PI-
CHINCHA: Sto. Domingo-Quevedo rd., km 45, Pan s
Pilar-24 de Mayo rd., km 12, 600 m, Apr. 1980, Dod-
son & Gentry 10337 (MO).

Passiflora montana Holm-Nielsen & Lawesson,
sp.nov. TYPE: Ecuador. Carchi: Tulcan-Mal-
donado rd., km 67, 2,600 m, 78°04'W,
pss. 21 May 1973, L. Holm-Nielsen, S.

eppesen, B. Lejtnant & B. @llgaard 6200

ei ciuis AAU). Figure 2.

Liana, ubique glabra. Caule bp atque tereto. Stip-
ulis valde magnis, 4-4.5 x m, latioribus quam lon-
gioribus, reniformibus mu incen ad basin obtusis,
margi ne glandulare- “serrata, Petioli is 3. 5-4 cm longis,

medium. peor ag basifixis, 3-lobatis, interdum
re lobatis, 9-11 x ys 5-13 cm, dimidium oon , lobi
3 cm lati, lanceolati, acu Iti; pedato-venatis, integris cum

glandibus in uus Bracteae verticillatae, ane
portatae 5 cm infra florem, ovatae, acuminatae, ad
basin cordatae, integrae, 2.5-3.0 x cm viridibus.
Flores solitarii; pedicellis 4.5—6 cm longis, teretis. Flo-
res 5-6 cm lati; tubo calycis breve campanulato, 5-6 x
7-8 mm; lobis calycis oblongo-lanceolatis, Meise
1.5 x 0.5 em, d manifeste carinatis, c
arista 6 mm longa quae apicem excedit; dein oblon
gis, subacutis, pallido- aaa 1-1.5 x 0.3-0.5 c
eriebus; externae A se-

versus operculum. ENDE uni epum recur-

HOLM-NIELSEN & LAWESSON — NEW ECUADOREAN PASSIFLORA SUBG. PASSIFLORA

499

vatum, modice plicatum, 4 mm longum, dun d
superioris filiformiter fissum. Limen is n base
phori affixum, margine lobata. Ova pr bro. p

pallido-virides cum maculis lilacinis. Pom non visi.

Liana, glabrous throughout. Stem striate, te-
rete. Stipules very large, 4—4.5 x 2 cm, reniform,
mucronate, obtuse at base, margin glandular-ser-
rate. Petioles 3.5—4 cm long, terete, striate, with
two alternate, stipitate glands above middle, 4
mm long; blades basifixed, 3-lobed, occasionally
4-lobed, 9-11 x 11.5-13 cm, lobed halfway; lobes
3 cm wide, lanceolate, acute; pedate-veined, en-

acuminate, cordate at base, entire, 2.5-3 x

cm. Flowers 5-6 cm wide, solitary, lateral; ped-
icels nins cm long, terete; hypanthium short
campanulate, 5-6 x 7-8 mm; calyx lobes ob-
c sq green, 1.5 x 0.5 cm, obtuse,
prominently carinate, with a 6 mm long awn
exceeding apex; petals oblong, cr light green,
1-1.5 x .5cm. Coronal filaments of several
series; outer three series of filiform elements 12
mm long succeeded by about 5 irregular series,
of slender filiform elements, 6 mm long, lilac,
extending towards operculum. Operculum mem-
branaceous, recurved, slightly plicate, 4 mm long,

bu . Nectar ring present, conspicuous.
deta absent. Ovary glabrous; styles light green
with lilac spots. Fruits not seen.

Passiflora montana as member of "series"
Lobatae is most clearly related to P. sprucei Mast.,
from which it differs by not having the leaves
divided below the middle, by having the stipules
twice as large and having two (vs. three or four)
petiolar glands 4 mm long and not sessile, and,
especially, by the floral composition. In P. mon-

ana the operculum is recurved and has a filiform
upper half and the filaments are 3-4 mm long,
whereas the operculum in P. sprucei is erect with
the lower quarter membraneous and the upper

two) outer series of long filiform filaments suc-
ceeded by four or five inner series of 4-5 mm
(vs. 2-3 mm) long filaments. Passiflora montana
is the highest growing member of "series" Loba-
tae found in Ecuador at altitudes of 2,000-3,000
m; the related P. sprucei and P. resticulata Mast.
& André usually do not occur above 2,000 m.

500 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

2cm

FIGURE 2. Passiflora montana Holm-Nielsen & Lawesson.— A. Flowering stem.— B. Longitudinal section of
flower.

1987]

Additional specimens examined. ECUADOR. PI-
CHINCHA: Niebly, Pululagua, Sodiro s.n. (S).

T dicas: Holm-Nielsen & Lawes-

, Sp. nov. TYPE: Ecuador. Napo: Baeza-

Tena rd., DEB 1 800-1, 900 m, 4 Nov.

1980, G. Harling & L. Andersson 16216 (ho-
lotype, GB; isotype, AAU). Figure 3.

a admodum glabra superficie foliarum. Caule

striato, tereto. Stipules reniformibus, 1 x 2 cm, lobus
nf obtusus, superior lobus cuspidatus, mucron
onga, margine glandulare serataque. Petiolae
teretae, 3 cm longae, 4-glan , 1-2 mm gis, al-
ternis; lamina basifixa, cordata, integra, cir scrip
tione deltoidea, 8 x 9-10 , 5S-7-n a, coriacea
Flores solitarii lateralesque; SU. i 5-6 c
longis; brac infr t erticilla-
tae, liberae, ovatae, acuminateae, , cu
margine glandulare serrata. Flores 7-8 b

longis, consistunt, tubu secunda usque ad
operculum obtegentibus. Operculum membranaceum,
erectum, dense plicatum, 8-9 mm lon ngum, MT

spathulato-ligulata

3 mm infra operculum. Limen arcte = Sessea “==
gynophori, margine lobata. Ovario glabro. Fructu
ovoideo. Semina non matura.

Liana, stems sparsely puberulent, leaves pu-
bescent at lower and glabrous at upper surface.
Stem terete, striate. Stipules reniform, 1 x 2 cm,
the lower lobe obtuse, the upper lobe cuspidate
with a mucro 2 mm long, the margin glandular

and serrate. Petioles terete, 3 cm long, with 4
stipitate, alternate glands 1-2 mm long; blades
basifixed, cordate, entire, deltoid, 8 x 9-10 cm,
5—7-nerved, coriaceous. Flowers solitary, lateral,

7-8 cm wide; pedicels terete, 5-6 cm long; bracts
verticillate, ee inserted 1 cm below flower,
ovate, acum , 0.7 x 0.3 cm, margin glan-

dub am NM onim short-campanulate,
1 x 2em; calyx lobes and petals ovate to oblong,
violet, 3-4 x 1-2 cm; calyx lobes carinate with-
out an awn. Coronal filaments, violet, in 5-8
series; outer 2 series of ligulate, 2.5-3 cm long
filaments with filiform apices, the second series
ongest; innermost 3-6 series irregular, of spat-
ulate to ligulate or tuberculate filaments 1-5 mm
long, covering the interior of the hypanthium
from the second series to the operculum. Oper-
culum membranaceous, erect, densely plicate, 8—
9 mm long, the upper part split into spatulate or

HOLM-NIELSEN & LAWESSON — NEW ECUADOREAN PASSIFLORA SUBG. PASSIFLORA

501

ligulate segments. Nectar ring a horizontal ridge
3mm below operculum. Limen tightly surround-
ing base of gynophore, margin lobulate. Ovary
glabrous. Fruit ovoid, 5.5 x 2 cm

Passiflora deltoifolia 1s related to P. menisper-
mifolia Kunth and P. crassifolia Killip in “series”
Menispermifoliae. The new species differs from
these two species by having leaves as broad as
long and not longer than broad. Passiflora men-
ispermifolia has three-lobed leaves, whereas P.
deltoifolia and P. crassifolia have unlobed or in-
conspicuously iiis leaves. The operculum is
plicate and there are no awns at the calyx lobes

of P. deltoifolia. The bracts of P. deltoifolia are
smaller, 7 mm long and 4 mm broad (vs. 1-2
cm long and 5-8 mm broad) and not cuani hh
or long acuminate, as in P. menispermifolia and
P. crassifolia. In addition, the new species lacks
the dense indumentum characteristic for P.
menispermifolia and P. crassifolia.

Passiflora pergrandis Holm-Nielsen & Lawes-

of Cumbaraza, 900 m, 20 Apr
Harling & L. Andersson 13771 (holotype,
GB; isotype, AAU). Figure 4.

Liana, ubique glabra praeter inflorescentiam atque
varium. Caule

,l€ ase; laminibus psi fixis, anguste-ova-

tis, SE 15-20 x 9-10 cm, obtuso-truncatis,
crasso-coriaceis € — lateralis, ee
non-terminalis, usque e ad 30 cm longa, c pluribus

15-16 cm latis floribus aus amet ieee:
11 x 1-4.5 cm. Inflorescentia interdum D modo
paucis foribus pedicellis teretis, 3-9 cm longis, puber-
nfra florem portatae, erin lide,
cm, obscure lacin-
ato- lobatae i in parte distale, obtusae in base, margo
Ingracum p puberu-
lis. Tubo inis campanulato 1x2 cm; lobi calycis
aes bs 6 x 3.5—4 cm, obtusi, carinati cum aris-
ta folia onga, quae apicem non excedit.
Petalis lanceolatis ovatis, 5.5-6 x 2 cm, alba. Fila-

a t
3 cm, cruciatim eim colore
i um

albidocanum tomentosum. Fructus non visi

Liana, glabrous throughout except inflores-
cence and ovary. Stem terete, striate, older parts

502 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

D A Š 5cm

FIGURE 3. Passiflora deltoifolia Holm-Nielsen & Lawesson.—A. Flowering stem.— B. Longitudinal section
of flower.—C. Operculum, densely plicate.—D. Fruit.

1987] HOLM-NIELSEN & LAWESSON—NEW ECUADOREAN PASSIFLORA SUBG. PASSIFLORA 503

5cm

FiGurE 4. Passiflora pergrandis Holm-Nielsen & Lawesson.— A. Leaf and reduced inflorescence. — B. Old
stem. — C. Inflorescence with several flowers and bracts.— D. Longitudinal section of flower with recurved
operculum.

504

2 cm diam. Stipules reduced. Petioles 2-3 cm,
with one pair of ovate, sessile, black glands 3 x
l mm, | cm from base; blades basifixed, narrow-
ovate, acuminate, 15-20 x 9-10 cm, obtuse to
truncate at base, Laban pinnate-veined,
entire, thick-coriaceous. Laminar nectaries ab-
sent. A distal bud pe to a short shoot,
this forming a conspicuous indeterminate inflo-
rescence to 30 x 15-16 cm with several flowers
(or sometimes reduced with few flowers), the
lowermost flowers developed first, the flowers
subtended by 4-11 x 1-4.5 cm reduced leaves;
pedicels terete, 3-9 cm long; bracteoles verticil-
late, free, inserted 5 mm below flower, ovate to
oblanceolate, 5 x 4 cm, obscurely laciniate-lobed
at distal part, obtuse at base, margin entire with
several stout black glands. Hypanthium cam-
panulate, 1 x 2 cm; calyx lobes oblong-ovate,
6 x 3.5—4 cm, obtuse, carinate with a foliaceous
awn 4 mm long, not exceeding apex; petals nar-
row-ovate, 5.5-6 x 2 cm, white. Corona fila-
ments of 3 series; filaments of outer series mi-
nutely setaceous, 2 mm long; filaments of second
series stout, ligulate, 5 x 0.3 cm, cross-banded
with white and dark violet; third series close to
operculum, minutely tuberculate 1-2 mm. Oper-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

culum membranaceous, recurved, the margin
with short fimbriate filaments. Limen tightly ad-
herent to androgynophore between the promi-
nent nectar ring and trochlea. Ovary densely
whitish-gray tomentose. Mature fruits not seen.

Passiflora pergrandis is a member of "series"
Laurifoliae and is related to P. ambigua Hems-
ley, which is also present in Ecuador. Passiflora
pergrandis differs in several aspects from P. am-
bigua, these being much larger inflorescence and
longer pedicels, larger leaflike bracts, calyx lobes
and petals only twice as long as broad (vs. three
or four times longer), and much shorter outer
coronal filaments. The awn in P. pergrandis does
not exceed the apex of the calyx lobe as it does
in P. ambigua. Passiflora ambigua is known from
Central America.

LITERATURE CITED

n L. B., P. M. JORGENSEN & J. E. La-
ESSON. Passifloraceae. In Harling and Anders-

ñoraceae Publ. Field Mus. Nat. Hist., Bot. Ser
19: 1-613.

THE GENUS ATTALEA (PALMAE) IN PANAMA!

GREGORY C. DE NEVERS?

ABSTRACT

The two Panamanian species of Attalea (Palmae) are described and illustrated, A. iguadummat for

the first time.

Attalea (Palmae) is a poorly collected neotrop-
ical genus of about 25 species centered in Am-
azonian South America and reaching its north-
western limit in Panama. When Bailey (1943)
treated the palms of Panama the genus was not
known from the isthmus. Attalea was last revised
by Glassman (1977), at which time one species
was recorded in Panama. Recent fieldwork has
revealed an undescribed species of Attalea there.
A circumscription of the genus in Panama is pro-
vided, and the new plant is described and named
A. iguadummat.

Attalea Kunth, Nov. Gen. et Sp. 1: 309. 1816.
TYPE: A. amygdalina Kunth, Nov. Gen. et
Sp. 1: 310. 1816.

Arborescent or acaulescent, monoecious.
Leaves pinnate, the pinnae inequilateral at the
tip, clustered in groups or evenly distributed along
the rachis. Inflorescences either androgynous or
staminate, bearing a pair of bracts, the pedun-
cular bract large, woody, sulcate, terminating in
a long or short rostrum, enclosing the inflores-
cence in bud; staminate inflorescence branched
to one order, the rachillae with many flowers,
these disposed in 1 or 2 rows, or spirally arranged
in dyads. Staminate flowers with 3 short, tri-
angular sepals, and 3 valvate, lanceolate-apic-
ulate, flat petals; stamens 6-10 (in Panama),
shorter than the petals, the anthers straight, de-
hiscing longitudinally. Androgynous inflores-
cence with sessile or short pedicelled pistillate
flowers. Pistillate flowers 2-4 cm long, with 3
sepals and 3 petals, the sepals and petals imbri-
cate; stamens reduced to a prominent stamino-
dial ring; ovary ovate; stigmas 3, apical. Fruit
with exocarp thin, fibrous; mesocarp pulpy and

fibrous; endocarp thick, hard, without fibers.
Seeds 1-3, irregularly shaped.

Attalea is characterized by its large size, sep-
arate staminate and androgynous inflorescences,

ng, ru

Attalea is closely related to Scheelea, Orbignya,
and Maximiliana, together comprising the sub-
tribe Attaleinae, the genera of which are distin-
guished from each other by characters of the an-
droecium.

Attalea allenii is abundant throughout San Blas
and many plants produce fruits annually, yet in-
florescences at anthesis are rarely seen. During
two years of collecting in San Blas, the androgy-
nous inflorescence was seen only once and the
staminate inflorescence only three times. When
Glassman (1977) prepared his preliminary treat-
ment of Attalea, the androgynous inflorescence
was unknown. Androgynous inflorescences of A.
iguadummat (described below) are equally rare,
while the staminate inflorescences, in contrast,

The pollination syndrome of the genus is un-
known (Henderson, 1986). The staminate inflo-
rescences of both Panamanian species produce a
strong musky odor at anthesis.

KEY TO THE SPECIES OF ATTALEA

la. Pinnae evenly ip along the rachis; mid-
dle pinnae 6.5-7 ; staminate inflo-
rescence 25.155. cm long; rachillae 12.5-

! The first year of fieldwork was made possible by a Smithsonian Fellowship; the second year was conducted

under the a

uspices of a contract with the Asociación de Empleados K

una. Additional support was provided by

thi Missouri Botanical Garden through its Flora of Panama project. Bruce Allen translated the Latin diagnosis.

Hecho deis assisted with fieldwork.
ri Botanical Garden, P.O. Box

9. St. Louis, Missouri 63166, U.S.A. Current address: California

iss 29
A pe of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A.

ANN. MissouRi Bor. GARD. 74: 505-510. 1987.

506

18.2 cm long; flowers spirally arranged in
dyads; stamens 8-10 iguadummat
. Pinnae in groups of 3 - 5 along the rachis;
middle pinnae 2-4 cm wide; staminate inflo-
rescence 15-25 cm pee rachillae 1-4.5 cm
long; flowers two-ranked; stamens 6 ... A. allenii

c

Attalea allenii H. Moore, Gentes Herb. 8: 191.

949. TvPE: Panama. Colón: Puerto Pilón,

10 m, Allen 4103 (holotype, MO; isotype,
BH). Figure 1.

Stem solitary, short or subterranean. Leaves
12-15, about 5 m long; petiole 60-80 cm long,
2.5 cm diam. at base, 2 cm diam. at apex; rachis
3.3-3.7 m long; pinnae 85-87 per side, arranged
in groups of 3-5, linear, glabrous, inequilateral
at tip, the margins ferruginous-lepidote; middle
pinnae 75-95 cm long, 2-4 cm wide; apical 9—
10 pinnae regularly spaced, separated basally,
coherent at the apex, forming 2 broad lobes about
10 cm wide at the terminus of the rachis. Inflo-
rescences interfoliar, produced at ground level,
erect, either staminate or androgynous; bracts 2;
prophyll 30 cm long, 4 cm wide, the apex short
and rounded; peduncular bract 35 cm long with
a rostrum 7 cm long. Staminate inflorescence 1 5—
25 cm long; peduncle 8 cm long, 8-10 mm wide,
brown furfuraceous at anthesis; rachis 23-26 cm
eae demi about 16-28(—50), 1-4.5 cm long,

—8 staminate flowers in 2 ranks. Sta-
ee flowers with sepals 3, deltoid, 1 mm long;
petals 3, valvate, lanceolate-apiculate, glandular,
9-13 mm long, 2-3 mm wide; stamens 6, fila-
ments 2-3 mm long, the anthers straight, 4—6
mm long; pistillode minute. Androgynous inflo-
rescence unbranched; peduncle 10-24 cm long,
1-1.4 cm wide; rachis 10 cm long; pistillate flow-
ers disposed in triads with 2 staminate flowers.
Staminate flowers of triads 6-7 mm long, with
3 sepals, these connate for less than 0.5 mm at
base, triangular, 2 mm long; petals (5—)6, 4—5
mm long; stamens 6, erect, dorsifixed, the an-
thers 1-1.5 mm long; pistillode minute. Pistillate
flowers sessile, crowded on the rachis, 3.2 cm
long, enveloped at the base by 3 triangular bracts
2 cm long and 1.6 cm wide; sepals 3, broadly
imbricate, 2.7-2.9 cm long, 1.5-2 cm wide, ir-
regularly lobed at apex; petals 3, imbricate, 2.5-
2.7 cm long, 1.8-2 cm wide, irregularly lobed at
apex; staminodial ring 6-7 mm deep, minutely
6-lobed; ovary conical, 2.3-2.5 cm long, 1.2-1.4
cm wide at base, densely brown tomentose; stig-
mas 3, arching, 8-11 mm long; ovules 3, basal.
Infructescence with 7-24 fruits, these obovoid,

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

6-7.5 cm long, 3.5-4 cm wide, the exocarp thin,
fibrous, densely ferruginous-lepidote, appearing
smooth; mesocarp 1.5-2 mm thick, fibrous; en-
docarp 3-5 mm thick, bony. Seeds 1-3, irregu-
larly shaped, conforming to the shape of the cav-
ity in which they develop.

Additional specimens examined. PANA BOCAS
DEL TORO: Río San Pedro, Gordon 15c (MO). CANAL
AREA: Coco Solo, Gentry 6298 (MO). COLON: Nombre
de Diós, July 1911, Pittier 4237 (US); Santa Rita Ridge,
Croat 15308 (MO); Santa Rita Ridge, 300-500 m, Gen-
try 6556 (MO). PANAMÁ: El Llano-Cartí Rd., 12 Jan.
1981, Read et al. 81-57 (US). COMARCA DE SAN BLAS:
El Llano—Carti Road km 16.7, 350 m, 9?19'N, 78?55'W,
*igua" (Kuna), “mange” (Spanish), 4 Nov. 1984, de
Nevers et al. 4152 (MO, NY); El uon Road km
19, 10 Aug. 1984, de Nevers 3639 (MO, PMA); same
locality, 18 June 1986, de Nevers & Herrera 7954 (CAS,

MO); same locality, 11 Mar. 1986, de Nevers et al.
7301 (MO); Cangandi, 30-150 m, 9°24’N, 79*8'W, 29
Jan. 1985, de Nevers et al. 4735 (MO); same locality,
*igua kaa” or “‘igua sai la let" (Kuna), 10 Feb. 1986,
de Nevers & Herrera 7191 (MO); Río Tiwar (R. Acla),
8^48' N, 77°40'W, 25-100 m, Sugden 624 (MO); Playón

Hydro Camp 14, 200 m, Duke 11377 (BH); 2 km from
Las Animas on rd. to Quibdó, 5?4'N, 76?47'W, King
et al. 664 (BH, NY). VALLE: Buenaventura Bay, Agua-
dulce Island, Moore et al. 9468 (BH); isi dise dank
Moore et al. 9460 (BH); km 14 marker between Bue
ventura and Bajo Calima, below 50 m, 3°56'N, 76°59' W,
Croat 57552 (MO); Dindo area, Bajo Calima, 100 m,
3°59’N, 76°58'W, Gentry & Monslave 48429 (MO); Bajo
Calima, Gentry et al. 40395 (MO); Río Calima, La
Trojita, 5-50 m, 19 Feb. 1944, Cuatrecasas 16397 (US);
Bahía de Buenaventura, Quebrada de San Joaquin, 0-
10 m, 20 Feb. 1946, Cuatrecasas 19948 (US). BOLÍVAR:
Mun. Morales, cgfo. Norosi, camino a Tiquisionuevo,
130-200 m, Cuadros 2194 (MO).

Attalea allenii is well known from the original
description and many collections. It ranges from
Panama to Colombia. In Panama it occurs in
tropical wet forest (sensu Holdridge et al., 1971)
on the Atlantic slope. The Kuna name is **igua,"
the leaves are used medicinally, and the imma-
ture fruits are eaten.

Attalea iguadummat de Nevers, sp. nov. TYPE:
n

4 350
79?45'W, 24 Feb. 1986, de Nevers 7197 (ho-
lotype, CAS; isotypes, K, MO, PMA). Fig-
ure 2

Species nova A. victoriana Dugand similis sed flo-
ribus masculis spiratim depositis, staminibus 8-10, fi-
lamentis 4-5 mm longis, antherisque 2-3 mm longis

1987]

FIGURE 1.

Stem solitary, short or subterranean. Juvenile
leaves 1.5-2 m long, 20-26 cm wide, undivided,
obovate, the margins dentate with triangular teeth
1.5-2 cm long. Mature leaves 9-17, arching; pet-
iole 55-82 cm long, 5.7-7.2 cm wide at base,
broadly channeled adaxially; rachis 6.75—7.23 m

DE NEVERS— ATTALEA IN PANAMA 507

dalli T €

P a^
Lh A

Infructescence of Attalea allenii, de Nevers 4152.

long; pinnae 104-109 per side, evenly spaced,
linear, glabrous, inequilateral at tip, the margins
ferrugineous-lepidote; middle pinnae 144-148
cm long, 6.5-7 cm wide, with midvein raised
adaxially and abaxially; apical pinnae 1.6-1.9 cm
wide, 55-59 cm long, free. Inflorescences inter-

A

508

foliar, produced at ground level, erect, either sta-
minate or androgynous, both inflorescence types
produced on the same plant; bracts 2; prophyll
20-25 cm long, encircling base of peduncular
bracts; peduncular bract woody, prominently
vertically sulcate, enclosing inflorescence in bud,
splitting and opening flat at anthesis, erect, 115-
203 cm long, 30 cm wide, with prominent non-
splitting rostrum 12-15 cm long. Staminate in-
florescence 125-155 cm long; peduncle 90-110
cm long, 1.5 cm wide, covered in a n of fur-
furaceous brown tomentum, this soon eroded
away; rachis 35-45 cm long: rachillae 40—50,
subtended by an acute, triangular bract 1-1.5 cm
long, this striate when dry; rachillae with basal
sterile portion 2.5-4.2 cm long, fertile portion
10-14 cm long, with scattered groups of minute
scales, these bright white in dried specimens. Sta-
minate flowers spirally arranged in dyads, each
subtended by a minute bract 0.5-1 mm long.
Sepals 3, 1-1.5 mm long, striate when dry; petals
3, valvate, 1.3-1.7 cm long, 1 mm wide, flat-
tened, curved to recurved at the tip, sometimes
straight or “S” curved, striate when dry, but not
when fresh; stamens 8-10, about ' the length of
the petals; filaments 2-3 mm long, separate and
free; adjacent stamens occasionally with the fil-

ments connate at the base for 1 mm or the
filaments connate completely, divergent just be-
low the anthers; tip of filament attached 'A-'^
way up the anther, the thecae united above the
point of attachment, separate or occasionally
united slightly below it; anthers 2-3 mm long,
straight, dehiscing longitudinally; pistillode 1—2
mm long, 3-lobed. Androgynous inflorescence at
anthesis not known; peduncular bract as in sta-
minate inflorescence; peduncle 60-70 cm long,
the rachis 35-45 cm long. Pistillate flowers short-
pedicellate or sessile; sepals 3, 22-35 mm long,
15-18 mm wide at base, broadly triangular; pet-
als 3, 40-45 mm long, acute; styles 3, apical,
exserted from the petals; perianth persistent in
fruit, the sepals chartaceous; petals chartaceous,
margin thin, undulate; the staminodial ring en-
larged, 1.1—1.8 cm deep, with 15-20 lobes, these
2-3 mm deep, 2-4 mm wide, commonly bifid.
Infructescence with (3—)45—65 fruits, these 7—
10 x 4.5-6.3 cm, obovate, brown tomentose,
with the styles and stigmas persistent; exocarp
1-2 mm thick, tough, fibrous; mesocarp 3-5 mm
thick, fibrous; endocarp 2-8 mm thick, bony,
smooth, without fibers; seeds 1-3, irregularly
shaped; endosperm homogeneous.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

The specific epithet is derived from the Kuna
name for the plant, *igua dummat.” It honors
the Kuna Indians of San Blas, Panama, who have
initiated a self-managed forest reserve and wild-
life sanctuary on their tribal lands.

Additional specimens examined. |. PANAMA. COLÓN:
Santa Rita Ridge, 8 miles from Transisthmica Hwy.,
in primary forest along small stream, 420 m, 9?25'N,
79°40'W, 2 Feb. 1986, Hammel, McPherson & Merello
14387 (MO); same locality, km 22, in forest on ridges,
slopes and in draws, 17 Feb. 1986, Hammel, Mc-
Pherson & Roubik 14507 (MO); same locality, 9 Mar.
1968, pid 3405 (US); same PEE km 13.8, 13
May 1986, de Nevers 7734 (BH, MO, PMA). COMARCA
DE SAN BLAS: Río Taindi, 9°25’ N, 79°11'W, 5 Apr. 1986,
de Nevers & Herrera 7656 (F, MO, PMA).

Attalea iguadummat is distinctive in its acau-
lescent habit, broad leaflets evenly spaced along
the rachis, large inflorescence, and dyads of spi-
rally arranged staminate flowers with 8-10 sta-
mens. Among acaulescent species of Attalea with
pinnae evenly distributed along the rachis, A.
iguadummat is similar to A. victoriana Dugand
and A. nucifera Karsten. It differs from A. vic-
toriana in its spirally arranged staminate flowers
(vs. the staminate flowers disposed in 2 rows on
one side of the rachilla), shorter staminate rach-
illae (12.5-18.2 cm vs. 25 cm), fewer stamens
(8-10 vs. 12-15), and sessile (vs. pedicellate) pis-
tillate flowers. Attalea iguadummat is distinct
from A. nucifera in its longer middle pinnae (144—
148 vs. 93-131 cm) and its glabrous staminate
petals (vs. staminate petals reddish glandular).
Attalea iguadummat may be most closely related
to A. tessmannii Burret, the only other species
with staminate flowers spirally arranged in dyads.
Attalea tessmannii is an arborescent species from
Amazonian Peru which differs from A. iguadum-
mat in its branched androgynous inflorescence
with dispu saben HOMES: Attalea pias-
sabossu Bondar
but they are o. in a single row, not spirally
arranged, and it is arborescent.

Attalea iguadummat is known only from the
extremely wet Atlantic slope of the mountains
between Colón and the western border of San
Blas, Panama. This may be the wettest area in
Panama (Myers, 1969; Anonymous, 1975). Fur-
ther collecting may reveal additional localities in
the Atlantic lowlands to the west in the Provinces
of Coclé, Veraguas, or Bocas del Toro, or the
palm may be truly endemic to the wet ocean
slopes between Colón and the Mandinga River.

The Taindi River locality of A. iguadummat

509

DE NEVERS— ATTALEA IN PANAMA

1987]

AT Un
bu li o.

FIGURE 2. Inflorescence of Attalea iguadummat at anthesis, Hammel 14387, photo B. Hammel.

lenii, *dummat" means big. When asked where
the “big Attalea” could be found the informant
mentioned the Iguagandi River, a Taindi tribu-
tary. In Kuna “‘-gandi”’ signifies place of, imply-
ing the Iguagandi River is the “river of Attalea.”

was discovered via a clue from a Kuna Indian.
During an ethnobotanical survey in San Blas in-
formants were asked to name wild edible plants
that they harvest. One man mentioned “igua
dummat.” **Igua" is the Kuna name for A. al-

510

Kuna botanists state that the Mandinga River is
the eastern limit of A. iguadummat, which has
not been found further east in San Blas during
two years of intensive palm collecting there. Im-
mature fruits of A. iguadummat are eaten oc-
casionally by the Kuna.

LITERATURE CITED

ANONYMOUS. 1975. Atlas Nacional de Panama. In-
stituto rein Nacional “Tommy Guardia,”
Panamá, Panam

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

BAILEY, L. H. 1943. Palmaceae. In Flora of Panama.
nn. Missouri Bot. Gard. 30: 327-396.
GLASSMAN, S. F. 1977. Preliminary taxonomic stud-
ies in the palm genus Attalea H.B.K. Fieldiana,
Bot. 38: 31-61.
HENDERSON, A. 1986. A review of pollinanenaty sayas
in the Palmae. Bot. Rev. (Lancaster) 52: 22
aes pic E ^ W. C. GRENKE, W. H. HATHEWAY,
& J. A. Tost, JR. 1971. Forest Envi-
ai in Tropical Life Zones. Pergamon Press,
rk.

w Yo
Myers, C. W. 1969. The ecological geography of cloud
forest in Panama. Amer. Mus. Novit. 2396: 1-52.

NOVELTIES IN MESOAMERICAN MISTLETOES
(LORANTHACEAE AND VISCACEAE)!

Jos KUIJT?

ABSTRACT

ive new species of Loranthaceae (Cladocolea primaria, Psittacanthus angustifolius, P. minor, P.

species of Viscaceae (Dendrophthora davidsei, D.

(including the var. wurdackii (Rizzini) Kuijt) is proposed for what has usually been called P. calyculatus
n in Mesoamerica. The latter name is now restricted to a different species endemic to

Mexico

The following novelties result from recent
studies in connection with the Flora de Nicara-
gua and Flora Mesoamericana.

1. Cladocolea primaria Kuijt, sp. nov. TYPE:
Panama. Panamá: Cerro Jefe, 2 km along
road to Altos de Pacora from junction with
road to peak, low cloud forest, 800 m, Syts-
ma & Knapp 4797 (holotype, MO; isotype,
LEA). Figures 1, 2.

Plantae glabrae, pauce ramosae; rami quasi teretes,

insignis, petioli ad 15 mm Io
bisexuales. I
triadis 3- vel 4-paribus, supra pari singulo, tunc pari

uno mon m ebracteolatarum et denique flore ter-
minali sequentibus; triadae basales ad axillas a -
gatae; inflorescentiae paribus nonnullis foliorum
squamiformium, c m, m s e. Flo-
res 4-partiti; petala dimorpha, 2- e onga; an-
therae perpar viora filamentis
brevissimis insertae ad ee ovarium 1.5 x 1 mm;

stylus rectus, stigma capitatum. Fructus 6 x 4 mm,
ruber, obscure violascens, ellipsoideus.

Plants sparsely branched, twining, glabrous.
Stems terete or slightly 4-ridged, often with con-
spicuous lenticels when older, straight and rather

apex mostly blunt or slightly apiculate; midrib
conspicuous and running into apex; petioles stout,
to 15 mm long. Inflorescences subtended by sev-
eral pairs of thick, brown scale leaves, solitary
in leaf axils, determinate, with 3 or 4 pairs of

triads below and a pair above, followed by a pair
of ebracteolate monads and a terminal flower,

morphic, 2-2.5 mm
sessile on the shorter petals and with very short
filaments on the longer ones; ovary 1.5 x 1 mm;
style more or less straight, the capitate stigma
reaching the petal tips. Fruit 6 x 4 mm, red,
becoming dark purple, elliptic in outline; calyc-
ulus inconspicuous; embryo dicotylous, slender,
the haustorial pole scarcely expanded.

Additional specimens examined. PANAMA. PANAMA:

Cerro Jefe, Clusia forest near radio tower, 900 m, D'Arcy
& Hamilton 14817 (LEA, MO); in forest near road to
Cerro Jefe near junction with road to Altos de Pacora,
Mori & Kallunki 72763 (LEA, MO); Cerro Jefe, 6.6
mi. above Goofy Lake, disturbed cloud forest, 850-
900 m, Sytsma et al. 2839 (LEA, MO).

Ait.

Cladocolea r ] 7 id bl

ficulties in generic assignment. When I mono-
graphed Cladocolea (Kuijt, 1975), I proposed the
notion that Struthanthus is polyphyletic, at least
many species being derived from a number of
independent sources within Cladocolea. Thus I
spoke of connecting bridges, these in some cases
characterized by species pairs, one member of
which was placed in Cladocolea, the other in
Struthanthus. With some very minor exceptions,
this left Cladocolea as a genus with determinate
spikes of monads, the flowers 4-, 5-, or 6-partite,
and either bisexual or the species dioecious. Stru-

panamensis, which has bisexual flowers a

i ive di pcs are Sie to Dr. Karel U. Kramer, Zürich, for the Latin descriptions, and to the Natural

Sciences and Engineering Res

arch Council of Canada for continuing financial suppo

? Department of Biological E. University of Lethbridge, Lethbridge, Alberta TIK 3M4, Canada.

ANN. Missounmi Bor. GARD. 74: 511—532. 1987.

512

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FIGURE 1. Cladocolea primaria Kuijt (Sytsma & Knapp 4797). Pendent branch.

bracteolate monads), its inflorescence made up
of mostly triads. The inflorescence of Struthan-
thus is generally indeterminate, but in a few
species it also bears two or four monads at the
tip, followed by a truly terminal flower.
Cladocolea primaria presents a predicament
from which there is no completely satisfactory
escape, for placement in either genus leads to the
need of significant modification of that generic
concept. As a species of Struthanthus it would
be the first four-partite species, and the second
one with bisexual flowers. If placed in Clado-

colea, however, it is the first species which is truly
triadic. I feel that placement in C/adocolea is
more acceptable, although I cannot deny an ele-
ment of arbitrariness in this regard. I continue
to hold the view that union of the two genera
would tend to obscure the complex relationships
between them, and that these difficulties concern
very few species. In fact, if the evolutionary re-
lationships are as I have proposed, such difficult
intermediate species would be expected.
Viewed in the above context, C. primaria is
closely related to C. /enticellata (Diels) Kuijt

1987]

1 ñ

KUIJT— MESOAMERICAN MISTLETOES 513

YW

b. Flower dissection.—c. Ma

ture

yt (S & Knapp 4797).—a. Inflorescence, the upper triads removed. —
fruit. —d. Embryo.

514

(which it greatly resembles superficially) and C.
roraimensis (Steyerm.) Kuijt, while in Struthan-
thus it is especially S. leptostachyus (Kunth) G.
Don and S. polystachyus (Ruiz & Pavon) Blume
that are related. Our species represents the sec-
ond Cladocolea reported for Panama and seems
to be limited to the Cerro Jefe area.

N

š ph ney page nios Kuijt, sp. nov. TYPE:
Costa Rica. ón: Cordillera de Talaman-
ca, Atlantic ue unnamed cordillera be-
tween the Río Terbi and the Rio Siní, 2,400-
2,750 m, elfin forest edge, Davidse et al.
28990 (holotype, MO). Figure 3.

Planta ad circ. 15 cm alta, monoeca, erecta, olivacea,
parvifoliata; cataphylla basalia nulla vel irregularia, ap-
pendices basales in plano medio. Folia (ob)lanceolata,
succulenta, ad 9 x 2 mm, acuta. Flores feminei ad
partes superiores internodium fertilium, plerumque

wes d “N 5 cm longae; internodia fertilia 2 vel

3, raro 4; flores uniseriati, ad 20 pro bractea eta
Fru ee albus, cllipsoideus ad .5 mm diam.,
petalis patentibus

Plants monoecious, to ca. 15 cm high, erect,
olive green, small-leaved; basal cataphylls absent
throughout in some individuals, irregularly pres-
ent in others; basal appendages oriented in me-
dian plane; young parts often with sparse, stiff,
white bristlelike hairs. Leaves (ob)lanceolate,
succulent, to 9 x 2 mm, acute. Spikes solitary
and axillary as well as terminal, to 5 cm long
including the peduncle ca. 7 mm long; fertile
internodes 2 or 3, rarely 4. Flowers uniseriate,
to 20 per fertile bract; female flowers in the upper
part of the fertile internodes, not generally out-
numbering the male flowers below. Fruit white,
ellipsoid to globose, 1.5 mm diam.; petals
spreading

Additional specimens examined. COSTA RICA.
LIMON: Cordillera de Talamanca, Atlantic slope, Valle

m, Davidse et al. 28755a (MO). HEREDIA: open road

side,
Lent 3826

Dendrophthora davidsei is the sixth known
ies

currently in this paper. The others are D. gua-
temalensis Standley, D. mexicana Kuijt, D.
squamigera (Benth.) Kuntze, and D. terminalis
Kuijt. No serious confusion is possible with those

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

species. The closest relative of D. davidsei, how-
ever, is D. paucifolia (Rusby) Kuijt, which ranges
from Bolivia to Venezuela. The major difference
between D. davidsei and D. paucifolia is that si
vegetative branches of the latter invariably
a prominent pair of basal cataphylls placed high
above the leaf axil (see Fig. 39 in Kuijt, 1986a).
This is not seen in any of the ten individuals
present in the two Davidse collections, but basal
cataphylls are present on some (not all) vegeta-
tive laterals of the Lent specimen. The latter looks
extremely slender but appears to belong to the
present species. A second and apparently con-
sistent difference is that leaf size in D. paucifolia
indles upwardly until the uppermost lateral
te are subtended by leaf scales; even though
perhaps a slight diminution takes place in D.
davidsei, none of the leaves on the main stem
ever reach scale size. The relationship of the two
species is certainly very close, but separation ap-
pears justified.

Terminal inflorescences were not seen in the
type but are represented in the otherwise iden-
tical collection Davidse et al. 287 55a. In the type,

s shown in Figure 3a, the tips of the larger
auctio have remained static, giving the
impression of buds with crowded small leaves.
Such shoot tips undoubtedly will expand into
new systems of inflorescences, and perhaps mark
a seasonal change. In fact, in the third collection,
Lent 3826, there are some branches where such
a new expansion seems to have taken place, de-
marcated from the older portion by several pairs
of very small leaves.

3. r. aC Kuijt, sp. nov.
TYPE: Costa a. Limón: Cordillera de Ta-
lamanca, od slope, Valle de Silencio,
along the Rio Terbi, 0.5-1.5 airline km W
of the Costa Rican-Panamanian border,
2,300-2,400 m, Davidse et al. 28755b (ho-
lotype, MO). Figure 4.

Planta tenuis, monoeca, erecta, ramosissima, circ.

30 cm alta, ad basem squamata; caules inflorescentias

gerentes haud ultra 1 mm crassi. Cataphylla basalia

ad dispersi et femineis multo pauciores; bacca alba,
m diam., ovoidea, sepalis clausis.

Plants monoecious, ca. 30 cm high, profusely

branched, slender, erect, squamate to the base.
Inflorescence-bearing stems | mm or less in

1987] KUIJT — MESOAMERICAN MISTLETOES 515

FIGURE 3. Dendrophthora davidsei Kuijt.—a. Habit (Davidse et al. 28990). —b. Same collection, old inflo-
rescence. — c. Terminal portion of compound inflorescence (Davidse et al. 28755a).

516

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FIGURE 4. Dendrophthora talamancana Kuijt (Davidse et al. 28755b). —a. Habit. — b. Fruiting inflorescence.

thickness; basal cataphylls absent throughout, the
basal appendages in a median orientation. Spikes
solitary, axillary only, to 15 mm long, the single
fertile internode slightly longer than the pedun-
cle, the apex acute. Flowers uniseriate, to 6 flow-
ers above each fertile bract; male flowers irreg-

ularly distributed and much outnumbered by the
female flowers. Berry white, 2 mm diam., ovoid;
sepals closed.

This distinctive plant represents the third
completely squamate Mesoamerican species of
Dendrophthora, the others being D. squamigera

1987]

(Benth.) Kuntze and D. terminalis Kuijt. It differs
from the former in its extremely fine, much-
branched stems and short fertile internodes, and
from the latter in its size, blunt scale leaves, and
lack of terminal spikes. It is conceivable that the
unusually slender Friedrichsthal s.n. specimen
from Guatemala, as cited in Kuijt (1961), will
turn out to be this species. I know of no similar
South American plants.

A

. Phoradendron fasciculatum Kuijt, sp. nov.
TYPE: Panama. Chiriquí: Jaramillo Arriba,
near Boquete, trail to Rio Palo Alto, 1,100
m, near paved road, hyperparasitic on Pho-
radendron undulatum Eichl., in turn para-
sitic on Psidium guajava, Churchill & Kuijt
5106 (holotype, MO; isotypes, BM, CR,
EAP, MEXU, NY, LEA, PMA). Figure 5.

Plantae erectae, glabrae, basi rami 4-6 e pulvino
communi orientes; internodia vetustiora bicarinata,

novella praecipue statu sicco leviter quadrangularia,
ad 10 cm longa; rami laterales paribus cataphyllarum
singulis. Rami solum novelli foliosi. Folia carnosa, 8 x
5 mm, apice rotundata, mox decidua. Monoeca; spicae
paribus cataphyllorum sterilium vel nullis; internodia
fertilia 5 vel 6. Flores masculi 1-3 ad apicem areae
floriferae supra bracteam fertilem, feminei usque ad
12 pro bractea et iis suppositae, bi- vel triseriati. Fruc-
tus n ovoideus, 3 mm diam.; petalis inconspicuis,
claus

Plants monoecious, erect, glabrous, fascicled
from the base with 4—6 stems from a common
cushion. Stems usually lacking basal cataphylls;
internodes 2-keeled when older, somewhat
quadrangular when young, especially when dry,
to 10 cm long, stout; lateral branches with 1 pair
of basal cataphylls ca. 4 mm above base, spread-
ing when dry. Leaves fleshy, soon deciduous, 8 x
5 mm, the apex rounded, sides parallel, the base
clasping. Spikes with 1 pair of sterile cataphylls
or without, 5—6 fertile internodes sometimes pro-
liferating terminally into a second series of youn-
ger fertile internodes. Male flowers 1—3 at the tip
of the flower area above each fertile bract, the
female flowers to 12 per bract below them, bi-
or triseriate. Fruit broadly ovoid, 3 mm diam.,
the petals very small, inconspicuous, more or less
closed.

This distinctive species undoubtedly belongs
to the P. dipterum group of species, in whic
hyperparasitism is the rule. Phoradendron fas-
ciculatum is no exception, in that all plants seen
of the type collection were growing on P. un-
dulatum Eichler. It is impossible to tell at this

KUIJT—MESOAMERICAN MISTLETOES

517

time whether this cluster of species is obligately
hyperparasitic; plants growing near the base of a
primary host may easily be mistaken for being
parasitic directly on the host tree. Another fea-
ture apparently held in common by these various
species is that several stems originate from a bas-
al cushion, as illustrated in Figure 5a (arrow). I
add a comparable illustration of a small plant of
Phoradendron dipterum Eichler from Nicaragua
(Fig. 6), which happens to be parasitic on a leaf,
the host again being a Phoradendron. A clear
basal cushion is visible (arrow). A third example
is Phoradendron aequatoris Urban from Ecuador
(Kuijt, 1986a, fig. 2), which also has basal
sprouting and is parasitic on a Phoradendron.
That this is not axillary branching from the nodes
ofa much shortened base is demonstrated by the
usual lack of basal cataphylls, in contrast to what
occurs elsewhere in the plant.

The explanation of the basal cushion almost
certainly lies in the original haustorial disk of the
seedling. It has recently been shown in an un-
related species of Viscaceae, Viscum minimum
Harvey, that the margin of the haustorial disk
regularly produces aerial shoots (Kuijt, 1986b).

he same is true for Viscum album L. (Tubeuf,
1923), and for Ixocactus hutchisonii Kuijt of
Loranthaceae (Kuijt, 1987). In Phoradendron, this
feature would appear to have some taxonomic
constancy in the group of species under discus-
sion.

Additional specimen examined. COLOMBIA.
ANTIOQUÍA: highway between Uramita and Canasgor-
das, on Phoradendron piperoides (H.B.K.) Trel., Bark-
ley & Gutierrez 535457 (US

5. Phoradendron molinae Kuijt, sp. nov. TYPE:
Nicaragua. Madriz: cut over cloud forest area
on Volcán Somoto, 10 km S of Somoto,
1,400 m, Williams & Molina 20270 (holo-
type, US; isotype, F). Figure 7.

Internodia compressa carinataque, ad 6 cm longa;
rami laterales

; pedunculus statu fructifero circ. 3
s; spica fructifera circ. 3 cm longa, interno-
diis fertilibus 2 vel 3, floribus pro bractea sions tribus,
uctus ovo-

ideus, laevis, 3 x 2 mm, petala clausa.

Plants dioecious (only the female seen), stems
with compressed, keeled internodes to 6 cm long,
basal cataphylls one very low pair on lateral

518 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FiGURE 5. Phoradendron fasciculatum Kuijt (Churchill & Kuijt 5106).—a. Base of young plant, ns
sprouting from basal cushion (arrow).— b. Habit of older plant.—c. Older inflorescence, the basal portion
fruit, the terminal portion proliferated and in flower

1987]

KUIJT—MESOAMERICAN MISTLETOES

519

FiGuRE 6. Phoradendron dipterum Eichler parasitic on Phoradendron sp., Nicaragua (Stevens & Montiel
17931, LEA). The hyperparasite is sprouting from a basal cushion (arrow) attached to a leaf of the primary host.

branches. Leaves to 10 x 4.5 cm; blade thin,
more or less palmately veined, ovate, the base
abruptly contracted into conspicuous, cuneiform
petiole to nearly 1.5 cm long. Female inflores-
cence solitary, less than 2 cm long, often with a
sterile pair of cataphylls; peduncle ca. 3 mm long
in fruit; fruiting spike ca. 3 cm long, with 2-3
fertile internodes and 3 flowers per fertile bract
just above the middle of the internode. Fruit
ovoid, smooth, 3 x 2 mm; petals closed.

6. Phoradendron nitens Kuijt, sp. nov. TYP
Costa Rica. Cartago: east side of sip

divide between Tres Rios and Cartago, on
Euphorbiaceae, Kuijt 2465 (holotype, CR;
isotype, UBC). Figure 19 in Kuijt (1964) and
Figure 5 in Kuijt (19862).

Planta magna, m osa, ramificatione saepe
furcata; innovationes gelu cataphyllis basalibus

Ramuli novelli aliquantum compressi,
que a

m dispositi; spica ad 4 cm
longa, cataphyllis nullis, pipi m fertilia 3 vel 4, flo-

520 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FiGure 7. Phoradendron molinae Kuijt (Williams & Molina 20270). —a. Habit. — b. Fruiting inflorescence. —

c. Basal cataphylls.

res pro bractea 6-9, biseriati. — flavus, 3 x 1.5
mm, ellipsoideus; petala claus.

Plants fleshy, monoecious, the branching
mostly forked, where percurrent perhaps with a
pair of intercalary cataphylls. Lateral shoots with

a large pair of basal cataphylls just above the
axil; stems somewhat compressed when young,
becoming terete. Leaves to 15 x 7 cm or larger,
thick and shiny when fresh, rigid, obovate to
nearly elliptical; base tapering or abruptly con-

1987]

tracted; petiole very stout and flat, to 6 mm wide.
Spike to 4 cm long, lacking cape fertile
internodes 3-4, with flowers 6-9 p act, bi-
seriate; male flowers very rare, iu in series.
Fruit yellow, 3 x 1.5 mm, ellipsoid; petals closed.

Additional specimens examined. NICARAGUA.

o desde “Santa Julia" A "La Él

95 1 m, es 680 (HNMN see jos spec-

imens listed under ee a. (Presl) Ei-
chler in Kuijt, 1964).

This species, which in Mesoamerica has been
erroneously referred to Phoradendron obliquum
(Presl) Eichler, is often extremely difficult to sep-
arate from vigorous specimens of P. robustissi-
mum Eichler if only herbarium material is avail-
able. In the fresh condition, its usually larger

robustissimum is also strictly dioecious, but the
rarity of male flowers in P. nitens can be very
misleading. Phoradendron obliquum is presently
placed in synonymy under Dendrophthora obli-
qua (Presl) Wiens (Kuijt, 1986a).

The species is now known from Panama, Costa
Rica (Kuijt, 1964), and Nicaragua, and has been
reported recently from Ecuador (Kuijt, 1986a, as
Phoradendron #5, fig. 5). We may thus antici-
pate that it will be found in Colombia.

N

. Phoradendron tardispicum Kuijt, sp. nov.
TYPE: Panama. Chiriquí: bridge over Río San
Felix, on Panama Hwy., 50 m, on Ficus along
river directly S of bridge, Churchill & Kuijt
5107 (holotype, MO; isotypes, BM, LEA,
MEXU, PMA). Figure 8

Plantae dioeca eae solum visae), vivae o
scure virides, glabrae. Caules deinde bs üternódiá
recta, 5-9 cm longa. "Ca taphylla basal

defoliata aliquot pro nodo, spica quaque par
bus cataphyllorum sterilium basalium internodiisque
fertilibus circ. ie flores biseriati, ad 10 pro e fer-
tili. Fructus ovoideus, luteo-viridis, 3 mm longus, pe-
talis clausis

KUIJT — MESOAMERICAN MISTLETOES

521

Plants dioecious (only the female seen), bright,
S m di

slightly flattened when young, soon becoming
terete; internodes straight, 5-9 cm long. Basal
cataphylls mostly 1 or 2 pairs, very inconspic-
uous, if 1 pair present nearly axillary, if 2 pairs
present the second pair to 20 mm above axil,
rarely to 4 pairs. Intercalary cataphylls present,
one pair between successive pairs of foliage leaves
but irregularly distributed along the branch, ab-
sent from some internodes, always inconspic-
uous. Leaves amplexicaul, cordate, shiny when
fresh, to 7.5 x 6 cm, rather thin; margin undu-
late; venation pinnate but obscure. Inflores-

2 pairs of sterile basal cataphylls less than 5 mm
above base, followed by ca. 7 fertile internodes.
Flowers biseriate, to 10 per fertile bract, yellow-
ish green, each fertile internode with stalk and
flowerless tip 2-4 mm long. Fruit ovoid, yellow-
ish green, 3 mm long, the petals closed.

This is a remarkable species for its irregularly
distributed intercalary cataphylls and for late de-
velopment of inflorescences. It is difficult to see
what known species might be related. As far as
I am aware, only in Phoradendron paradoxum
Urban from Venezuela do intercalary cataphylls
alternate in occurrence, but there this pattern
seems to be regular (Trelease, 1916). That species
and the closely related P. fendlerianum Eichler,
however, have triseriate flowers and long-peti-

ut there 0-3 cataphylls may be present on an
“internode”; nor does this species seem to be
closely related to P. tardispicum.

8. Phoradendron zelayanum Kuijt, sp. nov. TYPE:
Nicaragua. Zelaya: N of abandoned airstrip
near Alamikamba, along tributary of Caño
Alamikamba, 10 m, gallery forest among sa-
vanna, on Symphonia globulifera L.f., Ste-
vens 21717 (holotype, MO; isotypes,
HNMN, LEA). Figure 9.

nta monoeca, dichotoma, apice abortivo. Caules
menus innovationes laterales cataphyllis € positis
binis. Folia late ovata vel orbicularia, la 8 x
8 cm, palmato-venosa; petiolus validus, planus, supra
expansus

522 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ficure 8. Phoradendron tardispicum Kuijt (Churchill & Kuijt 5107).—a. Habit. — b. Inflorescence.

[VoL. 74

1987]

KUIJT—MESOAMERICAN MISTLETOES

523

TS
cm

FIGURE 9. Phoradendron zelayanum Kuijt (Stevens 21717).—a. Habit.—b. Inflorescence.

ternodia fertilia tria; flores 13-15 pro bractea fertili,
biseriati; spica 4 cm longa; spicae hermaphroditae.

Plants monoecious, forking, the apex aborting,
the inflorescences and young shoots dull golden-
yellow. Stems terete, stout; internodes to 8 cm
long; the lateral shoots with one low pair of cata-
phylls. Leaves broadly ovate to orbicular; blade
to 8 x 8 cm, with 5 or 7 very conspicuous pal-
mate veins running far towards the apex; petiole
stout, ca. 8 mm long, flat and expanding distally.
Inflorescences bisexual, lacking sterile cata-
phylls, the peduncle 3 mm long, this followed by
3 fertile internodes each with 13-15 biseriate
flowers per fertile bract, the entire spike 4 cm
long

9. Psittacanthus eo Kuijt, sp. nov.
TYPE: Nicaragua. Ma .5 km al S de San
José de Cusmapa, X m, parasitando en
un Pinus, Moreno 24419 (holotype, MO;
isotype, HNMN, LEA). Figure 10.

Caules acute quadrangulares. Folia bina, anguste fal-

cata, tenua, ven natione pinnata; lami

sistens. Pedunculi triaderum circ
bracteis foliaceis ad 2 cm

cm longa, antherae 6 mm longae. Stigma antheras su-
perans, capitatum.

524 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ficure 10. Psittacanthus angustifolius (Moreno 24419).—a. Habit.—b. Tip of petal.

[Vor. 74

1987]

Stems sharply quadrangular. Leaves paired,
narrowly falcate, thin; the blade to 17 x 2.5 cm,
venation pinnate, the base acute, the apex slen-
derly attenuate; petiole to 5 mm long. Inflores-
cences terminal, consisting of 4 or 6 triads; triad
peduncles ca. 1 cm long, the lowest ones with
foliaceous bracts to 2 cm long; pedicels 1.5 cm
long, with conspicuous terminal cupule, the ca-
lyculus smooth. Bud stout, more or less straight
or somewhat curved; base and tip 5 and 4 mm
wide, respectively, the latter blunt. Petals orange,
7.5—8 cm long; inner part of flower hairless, the
petal base 5 mm wide, without ligule; petal apices
4 mm wide, blunt, each with a fleshy, ligule-like
median crest extending inwards. Stamens di-
morphic; filaments attached at ca. 2.5 cm above
petal base, 5 cm long; anthers 6 mm long. Ovary
5 x 6.5 mm. Stigma placed above anthers, very
small, capitate. Fruit unknown.

Additional specimen examined. Same data as type,
Soza et al. 155 (HNMN, MO).

Thi cican t dicti ti , known

from what is essentially a single collection.
Whether the species is restricted to Pinus, as ap-
parently is Psittacanthus pinicola, can only be
shown by further fieldwork. The two species can-
not be confused, as P. pinicola is dyadic and has
irregularly whorled, blunt, more leathery leaves.
The type locality suggests that the species may
well be present in neighboring Honduras.

10. Psittacanthus minor Kuijt, sp. nov. TYPE:
Nicaragua. Matagalpa: SW slopes of Cerro
El Pilón and adjacent Laguna Tecomapa,
roadside, low thorn scrub and pastures on
rocky slopes, on ant acacia, Stevens 9466
(holotype, MO; isotypes, HNMN, LEA).
Figure 11

Plantae parva, PE teretibus, foliis binis. Folia
ad 5 cm, ovata,

gae, series duae vix
longa; stylus petalis fere aequilongus; stigma aegre dis-
tinguendum.

Stems terete, phyllotaxy paired. Leaves thin,
the blade to x 3.5 cm, ovate, the apex and
base obtuse or nearly so; venation more or less
palmate; petiole 3-5 mm long. Inflorescences ter-
minal, consisting of 4—6 pairs of triads on pe-
duncles ca. 12 mm; pedicels 10 mm long. Buds

KUIJT—MESOAMERICAN MISTLETOES

525

straight, not dilated at base. Petals ca. 3.7 cm
long, red-orange; ligular area weakly differen-
tiated. Stamens dimorphic; the anthers 3 mm
long, dorsifixed, the 2 series dpi overlap-
ping, the filaments attached ca. 21 mm above
the base, some 16 mm long. mel more or less
cylindrical, 4 x 2 mm; style nearly as long as the
petals; stigma weakly differentiated; calyculus
smooth. Fruits ovoid, 1.5 x 1 cm, with con-
spicuous calyculus, blackish.

Additional specimens examined. NICARAGUA. MA-
TAGALPA: Puertas Viejas, 2 km al N sobre la Carretera
Panamericana, “San Vicente," 600 m, on Acacia, Mor-
din 18288 (HNMN, LEA, MO); same, San Juanillo, 8

al SE de Ciudad Dario, 500 m,
pos 2618 (HNMN, LEA
de Carretera, quebrada, 460-480 m
Moreno 1669.
ino o Dari TiO- Presa Santa

Dario, on Leguminosae, Grijalva 2693 (HNMN, LEA,
MO).

Psittacanthus minor is closely related to P.
mayanus Standley & Steyerm., which appears to
be limited to the Yucatán region, has quadran-
gular stems, and fruits which are about half as
large as those of P. minor. Psittacanthus ma-
yanus is much smaller in general.

11. Psittacanthus pinicola Kuijt, sp. nov. TYPE:
Belize. Belize: Manatee Pine Ridge, on pine,
1931-32, Gentle 82 (holotype, GH; isotype,
MO). Figures 12, 13.

Caules plus minusve teretes; folia symmetrica, ter
verticillata, ad 11 x 2.5 cm, anguste elliptica vel lan-
ceolata, apice rotundata, basi in petiolum circ. 5 mm
longum angustata. Inflorescentiae laterales, ad nodos,
umbellulas e gas 2 vel 3 formantes. Petala circ. 4
cm longa, rubra, apice luteo- viridescentia, medio au-

uantum curvatum, ad latitudinem
circ. 5 mm dilatatum, ad apicem tenuissimum , leviter
curvatum, circ. 1.5 mm latum angustatum; beep

circ.
laevis. Stamina dimorpha; antherae dorsifixa
paulo pilosae. Ovarium plus minusve merida 4.5
mm longum; stylus longus, basi aliquantum torsus;
stigma ellipsoideum, subtiliter papillosum. Fructus el-
lipsoideus, calyculo inconspicuo, 13 x 5 mm, saturate
purpureus.

Stems more or less terete, becoming coarsely
fissured and blackish when old; leaves symmet-
rical, in (often somewhat irregular) whorls of 3,
to 11 x 2.5 cm, narrowly elliptical to lanceolate;
apex rounded; base tapering into petiole ca. 5
mm long. Inflorescences lateral, axillary, often
also on older, leafless stems, each being an umbel
of 2 or 3 dyads; inflorescence peduncle to 13 mm
long; dyad peduncles and floral pedicels 5-7 mm

526 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FiGurE 11. Psittacanthus minor Kuijt (Stevens 9466). —a. Habit, with immature inflorescence. — b. Mature
ud.— c. Base of petal. —d. Fruit (Moreno 18288).

FiGure 12. Psittacanthus pinicola Kuijt.—a. Habit with immature inflorescences (Pipoly 4013).—b. Same
collection, portion of inflorescence with mature buds, a single petal shown separately.—c. Complete inflorescence
just after anthesis (Gentle 82).—d. Mature fruit (Stevens 7600).

527

FiGURE 13. Psittacanthus pinicola (Gentle 82).—a. Leafy innovation.— b. Mature flower, the style and anther
shown separately to the right.—c. Relative positions of anthers and style.

528

1987]

long, the latter scarcely expanded at the tip. Pet-
als ca. 4 cm long, red with yellow-green tip, or-
ange in the middle, prominently ligulate at the
base. Buds somewhat curved above, inflated at
the ovary to a width of nearly 5 mm, tapering
to a very slender, slightly curved tip ca. 1.5 mm
wide. Stamens dimorphic

nearly trimorphic (Fig. 130); filaments of the
longer type attached 1.5 cm above the base, 1.5
cm long; anthers 3-4 mm long, dorsifixed,
sparsely pubescent on back. Ovary more or less
cylindrical, green, 4.5 mm long, 2 mm diam.
below, expanding slightly above; calyculus
smooth. Style 4.6 cm long more or less straight,
but the base somewhat twisted; stigma ellipsoid,
finely papillate. Fruit ellipsoid, with inconspic-
uous calyculus, 13 x 5 mm, “deep purple."

Additional specimens examined. BELIZE. BELIZE:
western highway, Mile 30, beside track, on pine, White-
a 2442 ies same, The de on pine, Whitefoord
562 (MO). DISTR. UNKNOWN: P. Cayo, in roadside
SA on ridge area a overlooking l 000 ft. falls, in area
ountain Pine Ridge area, Huston

ZELAYA: Rio Troncera at junc-

gallery RT in savanna, on Pinus caribea, Pipoly 4013

(HNMN, LEA, MO); near Tala Has and Puente Mango
(over 1 0—60 m, pine savanna, on Pinus
caribea, Stevens 7600 (LEA, MO); Comarca del Cabo,

Kornuk Creek above Puente Pozo Azul, old bridge,
Robbins 5831 (LEA, MO). NUEVA SEGOVIA: El Jicaro,
7 km sobre la carretera a Murra entrada al Quebracho,

00-620 m, on pine, Moreno 8305 (HNMN, LEA,
MO).

Other dyadic species north of Panama are P.
sonorae (Watson) Kuijt, P. palmeri (Watson)
Barlow & Wiens, P. nudus (Molina) Kuyt &
Feuer, and P. ramiflorus (DC.) G. Don. Psitta-
canthus pinicola is similar to the last species but
seems more closely related to P. dichrous (Mar-
tius) Martius (see Eichler, 1868, especially Pl. 5).
Not only inflorescence structure and general ap-
pearance indicate this affinity, but even the pe-
culiar curvature of the stylar base is seen in both.

12. Psittacanthus rhynchanthus (Bentham)
Kuijt, comb. nov. Loranthus rhynchanthus
Bentham, Bot. Voy. Sulphur 102-103. 1845.
TYPE: “Dr. Sinclair," Tiger Island (Hondu-
ras, Bay of Fonseca) (K). Figure 14d-f.

Psittacanthus chrismarii Urban, Bot. Jahrb. 24: 331.
1897. TvPE (here designated): Costa Rica. Foréts
de Nicoya, Tonduz 13706 Fir dn. US; iso-
lectotypes, CR, GH).

Psittacanthus calyculatus auct., non (DC.) G. Don, Gen.

KUIJT—MESOAMERICAN MISTLETOES

529

. 1834. TYPE: Mexico. Cuernavaca:
UE i 150 (G-DO).

An attractive Psittacanthus in which the buds
are distinctively curved and beaked occurs
throughout Mesoamerica, at low elevations from
southern Mexico to Venezuela. In the past, this
species has been called P. calyculatus (DC.) G.
Don or, earlier, P. chrismarii Urban. After study-
ing the types of both Loranthus rhynchanthus
Bentham and L. calyculatus DC., I conclude that
these are two distinct species. Consequently, the
name P. rhynchanthus must be applied to the
wide-ranging species mentioned above. True P.
calyculatus seems limited to Mexico, the type
originating from the area of Cuernavaca, further
collections having been seen from Puebla and

orelia.

Notwithstanding their general similarity, the
two species may be consistently separated mostly
on the basis of floral features. The mature, un-
opened bud of Psittacanthus calyculatus is very
nearly straight and has a rather blunt tip; that of
P. rhynchanthus shows a distinctive curvature in
the distal portion, the apex being sharply acute,
and more beaklike. Psittacanthus rhynchanthus
has smooth pollen sacs behind which are borne
long, conspicuous, reddish stamen hairs; the pol-
len sacs of P. calyculatus are distinctly lobed, and
stamen hairs are lacking. Furthermore, the stylar
base in P. rhynchanthus bears low protuberances,
and each adjacent petal base shows a ligule con-
sisting of a low, V-shaped ridge; the stylar base
in P. calyculatus is smooth, and ligules are ab-
sent. Leaves of P. calyculatus tend to be smaller
(to 8 x 4 cm), mostly less than twice as long as
wide, and approximately symmetrical, while
those of P. rhynchanthus are usually larger (to
12 x 4 cm), more than twice as long as wide,
and strikingly falcate.

In Venezuela, at least some individuals of the
latter species have extremely narrow leaves; these
plants belong to Psittacanthus rhynchanthus var.
wurdackii (Rizzini) Kuijt, comb. nov. (P. calyc-
ulatus (DC.) G. Don var. wurdackii Rizzini, Rod-
riguésia 41: 15. 1976). I have not yet encountered
the species from the Caribbean lowlands of Co-
lombia, but it would be surprising if it were not
present there.

13. Lione subtilis Kuijt, sp. nov. TYPE:
ds nama. Coclé: near continental divide along
is. road, 2.2 km beyond sawmill in
forest above El Copé, 900 m, Hammel 998

530 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

.
w— - — <...

ne
oe wget:

weit
ge

we — -—

RE 14. Psittacanthus calyculatus (DC.) G. Don (a—; Mexico, Puebla, slopes of Popocatepetl, Ugent et
i 9, WIS) and P. rhynchanthus (Bentham) Kuijt (d- á MSN. Managua, Cuatro Esquinas, Moreno 4400,
LEA). —a, d. Mature buds.— b, e. Petals.— c, f. Stylar ba

(holotype, MO; isotypes, LEA, PMA). Fig- lamina ae E ad late ovata, 4-12 cm longa,
ure 15. 2-6 ata, ap l

mm.
Plantae inconspicuae, subtiles; caulis gracilis, plus darum 2 vel constitutentes; pedunculus c
minusve teres. Folia tenua; venatio manifesta, pinnata; ad 8 mm longus, triadarum ad 5 mm, uterque 0.5 mm

.,
3
oa
nd
E
$
ue
TE
R
Ms
E d
H
H

FIGURE 15. Struthanthus subtilis Kuijt.—a. Leafy shoot (Croat 49004) and leaves (Croat 44584).— b. Inflo-
rescence (Folsom & Lantz 1894). —c. Same collection, petals and style, male flower. —d. Same collection, petals
and style, female flower.— e. Mature fruits (Hammel 998).

531

532

vel minus crassus; flores laterales paces pid 0.5
mm longis; ipsis bracterolaeque m e, caducae;
petala 2-2.5 mm longa; stigma cristis indistinct pa-
pillosis 6. Siete aurantiacus, subglobos

diam., calyculo inconspicuo; pedicelli Puch later-
aliu m ad 3 mm elo ongati.

Plants inconspicuous, rather delicate, branched.
Stems slender, more or less terete; stem roots
occasional, thin. Leaves thin, blade narrowly lan-
ceolate to broadly ovate, always with caudate tip,
2-6 x cm, the evident venation pinnate;
petiole ca. 3 x 1 mm. Inflorescences pale green,
solitary in leaf axils, subtended by 4 minute,
probably caducous bracts, consisting of a raceme
of 2 or 4 triads; inflorescence peduncle to 8 mm
long; triad peduncles to 5 mm, both 0.5 mm or
less thick. Lateral flowers on pedicels 0.5 mm
long at anthesis; bracts and bracteoles minute,
caducous. Petals 2-2.5 mm long. Stamens di-

orphic; upper portion of sterile stamens pa-
pillate; anthers 0.4 mm long; style 2 mm long,
the stigma with 6 indistinct papillate crests. Fruits
orange, nearly spherical, 5 mm diam.; calyculus
inconspicuous; pedicels of lateral fruits elongated
to 3 mm.

Additional specimens examined. PANAMA. COCLE:

near continental divide along lumbering road, 1.5 mi.

Copé, Croat 44584 (CR, LEA, MO); El Copé-

El Potrosa, Atlantic slope of Alto Calvario, 700-850

m, Folsom & Lantz 1894 (LEA, MO, PMA); along road
f à

and Coclesito (N of Pintada), 4 mi. N of Llano Grande,
600 m, Antonio 3575 (LEA, MO); trail from Cano
Blanco del Norte to continental divide N of El Copé,
on Hedyosmum, 400 m, Davidse & Hamilton 23654
(BM, LEA, MEXU, MO, PMA); El Copé, Atlantic side,
1,200 m, Antonio 1153 (LEA, MO); between conti-

2409 (EAP, LEA, MO); El Copé, along gravel road to

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

the right before sawmill, 800 m, Antonio 2207 (LEA,
; Alto Calvario cloud forest, 5.3 km
above El Copé, continental divide, above sawmill, 930
m, Antonio 3044 (LE , PMA); above El Potroso
sawmill at gare divide, 1,200-1, Ly m, d
1820 (LEA along
bora road un beris Agricola Alto P Piedra, aln
first major stream, 3 mi. from fork in road at school,
700 m, Croat 49004 (LEA, MO)

This species was previously listed as S. aff.
dichotrianthus Eichler (Kuijt, 1978). Struthan-
aL JS L . L í 4 ç L 1). J F 1

Blume) are indeed related to S. subtilis, as is S.
quercicola (Cham. & Schlecht.) Blume of western
Panama to Mexico, but the extreme slenderness
and small racemes of our plant, the leaf size, and
especially the consistently caudate to acuminate
leaf apex, leave little doubt that this is a distinct
species. Struthanthus subtilis appears to be en-
demic to the Coclé-Veraguas region.

LITERATURE CITED

EICHLER, A. W. 1868. ipid In C. F. P. Mar-
tius, Flora Brasiliensis 30 y 1
Kuur, Jos. 1961 revision of Pos NR (Lo-
ranthaceae). Wentia 6: 1-145.
964. A revision of qs Loranthaceae of Cos-
ta Rica. Bot. Jahrb. 83:
75. The genus C m (Loranthaceae).
J. Arnold Arbor. 56: 265-335.
. 1978. Co mmentary on the mistletoes of Pan-
ama. Ann. Missouri Bot. Gard. 736—763.
1986a. Viscaceae. Jn Flora of hon 24:

6b. Observations on establishment and
of Viscum minimum (Vis-
caceae). Acta Bot. 6.
987. Mi scellaneous mistletoe notes, 10-19.
Brittonia (in press
TRELEASE, 19
llinois Press Urba
TUBEUF, 23. Vus EE der Mistel. Olden-
bourg, Miei

; The, Genus Phoradendron. Univ.

A REVISION OF DILODENDRON (SAPINDACEAE)!

A. H. GENTRY? AND J. STEYERMARK?

ABSTRACT

Dilodendron of Brazil and adjacent regions and Dipterodendron of Central America and northern

discuss their relationships, present a key to the three species
Dilode

t
costaricense (Radlk.) Gentry & Steyerm. and D. elegans. (Radlk. ) Gentry & Steyerm.

Dilodendron Radlk., a monotypic genus of the
dry areas of subequatorial South America, is
closely related to Cupania L. and Matayba Aubl.,
from both of which it differs strikingly in having
bipinnate leaves. According to Radlkofer (1892-
1900, 1895) the other main differentiating fea-
tures of subtribe Cupanineae are that the petals
of Cupania and Matayba (as well as closely re-
lated monotypic Vouarana Aubl. have two
squamae, whereas those of Dilodendron lack
squamae. Two other monotypic genera, Brazil-
ian Scyphonychium Radlk. and Guianan Pen-
tascyphus Radlk., have an intermediate bifid or
emarginate petal scale. The final genus of sub-
tribe Cupanineae, Tripterodendron Radlk., like-
wise monotypic and restricted to Brazil, is unique
in having tripinnate leaves and the small sub-
cupular calyx and bisquamate petals of Matayba.

eneric limits in subtribe Cupanineae are gen-
erally not Geary defined, and C “paid and Ma-
tayba, the only significant deed the only
nonmonotypic genera) recognized by Radlkofer
(1892-1900), are notoriously difficult to tell apart.

When Radlkofer (1892-1900) published his
Flora Brasiliensis treatment, Dilodendron was
known from Brazil and Bolivia, and from a single
sterile collection from Costa Rica. Dilodendron
bipinnatum has also since been collected in Par-
aguay and disjunct in the dry part of the Rio
Urubamba Valley in Convencion Province of
Cuzco Department, Peru (Macbride, 1956), but
the Costa qepa collection was subsequently de-
termined not to be congeneric.

" Dipterodendron Radlk. is a small genus of three

ly reported from Costa
Rica, Panama, and northwestern Venezuela
(Radlkofer, 1933; Steyermark, 1952; Croat,
1976). Dipterodendron was described by Radl-

ndron

kofer (1914) on the basis of three Costa Rican
collections. He recognized two species separated
by rather tenuous differences: leaflets smooth and
drying bright green in D. costaricense Radlk. vs.
leaflets dark green and papillose and appressed
puberulous below in D. elegans (Radlk.) Radlk.
The sterile Oersted collection now recognized as
D. elegans had originally been described as a
variety of Dilodendron bipinnatum in the Flora
Brasiliensis by Radlkofer.

A third species, D. venezuelense Steyerm. was
described in 1952 from Merida State in north-
western Venezuela, representing the first report
of Dipterodendron for South America. Like that
of D. elegans before it, the type of D. venezuelense
was sterile. The Venezuelan plant was distin-
guished from D. elegans by larger, more coarsely
toothed leaflets 2.3—4.5 cm long and 0.6-1.5 cm
wide, and because of its geographic disjunction.
However, recent collections from Costa Rica also
have leaflets reaching 4 cm long and 1.5 cm wide.
Some Costa Rican collections have leaflets with
coarse teeth and others with fine teeth. Coarse
teeth appear to reflect juvenile state rather than
a consistent specific difference. Moreover, we
have recently closed the geographic gap by col-

cense and D. elegans. While the vegetative dif-
ferences— mainly a more strongly appressed-
puberulous leaflet undersurface in D. elegans—
might be inadequate to justify maintaining D.
costaricense as a species separate from D. ele-
gans, there are also previously unreported fruit

! We thank USAID (DAN-5542-G-SS-1086-00 from the Latin America and Caribbean Bureau Office of
2 oe Resources) and the Smithsonian Institution for support of the Peruvian fieldwork that led to this

revisio

2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

ANN. Missounmi Bor. GARD. 74: 533-538. 1987.

534

differences. Thus, we refer all Central American
and northern South American material of Dip-
terodendron to two rather than three very closely
related pas both of which range from Costa
Rica to northern Venezuela.

The dA between Dipterodendron of
northern South America and monotypic Dilo-
dendron of subequatorial South American dry
forests suggest that even more merging is in or-
der. This study was initiated when one of us
recently collected a bipinnate-leaved Sapinda-
ceae tree as a tree plot voucher at the Tambopata
Nature Reserve in geographically intermediate

of the genus. However, the equally striking veg-
etative similarity between the Peruvian plant and
Dilodendron was subsequently discovered in the
herbarium when an attempt was made to identify
it to species. This led to an examination of the
taxonomy ofthe entire complex, which has never
een menogrepheg except for a _tecopying of
Radlkofer 'searlie I
in his posthumous (1933) Pflanzenreich treat-
ment. It turns out that Dipterodendron was never
adequately differentiated from Dilodendron in the
first place. Radlkofer (1914), who had seen no
flowers, suggested that Dipterodendron is inter-
mediate between Dilodendron and Tripteroden-
dron but differentiated it only from the latter
(which has tripinnate leaves and a thick oily aril,
and lacks saponiferous cells in the embryo) rather
than from the former even though he had earlier
referred the first Dipterodendron collection to Di-
lodendron. Later (1933) he emphasized slight dif-
ferences in radicle position. Aristeguieta (1973)
questioned the validity of separating Diptero-
dendron from Dilodendron, noting that according
to the literature Dipterodendron usually lacks
petals and has the radicle on the margin of the
cotyledon, while Dilodendron has 3-5 petals and
the radicle descending down the middle of the
dorsal side of the cotyledons. He ultimately as-
signed the Venezuelan material to Dipteroden-
dron essentially on geographic grounds. Later,
Steyermark (in herb.) identified sterile collec-
tions from Bolivar State in eastern Venezuela as
Dilodendron bipinnatum, which would virtually
eliminate the geographic discontinuity.
Although Dipterodendron might be retained as
distinct on the basis of its apetalous flowers, ves-
tigial petals are sometimes present. Moreover, in
Dilodendron the very small petals are variable in

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

number, mostly 3-4 but sometimes making up
a full complement of 5, and sometimes vestigial
(Radlkofer, 1892-1900). In fact, male flowers can
lack petals altogether, just as in Dipterodendron.
Since number of petals and even their presence
or absence is variable in the single species D.
bipinnatum, there seems no compelling reason
to separate Dipterodendron from Dilodendron on
this basis. Th pl
in the same genus otherwise similar species s
differ in presence or absence of petals, e.g.,
Swartzia, Licania, or Combretum; moreover,
other like Alectryon and Mis-
chocarpus have both petaliferous and apetalous
species. Although interpretation of floral sexual-
ity in Sapindaceae from herbarium specimens is
very tricky, in the case of Dipterodendron loss of
petals (only in male flowers?) might reflect a shift
to full dioecy. We conclude that Dipterodendron
should be united with Dilodendron to reflect best
their extreme similarity. Indeed we are suspi-
cious that monotypic Tripterodendron, of which
we have examined only sterile material, might
also be congeneric with Dilodendron. At any rate,
some collections of Dilodendron have incom-
pletely tripinnate leaves or squamellate petals,
and the other distinguishing characters of disk
margin, aril consistency, and lack of

Z

cells in the embryo seem weak.

pi eceden t for incl uding

Dilodendron Radlk., Bees Math.-Phys.
l. Konig]. Bayer. Akad. Wiss. München 8:
355. 1878; TYPE: D. nete

Dipterodendron Radlk., Smithsonian Misc. Collect.
61(24): 5. 1914. TYPE: D. costaricense.

Medium to large dioecious trees. Leaves al-
ternate, bipinnate (rarely in part tripinnate), mul-
tifoliolate, the leaflets sessile or subsessile, serrate
or dentate. Inflorescence a narrow (often almost
subspiciform) panicle, usually borne clustered
near the end of a branch and flowering preco-
ciously or with the newly expanding leaves.
Flowers tiny, the sepals 5, the petals smaller than
sepals, sometimes absent, when present variable
in number and often in part vestigial, the stamens

in minute 2-3-lobed stigma. Capsule 2-3-lobed,
loculicidally (22)3-valved, the valves woody or
subwoody, the 1—2(-3?) seeds ellipsoid, with a
thin, shiny, dark brown testa, scarious aril (fide

1987]

Radlkofer), and basal hilum. Embryo (fide
Radlkofer) subcircinately curved, the thick car-
nose cotyledons saponiferous, the radicle dorsal.

KEY TO THE SPECIES OF DILODENDRON

la. Outer margin of leaflet teeth convex, the leaf-
midrib

often opposite pinnae; Brazil to southern
Peru l. D. bipinnatum
. Outer margin ofleaflet teeth dg the leaf-
lets not ciliate- -margine ed, the lower midrib
and lateral nerves s or strigillose with
a few appressed hairs on midrib nerves; flow-
ers apetalous; calyx lobes acute; leaves with
8-16, usually ri due pinnae; Costa Rica to
Venezuela and P
2a. Fruit trigonal-globose, at dehiscence the
or re-

c

the outer surface dull, puberu ous, and
the

tertiary venation visibly iig ue EN
2.

costaricense
2b. Fruit laterally compressed, at .
the valves splitting to the base and r

cellate, drying black and shiny, bed scat-
(CIC IMINULW visit n
the verruculose surface; leaflets densely
strigillose beneath, the tertiary venation
usually not evident 3. D. elegans

1. Dilodendron bipinnatum Radlk., Sitzungsber.
Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss
München. 8: 355. 1878. svNTYPES: Brazil.
Minas Gerais: St. Hilaire 1586, Martius s.n.,
Riedel 1090, Warming s.n. (US).

Tree 8-20 m tall, to 40 cm dbh, the branchlets
usually somewhat angled and/or longitudinally
ridged, puberulous with both long and short (in
part gland-tipped) hairs when young, becoming
glabrescent, lenticels absent or minute and in-
conspicuous. Leaves bipinnate with 5-10 subop-
posite or alternate pinnae, rarely the basal leaflets

foliolate, the usually alternate leaflets narrowly
ovate or oblong-ovate, obtuse to narrowly acute,
1.5-9 cm long, 0.5-3 cm wide, smaller at base
and apex of each pinna, the margin rather ciliate
and deeply toothed, the outer tooth margin

GENTRY & STEYERMARK—REVISION OF DILODENDRON

535

strongly convex, sometimes with 1 or 2 marginal
notches (= doubly toothed), puberulous with erect
or suberect (sometimes in part gland-tipped) tri-
chomes, above glabrescent except the main veins,

elow + persistently pubescent over surface but

rtiary

tion somewhat prominulous below, the petiole
6-12 cm long. Inflorescence a terminal fascicle
of narrow panicles, usually borne in the axils of
fallen leaves at apex of a leafless branchlet, 3-36
cm long, tannish puberulous with trichomes of
different lengths, the flowers sessile or subsessile,
borne singly or in widely spaced, few-flowered
clusters along and at tips of the lower branches
(only at apex of the much-reduced upper lateral
branches), subtended by bracteoles. Flowers
greenish to cream or yellowish, 2—3 (male) to 5
(female, fide Radlkofer) mm long; sepals 5, ovate,
unequal, sparsely appressed-puberulous and lep-
idote, the margins + ciliate; petals reduced,
shorter than sepals, 3—4(-5), sometimes com-
pletely lacking in male flowers, broadly obovate,
contracted to basal claw, puberulous at least on
2 tiny lateral projections (= scales) near apex of
basal stalk; disk glabrous pei for tuft of hairs
between filament bases; stamens exserted, (6-)
8(-9, fide Radlkofer), idm from center of
disk, the filaments ca. 3 mm long, much narrower
at apex, the anthers 1-1.5 mm long, reddish; the
female flowers (not seen, fide Radlkofer) with
puberulous disk, short thick style, and obtuse
3-lobed stigma. Capsule trigonal-obovoid, 1.5-2
cm long, splitting open somewhat unequally to
near base, 3-valved, the valves woody, 3-4 mm
thick, pubescent inside, glabrous or glabrate out-
side, drying black, the surface rugulose.

aa so kaa examined. BRAZIL. BAHIA: 10
“aen iras, 500 m, (fr), Irwin et al. 31317 (F,
NY, ire RITO FEDERAL: Bacia do Rio Sao Bar-
tolomeu, Brasilia, (fl), Heringer et al. 4530 (MO, NY),
(fr), Heringer et al. 5224 (NY), (fl), chile d et al. 7011
N of Veadeiros, (fl), Prance &
Silva 58264 (MO, NY, X5 Zona ina Ap AEA corrego
Maranhão, (fl), Pires et al. 9472 (F). MATO GROSSO:
Campinapolis, (st), Haridasan 72 (F); 270 km N of
Xavantina, 8 km E of base camp, (fl), Ratter et al.
1874 (NY). MINAS GERAIS: Ituiutaba, (fl), Macedo 761
(MO, US); 15 km de Grão Mogol, estr. Montes Claros-
Grão Mogol, (fl), Pirani et al. s.n. (CFCR 880) (NY);
Lagoa Santa, (st), Warming s.n. (US).
prope Concepcion, (fl), Hassler 7393

BoLiviA. BENI: Lake Rogagua, 300 m, Rusby 1686
(NY). SANTA CRUZ: Río Yapacani, (fl), Kuntze s.n. of
June 1892 (NY, US); Velasco, (fl), Kuntze s.n. of July
1892 (NY, US); Provincia del Sara, Montes de Dolores,
Cantón Buena Vista, 450 m, (fl), Steinbach 2515 (NY);

536 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

0 80 ^70 : 60 50 m)

FIGURE 1. Distribution of Dilodendron. Squares = D. bipinnatum, stars = D. costaricense, circles = D. elegans.

Buenavista, Prov. Sara, 450 m, (fl), Steinbach 6479 (F, moniha, pao pobre, farinha secca. Bolivia: cuta.

ERU. CUZCO: Santa Ana, 900 m, (fl), Cook & Gilbert Peru: sumbayllo hembra, sumbaillo.
1468 (US), 1617 (US); Potrero, Prov. Convención,
1,500 m, (fl), Vargas 8228 (US); Santa Ana, Prov. Con- 2, Dilodendron costaricense (Radlk.) A. Gentry
vención, (fl), Weberbauer 5020 (F), (fr), Weberbauer & Steyerm., comb. nov. Dipterodendron cos-
I tari Radlk., Smit! ian Misc. Collect.
Vernacular names. Brazil: Maria pobre, ma- 61(24): 7. 1914. TYPE: Costa Rica. Alajuela:

1987]

La Balsa de Rio Grande, Pittier 3645 (ho-
lotype, US; isotype, US)

Tree 14-35 m tall, to 45 cm dbh, with but-
tresses and flat spreading crown, the branchlets
glabrescently finely puberulous, minutely and in-
conspicuously lenticellate. Leaves bipinnate with

-14 i

n
the pinnae 5-32-foliolate (or the leaflets a the
uppermost incompletely differentiated), the al-
ternate leaflets elliptic to oblong-elliptic, obtuse
to acutish when mature, often subacuminate when
young, 1-4.5 cm long, 0.3-1.5 cm wide, slightly
smaller at base and extreme apex of each pinna
(or the terminal leaflets incompletely differen-
tiated when young), the margin not ciliate, ser-
rate with relatively even teeth (occasionally al-
most entire and typically more deeply dentate
when young), the outer tooth margin nearly
straight, above glandular-punctate, puberulous
on midvein, below glabrous or with very few
inconspicuous trichomes near base, the tertiary
venation visible and often + prominulous below;
petiole 2-9 cm long. Inflorescences several per
branch, 5-15 cm long, each a few-branched, nar-
row panicle arising from axil of fallen leaf, borne
below leaves near branch apex in fruit (presum-
ably as in D. elegans when in flower), puberulous
with crisped and subappressed trichomes. Flow-
ers unknown, presumably 8-staminate from the
vestiges at base of young fruits (fide Radlkofer).
Capsule trigonal-globose (even when only 2 fer-
tile seeds), 2-3.4 cm long and diam., splitting
open only partially, 3-valved, the valves woody,
2—4 mm thick, densely pilose inside, puberulous
and minutely raised lenticellate outside, drying
dull brownish with tannish lenticels, the seeds
bean-shaped, 1.8-2 cm long with a shiny brown
testa and a basal hilum.

Additional specimens examined. Co

ALAJUELA: La Balsa de Río Grande, h Pittier py
(US); El Coyolar, near San Mateo, 100 m, (fr), od
3681 (coll. Werckle) (US); El Coyolar, 240 m
Standley 40056 (US); vic. of Capulin, Rio Grande d
Tarcoles, 80 m, (st), Standley 40159 (US). GUANA-
CASTE: Santa Rosa National Park, 200-300 m, 10*51'N,
85°37'W, (st), Janzen 10683 (MO), Liesner 4236 (MO).
PUNTARENAS: Palmar Norte de Osa, 0 m, (fr), Allen
5738 (US

ANAMA. CHIRIQUI: Progreso, (st), Cooper & Slater
280 (NY, US); W of San iU Limite, (st), Croat
22159A (MO). DARIEN: Cerr i, Rio Coasi, (seed),
Duke 15629 (MO); Yaviza, (fo) pun 6589 (US). PAN-
AMA: El Llano, (fr), Duke 5818 (MO); Río Tapia, (st),

Kan

GENTRY & STEYERMARK—REVISION OF DILODENDRON

537

Standley 28087 (US), 28282 (US); Juan Diaz, (st),
Standley 30574 (US); Rio Tapia, (st), Standley 41186
(US).

OLOMBIA. BOLIVAR: San Juan Nepomuceno, 200 m,
dou 75°10'W, (fr), Cuadros & Gentry 3617 (MO).
oco: Municipio de Riosucio, Peye, 60 m, (st), Forero
1781 (COL, MO).
Noir BOLIVAR: 48 km NE del caserio Los Ro-
s, 17 km de Upata, (st), Blanco 334 (MO, NY, VEN);
Altiplanicie de Nuria, ESE of Villa Lola, 315 m, (st),
Steyermark 86364 (NY, VEN). MERIDA: El Vigía-Pan-
americana, 100-120 m, (st), Bernardi 2093 (NY). ZULIA:
Dtto. ColóN, 14—25 km NO de Pto. Chama, (fr), Bunt-
ing & Drummond 6324 (VEN); El Toro, 8 km SSO de
El Consejo, (fr), Bunting & Alfonzo 7054 (V EN); Misión
de Tucuco, 105-250 m, (fr), Ijjasz 88 (NY); La Cocha,
Mun. Uribarri, (fr), Trujillo 12211 (F).

Vernacular names. Costa Rica: /upinsacca.
Panama: guavino. Venezuela: tamarindo de
monte, machirio tamirindo.

The seeds are said to be edible (Bernardi 2093).

3. Dilodendron elegans (Radlk.) A. Gentry &

elegans Radlk., in Mart. Fl. Bras. 13(3): 597
1900. Dipterodendron elegans (Radlk.)
Radlk., Smithsonian Misc. Collect. 61(24):
7. 1914. svNTYPES: Costa Rica. Alajuela:
prope Alajuela, Oersted 4, 5 (C, not seen).

Dipterodendron venezuelense uir. Fieldiana, Bot.
28: 346. 1952. TvPE: Venezuela. Merida: ara
Mes Isidro Alto and Sani poses de Mora, 760-

1,800 m, Steyermark 56569 (holotype, F; dH

VEN)

Tree 8-25 m tall, the branchlets longitudinally
striate-ridged or slightly angled, minutely pu-
berulous with erect or subappressed trichomes,
glabrescent, tl
scattered or essentially lacking. Leaves bipinnate
with 10-16 frequently opposite or subopposite
pinnae, the rachis puberulous with crisped tri-
chomes, grooved above; pinnae (5-)9-23-folio-
late, the alternate to subopposite leaflets oblong-
elliptic, obtuse to acute, 1-6(-7) cm long, 0.3-2
cm wide, smaller at apex and base of each pinna
(or the terminal leaflets incompletely differen-
tiated), the margin not ciliate, serrate with rela-
tively even teeth, the outer tooth margin nearly
straight, the upper surface rather glandular and

venation usually not evident; petiole
long. Inflorescences few-branched, very narrow,
subspiciform panicles, typically arising in clus-
ters at the end of a leafless branch from the axils

538

of fallen leaves and often accompanying a cluster
of unexpanded new leaves, in fruit + clustered
at the base of the now fully expanded leaves, 3-
26 cm long, tannish puberulous with + ap-
pressed trichomes, the flowers mostly in subses-
sile or short-stalked clusters along it. Flowers
reddish, apetalous, the sepals 5, ovate, less than
1 mm long, puberulous; disk flat, densely pu-
berulous; stamens exserted at anthesis, 6—7(—8?),
the expanded filaments ca. 2.5 mm long, the short,
thick anthers ca. 1 mm long; female flowers sim-
ilar to immature male flowers, with the 1 mm
long densely puberulous ovary tapering ap a
ong, narrow style and surrounded by ca. 6-8
subsessile sterile stamens, the stigma saucy
2-lobed. Capsule compressed-obovoid, 1-1.8 cm
long, 1-2 cm wide, splitting to base at dehiscence
with the valves reflexed, 2-valved, the valves
subwoody, ca. 2 mm thick, sparsely pilose or +
glabrate inside, outside with sparse and incon-
spicuous scattered trichomes, drying black (red
when fresh) with a minutely wrinkled-verrucose
surface, elenticellate; seeds mostly 1 per fruit,
flattened ovoid, 1 cm long, with shiny dark brown
testa and a tan basal hilum.

Additional specimens examined. COSTA RICA.
ALAJUELA: Camino de San Ramón, is Brenes 435la
(NY); La Palma (San Miguel) de San Ramón, 900-
l ,000 m, (fl), Brenes 5351 (F, NY); San Pedro de San

JOS n o
675-900 m, (fl), Skutch 4850 (F, MO, NY), (fr), 4876
(MO, NY, US).

VENEZUELA. BARINAS: Barinitas, (st), Bernardi 3337
(VEN). BoLIvAR: between Tumeremo and El Dorado,
29 km N of El Dorado, 220 m, (st), Steyermark 86570
(NY, VEN); savanna de los Chacharros, 4 km upstream
from Raudal Cotua, É jp ga pad ais 86773
(NY, VEN); 2 km SE os , 30 km S of El
deed 365 m, (st), a sma siai 8695 7 (NY, US,

VEN). CARABOBO: : carretera Maracay-Magdaleno-Gui-
gue, Cuesta de Yuma, 450
TRUJILLO: Cerro Go rdo, sand oil on ridge, 9?45'N,
70?15' m. 1,000 m, (fr), Steyermark & Carreno 111646
(MO, NY, US). zuri4: Dtto. Colón, carretera Ma-
chique a Fria entre La Redoma y Placita, (D. Bun-
ting & Alfonzo 6930 E

PERU. MADRE DE DIO Tambopata, 12*49'S, 69°18'W,

280 m, (st), Gentry et P 46217 (AMAZ, MO, USM).

Vernacular names. Costa Rica: lorito, galli-
nazo. Venezuela: caro montañero.

As thus constituted, Dilodendron is a small
genus of three species with one species (D. bi-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

pinnatum) mostly in the subequatorial dry areas
of the Brazilian shield and adjacent regions, a

Central America, and a third (D. elegans) wide-
spread from Costa Rica to Venezuela and Am-
azonian Peru. Dilodendron elegans and D. cos-
taricense are apparently ecologically separated.
In Central America D. elegans occurs in wet for-
est, whereas D. costaricense occurs in moist for-

°
=
B
g:
5

Venezuela, D. costaricense occurs mostly in drier
forests and D. elegans in wetter ones.
Dilodendron provides an excellent example of
the importance of sterile collections. Not only
was the original Peruvian collection that led to
this entire revision sterile, but so are most of the
other collections of the former Dipterodendron.
Of the 22 collection numbers of D. costaricense,
all but six are sterile and six of the ten state
records from which it is known, including the
only report from Colombia, are based only on
sterile collections; the flowers are still unknown.
Described from a sterile collection, D. elegans is
now known from 17 collection numbers from
eight different states in three countries; however,
only nine of the collections are fertile and the
only records for several states as well as for the
country of Peru are based on sterile material.

collecting institutions) get over their prejudice
against sterile collections.

LITERATURE CITED

ARISTEGUIETA, L. 1973. Familias y Generos de los
Arb de Venezuela. Instituto Botanico, Min-
i ra y Cri

raca

CROAT, T. B. 1976. Sapindaceae. /n Flora of Panama
nn. Missouri Bot 19-5

MACBRIDE, J. F pindaceae. Fi Flora of Peru,

Fieldiana, Bot. XIIIA): 291-
RADLKOFER, L. 1892-1900. SERSA In Martius,
mon Bras. a 225-654.
Sapindaceae. In A. Engler & K. Prantl,
Die ce SA Pfanzenfamilien 3(5): alin 366.
New Sapindaceae from Panama an
Costa Rica. Smithsonian Misc. Collect. 6104):
1-8.

Sapindaceae. In A. Engler, Pflanzen-
reich IV. 165: 1019-1274 (Heft 98e).
STEYERMARK. J. Contributions to the Flora of
Venezuela II. Fieldiana, Bot. 28: 346.

UNA NUEVA ESPECIE DEL GENERO DIOSCOREA
(DIOSCOREACEAE) DEL ESTADO DE
QUERETARO, MEXICO!

O. TELLEZ V ? v B. G. SCHUBERT?

RESUMEN

Se describe Dioscorea matudae, una nueva especie del estado de Queretaro, México. Se discuten
sus características en relación al sentido seccional dado por Knuth en 1924.

Como parte del proyecto “Desarrollo del Her-
bario Nacional" en el Instituto de Biología,
.M., se han hecho colecciones por dife-
rentes regiones de México, una de éstas es en el
estado de Queretaro, en donde fue encontrada
una especie del género Dioscorea que después de
un análisis, se determinó como nueva especie
para la ciencia y la cual se describe a continua-
ción.

Dioscorea matudae O. Téllez & B. G. Schubert,
sp. nov. TIPO: México. Queretaro: 4 km al
E del poblado Arroyo Seco, carretera a Jal-
pan, alt. 840 m, selva baja caducifolia, 20
Oct. 1982, P. Tenorio L. & C. Romero de T.
2265 (holotipo, MEXU; isotipos, A, ENCB,
F, MEXU, MO, XAL)

Herba tenella. Caules sinistrorsum volubiles. Folia
flores 2-4(-6) quoque Jen pate 6, introrsa. Inflo-

me ala abortiva
vel vestigialiter semen circundanti.

Herbácea trepadora glabra. Tallo sinistrorso
1-1.5 mm, ligeramente angulado a lineado. H
jas (2.7-)3.5—5.8. cm de largo, (1.5—)2-4.3 cm de
ancho, alternas, ovadas, la base cordada, el ápice
agudo a larga y abruptamente acuminado; 7-ner-
vias, prominentes en el envés, ocasionalmente
escasa y cortamente serruladas, las más externas
bifurcadas; peciolo (0.7—)1—2 cm de largo, lige-
ramente angulado, cortamente serrulado. Inflo-
rescencia estaminada 1 ó 2 racimos de cimas o
panículas de cimas de 4-12 cm de largo por axila;

t ; flo-
res 2- 4(-6) porci cima; pedicelo ca. 1 mm de Lien:
rrulado; bráctea ex-
terior r1-2 mm de largo, 1 mm de ancho, ovado-
lanceolada, acuminada; bráctea interior 1 mm
de largo, 0.5-1 mm de ancho, ovada. Perianto
1-1.5 mm, verdoso a amarillento; tépalos 1—
1.3 mm de largo, 0.5 mm de ancho, oblongos a
elipticos; estambres 6, 1 mm de largo, insertados
en la base de los tépalos; anteras introrsas, las
tecas coherentes; pistilodio 0.2-0.4 mm de alto,
cónico a triangular, inconspicuo. Inflorescencia
pistilada 1 racimo de 2.4—11 cm de largo por
axila; raquis angulado, conspicuamente serrula-
do; flores solitarias; Delicato 1-2(-3) mm de lar-
go; bráctea exterior
mm de ancho, ovado-lanceolada, acuminada;
bráctea interior 1-1.5 mm de largo, 0.5 mm de
ancho, lanceolada. Perianto 1-1.3 mm, verdoso
a amarillento, los tépalos 1-1.3 mm de largo,
0.5 mm de ancho, oblongos a elipticos; estami-
nodios 6, ca. 0.5 mm de largo, insertados en la
base de los tépalos, anteríferos o no; columna
estilar 0.8-1 mm de alto, los estilos bifidos, ro-
llizos, delgados. Cápsulas 9-12 mm de largo y
ancho, con dos de los lóculos abortados, en al-
gunos casos incompletamente y no se producen
semillas (con los dos lóculos completamente
abortados 7-8 x 2-5 mm, o con los lóculos de-
sarrollados 9-12 x 6-9 mm), suborbiculares a
orbiculares, membranosas; pedicelo acrescente
en el fruto 4-6 mm de largo, cortamente serru-
lado; semilla 4 mm de largo, 3 mm de ancho,
suborbicular, parda, solo una semilla en el lóculo
desarrollado, el ala casi completamente abortada
u ocasionalmente presente como vestigios cerca
al hilo.

! Agradecemos al Biol. Pedro Tenorio por facilitar el material para el pua NONO: A] Dr. Fernando Chiang

por su ayuda con la diagnosis latina. A Elvia

Esparza por su magnífica ilust

š en de Botánica, Instituto de Biologia, U.N.A.M., Apartado Postal ' 70- 367, 04510 México, D.F.,

Mex
3 Arnold Arboretum, Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A.

ANN. Missouni Bor. GARD. 74: 539-541.

1987.

FIGURA 1. Dioscorea matudae (P. Tenorio L. & C. Romero de T. 2265).—a. Planta estaminada. — b. Cima. —
c. Flor estaminada. —d. Planta pistilada.—e. Flor pistilada solitaria. —f. Flor pistilada. — g. Cápsula. —h. Semilla.

540

1987]

Especie interesante y a la vez dificil de rela-
cionar con las especies cercanas en el sentido
seccional dado por Knuth (1924). Un conjunto
de caracteristicas permiten distinguirla facil
mente de cualquier otra especie. Estas son la pre-
sencia de seis estambres introrsos insertados en
la base de los tépalos; pistilodio cónico a trian-
gular, conspicuo, semejante a los observados en
la sección Macrogynodium Uline; el habito her-
báceo delicado; los tallos sinistrorsos; las hojas
pequenas y glabras, semejantes a las de algunas
especies con tres estambres y tres estaminodios
encontradas en varias secciones del subgénero
Dioscorea Pax.; su cápsula con dos de los lóculos
abortados; la semilla con el ala completamente
a casi completamente abortada; y el pedicelo
acrescente en el fruto. Estos caracteres hasta aho-
ra conocidos de D. cyphocarpa Robinson ex R.
Knuth y D. tacanensis Lundell de la sección Poly-
neuron Uline, pero que difieren de D. matudae
por sus tres estambres formando una columna
estaminal insertada en el centro del toro

erior evidencia con gran punbahilidad
la presencia de estructuras homólogas en el gru-

TELLEZ V. Y SCHUBERT—NUEVA DIOSCOREA

541

po. Burkill (1960) y Téllez et al. (en prep.) hacen
referencia a estas estructuras, interpretándolas
como paralelismo en diferentes líneas dentro del
género, lo cual probablemente ha dificultado en
gran medida determinar las relaciones de sus taxa
en una forma objetiva.

Con gran probabilidad ésta especie podria dar
origen a una nueva sección por si misma o quedar

El epiteto específico está dedicado al cud Eizi
Matuda, quien contribuyó en forma sustancial al
conocimiento de las especies mexicanas de éste
género.

BIBLIOGRAFIA

BURKILL, I. H. 1960. The organography and the evo-
luti tion of Dioscoreaceae. The family of the yams.

Bot. 56: 319-412.

I Dioscoreaceae. Pflanzenreich IV.
43:1-387 (Heft 87).

TELLEz, V. O., R. MEDINA L., E. MARTÍNEZ O. & P
HIRIART V La Relevancia de la Pali-
nología en la Sistemática de Dioscorea (Diosco-
reaceae

SYSTEMATICS OF THE SOUTHERN AFRICAN GENUS
HEXAGLOTTIS (IRIDACEAE—IRIDOIDEAE)'

PETER GOLDBLATT?

ABSTRACT

Hexaglottis is a genus of six species occurring along the west and south coast of southern Africa,

an area of predominantly winter rainfall. It is a m btribe Ho ae, whic
is characterized by a cormous rootstock and secondarily bifacial leaves. Hexaglottis is defined largely

y an unusual flower structure with shortly clawed subeq d completely divi filifo
style arms. This revision includes three new species, H. na den sa H.

genus Rheome. The ese scr di and history o ottis are discussed, and, owing a detai

presentation of taxonomically important characiers includin chromosome cytology, Hexaglottis is
analyzed cladistically Sm namaquana is suggeste nc an isolated and primitive relict and
the sister species of the remainder of the genus. The so frican genus Homeria is probably the

to a group of species of Moraea section Moraea. B

while H. nana has x

Hexaglottis is a small genus of Iridaceae tribe
Irideae restricted to the winter rainfall area of
southern Africa (Fig. 1). It has linear, bifacial,
and usually channeled leaves; corms of the Mo-
raea type composed ofa single swollen internode
and apically rooting bud; and umbel-like inflo-
ipidia) enclosed in large, op-

"3
°
e

ace
of eee establishes its vite po-
sition in the predominantly southern African
subtribe Homeriinae OMA pu
1980) of the Old World tribe Irideae. The ñoral
structure is distinctive and, although not unique,
defines the genus. The subequal tepals have short
erect claws and horizontally extended limbs, and
the styles are short with branches divided almost
to the base into paired filiform arms that extend
outwards on either side of the subtending anther.
The flowers are yellow and fugacious, lasting only
a few hours. Additional features are firm, brown-
to blackish- dines corm tunics and a basic
chromosome nu

Moraea nins (Conn, 1986) has a
similar flower structure, but the flowers are blue,

romosome n r in Hexaglottis is x = 6,

= 10, a number shared with Rheome and basic "y Morae

unlike Hexaglottis, and the basic chromosome
number is x — 10. A second species, described

y Bolus as H. nana, has flowers es-
sentially identical to those of other species of
Hexaglottis, but it has dark brown, unbroken
corm tunics, unusual fasciculate rhipidia, and a
base number of x = 10, B is here excluded from
Hexaglottis. C
phology (detailed below), and vegetative mor-
phology indicate that it is related to the small
genus Rheome, comprising R. maximiliani and
R. umbellata, and probably also to Moraea lin-
deri and M. margaretae, which recent unpub-
lished investigation has indicated are closely al-
lied to Rheome.

The peculiar dpa eer type of flower is thus
believed to have evolved independently three
times. It defines H E only in combination
with the vegetative and chromosomal features
mentioned above.

RELATIONSHIPS

Hexaglottis is probably most closely allied,
within the Homeriinae, to Homeria, and avail-

! Support for this research from the U.S. National Science wasan grants DEB 78-10655 and DEB 81-
fric

Compton Herbarium
iid woe for the illustrations publishe
ff C

sch, in the sania of my

ment of Enviro

n Rourke and his staff,
eldwork is acknowledged with gratitude. I also thank

urator of African An. ibn Botanical Garden, P.O. Box 299, St. Louis, Missouri

S U. Eg A.
ANN. Missouni Bor. GARD. 74: 542-569. 1987.

1987]

BZ | S GAN
Bs AW
E 1

H. lewisiae

H. virgata

H. riparia

H. longifolia

GOLDBLATT —SOUTHERN AFRICAN HEXAGLOTTIS

URNI OE E TELE UG AZ MA

S f
ON A P
D

543

nll

y

NU

QUT

22

s S
Ra eak:

| <: T

u. s N ES
AL. D
ZZ — — P

DATES

FIGURE 1. Geography of Hexaglottis.

able morphological and cytological data suggest
that the two genera have as a common ancestor
(Fig. 5) probably a species or group of species of
Moraea allied to M. flexuosa (Goldblatt, 1982).
The characteristics that Homeria and Hexaglot-
tis share include subequal tepals, the claws of
which cup the lower part of the filaments, and
style branches reduced from the elaborate flat-
tened structures basic for Homeriinae (Gold-
blatt, 1980, 1986) and probably for the entire
tribe Irideae (Goldblatt, in prep.). The two gen-
era also share a similar and derived karyotype
with x = 6 comprising strongly acrocentric to
subacrocentric chromosomes. Genome size
(Goldblatt et al., 1984) is similar, 22-29 pg DNA
in Homeria and 20.6 pg DNA in Hexaglottis
namaquana, the only species of the genus for
which this is known. Moraea flexuosa has a com-
parable karyotype. Other members of Moraea

that appear less closely related include a part of
the heterogeneous section Moraea, x — 10, and
section Polyanthes, x = 6, the latter distinguished
by having blue to violet flowers (a derived con-
dition in Moraea). The karyotype in section
Polyanthes is also somewhat different in com-
prising acrocentric and submetacentric chro-
mosomes (Goldblatt, 1980).

HISTORY OF HEXAGLOTTIS

The taxonomic history of Hexaglottis has been
described by G. J. Lewis (1959) in detail, and it
is reviewed here briefly. The first of the species
now admitted to Hexaglottis was described by
Nicholas Jacquin in 1776 as Ixia longifolia. The
excellent figure that now serves as the type of the
species is unmistakably this Cape Peninsula
species, often confused with the more common
and widespread H. /ewisiae. Shortly afterward,

544

the younger Linnaeus described a second species
as Moraea flexuosa (Linnaeus fil., 1781). Thi
name is now regarded as nomenclaturally su-
perfluous and illegitimate, and a new name, H.
lewisiae, was proposed for the species in 1971
(Goldblatt, 1971a) (see discussion under this
species).

The very distinctive, late-flowering Hexaglot-
tis virgata was described in 1791 by Jacquin, this
species also being assigned to Moraea. As with
Jacquin's earlier species of Hexaglottis, a fine
illustration leaves no doubt about its identity.
Thus, all three common southwestern Cape
species of the genus were known and described
by the beginning of the nineteenth century when
E. P. Ventenat erected the genus in 1808. Ven-
tenat made no transfers to his new genus, men-
tioning only Ixia longifolia Jacq. by name, “Ixia
longifolia Jacq. etc.," which leaves one wonder-
ing whether he had further un in mind, The
acceptanc

Nn

OPN Ic cnnn oa1mo.

were made ‘for H. ut ayay by R. A. Salisbury
(1812)and for H. virgata and the illegitimate H.
flexuosa by Sweet (1830). One more species was
collected in the nineteenth century, H. riparia,
discovered by C. F. Ecklon & C. L. Zeyher (their
Irid. 30), but E iia was consistently as-
signed to H. flex

The three PONENS, species of Hexaglottis were
regarded as a single taxon by Klatt (1866) under
the name Homeria spicata (Ker) Sweet, the type
of which is conspecific with the earlier Homeria
elegans (Jacq.) Sweet (Goldblatt, 1981). Later,
Klatt (1882: 52, 1895: 159) recognized Hexa-
glottis with H. longifolia (including H. lewisiae)
and H. virgata. Baker’s (1896) definitive nine-
teenth century floristic treatment of the Iridaceae
in Flora Capensis is identical, but he understood
Hexaglottis so inadequately (Lewis, 1959) that

is work on the genus must be disregarded.

Louisa Bolus added one more species to Hexa-
glottis in 1932, the west coast H. nana, which,
although common, was apparently only discov-
ered in the 1920s. Hexaglottis nana as already
outlined differs markedly i in its vegetative mor-

Hexaglottis and is now excluded from the genus
(for comparison of H. nana with H. lewisiae see
Fig. 2).

Lewis’s (1959) revision of Hexaglottis admit-
ted four species to the genus and two new vari-
eties, H. virgata var. lata and H. longifolia var.
angustifolia, neither of which is recognized here.

Collecting since the publication of Lewis’s re-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

vision, especially in the arid country to the north
of the Cape Floristic Region, has substantially
expanded the knowledge of Hexaglottis. I dis-
covered a new species, H. namaquana, in the
Spektakel Mountains of northern Namaqualand
in 1974 and extended the range of H. lewisiae
into these mountains west of Springbok. The dis-
tinctive new H. brevifolia of northern Nama-
qualand and the Richtersveld is now recognized,
specimens previously having been placed in H.

described from the Roggeveld Escarpment in the
western Karoo. Lastly, plants treated as H. /on-
gifolia var. angustifolia by Lewis (1959) have
been re-collected and are regarded as a distinct
species described here as H. riparia.

The picture in 1959 of Hexaglottis as a small
genus essentially of the Cape Floristic Region
with minor extensions of H. lewisiae and H. vir-
gata to the north into semi-arid Namaqualand
has changed fundamentally. Hexaglottis must
now be viewed as centered along the interior
Cape West Coast with extensions south and east
into the Cape Floristic Region.

MORPHOLOGY

Rootstock. Species of Hexaglottis have a corm
ofthe Moraea type (Goldblatt, 1976b: 670, 1981:
428) consisting of a single swollen internode with
an apical primordium from which both shoot
and roots are produced. This organ is one of the
two major specializations defining subtribe
Homeriinae. The corm originates from an axil-
lary bud near the base of the flowering stem. The
corm tunics are basically like those found in
H I d several species of M. l

L

Moraea and consist of a coarse open network of
hard, wiry, dark brown to black fibers. In Hexa-
glottis the mealy substance between the fibers
often persists and clings to the fibers, imparting
a lighter color to them. The outer tunic layers
are usually paler in color and characteristically
medium brown in many collections of H. virgata
and H. lewisiae. The two moisture-loving species,
H. riparia and H. longifolia, have softer-textured
tunics, the outer layers of which become light
brown and the fibers are relatively fine

Leaves. The leaves are bifacial witha a sheath-
ing base and more or less linear and channeled,
this being the basic leaf type for Homeriinae. The
leaves of Hexaglottis namaquana are the most
distinctive, being relatively broad, almost pros-
trate, strongly undulate, lightly twisted with the

1987]

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

Habit and flowers of Hexaglottis lewisiae (A) and for comparison the vegetative and floral mo
Bises of ‘Hexaglottis’ nana (B). Compare for example, Figures 6 and 11. Habits x 0.5; single flowers full size;
separated stamens and style branches x2.

margins sometimes somewhat crisped. The leaf
condition in H. namaquana must be regarded as
derived from the much more common erect lin-
ear leaf with straight margins found in the other
species of the genus and thus represents one or
possibly two autapomorphies. The leaves of H.
virgata may be lightly coiled distally, a feature
obscure in herbarium material. There are usually
two to three foliage leaves per plant, but the num-
ber depends on growing conditions, so that fewer

leaves are produced in drier seasons. Plants al-
ways produce more leaves in the greenhouse than
in the wild. Under optimal conditions H. /on-
gifolia, H. riparia, and H. virgata subsp. karooica
have four or five leaves. The leaves are inserted
fairly close together near or slightly below ground
level, but in H. lewisiae subsp. secunda, the leaf
or leaves may be inserted some distance above
the groun

Flowering stem, sheathing bract leaves, and

546

branching patterns. The flowering stem is more
or less erect, ee so in Hexaglottis vir-
gata, but willowy and nodding in H. longifolia
and H. riparia. There may be up to three or four
major branches, each stalked and bearing a few
to several sessile lateral rhipidia (the inflores-
cence units). The exception is H. namaquana in
which all the rhipidia are stalked, presumably
the ancestral condition, and thus terminal on the
main or lateral branches. The sessile lateral rhi-
pidia are a derived feature, and a synapomorphy
separating the main group of species of Hexa-
glottis from H. namaquana.

A sheathing bract leaf with a closed sheath
subtends each branch or lateral rhipidium. In the
latter, the sheathing bract leaf resembles the in-
florescence spathes, which it may completely
conceal. Each sheathing bract leaf generally over-
laps the one above, except in H. /ongifolia, the
cauline internodes of which are comparans
long. Lewis (1959) used this feature as an im
portant character for distinguishing H. pee

Rhipidia. The inflorescence units are of the
basic type for Iridaceae, consisting of compressed
cymose umbels enclosed in two large, opposed,
sheathing spathes, the inner of which exceeds the
outer except in the lateral rhipidia of H. longi-
folia, where they are nearly equal, a presumably
specialized condition. Individual flowers are
pedicellate and subtended by a single membra-
nous bract contained within the spathes. The
flowers are produced serially, a few days apart,
at which time the pedicels elongate to raise the
flowers out of the spathes. In Hexaglottis brevi-
tuba and H. virgata the pedicels are short and
the ovaries are included in the spathes, but the
flowers have a perianth tube that serves the same
function as the pedicel in extending the flowers
beyond the spathes. The spathes are initially her-
baceous with dry attenuate apices, but towards
the middle of the flowering season they begin to
dry out and become light brown and chaffy to-
wards the middle. The short pedicels, included
ovaries, and perianth tube are important syn-
apomorphies separating H. brevituba and H. vir-
gata from the rest of the genus.

Flower. The flower is almost uniform
throughout Hexaglottis, except for the presence
of a tube in H. brevituba and H. virgata, and is
unusual in Iridaceae in the structure of the style
and style branches and the relationship of the
latter to the stamens. The shortly clawed tepals
are pale to deep yellow and subequal, or those
ofthe inner whorl are slightly smaller. The claws,
1-2 mm long, are erect and form a cup around

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

the base ofthe filaments, while the relatively long
limbs spread horizontally. The filaments are
united below for 1-3 mm into a column, at the
apex of which the style divides into three branch-
es, each opposite a stamen. The branches divide
almost immediately to form two long, filiform,
apically stigmatic arms which extend outwards
more or less horizontally to either side of the
subtending stamen. The flowers are short-lived,
opening in the mid to late afternoon and fading
about three hours later. The unusual structure of
the style of this flower has led to the placement
of all species with this character together in a
single genus, but data presented in this revision
indicate that H. nana, described by Louisa Bolus
in 1932, is distantly related to the other species
of Hexaglottis and must have acquired its Hexa-
glottis-like flower by convergence. A similar
flower has also evolved in Moraea hexaglottis
(Goldblatt, 1986). The divided style branches
and filiform ascending arms are the primary
characters separating Hexaglottis from other
genera of Homeriinae, to which should perhaps

cally free filaments as in the putative relatives
Moraea flexuosa and Homeria (Fig. 5). The free
part of the filaments allows the long anthers to
be displayed prominently and also well separated
from the style branches which almost always in
Homeriinae divide at the top of the united part
of the filaments.

As already mentioned in the paragraph dealing

ovaries. In other species the pedicels are about
as long as the spathes, and at anthesis the ovary
is almost always exserted.

Fruit. Capsules of Hexaglottis species vary
considerably and are important in recognizing
species and in assessing phylogenetic relation-
ships. The capsules are typically exserted from
the spathes and are basically ellipsoid in shape,
as in H. namaquana and H. lewisiae subsp. se-
cunda. In the latter, the capsule has a short beak,
a feature not always evident in populations from

dric capsules, while obovoid to clavate capsules
distinguish H. riparia and H. longifolia. The cap-
sules of H. longifolia are relatively large, 12-16
(7-23) mm long, but only 6-10(-12) mm long in
H. riparia, the smaller size presumably basic and
consistent with capsule dimensions of other

1987]

TABLE 1.

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

547

Chromosome numbers in Hexaglottis and Rheome. Original counts are marked with an asterisk.

Previous counts were reported by Goldblatt (1971b, 1980).

Haploid
Species Number Collection Data
HEXAGLOTTIS
H. namaquana Goldbl. 6* Spektakel Pass, west of Springbok, Goldblatt 3059 (MO).
H. lewisiae Goldbl.
subsp. /ewisiae 6* Tulbagh Cemetery, Goldblatt 5224 (MO); Cape Town, Kir-
stenbosch Gardens (wild plants), Goldblatt 5104 (MO).
subsp. secunda Goldbl. 6 Loeriesfontein road, north of Nieuwoudtville, Goldblatt 108
(J).
6* Spektakel Pass, west of Springbok, Goldblatt 6513 (MO); gran-
ite outcrops SW of Skuinskraal, near Hondeklipbaai, van
Berkel 453 (MO); near Nieuwoudtville, Goldblatt 6535
(MO).
H. riparia Goldbl. 6* Olifants R. bank at Citrusdal, Goldblatt 6555 (MO).
H. longifolia (Jacq.) Sweet 12* Cape Town, Kirstenbosch Gardens (wild plants), Goldblatt
5934 (MO), Malan 120 (NBG).
H. brevituba Goldbl. 12* Near the Kosies road, NW of Steinkopf, Goldblatt 5748 (MO).
H. virgata (Jacq.) Sweet
subsp. virgata 5 Signal Hill, Cape Town, Goldblatt 71 (J).

5” Slopes near parking area, Signal Hill, Cape Town, Goldblatt
6747 (MO); Signal Hill, near Sheik’s tomb, Goldblatt 6768
(MO); lower slopes of Devils Peak, Goldblatt 6717 (MO).

6* Hills west of Riversdale, Goldblatt 5436 (MO); Franskraal,

near the coast, Goldblatt 5368 (MO); near Misgund,

Gold-

blatt 6792 (MO); between Bredasdorp and Napier, Goldblatt
6937 (MO); between Doorn River and Bidouw road, Gold-
blatt 5941 (MO)

subsp. karooica Goldbl. 7*

Roggeveld, near Voelfontein farm, Goldblatt 6336 (MO);

Blomfontein farm west of Middelpos, Snijman 765 (MO,
NBG)

Olifants River valley near Alpha, Goldblatt 5120 (MO); Pak-

‘Hexaglottis’ nana L. Bolus 10*
huis Pass, near Soldaat Kop, Goldblatt 5158 (MO).
RHEOME
R. maximiliani (Schltr.) Goldbl. 10 Brandewyn River near Travellers Rest, Goldblatt 3884 (MO).
R. umbellata (Thunb.) Goldbl. 10* East end of Du Toits Pass, Goldblatt 5907 (MO).
15 Paarl Golf Course, Goldblatt 4414 (MO).
15* Piketberg, top of Versveld Pass, Goldblatt 5163 (MO); foot of

the Elandskloof Mts. at Elandsberg farm, Goldblatt 5853
(MO)

species. The capsules of H. virgata and H. brevi-
tuba are included in the spathes and are narrowly
fusiform. They remain enclosed by the spathes
through ripening and dehisce only in the upper
part

The seeds are brown and basically angular, but
elongate in H. virgata and H. lewisiae subsp. lew-
isiae, both of which have narrow capsules (seeds
of H. brevituba are not known). Seeds of H. lon-
gifolia are unusually large, a feature possibly re-
lating to its polyploid state. Hexaglottis lewisiae

subsp. secunda can usually be distinguished by
the raised and winglike angles of the seeds. The
seeds are known from only a few populations of
this subspecies, and I hesitate to regard the wing-
like angles as characteristic of the taxon until
more is known about their occurrence.

CHROMOSOME CYTOLOGY

The cytology of Hexaglottis was investigated
extensively for this study. The method followed
here is the same as that outlined for similar in-

548

subsp. /ewisiae.—C. H. lewisiae subsp.
kraal, Goldblatt 5368). —F. subsp. Pb us =12 (Doorn
10 (Signal Hill, Goldblatt 674
J. ‘Hexaglottis’ nana, 2n = fed Scale = 10 um

vestigations in Iridaceae (Goldblatt, 1979, 1980).
The results are presented in Table 1. Base num-
ber in Hexaglottis is x = 6, this originally sug-
gested on the basis of one count for H. lewisiae
(as H. flexuosa), 2n = 12 (Goldblatt, 1971b),
while a single count for the specialized H. virgata,
2n — 10, suggested that this species was a derived
aneuploid. Several additional counts for Hexa-
glottis have confirmed x = 6 as basic. Hexa-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

FIGURE 3. Mitotic ns cla ne sa of j ge dia species. —
unda. — D. H. riparia. — E. H.

H. virgata subsp. karooica, 2n = 14

[Vor. 74

C
——
N CS
< ! L NT

—A. H. namaquana.—B. H. lewisiae
virgata subsp. virgata, 2n = 12 (Frans-
R. BAUN. Goldblatt 5941).—G. subsp. virgata 2n =
—I. Rheome umbellata, 2n = 20.—

glottis lewisiae, H. namaquana, H. riparia, and
three populations of H. virgata all have 2n = 12.
The two known populations of H. virgata subsp.
karooica both have 2n = 14, while H. longifolia
is tetraploid with 2n = 24.

The two populations of Hexaglottis nana ex-
amined cytologically were found to have a dip-
loid number of 2n = 20 and a karyotype exactly
like that of the diploid Rheome maximiliani

1987]

(Goldblatt, 1980). Basic chromosome number in
Rheome, already established (Goldblatt, 1980)
as x = 10, has been confirmed here, with addi-
tional counts for R. umbellata. This species was
originally thought to be triploid, 2n =
one diploid population has been discovered, as
well as two more triploid populations (Table 1).

The chromosomes of Hexaglottis species are
fairly large, ranging in size in the basic karyotype
from 5 to 9 um with the method used here. The
basic karyotype as exemplified in the least spe-
cialized species, HE wamaqa, consists Bra eid

Ww
eo
+

a satellite on the distal end of the short arm ane
the third or fourth longest pair (Fig. 3A).

Hexaglottis lewisiae subsp. lewisiae and H. ri-
paria (Fig. 3B, D) have a similar karyotype, but
the satellite is located on one of the longest chro-
mosome pairs. Size differences are relatively
small, and the shortest chromosomes are only
+35% smaller than the longest (Fig. 3A-C; see
also Goldblatt, 197 1b: 364, fig. 14E). Hexaglottis
A is tetraploid, 2n = 24, but otherwise
has a karyotype comparable to that of G. riparia
and G. lewisiae subsp. lewisiae.

The Namaqualand populations of Hexaglottis
lewisiae subsp. secunda (Fig. 3C) have an ap-
parently derived karyotype. The longest chro-
mosome pair is metacentric and about 12.5 um
long, nearly twice as long as the next in size, an
acrocentric pair. The third or fourth pair is sub-
metacentric, while the smallest pair has a large
satellite (Fig. 3C) and is only 5 um long, about
one-third as long as the long metacentric. This
karyotype has been found in two widely sepa-
rated Namaqualand populations of this poorly
sampled subspecies (Table 1), but a population
from Nieuwoudtville, well to the south, has a
karyotype of acrocentric chromosomes, unusual
only in having a satellite on the end of a long
arm ofa long chromosome pair. The single plant
that I examined was structurally heterozygous,
having only one satellite present.

In the specialized Hexaglottis virgata, there is
unexpected intraspecific variation in the karyo-
type. The presumed basic karyotype (Fig. 3E) as
found in southern Cape populations of subsp.
virgata consists of six pairs of acrocentrics, the
first and third of which have a distinctly longer
short arm. The satellite is located on the second
longest and strongly acrocentric pair. A northern
population of H. virgata (Goldblatt 5491) can be
distinguished cytologically by having satellites
on the distal end of the long arm of the longest

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

549

and strongly acrocentric pair (Fig. 3F). The
northern populations may be a separate cytolog-
ical race, but more material needs to be exam-
ined. Cape Peninsula populations of H. virgata
are aneuploid, 2n = 10 (Goldblatt, 1971b), and
have a karyotype exhibiting considerable struc-
tural rearrangement. The longest pair (Fig. 3G)
is metacentric and about 12 um long, while un-
usually large satellites are located on the shortest
and acrocentric pair, the satellite being longer
than the short arm. Robertsonian fusion of two
acrocentric, medium-sized pairs and the trans-
location of the satellite to the smallest pair would
account for the modified karyotype.

Hexaglottis virgata subsp. karooica is unusual
in the genus in its diploid number of 2” = 14
(Fig. 3H). Three individuals of both known pop-
ulations were examined. The chromosomes are
more strongly acrocentric than in the basic 27 =
12 cytotype of subsp. virgata and there are two,
rather than a single, small pairs. The origin of
the extra small pair is unknown. Satellites in the
subspecies are located on the ends of the short
arms ora a large Wa dom pair.

Kar lottis appears to
have ded from a en rather uniform,
acrocentric set of chromosomes to increasing
mmetry with the development of greater size
ieee and, in H. lewisiae subsp. secunda and
the Cape Peninsula populations of H. virgata
subsp. virgata, the evolution of large metacen-
trics, in the latter with a decrease in base number
o x = 5. Hexaglottis virgata subsp. karooica
seems to be one ofthe rare examples in Iridaceae
of an increase in base number, as the karyotype
of this specialized and rare taxon is almost cer-
tainly derived from ancestors with x — 6. The
origin of the extra pair of small chromosomes is
problematic. The tetraploid H. /ongifolia may
have evolved by amphipolyploidy, and its large
size, especially in vegetative and fruit characters,
may be a direct result of its polyploidy. Hexa-
glottis brevifolia has the same base number as
the genus but details of its karyotype were not
seen in the poor material available.

2

c

HYBRID STUDIES

A crossing program involving three species of
Hexaglottis, H. nana, and Rheome maximiliani
in the spring of 1982 produced results (Fig. 4)
that confirm the indications from cytology that
H. nana is allied to Rheome rather than to Hexa-
glottis. Rheome maximiliani could be crossed
readily to emasculated flowers of H. nana, while

550

HEXAGLOTTIS

H. virgata

RHEOME

GURE 4. panies relationships in Hexaglottis, ‘H?
nana, and me maximiliani. Heavy lines indicate
ncc oe dotted lines indicate crosses were
attempted but failed.

repeated attempts to cross both of these species
with three species of Hexaglottis used in the study
failed. Attempts to make interspecific crosses be-
tween other species o

quana, but all attempts to cross H. virgata with
these species failed. The study was not extended
to the species of Moraea that are most similar
to Rheome, namely M. linderi and M. marga-
retae, as neither was available in cultivation.

REPRODUCTIVE BIOLOGY AND POLLINATION

The Hexaglottis flower is relatively small and

times depending on the species or population, in
mid to late afternoon and closing in the early
evening. All species except H. longifolia are
strongly self-incompatible (H. brevifolia un-
known). Very rarely a few undersized capsules
are produced on plants by autogamy late in the
flowering season, but normally flowers do not set
seed by their own pollen, even though a small

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

quantity of pollen is usually deposited on the
stigmas while the flowers are open. The excep-
tion, H. longifolia, is self-compatible and autog-
amous. Plants produce numerous full capsules
with fertile seed without cross-pollination.

The pale to deep yellow stellate flowers of Hex-
aglottis are typically pollinated by bees. The small
nectaries, located at the base of the outer tepals,

marily to the pollen, a large amount of which is
produced in the relatively long anthers.
Although the flowers of all species are very
similar, there is a considerable difference in the
response of bees to the flowers ofthe three species,
H. lewisiae, H. virgata, and H. lon

longifolia is totally unattractive to bees, which
ignore open flowers even when they pass close
to them. The same bees, however, visit the open-
ing flowers of H. lewisiae subsp. lewisiae growing
within a few meters of H. longifolia. Bees gather
around populations of H. lewisiae in the middle
of the afternoon about the time its flowers nor-
mally open, and den begin to gather pollen as
soon as the flow pen.

Hex aglottis bris is similarly attractive to

and species of Anthophoridae were observed
pollinating Hexaglottis flowers.

PHYLOGENY

Cladistics affords the most objective and crit-
ical method of assessing the phylogeny of a group,
and the results of a cladistic analysis of Hexa-
glottis and its immediate allies are presented be-
low. The cladogram (Fig. 5) was constructed
manually following concepts of clustering by
shared derived characteristics (synapomorphies)
and parsimony established by Hennig (1966) and
adapted by several botanists recently (Bremer,
1976; Humphries, 1981; Funk, 1982; Goldblatt,
1985). As discussed in the preceding pages, Hexa-
glottis (excluding H. nana) is believed to be a
natural (monophyletic) assemblage distin-
guished by a number of specialized features, their
polarity determined by outgrou
by generally accepted trends in Iridaceae. The
characters used for the cladistic analysis are pre-
sented in Table 2, and most of them are discussed

camnari an

1987]

D
> AC
E Qo
AM? yt
SU VU

2 S ES
"Pdl an A
c < c S x
e e MU. x 2 Cd SP
9? e? Qo? qv < S Ri SU ae ui a?
pU um ah < e "i eU e e tuy am
SUO M" WT X9 we es eo BB Ww wi

RE 5. Cladogram of Hexaglottis and its rela-
Homeriinae indicating the possible

(par allelisms) are — by double lines and a prob-
able reversal by a

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

551

in more detail in the pages dealing with mor-
phology and cytology of Hexaglottis. The basic
structure of the Moraea flower and the reduction
and SESE 2 Homeria and some species
of a hav n discussed at length else-
oye (Goldblatt 1980, 1986). Reasons for con-
sidering x — e basic chromosome number
in Moraea and "s allies have been presented in
two studies (Goldblatt, 1971b, 1976a) dealing
primarily with chromosome cytology.

The immediate sister group of Hexaglottis is
probably Homeria, and the genera share a series
of derived and reduced floral features as well as
the same chromosome number and karyot

genera that stand out phenotypically in a variety
of unusual features. A study in progress suggests
that each of these segregates 1s related to a species
or section of Moraea. Homeria and Hexaglottis
together are probably related to M. flexuosa of
the monotypic section Flexuosa, in turn most
likely derived from species at present placed in
section Moraea. The sole synapomorphy that

LE 2. Characters used in the cladogram (Fig. 5), the derived (apomorphic) state listed first, followed by

the presumed ancestral (plesiomorphic) condition.

Corm tunics composed of hard, wiry, blackish fibers forming an open reticulum —tunics composed of straw-

colored fibers forming a fine reticu um.

. Karyotype comprising only acrocentric odo mos Gates comprising submetacentric pairs as weil

id.
dtu and nectar guides present on inner and outer tepals— nectaries and nectar guides present on outer

e branches ned iren arn above the stigma lobe.

d

2. Basic chromosome number x = 6— number x
3
as acrocentri
4. Style branches ` narrow and not petaloid—style branches broad and petalo
5.
tepals o
6. Filaments united entirely (or free near the apex)— ae united in the lower half.
7. Flowers shades of blue to v UN shades of yello
8. Stem flexuose—stem more mc
9. Tepal claws longer than the limbs- tepal p about as long as or shorter than the limbs.
10. Style branches divi to
11. Armsofthe style branches 1. apically stigmatic
12. Lateral rhipidia sessile and
eeding the subtending
13. Hypanthium tube present, E least 1 mm a tepals free from the base.
14. Hypanthium tube at least long—tub m long.
15. Ovary enclosed in the iria es—ova sert om the spath
16. Capsule narrowly obovoid to clavate-truncate — c psule ellipsoid
l psule more or less cylindric-trigo patra elli

Capsule iro i ellipsoid and iari sed in t
u ed— — not bea
Plants polyploid (2

Nee

6230.99.54
fo
"a
a

psoid.
he s nee uis ellipsoid and exserted.

= 24)— ae diploid ei = bns

. Leaves spreading and undulate — leaves ascending to erect and more or less straight.

21. Basic c some number x = 7— basi

22. Plants self-compatible and Ben DH ai tneonieedbln
23. Seeds angular-fusiform — seeds broadly angular.

24

25. Leaf margins undulate to lightly crisped —leaf margins straight.

552

unites these species is the corm tunic which con-
sists of coarse, dark brown to black, netted fibers.
Within the group with coarse black corm tunics,
Hexaglottis, Homeria, Moraea section Flexuosa,
and Moraea section Polyanthes appear to form
a monophyletic alliance that shares the derived
basic chromosome number of x = 6 and a karyo-
type of predominantly to exclusively acrocentric
chromosomes. Section Polyanthes has blue flow-

guides on both inner and outer tepals and have
style branches reduced from the basic petaloid
condition. The species of section Moraea be-
longing to this alliance include M. namaqua-
montana, M. serpentina, M. tortilis, and their
close allies, all except M. namaquamontana being
united by having included ovaries and capsules.
The detailed relationships of the species in sec-
tion Moraea are not dealt with further and will
be the subject of a future study.

Hexaglottis itself stands out in having spe-
cialized style branches and having a possible re-
versal in the filaments being free in the upper
half (see discussion under Flower in the section
dealing with Morphology). Hexaglottis nama-
quana stands out in the genus as unspecialized
and taxonomically isolated. It is probably close

species. It is the sister species to the other species
of the genus, which all have sessile lateral rhi-
pidia, an important synapomorphy in Hexa-
glottis. The included ovary and hypanthium tube
are synapomorphies uniting H. brevifolia and H.
virgata, the latter distinguished by its longer tube
and particularly short pedicel. Hexaglottis vir-
gata subsp. karooica has unusually large flowers
(probably a specialized condition but not reflect-

[o
m 4. Hexaglottis riparia
and H. longifolia form another species pair, linked
by the derived capsule shape. Hexaglottis lon-
gifolia stands out here in having large capsules,
autogamous reproduction, and in being poly-
ploid. The two subspecies of H. lewisiae appar-
ently share no synapomorphy, or at least none
that I have been able to identify. However, they
are too similar morphologically to be regarded
as separate species. Further study may throw more
light on their relationship and will perhaps in-
dicate the presence of specialized features linking
them

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

The parallelism shared by Hexaglottis lewisiae
subsp. /ewisiae and the H. virgata-H. brevituba
clade, narrow fusiform seeds (character 23), sug-
gests a possible alternative phylogeny with these
taxa forming a single clade. This is an attractive
hypothesis, suggesting as it does the derivation
of the species with a perianth tube and cylindric,
included ovary (i.e., H. virgata and H. brevituba)
from an ancestor like subsp. /ewisiae, which at
least has an unusually narrow ovary. The sepa-
ration of H. lewisiae subsp. secunda from subsp.
lewisiae that would result from such an inter-
pretation is difficult to accept given their other-
wise similar morphology.

One significant conclusion that is evident from
the cladistic analysis is that Moraea as presently
defined is shown to be paraphyletic, consisting
of a number of discrete lineages, some of which
are treated as distinct genera. The analysis of the
relationships of the main species groups in the
whole alliance is in progress and there will likely
be some changes to the taxonomy of Moraea and
its segregates. Such changes may include Hex-
aglottis but it seems preferable, nevertheless, to
publish the revision according to the present tax-
onomy. The species, H. nana, is not reassigned
to any genus, as this would be premature given
the prevailing uncertainty about the relation-
ships of species in Moraea itself.

SYSTEMATIC TREATMENT

Hexaglottis Ventenat, Decades Generum No-
vorum 6, no. 3. 1808. TYPE SPECIES: H. lon-
gifolia (Jacq.) Salisb., lectotype, designated
by Lewis (1959: 219-222)

Important references: Baker, Handbook Irid. 75-
76. 1892 et Flora Cap. 6: 31-32. 1896; Lewis,
Flora Cape Peninsula 225. 1950; J. S. African
Bot. 25: 215-230. 1959

? — Herbert, Edwards Bot. Reg. 30: Misc. 89

. TYPE SPECIES: P. flava Herb. [The identity

of E flava, treated by Lewis (1959) as conspecific
with H. virgata, is uncertain. N

known. The protologue seems to ma

ae and snae leafed species of Homeria sec-
Hom and Conanthera equally we ell.
Plantia is iir congeneric wit

Hexaglottis.]

Plants variable in size, 12-120 cm high. Corm
globose, + symmetric, 10-20 mm diam. or larger
if surrounded by accumulated tunic layers, the
tunics consisting of dark brown to blackish layers
of thick vertical ribs connected by fine cross-

1987]

fibers, the older layers increasingly fibrous and
dissected. Leaves usually 2-3, occasionally only

, linear, usually ascending and longer than the
stems, often trailing distally, or + prostrate, flat
or channeled, usually inserted towards stem base
or at some distance above the ground, the mar-
gins plane or undulate. Stem erect, straight to
slightly flexuose, branched, with only one main
axis or with 2-6 diverging secondary axes; lateral

o

amaqu
partly enclosed by the subtending stem bract, this
usually as long or longer than spathes. Rhipidia
stalked (H. namaquana) or sessile except the ter-
minal, 2—several-flowered; spathes herbaceous,
or partly to entirely pi and pale at flowering
time, attenuate, except in H. namaquana, the
outer often d E the sheathing stem bracts
and membranous below, !^—/ as long as the inner
(subequal in e lateral rhipidia of H. longifolia);
in sessile rhipidia the upper part of inner visible
above bracts except in H. longifolia. Flowers stel-
late, upright or facing sidewards, yellow, scented
or not, the nectar guides deeper yellow and usu-
ally surrounded by small dark spots, larger on
the outer tepals, located near the base of the limbs,
the tepals free or united below as a closed tube;
perianth tube (when present) cylindric, 1-7 mm
long, narrow, usually curving slightly outward,

GOLDBLATT —SOUTHERN AFRICAN HEXAGLOTTIS

553

partly enclosed in the spathes; tepals with short
erect claws 1-2 mm long, forming a narrow cup
enclosing base of filaments or filament column;
the outer slightly larger than the inner and with
a small nectary on the claw; limbs extended hor-
izontally, the outer + ovate to oblong, the inner
oblong to cuneate. Filaments 4—6 mm long, mon-
adelphous, united for 1-2(-3) mm, weakly di-
verging above; anthers 3-9 mm long, linear, ini-
tially erect, curling inwards and partly collapsing
after anthesis.

indric or wider
the spathes; style dividing at apex of the united
part of the filaments into 3 short branches, each
divided almost to the base into 2 filiform (mi-
croscopically grooved) arms, extending outwards
on either side of the subtending filaments, ciliate
and stigmatic only at the apex. Capsule ellipsoid,
obovate to clavate, or + cylindric, exserted or
included in the spathes, usually only 1 per inflo-
rescence in H. virgata; seeds angular, sometimes

Ó

ud hur basa 2n = 12, 14,

Distribution. Winter rainfall parts of south-
ern Africa, from Port Elizabeth in the east, to the
Cape Peninsula and north throughout Nama-
qualand, also locally on the Roggeveld Escarp-
ment in the western Karoo.

KEY TO HEXAGLOTTIS AND OTHER AFRICAN IRIDACEAE WITH A HEXAGLOTTIS-TYPE FLOWER

la. Rhipidia 2-several, arranged in a fascicle, each on a short stalk; outer inflorescence spathes usuall
‘Hi

not entirely sheathing, but with a diverging apex

exaglottis nana

—

outer inflorescence spathes entirely sheathing.
2a. Lateral rhipidia stalked.

3a. Flowers blue-violet; style arms 2.5- - mm lon

ie

mm wide, spreading on the ground nd twisted, the margins undulate and often cri a.
1. Hin

3b. Flowers yellow; style arms about 4

b. Rhipidia many, not fasciculate but in spicate or racemose arrangement, each either sessile or stalked;

Moraea bir d dn
leaves comparatively broad and short, to 11

namaquana

N
c

4a. Tepals free to base;

point, plants of open a

wn
c

habitats, streamsides, an
6a. ee apsules 12-16(-

shorter than outer and concealed by the sheathing bract leaves ..
nd ca. 4 mm wide; inner spathe of the lateral rhipidia
H. ri

ually
6b. Capsules 6—10(-12) mm long an

sually longer than outer and not concealed

ry
Capsules obovoid to clavate, truncate above; 4-8 mm at the widest point; plants of moist

. Lateral rhipidia sessile, ed arranged on the main axes.

sule partly to well pom from the spathes

5a. Capsules psa dca ellipsoid to | cylindric- trigonous; rarely more than 3 mm at the pus
2.

H. lewisiae

23) mm long and 6-8 mm wide; inner spathe of lateral fpa

H. longifolia

riparia

4b. Tepals united into a tube below the claws; ovary and capsule enclosed within the spathes.
7a.

Perianth tube 1-2 mm long; upper part o

plants of northern Namaqualand and the Richtersveld

7b. Perianth tube (3-)4-9 mm long; inh entirely included; plants of the northwest, p
and 6.

southern Cape and western Karo

of the ovary often emerging from the spathes;
5. H. brevituba

H. virgata

554

—
`

Hexaglottis namaquana Goldbl., sp. nov.
TYPE: South Africa. Cape: Namaqualand, top
of Spektakel Pass, stony clay soil among
patches of quartzite, Goldblatt 3059 (holo-
type, MO; isotypes, K, NBG). Figure 6.

Plantae 15-30 cm altae, foliis 2-3 prostratis undu-
latis, marginibus undulatis vel crispis, omnibus rhi-
pidiis pedunculatis ex bracteis vaginantibus caulium
exsertis, tepalis liberis, ca. 2 cm longis, ovario o 4-5 mm

a
ellipsoideis

ca. 8-10 mm longis.

Plants 15-30 cm high. Corm 1-1.5 cm diam.,
the tunics fibrous, dark brown to black, extend-
ing above into short stiff bristles. Leaves 2-3,
all + basal, 8-15 cm long, to 11 mm wide, +
prostrate, irregularly undulate or twisted, the

nches (o
bloom) stalked, the branches subtended by dry,
sheathing bracts 17-21 mm long. Rhipidia
stalked, exserted from the subtending bracts;
abov e, acute,
the i inner 2.5-3 cm long, the outer about half as
long. Flowers yellow, stellate with free, spreading
tepals; tepals about 2 cm long with claws about
2 mm long, the outer tepal limbs to 5 mm wide,
the inner narrower. Filaments united only at very
base (seemingly free), 3—4 mm long; anthers about
3 mm long, straight and suberect before dehis-
cence. Ovary 4-5 mm long, usually just exserted
from spathes, the style arms spreading, about 4
mm long. Capsules narrowly oblong-ellipsoid, 8—
10 mm long; seeds Ric about 1 mm diam.
Chromosome number 2
Flowering time. Lais Sepieribar to October;

flowers opening in the mid afternoon, after 3:00
P.M. and fading near sunset.

Distribution. Hexaglottis namaquana is
known only from the eastern slopes of the Spek-

wards the top of Spektakel Pass, in hard, stony,
clay soil, sometimes covered with white quartzite
pebbles of the Nama System. Outcrops of Nama
shales and quartzites are rare in Namaqualand,
where granites and granitic sands are the rule.
Hexaglottis namaquana may be found in other
parts of Namaqualand where there are similar
outcrops of the Nama System. The species is
sympatric with H. /ewisiae, which on Spektakel
Pass is a tall slender, narrow-leafed plant, bloom-
ing very late in the day. Its flowers open at about
5:30 P.M. and last about three hours. The flowers

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

of H. namaquana open at about 3:00 P.M. and
begin to fade at 5:30. They are further isolated
reproductively by a shift in flowering season.
Hexaglottis namaquana blooms from late Sep-
tember into mid October, whereas H. /ewisiae in
this area blooms from mid October to late No-
vember.

Diagnosis and relationships. Hexaglottis
namaquana, discovered only in 1974, is a re-
markable member of the genus. It has the small

ellow flowers that characterize Hexaglottis, but

are borne on long s rather than being
sessile. The former feature must be regarded as
primitive for the genus and separates H. nama-
quana from the other species. Members of this
species are fairly short and are unusual also in
having the leaves strongly undulate to crisped.
The capsule is unspecialized in being well exsert-
ed from the spathes and in being ellipsoid. In
other species of Hexaglottis the capsule is either
elongated and sometimes linear or is shortly
stalked and enclosed within the spathes

Fendi s specimens s. SOUTH AFRICA.

—29.17 (Springbok): Spektakel pec, west of
Springbok (DA), Goldblatt 5172 (MO, NBG); rocky
eastern the top of Spektakel Pass, Goldblatt
6672 (MO).

2. Hexagon lewisiae Goldbl., J. S. African Bot.
: 234. 1971. T

the lectotype of the invalid M. /lexuosa)].
Figure 7.

Moraea flexuosa L. f., Suppl. Pl. 100. 1782; Ker, Bot.
Mag. 19: tab. 695. 1803, nom. illeg. superf. pro
Ixia longifolia Jacq. T flexuosum (L.

1825, nom. illeg.

27, nom. illeg. bas. illeg. Hexa-
glottis flexuosa (L. f.) Sweet, Hort. Brit. ed. 2: 498.

: s, J. S. Afr. Bot. 25: 223. 1959 et FI.
Cape mih 225. 1950, nom. illeg. bas. illeg.
he as for Ixia longifolia (= Hexaglottis longi-

folia ia
)Salisb. sensu Baker, Flora
Cas, 6: 32. 1896, pro parte (excluding the type of
H. longifolia).
Homeria spicata (Ker) Sweet sensu Klatt, Linnaea 34:
H

J 4 25 í Y

we ]. s
(1866) concept of H. spicata included Hexaglottis
virgata, H. longifolia, and H. lewisiae.

1987]

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

555

E 6. Morphology and distribution of Hexaglottis namaquana. Habit x0.5; flower full size; side and
x2.

FIGU
top view w of the stamens and style branches

Plants variable in size, (12-)20-60 cm high.
Corm 15-20 mm diam., symmetric, the tunics
(pale-)dark brown, fibrous, occasionally pro-
duced upwards into a neck. Leaves 1—3(—-4), in-
serted towards the base (sometimes shortly above
the ground), ascending, linear, channeled, the
margins sometimes inrolled or rarely undulate
and the leaves rather short, normally much ex-
ceeding the stem and trailing above. Stern usually
bearing 1—3 secondary branches near the base,
often flexuose, the lateral rhipidia sessile, usually
overlapping the rhipidium above, subtended by

a sheathing stem bract concealing at least the
lower part of the spathes, and usually about two-
thirds their length. Rhipidia sessile except the
terminal; spathes herbaceous, dry above, (2.5-)
3-4.5 cm long, attenuate, the inner slightly longer
than the outer. F/ower golden yellow with a strong
sweet scent, stellate with free tepals, the outer
often feathered brownish on the reverse, the nec-
tar guides deeper yellow, usually surrounded by
several small dark greenish spots; tepals 19-24
mm x 7-10 mm (subsp. /ewisiae), 24-30 mm x
10-13 mm (subsp. secunda), the claws 1.5-2

556 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

ZZ:
UP Sg Qa
: p ny ete Hep ur tem

L °.
= M
1500
7 ALL NM
|
l
Yi PFA
, AD
ea N
Tad as
edi
uw 4.

FIGURE 7. Morphology of Hexaglottis lewisiae subsp. lewisiae and the distribution of both subspecies of H.
lewisiae. Habit, corm, and fruiting branch x0.5; flower full size; seeds much enlarged.

1987]

mm long, the limbs horizontal, the outer ovate,
the inner more or less cuneate, about as long to
2 mm shorter than the outer. Filaments 4-6 mm
long, united for 1-1.5 mm; anthers 5-7 mm long
before e Ovary (5.5-)6-12 mm long,
usually partly to entirely exserted at flowering,
rarely eel included (Barker 2558); style about
ong, the style arms spreading and 6
mm long. Capsule triangular in section and nar-
rowly cylindric, 11—16(-20) mm long (subsp.
lewisiae) or terete and ellipsoid, 8-13
(-15) mm long (subsp. secunda), then often shortly
beaked, dehiscing in the upper third to half only;
seeds angular, sometimes narrowly so and ta-
pering at both ends, 1.2-2 mm long, 0.7-1 mm
at the widest, winged in northern populations.
Chromosome number 12.

Flowering time. (September-)October-No-
vember; flowers opening between 3:00 and 4:00
P.M. in the south and beginning to fade towards
7:00 P.M., only opening towards sunset in the
Namaqualand populations and fading at about

Distribution and biology. Hexaglottis lewi-
siae is widespread in the southern African winter
rainfall area (Fig. 7). It extends from Springbok
in the north through the southwestern Cape as
far east as the Humansdorp district. It is com-
mon only in the western Cape and has a scattered
distribution to the north of Piketberg and east of
Bredasdorp. It is found in dry and exposed sites,
often on stony ground, and occurs on sandy and
clay soils.

Like most species of Hexaglottis, H. lewisiae
is pollinated by bees, but pollination has been
studied in most detail in this species, and my
observations are summarized here. The rela-

tively
appear to be very attractive to small bees. When
the flowers open in mid afternoon, bees rapidly
appear around the plants and begin to gather
pollen. They visit the same flowers several times,
collecting pollen as soon as it is exposed in the
anther sacs, which dehisce slowly from the apex.
Pollen is the primary reward and only occasion-
ally is some nectar also taken. The small quantity
of nectar produced seems to be of minor interest
to the pollinators. Fruit and seed production is
very successful in H. /ewisiae, and plants develop
several full capsules from each of the many in-
florescences.

Diagnosis and relationships. Hexaglottis
lewisiae is a diploid and self-incompatible. Its

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

337

yellow flowers are almost identical to those of all
other species of Hexaglottis, except that unlike
H. virgata and H. brevituba, the tepals are free
to the base. It shares with all but H. namaquana
the similar vegetative feature of sessile lateral
rhipidia. It is most easily confused with H. ri-

or clavate capsules, thus always broadest in the
upper third and markedly flat-topped, unlike the
cylindric or ellipsoid, and sometimes beaked,
capsules of H. lewisiae. Hexaglottis riparia and
H. longifolia grow in moist situations and, pos-
sibly as a consequence, have more and longer
leaves than is usual in H. /ewisiae. The combi-
nation of init three or = leaves; obovate: cla-
vate oist habit

smaller RR makes it ol that H. riparia
and H. longifolia will be confused with H. /ew-
isiae.

History. Hexaglottis lewisiae has been known
since the younger Linnaeus (1782) described the
species as Moraea flexuosa based on material
collected by Carl Peter Thunberg a few years
earlier. The epithet is regarded today as super-
fluous and illegitimate since Linnaeus cited as a
synonym /xia longifolia Jacq., now H. longifolia.
The combination * ee flexuosa was made
by Sweet in 1830, and the species, usually in-
cluding H. Tonerli, was known by this name
for several years. The later nineteenth century
botanists generally did not recognize H. flexuosa.
F. W. Klatt included it, together with H. virgata
and H. longifolia, in Homeria spicata, this a syn-
onym of Homeria elegans (Goldblatt, 1981). Lat-
er, Klatt (1882) recognized Hexaglottis, includ-
ing H. virgata and H. longifolia (presumably but
not explicitly including H. lewisiae). J. G. Baker
(1896) included H. flexuosa in H. longifolia, and
the distinction between the two species was only
reestablished by Lewis in 1950. The new name
H. lewisiae was proposed in 1971 by the present
author for the species that until this time was
known by the illegitimate name H. flexuosa.

Variation. There appear to be two major
forms of Hexaglottis lewisiae, the southern and
typical, treated here as subsp. /ewisiae, which
extends from the western Cape coast eastwards
through the southern Cape to Humansdorp. It
has medium-sized flowers with tepals 19-24 mm
long and distinctive long slender capsules 1 1-20
mm long that tend to dehisce only in the upper
part. The seeds are also comparatively small as
a result of the need to be accommodated in the

558

narrow locules. The karyotype in this form ap-
pears to be uniform and consists of four strongly
acrocentric chromosome pairs and two acro- to
submetacentric pairs.

Populations to the north of the Olifants River
mountains, from Clanwilliam north to Spring-
bok, treated as subsp. secunda, comprise plants
with larger flowers, the tepals 24-30 mm long,
somewhat shorter ellipsoid capsules 8-13(-15)
mm long, and larger seeds. The chromosome cy-
tology of this series of populations is not as well
known, but two Namaqualand populations ex-
amined have karyotypes with a large metacentric
chromosome pair. This northern form is mor-
phologically variable. Namaqualand plants have
a single leaf in the wild, but two leaves in cul-
tivation, and slightly smaller flowers than those
from the northwest Cape. The extensive popu-
lations from the western Karoo near Nieuwoudt-
ville have capsules with a beaklike apex, a feature

n
Clanwilliam and Y nrhyn
and karyotypes din been determined for one
northwest Cape population. The flowers of the
Namaqualand populations have a different phe-
nology, opening between 5:30 and 6:00 P.M.,
whereas all other forms of H. /ewisiae open be-
tween 3:00 and 4:00 P.M. and fade at about 6:00
P.M

KEY TO THE SUBSPECIES OF
HEXAGLOTTIS LEWISIAE

la. Capsules ellipsoid, 8-13(-15) mm long, often
distinctly beaked; outer tepals 24-30 mm long,
flowers usually secund ........

lb. pA cylindric or nearly
m long, not beaked; outer tepals 19-24 mm

pen flowers usually upright aaa.

2B. subsp. /ewisiae

2A. subsp. secunda Goldbl., subsp. nov. TYPE:
South Africa. Cape: stony east-facing slopes
near the top of Spektakel Pass, west of
Springbok, Goldblatt 6673 (holotype, PRE;
isotypes, K, MO, NBG, S, US, WAG)

Planta 30-60 cm alta, floribus usitate hus te-

end 24-30 mm longis 1 rio 5.5-9

m longis, APS ellipsoideis 8- 1315) m mm longis
ipia rostratis

3

Plants 30-60 cm high. Flowers usually secund;
tepals 24-30 mm long, 10-13 mm wide. Ovary
5.5-9 mm long, usually at least partly exserted
at flowering. Capsule terete and ellipsoid, 8-13
(-15) mm long, dehiscing for at least half its

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

length; seeds angular, narrowly winged on the
ridges, about 2 mm long, ca. | mm at the widest.

Distri bution. Subspecies secunda has a scat-

william north through the Olifants River Valley
to Vanrhynsdorp, to the Karoo north and east
of Nieuwoudtville, and into Namaqualand, where
it has been recorded west of Garies, in the Ka-
mieskroon district, and on the Spektakelberg, west
of Springbok. Plants grow in rocky ground, either
in clay soil, as on the Spektakelberg and in the
Karoo, or in sand.

Additional specimens examined. SOUTH AFRICA.
Se 17 (Springbok): stony east-facing pus near
p of Spektakel Pass, west of Springbok (DA),
Goldratt 6613 (MO).
30.17 (Hondeklipbaai): Klip Vlei, between Kamies-
A 9

of Skuinskraal farm, Hondeklipbaai
road (DB), van Berkel 453 (M B
31.18 (Vanrhynsdorp):
EET byi (K), Barker s665 (NBG), Snijman s
G); flats below the erg, 12 km south of
in veri Goldblatt 6992 (MO, PRE); slopes of
the Olifants River Bridge, south of Klawer, rocky sand-
stone soil (DC), M ae (K, MO, NBG, PRE).
31.19 (Calvinia): Loeriesfontein road, ca. 2 mile
north of the Waterfall (AC ), Goldblatt 1 08 (J); karroid
hills anges of the Klip Koppies, Nieuwoudtville,

ndkraal sandveld (D

farm, Nieuwoudtville, Goldblatt
y; Nieuwoudtville rocks on top of kopies,
Galpin 11137 (K
32.18 (Clanwi is north of Clanwilliam,
rocky sandstone soil (BB), Goldblatt 6990 (MO, PRE,
, US).

2B. Subsp. lewisiae.

Plants (12-)20—-60 cm high. Flowers bur
upright; tepals 19-24 mm long, 7-10 mm wide.
Ovary 8-10 mm long, usually partly to nes
exserted at flowering, rarely entirely included.
Capsule terete to triangular in section, narrowly
cylindric, 11-15(-20) mm long, dehiscing in the
upper third to half only; seeds angular-ellipsoid,
tapering at both ends, 1.2-1.4 mm long, ca. 0.7
mm at the widest.

Distribution. Subspecies /ewisiae extends
from the Cape west coast east through the south-
ern Cape to Humansdorp. It appears to be com-
mon only in the west and records are very scat-
tered east of Bredasdorp. Figure 7.

Specimens examined. SOUTH AFRICA. CAPE-32.1
(Clanwilliam): NE edge of Verlorenvlei (AD-BC), m

1987]

lans 7803 Do De Hoek, Piketberg (DD), Barker
2558 (NBG

33.18 (Ca own): Mamre hills (AD), Compton
9828 (NBG); d (BB), Loubser 466 (NBG); ca.
10 km north of Malmesbury (BC), Goldblatt 6173 (MO,
S, US); mountains around Cape Town vy Ecklon
& Zeyher s.n. (84) (S); foot of Lions Head, Pappe s.n.
(SAM 70674); Oudekraal, Cape Peninsula, Goldblatt
163 (J); hang Lewis 97 1 (SAM); Camps Bay, Moss
13403 (BM); Table Mountain (CD), Ecklon 536 (BM,
K, MO, PRE); Wynberg Hill, Ee 10198 (MO, US);
Kirstenbosch, Lewis 672 (N SAM); Kirstenbosch,
slopes near the herbarium, Lei 5104 (MO); Kir-
stenbosch, near kd npud offices, Goldblatt 6634
(BR, M AG); below Pearson House, Kir-
stenbosch, i. 6634 (BR, MO, PRE, WAG); near
Bishopscourt, Salter 9002 (BOL); behind Groot Schuur,
Wolley Dod 360 (BM, BOL, K); Rosebank, H. Bolus
3801 (BOL, K); Tygerberg Nature Reserve (DC), Loub-
ser 3059 (MO); Langverwacht, above Kuils River, Oliver
4803 (K, MO, PRE, STE); Stellenbosch Flats (DD),
Garside 66 (K); between Klapmuts and Paarl, Acocks
3677 (S); Berg 2 near Paarl, Drége s.n. in 1840 (K,
S), Barker 8797 (N

33.19 ocu. Gydo Pass (AB), Wall 705 (S);
orcester, beim Wasserfall (AC), Ecklon & Zeyher
(LD, MO); near Tulbagh, Leighton 1317

(BOL); Tulbagh plains, Marloth 9575 (PRE); Tulbagh

).
21 (Ladis a 5 mi. west of Ladismith (AC),
rat 3030 (N
E ot Ruigtevlei,
eb C), Fourcade 1525 (BOL).

33.24 T aaa Essenbosch hills (CD), Four-
cn 4420

4.18 i i UE Simons Bay (AB), Wright 269
u ene Bay, Wall s.n. (S); s Barker 2728
(BOL, NBG); Bergvliet Farm, Purcell 124 (SAM).

34. 19 (Caledon) Elgin (AA), Dd 1164 iei
Dwarsberg-Somerset Sneeukop, Stokoe s.n. (SAM
55728); Genadendal (AB), Prior s.n. (K); between Houw
Hoek and Kleinmond (AC), Werdemann & Oberdiec
678 (B, K, PRE); Hemel en Aarde (AD), Gillett 90
(STE); 8 mi. from Stanford on the road to Elim, Gillett
4506 (BOL, K); SEV: Vogelklip, Sapa S.
Williams 873 (C, M AG); near Napier (BD), Lei-
poldt 3551 (BOL); mb Baardscheerdersbos and
Elim (DA), Goldblatt 7107 (MO, PRE).

34.22 (Mossel Bay): ps the river at Great Brak
River (AA), Young s.n. (BOL 5541); between (ado
and Great Brak River (?AB), y eun 6151 (K).

34.24 (Humansdorp): Oudebosch flats (AA), Four-
cade 960 (BO L).

Without precise locality: Cape of Good Hope (CBS),

near Swart R.,

Swartz"); rock crevices above forest plantation, Clan-
william, Galpin s.n.

Introduced: Réunion, Trou aux Cerfs, Vaughan 3255
(SAM), Lorence 15764 (K).

3. Hexaglottis riparia Goldbl., sp. nov. TYPE:
South Africa. Cape: along the Olifants River

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

559

at Citrusdal campsite, after fire, Goldblatt
6706 (holotype, NBG; isotypes, K, MO,
PRE, STE). Figure 8

sa aa ee aor var. angustifolia Lewis, J. S. Af-
2. 1959, pro parte (excluding the
type ES var. annal e placed in H. longi-
n the present pap
(Plantia flava Herb., Edwards Bot. R eg. 30: misc. 89.
1844 1. possible synonym. See discussion below
der tory.)

Planta 45-90 cm alta, foliis (2-)3 linearibus rhipidiis
terius sessilibus, bracteis caulis imbricatis lon-
gioribus m internodis, spathis onus ide
in dudes caulis T tepalis liberis 16-21 mm
longis 6-8 mm latis, ovario 5-7 mm longo exserto,
capsulis ue do 6-10C1 2) mm longis.

Plants 45-90 cm high. Corm 13-20 mm diam.,
the tunics of fine, light brown fibers. Leaves

long as the stem, the upper decreasing in length
and width. Stem straight, simple or 1-3-branched
from the lower nodes, the lateral rhipidia sessile
at each node, subtended by a sheathing stem qas

ftan

these as ines or sontes hài longer than the inter-

node and overlapping the next bract. Rhipidia
sessile, except the terminal; spathes herbaceous,
attenuate, dry apically, 3.3-3.8 cm long, about
as long as the subtending stem bract, the outer
shorter than the inner, often hidden. Flowers stel-
late with free tepals, deep yellow, strongly scent-
ed; tepals 16-21 mm long, 6-8 mm wide, with
claws about 2 mm long, the limbs spreading, the
inner slightly shorter than but as wide as the
outer. Filaments 4-6 mm long, united for 1.5-
2.5 mm; anthers 5-6 mm long. Ovary 5-7 mm
long, exserted; style arms 5-6 mm long, extended
horizontally. Capsules obovoid-clavate, some-
what truncate, 6—10(-12) mm long, about 4 mm
wide; seeds angular, 1-1.5 mm long, 1 mm at the
widest diam. Chromosome number 2n = 12.

Flowering time. October-November; flow-
ers open at about 5:00 P.M. and begin to fade
after 7:30 P.M.

Distribution. Hexaglottis riparia has a lim-

south and Clanwilliam in the north (Fig. 9). It
appears to be restricted to streambanks and pos-
sibly edges of marshes. Such areas are usn)
overgrown with tall vegetation, and H. ripar
accordingly blooms only after fires or heavy graz-

560

f£ II

vias Morphol Habit
flowering, and trating branches x0. 5; Fendi flower and
eds full siz

ing when the habitat has been opened up con-
siderably.

Diagnosis and relationships. Hexaglottis ri-
paria has flowers typical of the genus. It is dis-
tinctive largely in its small obovoid to clavate
fruits and in its slender, often willowy stems. The
capsules are 6-10 mm long, or occasionally in
robust plants up to 12 mm. Hexaglottis longifolia
has similarly shaped capsules but they are much
larger, usually 16-23 mm long. The similarity in

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

capsule shape prompted Lewis (1959) to include
the only collection of H. riparia known to her in
H. longifolia var. angustifolia. The type of the
latter has unusually narrow leaves but in other
respects the type corresponds well to typical H.
longifolia. This species differs further from H.
riparia in having pale yellow, unscented flowers
and in being self-compatible, autogamous, and
tetraploid with 2n = 24. The flowers of H. riparia
are deep yellow and scented, and, like other
members ofthe genus, it is self-incompatible and
diploid with 2n = 12. The similar capsule shape
in H. riparia and H. longifolia, a derived feature,
probably indicates that they are closely related.

History. This species was apparently collected
first by Ecklon & Zeyher in the Tulbagh district
in the 1820s, and rarely since then. It was initially
assigned to Hexaglottis longifolia and subse-
quently cited by Lewis (1959) under H. longifolia
var. angustifolia. There is a possibility that the
species described as Plantia flava may be the
same as H. riparia. Baker (1896) and Lewis (1959)
treated the monotypic Plantia as congeneric with
Hexaglottis. The type is, however, unknown, and
the description is ambiguous and could apply as
well to some species of Homeria as to Hexa-
glottis. Plantia flava was grown and flowered in
England from corms collected by George Synnot,
who lived in the Clanwilliam district from 1821
to 1825 (Gunn & Codd, 1981). If Plantia flava
is a species of Hexaglottis, it is probably con-
specific with H. riparia, the only species of Hexa-
glottis in the northwest Cape with capsules that
correspond with Herbert’s description of the cap-
sules as obovate.

Specimens examined. SOUTH AFRICA. CAPE-32.18
(Clanwilliam): river banks, Clanwilliam (BB), Galpin
11483 (BM, K, PRE, UPS); Clanwilliam, Leipoldt 376
(S

32.19 (Wuppertal): along the Olifants river at Cit-
rusdal campsite (CA), iege 6555 (K, MO, NBG),
P (K, MO, NBG, PRE, STE).

3.19 (Worcester): cu Tulbaghskloof, etc.,
; & Zeyher Irid. 30 (77.9) (K, MO).

4. Hexaglottis longifolia (Jacq.) Salisb., Trans.
Hort. Soc. 1: 313. 1812; Baker, Flora Cap.
6: 32. 1896, in part excl. H. lewisiae (as H.
flexuosa); Lewis, Flora Cape Peninsula 225.
1950 et J. S. African Bot. 25: 223-225. 1959.
Ixia e. Jacq., Hort. Vindob. 3: 47 &
tab. 90. 1776. Moraea longifolia (Jacq.) Pers.,
Syn. Pl. 1: 49. 1805. Moraea flexuosa L. f.,
Suppl. Pl. 100. 1782, nom. illeg. superf. pro

1987]

Ixia longifolia Jacq., applied to H. lewisiae.

Sisyrinchium flexuosum (L. f.) Spreng., Syst.

Veg. 1: 167. 1825, nom. illeg. bas. illeg.

Homeria flexuosa (L. f.) Sweet, Hort. Brit.
95.

223. 1959, nom. illeg. bas. illeg., applied to
H. lewisiae. TYPE: South Africa. Cape: illus-
tration in Hort. Vindob. 3: tab. 90. Figure 9.

j ge pone wht es var. angustifolia Lewis, J. S. Af-

9. TYPE: South Africa. Cape:

Du pim Rs ‘Pillans 8436 (BOL, holotype),

pro parte (including only the type, other speci-
mens here treated as H. riparia).

Plants 60—150(—200) cm high. Corm 15-20 mm
diam., the tunics of fine, light brown fibers. Leaves
3-4(—5), linear, ascending, the lower longest, 50-
100 cm long, the upper decreasing in size, 6-12
(-20) mm wide, flat or channeled, usually bent
and trailing distally. Stem straight, simple or 1—
3-branched from the lower nodes; lateral inflo-
rescences sessile at each node, subtended by a
sheathing stem bract often entirely concealing
the spathes, as long, longer, or shorter than the
internode. Rhipidia sessile, except the terminal;
spathes herbaceous, attenuate, dry at apex, 3.5-
5.5(-7.5) cm long, the inner about as long as the
subtending stem bract, the outer shorter, often
hidden. Flowers pale yellow, odorless or with a
very faint odor, stellate, with free tepals, the nec-
tar guides usually only on the outer tepals, in-
conspicuous; tepals with claws about 2 mm long,
the outer with limbs 19-27 mm long, 8-11 mm
wide, the inner 16-23 mm long, 6-9 mm wide.
Filaments 4-6 mm long, united for 2-3 mm;
anthers 5-6 mm long before anthesis. Ovary 8—
12 mm long, exserted, the style arms compara-
tively short, 4-6 mm long, ascending, reaching
only to the base of the anthers. Capsules well
exserted, obovoid-clavate, truncate, 12-16(-23)
mm long, 6-8 mm wide, dehiscing for /2— their
length; seeds angular, 2-2.3 mm long and up to
1.5 mmatthe widest. Chromosome number 2n =
24.

Flowering time. Mid October-November;
flowers opening (2:00—)2:15-2:45 P.M. and be-
ginning to fade ca. 6:00 P.M.

Distribution and biology. Hexaglottis longi-
folia is restricted to the Cape Peninsula and a
few valleys in the mountains between Somerset
West and Paarl. It grows along streams and in
marshes or shady and damp sites (Fig. 9).

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

561

It is the only polyploid species in the genus
and further unusual in Hexaglottis in being self-
compatible and normally autogamous. Although
the flowers appear to the human eye similar to
those of the bee-pollinated H. /ewisiae, H. lon-
gifolia is seldom visited by insects. Plants of H.
longifolia growing a few feet away from H. lew-
isiae are ignored by bees gathering pollen from
the latter species. Flowers of H. longifolia placed
among those of H. lewisiae are consistently
avoided by bees. Clearly to these insects, the
odorless flowers of H. longifolia are very different
to those of H. lewisiae, which are strongly scent-
ed. Possibly there are significant differences in
ultraviolet patterning as well.

Diagnosis and history. The good illustration
of Jacquin's Ixia longifolia accompanying the
protologue and now regarded as the type of the
species corresponds unmistakably to the tall, pale
yellow-flowered Hexaglottis of damp shady sites
on the Cape Peninsula. The painting shows clear-
ly the characteristic pale yellow flowers, six rath-
er short style branches, and the large, obovoid
to clavate capsules. Despite this clear illustration,
H. longifolia has often been associated with the
very different H. lewisiae (as H. flexuosa) (Klatt,
1895; Baker, 1896), which has deep yellow,
scented flowers, long style branches, and linear
cylindric capsules. Lewis (1959) has explained in
detail this historical confusion and has elabo-
rated the several differences between these two
quite distinct species. No further discussion seems
necessary here.

Lewis (1959) recognized two varieties of Hex-
aglottis longifolia in her revision, the typical, and
var. angustifolia, the latter distinguished by nar-
rower, stiff, and strongly channeled leaves, short-
er spathes, and smaller capsules. Her decision to
treat this somewhat smaller form as a variety of
H. longifolia was evidently based on similarity
of general appearance, including the shape of the
capsules, which, although smaller, are obovoid
and truncate, and by the habitat: moist places
along streams and rivers. The type, from du Toits
Kloof, seems to me merely a slightly smaller
specimen of H. longifolia and can be matched in
size by some collections from the Cape Penin-
sula. It does not appear to warrant taxonomic
recognition. However, the other collection cited,
Galpin 11483, from the Olifants River Valley, is
a different species treated here as H. riparia. It
has smaller capsules and the flowers, examined
live from this area, are unlike those of H. /on-

562

ANNALS OF THE MISSOURI BOTANICAL GARDEN

A H. riparia

Zm
mi

Nau

|

E 9. Morphology of Hexaglottis longifolia and distribution of H. ol ^d H. riparia. Habit,
eben and fruiting branches, and corm x0.5; flower + full size; seeds much enlarg

gifolia. They are bright yellow and strongly scent-
ed, in both characteristics corresponding better
to H. lewisiae and in conflict with H. longifolia
with its pale yellow, scentless flowers. In addi-
tion, H. riparia is diploid, 2n = 12, and self-
incompatible, in contrast to the tetraploid and
autogamous H. longifolia.

Specimens examined. SOUTH AFRICA. CAPE-33.18
(Cape Town): Liesbeek River, below Fernwood (CD),
Salter 8973 (NBG), 8775 (NBG, SAM), 8999 (NBG);
Kirstenbosch, Lewis 673 (NBG, PRE, SAM), Goldblatt
5934 (MO); Kirstenbosch, below Pearson House, Ma-
lan 120 = “a Goldblatt 6635 (K, MO, PRE, WAG);

low Fern , Salter 9377 (BM); above Rhodes
Drive, Salter 9154 (BM); Table Mt., east base in damp

soil, Pillans 10262 (BR, MO); ee Wer-
dermann & Oberdieck 722

33.19 (Worcester): du ci ae (CA), Pillans 4836
ou

ou

Simonstown): Orange Kloof, swamp (AB),
Wolle Dod 3479 (BM, BOL, K, PRE); shady roadside
ing below Constantia Nek on the road to Groot
Ca Goldblatt 6640 (MO, PRE, S).
Without precise locality: Thunberg s.n. (S "Herb.
Casstrom").

5. Hexaglottis brevituba Goldbl., sp. nov. TYPE:
South Africa. Cape: Richtersveld, Sabiesies,
on the a to Cornelsberg, Viviers s.n. in
1983 (holotype, NBG; isotypes, K, MO,
PRE). Figure 10.

[VoL. 74

1987]

Planta 40-55 cm alta, foliis 2-4, rhipidiis : apen

exterioribus ex bracteis caulis “ra nalis [aD
bonus tubo 1-2 mm longo, ovario 8-1 m longo
supra ex bracteis exserto, pedicellis 4-10 mm mg a

Plants 40-55 cm high. Corm 12-18 mm diam.
or larger if surrounded by accumulated tunic lay-
ers, the tunics of coarse, dark brown to blackish
fibers. Leaves 2—4, linear, ascending, longer than
the stems and trailing distally, seh ie with
margins incurved, inserted towards stem base.
Stem with 1 main axis or with 23 diverging
secondary axes, straight or rarely slightly flex-
uose; lateral rhipidia sessile, partly enclosed by
the somewhat shorter subtending stem bracts.
Rhipidia sessile except the terminal, 3-4-flow-
ered; spathes herbaceous, or partly to entirely dry
and pale at flowering time, attenuate, (20-)25-
30 mm long, exserted from the stem bracts, often
membranous below, the outer half to two-thirds
as long as the inner. Flowers stellate, upright or
secund, pale yellow, evidently odorless; tepals
united below into a short closed tube; perianth
tube 1-2 mm long, exserted from the spathes;
tepals 16-19 mm long, the claws about 1 mm
long, the limbs extended horizontally, 5-8 mm
wide, the outer larger, narrowly ovate, the inner
smaller. Filaments 3—4 mm long, united for about
1 mm; anthers 7-8 mm long. Ovary 8-11 mm
long, cylindric, often curved outwards, the upper
1 mm narrow and sterile, the apex often exserted
from the spathes; pedicel 4-10 mm long, the style
arms about 6 mm long. Capsule a ep seeds not
known. Chromosome number 2n

Flowering time. September to mM flow-
ers opening mid afternoon and fading after 7:00
P.M.

Distribution. Hexaglottis brevituba is known
t

10). There is only a single record from the central
part of Namaqualand, at Stinkfontein south of
Garies, and other collections are from the Spring-
bok area and north to the Richtersveld, where
the type collection was made. Hexaglottis brevi-
tuba is probably more common than the present
record indicates. It is inconspicuous except when
in bloom, and the flowers are open for only a few
hours in the mid afternoon of a few weeks, usu-
ally towards the end of spring.

Diagnosis and relationships. Hexaglottis
brevituba is clearly allied to the widespread H
virgata, which has similar vegetative and floral

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

563

morphology but is readily distinguished by its
flowers with a well-developed perianth tube some
4—7 mm long and an entirely included ovary with
a pedicel 4-6 mm long. In H. brevituba the peri-
anth tube is only 1-2 mm long; the ovary is often
apically exserted and the pedicel (4-)7-10 mm
ong

History. The first record of H lottis brevi-
tuba was made by Rudolf Schlechter i in 1897 and
the collection was identified only as Hexaglottis.
In Lewis's revision of the genus she placed it in
H. virgata var. lata, the type and only other col-
lection being from the Biedouw Valley. A third
collection, made by G. J. Lewis near Springbok,
was assigned to H. lewisiae (as H. flexuosa) by
Lewis, who did not notice the characteristic short
perianth tube in the poorly preserved flowers.
The perianth tube can, however, be seen in this
collection when buds are examined carefully. The
range of the species was substantially extended
to the Stinkfontein Mountains in the Richters-
veld by Mike Viviers in 1983. This, the only
adequate collection of H. brevituba, has been se-
lected as the type.

Specimens examined. TH AFRICA. CAPE: 28.16
(Vioolsdrif): Richtersveld, Sabiesies, on the road to
Cornelsberg (CA), Viviers s.n. (K, MO, NBG, PRE),
1337 (NBG).

29.17 (Springbok): north of Steinkopf, near Kosies
road in kloof northeast of Rabas (BA), Goldblatt 5746

~
3
Am
Nn
a
-
xí

30. g): Stin
Doorn R.) (CC), Schlechter 114876 (BOL).

6. Hexaglottis virgata (Jacq.) Sweet, Hort. Brit.
ed. 2: 498. 1830; Baker, Flora Cap. 6: 32.
1896, pro parte; Lewis, Flora Cape Penin-
sula 225. 1950 et J. S. African Bot. 25: 225.
1959. Moraea virgata Jacq., Ic. Pl. Rar. 2:
tab. 228. 1791 et Coll. Bot. 3: 194. 1791.
Ixia virgata (Jacq.) Willd., Sp. Pl., 1: 202.
1798. Homeria virgata (Jacq.) Sweet, Hort.
Brit. ed. 1: 395. 1827. TYPE: South Africa.
Cape: cultivated in Vienna, illustration in
Jacq., Ic. Pl. Rar. 2: tab. 228. Figure 11.

Plants variable in size, 12-85 cm high. Corm
13-18 mm diam., or larger if surrounded by ac-
cumulated tunic layers, the tunics of coarse, dark
brown to blackish fibers. Leaves — 2-3, oc-
casionally only 1, rarely 4, linear, ascending,
longer than the stems and trailing talis. chan-
neled with margins curving inward, occasionally
loosely coiled above, inserted towards the stem
base. Stem with 1 main axis or with 1—3(-6) fairly

564

ANNALS OF THE MISSOURI BOTANICAL GARDEN
"X
Si

[Vor. 74

A B H. brevituba
H. virgata
subsp. virgata

>
HER. subsp. karooica JN
"r.i 4 D>
| — N
QUAE
id As

FIGURE 10. Morphology of Hexaglottis brevituba and the distribution of H. brevituba and H. virgata. Habit
x 0.5; flower and corm full size; detail of stamens, ovary, and style branches x

1987]

long, strongly diverging secondary branches,
straight to slightly flexuose, the lateral rhipidia
sessile, each partly to almost entirely enclosed
by the subtending stem bract, this usually as long
or longer than spathes. Rhipidia sessile except
the terminal, 2—3-flowered; spathes herbaceous,
or partly to entirely dry and pale at flowering
time, attenuate, 22-30(-37) mm long, the outer
concealed by a stem bract, often membranous
below, !1⁄— as long as the inner, upper part of
the inner extending above the bracts. Flowers
stellate with the tepals united below into a closed
tube, upright or secund, pale yellow, evidently
odorless, the nectar guides deeper yellow; peri
anth tube cy narrow, usually slightly curv-
ing outward, (3-)4—6(-9) mm long, usually at least
partly enclosed in the spathes; tepals 14—25
(subsp. virgata), 23-32 (subsp. karooica) mm
long, with short claws 1-1.5 mm long, the limbs
extended horizontally, 4-8(-11) mm wide; the
outer larger, lanceolate to ovate, the inner small-
er, lanceolate to cuneate. Filaments 3.3-6 mm
long, united for 1-2 mm; anthers initially 6—9
mm long. Ovary 8-16(-20) mm long, the upper

in the spathes, the pedicel short, 5-6 mm
style arms 5-6 mm long. Capsule narrowly spin-
dle-shaped, included in the spathes, 9-13 mm x
2-3 mm (subsp. virgata), or 16-22 mm x 4mm
(subsp. karooica), only 1 (rarely 2) developed in
each rhipidium; seeds narrowly angular, 0.7-1
mm wide and 1-2 mm long. Chromosome num-
ber 2n = 12, 10 (subsp. virgata), or 2n = 14
(subsp. karooica).
Flowering time. (Late September-)October-
December(—mid January); flowers opening 3:00-
:30 P.M. (or up to 4:00 P.M. on cooler days),
beginning to fade ca. 6:30 P.M., usually collapsed
by 7:00 P.M

Distribution. Hexaglottis virgata is wide-
spread in the southern African winter rainfall
area, extending from the Nieuwoudtville district
in the northwest through the western and south-
ern Cape to Port Elizabeth in the east (Fig. 10).
It is rather scattered in the west of its range, but
common from Malmesbury and the Cape Pen-
insula eastwards through the southern Cape. It
is found more often on heavier soils, particularly
shales of the Malmesbury System, but also occurs
on granitic substrates. Occasionally H. virgata is
found on sandy soils of the Cape System. Sub-
species karooica occurs inland on the Roggeveld

GOLDBLATT —SOUTHERN AFRICAN HEXAGLOTTIS

565

Escarpment, where it favors sheltered and damp
situations.

Diagnosis and relationships. Hexaglottis vir-
gata is a distinctive species, unmistakable in its
slender, relatively long perianth tube some 4-7
mm in length and very straight stems with
strongly diverging lateral branches. The historic
confusion about the identity of this species has
been due to the poor observation that resulted
in overlooking the perianth tube or confusing it
with the slender ovary (Lewis, 1959). In fact the
upper part of the ovary is narrow, sterile, and
tubular, but this is clearly different from the peri-
anth tube from which it is separated by an ab-
scission layer. The only other species of Hexa-
glottis with a perianth tube is H. brevituba,
described in this paper. Hexaglottis brevituba has
a much shorter tube, only 1-2 mm long, com-
pared with a tube (3-)4-7 mm long in H. virgata.
The two species differ in several other features.
Hexaglottis virgata has an ovary entirely includ-
ed in the spathes on a short pedicel up to 5 mm
long, while H. brevituba has a longer pedicel 7—
10 mm long, and the ovary is curved and usually
just exserted from the spathes.

Two subspecies of Hexaglottis virgata are rec-
ognized here. The typical and most common,
subsp. virgata, has relatively small flowers and
capsules. It has a wide distribution extending
over almost the entire range of the species. Subsp.
karooica, known only from two isolated sites on
the Roggeveld Escarpment, has unusually large
flowers, spathes, and capsules.

History. The earliest existing records of Hexa-
glottis virgata are those made by the Swedish
botanists Carl Peter Thunberg and Anders Sparr-
man in the 1770s, but this common southwest-
ern Cape species must surely have been collected
earlier than this. However, it was Nicholas Jac-
quin who first described the species in 1791, based
on plants grown in Vienna, and probably sent to
him some years earlier by Franz Boos and Georg
Schol, the collectors who provided Jacquin with

many of the Cape plants that he illustrated and
described. The painting that accompanies the de-
scription and serves as the type in the absence
of preserved material is excellent and leaves no
doubt about the identity of what Jacquin called
Moraea virgata. After the genera Homeria and
Hexaglottis tinct from Mo-
raea by E. P. Ventenat in 1808, M. virgata was

566 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

SUBSP. KAROOICA

A

SUBSP. VIRGATA

FIGURE 11. Morphology of sre quia Is —A. Subsp. virgata. —B. _Subsp. karooica. Habits x 0. 25;
corms x 0.5; flowers, fruiting branch (su
and gynoecium full size; single capsule ale pn es virgata) + x 2:

1987]

transferred to Homeria (Sweet, 1827) and shortly
afterward to Hexaglottis (Sweet, 1830). J. G.
Baker included H. /ewisiae (as H. flexuosa) and
H. longifolia in H. virgata in Flora Capensis
(1896). This unsatisfactory treatment was cor-
rected by G. J. Lewis in 1950.

KEY TO THE SUBSPECIES OF
HEXAGLOTTIS VIRGATA

la. Outer tepals 14-22(-25) mm long x 5-7
(711) mm wide; spathes (at least the terminal)
ERU mm long; foliage leaves 1—3, rare-

E subsp. virgata

0.5

1b. i dien 23-32 mm long x 7-1
wide; spathes (at least the zn 32-37
mm long; foliage leaves (2—

ns subsp. karooica

6A. Subsp. virgata. Figure 11A.

Hexaglottis virgata var. lata G. Lewis, J. S. African

t. 25: 228-229. 1959. TYPE: South Africa. Cape:

Welbedacht, Bidouw Valley, Lewis 2514 (holo-
type, SAM 61821; isotypes, BOL, PRE, SAM).

Plants 12—50(—70) mm high. Leaves 1—3(-4).
Spathes 22-28(-32) mm long. Flowers with peri-
anth tube (3—)4-5.5(-7) mm long; tepals 14-
22(-25) mm long. Filaments 3.3-6 mm long; an-
thers 5-8 mm long. Ovary 8-12 mm long; cap-
sules 9-13 mm long. Chromosome number 2n =
12, 10.

Distribution. The distribution of subsp. vir-
gata is the same as that for the species, except
that it does not occur on the Roggeveld Escarp-
ment (Fig. 10)

Variation. Ascircumscribed here, subsp. vir-
gata includes the variety /ata, described by G. J.
Lewis for somewhat larger-flowered plants with
comparatively wide tepals, from the Biedouw
Valley. When this was described in 1959, no oth-
er population of H. virgata was known from the
northwestern Cape, and only one was known from
Namaqualand, a poorly preserved specimen,
which Lewis included in var. /ata despite the
distorted flowers that could not be accurately
measured. Hexaglottis virgata is now known from
several sites in the northwest Cape, from the Oli-

a
tion in flower dimensions in these jomdan
is considerable and covers the whole range from
typical rather small-flowered subsp. virgata (te-
pals 14-20 x 5-8 mm; spathes 22-28 mm long),
which occurs in the Cape Peninsula and southern
Cape, to the type of var. lata and very similar
plants from Nieuwoudtville (tepals 20-25 x 9-

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

567

11; spathes 23-30 mm long). The rationale for
recognition of the variety has thus disappeared.
These northwestern populations do, however,
appear to represent a distinct race. Plants from
two populations from the northwestern area have
a karyotype with strongly acrocentric chromo-
somes (Fig. 3F) that contrast with the presence
of acrocentrics and submetacentrics in the most
common southern cytotype (Fig. 3E

Populations examined from the Cape Penin-
sula have n = 5, and a very different karyotype
from those Si n = 6 (Fig. 3G) (see discussion
under Cytology). There seems to be no corre-
sponding ipic grid difference in the Cape
Peninsula plants.

Specimens examined. SOUTH AFRICA. CAPE-31.19
(Calvinia): Grasberg road northwest of Nieuwoudt-
ville, renosterveld (AC), Goldblatt 707 1 (K, MO, NBG,
PRE, US); 2-3 km from Nieuwoudtville on the north
side of hos road to the escarpment, Goldblatt 7411
(MO); 8 km south of Nieuwoudtville on sandstone
on oo 7395 (MO

8 (Clanwilliam): 10 km south of Clanwilliam,
j a bank (BB), Goldblatt 6705 (MO, PRE); 12
km south of Clanwilliam, stony clay bank, Goldblatt
6989 (K, MO, de PRE, WAG); clay hillside wi
south of the Alp rnoff, on the National Road t
Citrusdal (BD), pone 3028 (MO).

32.19 (Wuppertal): between Doorn River and Bi-
douw Valley turnoff, stony clay (AA), Goldblatt 5941
(MO); Welbedacht, Bidouw Valley, Lewis 2514 (BOL,
PRE, SAM

33.18 (Cape Town): Table Mountain (CD), Tyson
2488 (SAM), Bayliss 3054 (UC); kloof between Lions
Head and Table Mountain, Burchell 252 (K); slopes of
Devils Peak, above de Waal Drive, Cape Town, Gold-
blatt 67 17 (MO); near Bishopscourt, Salter 9001 (BOL);
Wynberg Hill, Pillans 10819 (MO, UPS), Salter 8978
(SAM), 9545 (BM); Signal Hill, Lewis 665 (SAM),
Goldblatt 71 (J), 6747 (MO, S), 6748 (MO), Marloth
7234 (PRE); Camps Bay, Moss 13403 (J); Observatory
grounds, Davis s.n. (SAM 61050); Tygerberg Nature
Reserve (DC), Loubser 3004 (MO); Stellenbosch (DD),
Boucher 3392 (PRE, STE); 4 miles from Faure on the
Stellenbosch road, Lewis 2340 T Groot Dra-
E und fuss ue POR Drége s.n

(Worcester): farm Waterval, near ‘Porterville
(AA), Loss 966 (NBG); Tulbagh Cemetery (AC),
Goldblat aged K, MO, S); 9 miles along the

Leeuwfon ad (AD) Pearson Deg (K); Bains Kloof

(CA) uri tie 9106 (BM, BOL B

west of (ox saa near the
G, PRE); Karoo Gar-

orcester, Compton 17849 (NBG), Lewis 5304
DEM Pok i Rawsonville, /e Roux s.n. (PRE); be-
tween Worcester and Robertson (DA-DB), Zinn s.n.

ice dea 3).
0 (Montagu): pasture pee 10 O’Clock Mt.,
Svellendam (CD), Wurts 486 (NBG).

3.21 (Ladismith): south dign to Attaquas Kloof
m 4 Thompson 1636 (PRE), Attaquaskloof, Mossel
Bay, Barker 7692 (NBG).

568

33.23 (Willowmore): hills near Uniondale (CA), H.
Bolus 2484 (Ky; Prince Alfreds Pass (CC), Wall 18
(LD); near Misgund (CD), Goldblatt 6792 (MO); be-
tween Misgund Net Nieuweplaats Pa Fourcade 5485
(PRE, STE); Kliprivier, Tsitsikamma Park, Bower 625
(PRE); hills hear Jouberti na (DD), Fourcade 2374 BOL,
K); Joubertina, B BOL)

33.25 (Port Elizabeth): Van Staadens Hoogte (CC),
MacOwan 2055 (BM, K); Greenbushes (CD), Holland
4051 (BOL); Baakens River Valley, Port Elizabeth (DC),
Olivier 1739 (WAG); Port Elizabeth, Long 502 (K),
—— sub Rogers 2414 (J).

4.18 (Simonstown): Steenberg (AB), Compton 1665
ER Bergvliet, Purcell s.n. (BOL 1638, SAM), 124
(SAM); Somerset West, on stiff clay soil (BB), Parker
4388 (BOL, K, MO, NBG).

34.19 (Caledon): Houw Hoek (AA), Penther 572 (K,
S); Greyton-Genadendal (AB), Lindeberg s.n. (S); Na-
pier Ruggens (BD), Marloth 10006 (PRE); between Na-
pier and Bredasdorp (CA), Goldblatt 6937 (MO);
Franskraal, along the coast above the beach (CD),
Goldblatt 5368 (MO); sandy soil on slopes near Avoca
(DA), Goldblatt 6939 (MO, PRE, S, WAG); Bredas-
dorp Poort (DB), Esterhuysen 19580 (BOL).

34.20 (Bredasdorp): Storms Vlei Kloof (AA), Lei-
poldt 3549A (BOL); Bontebok Park, Swellendam (AB),
Liebenberg 6710 (STE); Zuurbraak (BA), Barker 5029
( ); Grootvadersbos, paths in wood (BB), Willems

shale hills west of Heidelberg, ren

7416 (MO); Potteberg (BC), David s.n. n BG);
=a distr., Leipoldt 3550 (B

21 (Riversdale): hill top 5 km west í Riversdale
Ps Goldblatt 5436 (MO); Still Bay, limestone hills
(AD), Esterhuysen 19538 (BOL, PRE); limestone hills
south of Albertinia, Stilbaai road on turnoff to Riet-
huiskraal, Goldblatt 7428 (MO); Onverwacht, Alber-
tinia (BA), Muir 1207 L).

34.23 (Knysna): Plettenberg Bay (AB), Rogers 28241
(K), 26762

Without precise locality: CBS, A/exander s.n. (BM,
K); Sparrman s.n. Iris edulis (S); Thunberg s.n. Iris
edulis (S)

6B. Subsp. karooica Goldbl., subsp. nov. TYPE:
South Africa. Cape: eastern border of farm
Blomfontein, 22 km from Middelpos to-
wards de Hoop, Snijman 765 (holotype,
NBG; isotypes, K, MO, PRE). Figure 11B.

Planta robusta, 30-85 cm alta, ws (2—)4—5, spathis
32-37 mm longis, ind) arag 1 6.5-9 mm longo,
tepalis 23-32 mm lon -11 mm S antheris 6—8

mm longis, capsulis TEN mm long

Plants robust and 30-85 cm high. Leaves usu-
ally 4-5, rarely 2-3. Spathes 32-37 mm long.
Flowers with perianth tube 6.5-9 mm long; tepals
23-32 mm long, 7-11 mm wide. Filaments 4—6
mm long; anthers 6-8 mm long. Ovary 15-20
mm long. Capsules 16-22 mm long. Chromo-
some number 2n = 14.

Distribution. Subsp. karooica is known from
two widely separated localities along the Rog-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

geveld Escarpment, near Blomfontein west of
Middelpos in the north, and on the slopes of
Sneeukrans on the farm Voelfontein northwest
of Sutherland, some 50 km to the south. The
habitat at both sites is moist, the plants growing
among large rocks near seepage zones.

Variation. The subspecies is variable in flow-
er size. Plants from the Middelpos area have very
large tepals 28-32 mm long while those from
Voelfontein have smaller flowers with the tepals
somewhat narrower and only 23-24 mm long.
Despite this variation, other features, including
large spathes and capsules, higher leaf number,
and the unusual karyotype with 27 = 14, in both
populations indicate that subsp. karooica is in-
deed a distinct geographical variant of Hexa-
glottis virgata.

Specimens examined. SOUTH AFRICA. CAPE-31.20
(Williston): Roggeveld Escarpment, 71 km SE of Cal-
vinia on Middelpos road via Blomfontein (CC), Gold-
blatt 4612 (MO, NBG); eastern border of farm Blom-
fontein, 22 km from Middelpos towards de Hoop,
Snijman 765 (K, MO, NBG, PRE).

32.20 (Sutherland): Roggeveld, Sneeukrans south of
Voelfontein farm, ca. 4,500 ft., in wet site (AD), Gold-
blatt 6336 (MO), 7126 (MO).

LITERATURE CITED

BAKER, J. G. 1892. Handbook of the Irideae. George
Bell & Sons, London
1896. Irideae. In W. T. Thiselton-Dyer (ed-
itor), dis DRUMS 7-171. Reeve & Co., Ash-
d, Ken

1932. Novitates Africanae. Ànn.
9.

Opera Bot. 40: 1-85.
. The systematics of Montanoa
(Asteraceae, Heliantheae). Mem. New York Bot.
Gard. 36: 1-133.
GOLDBLATT, E 1971a. A new species of Gladiolus
and some nomenclatural changes in the Iridaceae.
1 S. African Bot. 37: 229-236.
1971b. Cytological and morphological pud
ies in the southern African Iridaceae. J. S. Afric
Bot. 37: 317—460.
976a. Evolution, cytology and subgeneric
classification in Moraea (Iridaceae). Ann. Mis-
souri Bot. Gard. 63: 1-23.
1976b. The genus Moraea (Iridaceae) in the
winter rainfall area of í Africa. Ann. Mis
ri Bot. Gard. 63: 657-
1979. Chromosome Am and karyotype
change in Galaxia (Iridaceae). Pl. Syst. Evol. 133:
61-69.

Redefinition of Homeria and Moraea
(Iridaceae) in the light of eat sui data, with
Rheome gen. nov. Bot. 9.
1981. Systematics aa biology of Homeria
pes Ann. Missouri Bot. Gard. 68: 413-

1987]

. 1982. A synopsis of Moraea (Iridaceae) with

new taxa, transfers and notes. Ann. Missouri Bot.

rue 69: 351-369.

Systematics of the southern African

idaceae — Ixioideae). Ann
77-447

6. Convergent evolution of the Homeria
flower type in six new species of Moraea (Irida-
ceae — Irideae) 7 eae Africa. Ann. Missouri
Bot. Gard. 73:
oe E. P ZIMMER. 1984. Estima-
tion of genome size (C-value) in Iridaceae by cy-
tophotometry. Ann. Missouri Bot. Gard. 71: 176-
l

, M. & L. E. Copp. 1981. Botanical se
of Southern Africa. A. A. Balkema, e wn.
66. Phylogenetic Systematics. Uni
Illinois Press, Urbana, Illinois. [English 1 tine
tion by D. D. Davis & R. Zangerl.]

HUMPHRIES, C. J. 1981. Cytogenetic and cladistic
studies in tue are (Compositae: Anthemideae).

Nordic J. Bot. 1: 83-96.

JACQUIN, N. 1776. Horus Botanicus Vindobonensis
3. Kaliwoda, Vien

91. oe tone virgata. In Icones Plantarum

Rariorum 2: tab. 228. Wappler, Vienna.

GOLDBLATT—SOUTHERN AFRICAN HEXAGLOTTIS

569

Krarr, F. W. 1866. Revisio Iridearum (Conclusio).
Linnaea 34: 537—689.

. 1882. Ergánzungen und Berichtigungen zu

Baker's Systema Iridacearum. Abh. Naturf. Ges.

Halle 15: 337-404

895. Irideae. In Th. Durand & H. Schinz

(editors), Conspectus Florae Africanae 5: 143-230.
Jardin Botanique de l'État, Bruxelles.

Lewis, G. J.

Cape Peninsula. Juta & Co., Cape Town.

South African Iridaceae. A revision of
Hexaglottis. J. S. African Bot. 25: 215-230.
LINNAEUS, C. (FiL). 1781 [1782]. Supplementum
Plantarum. Orphanotropheus, Braunschweig

A. 1812. On the cultivation of rare
plants. Trans. Hort. Soc. London -366.
1827. Hortus Britannicus,

Ist edition.
Ridgway, London
pi. _ Hortus Britannicus, 2nd edition. Ridg-

y
VENTENAT, E P. "1808. Decades Generum Novorum
Dufarb, Paris.

NOTES ON THE VARIATION AND TAXONOMY OF
WATSONIA BORBONICA (W. PYRAMIDATA, W. ARDERNEI)
(IRIDACEAE) IN THE SOUTHWESTERN CAPE,
SOUTH AFRICA!

PETER GOLDBLATT?

ABSTRACT

Lomenia borbonica Pourret (1788) is an earlier name for the common Cape species formerly called

Watsonia pyramidata, and the new combination in Watsonia is made here. Significant
either declinate or arcuate, characterizes southern and
=e ble. The whi red W. ar
uced to synonymy. DUE synonyms of W. borbonica
ted.

Melee in the stamen and style orientation,
rthern races which otherwise seem indist
an n albino sport of the northern an

t and unusual

ite-flowe ernei is considered

are discussed and the complete Musicien of the species is presente

The name Watsonia borbonica is a new com-
bination based on Lomenia borbonica Pourret
(1788), only species of Lomenia Pourret. The
epithet was given to a plant believed to have been
collected on the Isle de Bourbon, now called Ré-
union. Lomenia has long been misunderstood
and was for a time considered a synonym of
Freesia Klatt, a southern African genus. During
a study of Freesia (Goldblatt, 1982), I discovered
that Lomenia was congeneric with Watsonia.
Based on the illustration in the protologue, I con-
sidered L. borbonica conspecific with W. mar-
ginata (L. f.) Ker, the basionym of which pre-
dates Lomenia. Later, I found the type specimen
of L. borbonica at the Paris Herbarium (P) in
1985. The specimen is in excellent condition,
and there can be no doubt that it is identical with
the species presently known as W. pyramidata
(Andrews) Klatt, and in the Flora Capensis as
W. rosea Ker (Baker, 1896). The type collection
is attributed to Philibert Commerson, who col-
lected in the Mascarene Islands in 1770-1774.

of this and a few other Cape plants attributed to
Commerson's collection remains unknown.

Nothing could be more inappropriate than for
a native southwestern Cape species to have a
specific epithet ‘borbonica’. However, there is no
alternative but to use the earliest valid and le-
gitimate name for a species as mandated by the
1983 Code of Botanical Nomenclature.

HABITAT AND VARIATION

Watsonia borbonica is a fairly common and
well-known species of Watsonia of the south-
western Cape, South Africa. It is usually found
in mountain habitats and is locally common and
conspicuous after fires on lower slopes, but it also
occurs on sandy flats and at relatively high al-
titudes up to 1,000 m. Its range extends from the
Cape Peninsula eastward to Bredasdorp and lo-
cally north through the du Toits Kloof moun-
tains to the Breede River floodplain near Ro-
mans River. Plants flower in the early summer,
from October to early December and at high
elevations into January. They prefer rocky sand-
stone soils but may be found on soils derived
from granite.

Although Watsonia borbonica is not particu-
larly variable, what may be called the northern
and southern races differ in one unusual feature,
the orientation of the stamens and style. The
southern and typical populations have declinate
stamens that lie against the lower (anterior) tepal,
and when the style is receptive it lies below the
stamens. Northern populations from Paarl
Mountain, du Toits Kloof, and the Breede River
valley have unilateral stamens that lie horizontal
to somewhat arched under the upper (posterior)
tepal, and the style lies between the stamens and
the upper tepal. In all other features the plants
appear to be morphologically identical and can-
not be distinguished even on close inspection.

1 oe by Grant DEB 81-19292 from the U.S. National Science Foundatio
. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri

eee U. S.A.
ANN. Missouni Bor. GARD. 74: 570—572. 1987.

1987]

The diagnostic features of Watsonia borbonica
are its large flowers with subequal, pink tepals
30-36 mm long; a short, flared upper perianth
tube 8-12 mm long; robust and branched habit
(1-2 m tall); oblong-ovate capsules 20-3
long; and seeds with two wings. Pink-flowered
specimens of the northern form have always been
included in W. borbonica (as W. pyramidata).
However, a white-flowered sport believed to have
come from the northern part of its range has been
recognized as W. ardernei, a plant that has been
in cultivation for close to 100 years and is the
best-known cultivated Watsonia species. Wat-
sonia ardernei is here reduced to synonymy in
W. borbonica.

The significance of the variation in stamen and
style orientation is uncertain. It is clear that dec-
linate stamens and style are unusual in Watsonia,
occurring in only six other species. Arcuate sta-
mens are more common and are found in all but
one of the remaining ca. 45 species of the genus
and in many other genera of Ixioideae. Presum-
ably the declinate condition, rare in Iridaceae, is
derived directly from the more common arcuate
state and not independently from an ancestor
with symmetrically disposed stamens. Despite
the apparently gross morphological difference in
stamen and style orientation between the north-
ern and southern races of W. borbonica, I believe
that they should be treated as a single species. I
defer recognition of intraspecific taxa until my
monograph of Watsonia, currently in prepara-
tion, is completed.

OTHER SYNONYMS

Among other species I regard as synonyms of
Watsonia borbonica is W. cooperi G. Lewis, based
on the illegitimate homonym Tritonia cooperi

aker. The type is a fragment comprising a lat-
eral branch of the spike and three poorly pressed
flowers. The leaf mounted with the flowering ma-
terial has three conspicuous veins and is clearly
misplaced, belonging either to a species of Ana-
palina N. E. Br. or Tritoniopsis L. Bolus. How-
ever, the flowering material seems to belong to
W. borbonica. The tube and tepals are much dis-
torted, but the stamens seem declinate, and the
dimensions of the tepals and perianth tube ac-
cord with W. borbonica.

The reasons for treating Watsonia ardernei
Sander and W. ardernei Hort. ex Mathews & L.
Bolus as conspecific with W. borbonica have al-
ready been outlined, but the existence of the

GOLDBLATT — WATSONIA BORBONICA

571

homonyms applied to the same species is puz-
zling. Possibly Mathews & Bolus believed that
the name as described by Sander in a nursery-
man’s catalogue was not valid, hence their formal
recognition of a name already in use. The white-
flowered form, originally collected at Romans
River near Wolseley, on which W. ardernei (both
of Sander and of Mathews & Bolus) is based is
no longer known in the wild. It probably com-
prised one or a few individuals that were intro-
duced into gardens and then multiplied. The plant
named W. iridifolia var. obrienii by N. E. Brown
is based on cultivated plants from the same source
as the type material of W. ardernei.

The complete synonymy of W. borbonica is
outlined below. All type specimens cited have
been seen.

SYNONYMY

Watsonia borbonica (Pourret) Goldbl., comb.
nov. Lomenia borbonica Pourret, Hist. &
Mem. Acad. Roy. Sci. Toulouse (Acta Acad.
Sci.) 3: 74 & tab. 5. 1788. TvPE: said to have
been collected on Réunion (Ile de France)
but clearly from the SW Cape, Commerson
s.n. (holotype, P).

Gladiolus pyramidalis Lam. egg Meth. 2: 726. 1786.
wis :

patens Aiton (Lewis et al.,

fete oo precise locality unknown, collector
n (holotype, P—Herb. Lamarc
G lado iridifolius varietas speciosa floribus roseis
. Pl. tab. 235. 1793.

A HAM a (Ker) Ecklon, Top. Verz. 37. 1827, nom.
illeg., genus sine descr.

Watsonia E (Andrews) Klatt, Durand &
S . Africa 5: 194. 1895; Stapf, Bot.
Mag. edm pes 0261. 1931; L. Bolus, Fl. Pl. Africa
25: tab. 974. 1946; Lewis, Flora Cape Peninsula
238.1950. Gladiolus pyramidatus Andrews, Bot.
Rep. 5: tab. 335. 1803. Watsonia Hae Banks ex
Ker, Konig & Sims, Ann. Bot. 1: 230. 1804, Bot.
Mag. tab. 1072. 1807; Baker, Handbk. Irid. 158.

Africa. Cape: precise locality unknown, 1
= in Bot. Rep. tab. 335 (lectotype here desig-
ated).
Watsonia striata Klatt, Abh. Nat. Ges. Halle 15: 352.
(
elieved Ses hd Identity determined

from descriptio
— iridifolia var. pom ii N. E. Br., Gard. Chron

se 0. 1889; Klatt, Durand & Sch inz, Con-
= L l Africae 5: 193. 1895. TYPE: South Africa.
Cap t known; authentic material present at
Her

. Kew (probably originally from Romans
River farm, Tulbagh Div., flowered at Kew from

plants sent from St. George’s Park, Port Eliza-
beth).

W. obrienii van Tubergen, Gard. Chron. ser. 3,
01. 1894, nom. nud. (not a valid pee A
based on W. iridifolia var. obrienii as suggested
by Marais, Kew Bull. 35: 172. 1980).

Tritonia cooperi Baker, Handbk. Irid. 192. 1892, Flora
Cap 122.1896, hom. illeg., non Klatt (1882),

which is Tritonia cooperi (Baker) Klatt. Watsonia
iL. Bo Bo :

cooperi lus, J 67: 135. 1929; Lewis, J
S. African Bot. 7: 55. 1941. Tritonia quinquener-
vata Foster, Contr. Gray. H 36,

comprises a flowering stem and poorly pressed
flowers of W. borbonica and a leaf of a species of
Anapalina or rii

Watsonia ardernei Sander w Plants for d ie
20 & fig. 18

Watsonia ardernei Hort. ex Mathews & L. Bolus, Ann.
Bolus Herb. 4: 25-26. 1925; Fl. Pl. Africa 19: tab.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

750. 1939, hom illeg., non W. ardernei Sander
Sane due South Africa. Cape: Romans River
farm, Tulbagh Distr. (cult. Kirstenbosch), Tred-
midan n. we an atra BOL 17839, here Mcd E
isolectotypes, BM (as Arderne s.n.), BOL).

LITERATURE CITED

BAKER, J. G. 1896. Irideae. /n W. T. Thiselton-Dyer
(editor), Flora Capensis 6: 7-171.

GOLDBLATT, P. 1982. Systematics of Freesia Klatt
(Iridaceae). J. S. African Bot. 48: 39-91.

Copp. 1981. Botanical Exploration

1788. Description de deux nouveaux
genres de la famille des Liliacées désignés sous le
nom de Lomenia & de Lapeirousia. Hist. & Mém.
Acad. Roy. Sci. Toulouse (Acta Acad. Sci.) 3: 73-
78.

NOTES ON THE FLORAL BIOLOGY, CYTOLOGY, AND
EMBRYOLOGY OF CAMPYNEMANTHE
(LILIALES: CAMPYNEMATACEAE)!

PORTER P. Lowry II,? PETER GOLDBLATT,? AND HIROSHI TOBE‘

ABSTRACT

Field observations of the endemic New Caledonian genus Campynemanthe (Liliales: Campynema-
Pisin indicate that at least C. neocaledonica and C. viridiflora are strongly p oe and have an

he filaments. A chromosome number

of n= 11 has been determined

for both species from meiosis in 1 pollen mother cells, these being first records for the genus and family.

Endosperm formatio
putatively related Melanthiaceae.

Campynemanthe Baillon comprises three
species restricted to the Pacific island of New

Campynem a Tabi. and
followed Dahlgren & Clifford due and Dahl-
gren & Lu (1985) in placing both genera in Cam-
pynemataceae. The family, which appears to oc-
cupy an unspecialized position in Liliiflorae, is
possibly most closely related to Melanthiaceae

mall greenish flowers with persistent and ac-
crescent tepals, partly inferior ovaries, and free
stylodia. The embryology of both genera of Cam-
pynemataceae is of the basic type for the mono-
cots (Dutt, 1970; Dahlgren & Lu, 1985), except
that the endosperm development is now known
to be nuclear, a derived condition.

We report here some observations on the floral
biology of two species of Campynemanthe, in-
cluding the occurrence of protandry and an un-
usual postanthesis behavior of the persistent fil-
am m

counts for the genus and family, em we observed
nuclear spect ofthe

r

n in C. neocaledonica is nuclear rather than helobial as in several genera of the

embryology of Campynemataceae previously
unknown.

FLORAL BIOLOGY

Recent field observations indicate that Cam-
pynemanthe neocaledonica (Rendle) Goldblatt is
strongly protandrous. This phenomenon was ini-
tially observed in December 1985, in a popula-
tion of about 100 individuals covering an area
of ca. 20 m? located at 900 m on the Plateau de

ers exhibited discrete male and female phases of
sexual expression. In the initial, male phase the
straight filaments are erect to ascending and are
usually twisted slightly counterclockwise so that
the oblong to ovate anthers alternate with the
tepals (Fig. 1a). During this phase the three small
stylodia are erect and closely appressed, formin

(Fig. 1b)
siderably and recurve to expose the whitish stig-
matic surfaces.

hile there appears to be no overlap in the
sexual phases within a single flower, flowers of

! We thank J. H. Beach for many helpful suggestions on the manuscript, and J. K. Myers for preparing the
illustration. Assistance was provided in New Caledonia by L. B. Thien, Ph. Morat, J.-M. Veillon, ORSTOM-

ouméa

, and the Service des Foréts et du Patrimoine Naturelle. Work by P

. Lowry was supported in part by

NSF Doctoral Dissertation Improvement Grant BSR83-14691, the Missouri Botanical Garden, the W. purs
Jones Foundation, and the Division of Biology and Biomedical Sciences of Washington University, St. Loui

? Missouri Botanical Garden, P.O. Box 299, St.
Phanérogamie,
? B.

63166- 0299.

Louis, Missouri 63166-0299, U.S.A.;
Muséum National d'Histoire Naturelle, 16, rue Buffon, 75005 Paris, France
Krukoff Curator of African Botany, Missouri Botanical Garden, P. O. Box 299, St. Louis, Missouri

and Laboratoire i

* Biological Laboratory, Yoshida College, Kyoto University, Sakyo-ku, Kyoto 606, Japan.

ANN. MISSOURI Bor. GARD. 74: 573-576. 1987.

574

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

2mm

Flowers of Campynemanthe neocaledonica (Lowry 3945).—a. Male phase.—b. Female phase.

FIGURE 1.

both phases frequently occur within individual
inflorescences. Geitonogamy thus appears pos-
sible, assuming at least some self-compatibility.
There did not appear to be synchrony of flow-
ering among the individuals in the population,
although observations were made on only a sin-
gle da

It has since been possible to confirm protandry
in another population of Campynemanthe neo-
caledonica from Haute Ouinné (Lowry & Suprin
3698) from photographs taken in the field. Sim-
ilarly, two distinct floral phases can be seen in
photographs of C. viridiflora Baillon taken at 950
m on the Montagne des Sources (Lowry 3762)
and at 1,350 m on Mt. Humboldt (Lowry 3812),
indicating that this species is dichogamous and
almost certainly protandrous. Unfortunately, we
have not been able to determine whether pro-
tandry occurs in the third member of the genus,
C. parva Goldblatt, or in Campynema lineare

anthers, as figured by
Dahlgren & Lu (1985) and reported by Goldblatt
(1986).

Bawa & Beach (1981) and Lloyd & Webb
(1986), who have reviewed the evolutionary as-
pects of dichogamy in general, and protandry in
particular, concluded that dichogamy in self-
compatible species permits pollen and stigmas
to be positioned close to one another without
high levels of self-pollination. They also pointed
out that protandry is by far more common than
protogyny in biotically pollinated species, which
is presumably the case for C. neocaledonica. We

did not observe floral visitors on species of Cam-
pynemanthe.

CHROMOSOME NUMBER

A haploid chromosome number of n = 11 was
determined from pollen mother cells in flower
buds of Campynemanthe neocaledonica and C.
viridiflora. Inflorescences of these species were
fixed in the field with 3:1 ethanol-acetic acid
and squashed with FLP orcein. The relatively
large chromosomes were ca. 3 um long at meiotic
metaphase but exhibited no distinctive details.

Chromosome numbers in Melanthiaceae are
diverse. The counts summarized in Table 1 were
taken from the /ndex to Plant Chromosome
Numbers series (Goldblatt, 1985, et praec.)
and unpublished work (Ambrose, 1975; Bodkin,
1978). Tofieldieae apparently have x = 15, the
only known base number in the monotypic gen-
era Pleea and Harperocallis, and the most com-
mon one in Tofieldia. Tofieldia coccinea Rich-
ards. has x = 16, a count confirmed by a number
of workers, but several other species have x —
15. Counts of n = 14 and n = 15 have been
reported for T. calyculata (L.) Wahlenb. Japan-
olirion, which perhaps belongs in Tofieldieae
(Utech, 1984), has n = 12

In Melanthieae x = 8 is the most frequent base
number and the only one in Melanthium,
Schoenocaulon, and Veratrum. There are two
distinct series in Zigadenus: n = 16 in Z. elegans
Pursh, Z. glaucus Nutt., Z. sibiricus (L.) A. Gray
ex S. Wats., Z. nuttallii A. Gray ex S. Wats., and
Z. volcanicus Benth.; and n = 11 in Z. fontanus

1987]

Eastw., Z. fremontii (Torr.) Torr. ex S. Wats., Z.
brevibracteatus (M. E. Jones) Hall, and Z. ve-
nenosus S. Wats. Amianthium has x = 16 (Utech,
1986), and Stenanthium has x = 8 and a puzzling
second base of x =
Narthecieae, the largest tribe in the family, ap-
pears to be paleotetraploid. Narthecium, Aletris,
Nietreria, an x = 13, while
Helonias and Heloniopsis, now usually segregat-
ed as a distinct tribe (Utech, 1984) or even fam-
ily, have x = 17. Lophiola, perhaps allied to Nar-
gi Sr P n = 21.
The s genus Chionographis, placed in
Chionographidea (Dahlgren et al., 1985), has
= n C. koidzumiana Ohwi and C. japonica
PO) Maxim. var. japonica, but C. japonica
var. kurohimensis Ajima & Satomi has n = 22,
and subspp. minoensis (Hara) Hara and hisu-
achiana (Okuyama) Hara have n = 21. The most
parsimonious interpretation here is a base num-
ber o = 12, with reduction to x = 11 and
polyploidy resulting in n = 22, then reduction to
= 21. The taxonomically isolated XeropAyllum
es ie has x =
Summarizing, the available counts suggest to
us a possible ancestral base number of x = 8 for
Melanthiaceae, persisting only in Melanthium,
Schoenocaulon, Veratrum, and Stenanthium (all
Melanthieae). Other genera and tribes are ap-
parently paleotetraploid, with Tofieldieae having
x = 15 l

°

= 17 reported in Helonias and Heloniopsis 1S
difficult to reconcile with other members of the
Narthecieae. The odd counts of n = 11 in Zi-
gadenus, which has several species with x — 16,
and n — 20 in Stenanthium, which also has n —
8, remain to be explained as well.

Our report of n = 11 for Campynemataceae,
which suggests a base number of x = 11, does
not fit with any assemblage of Melanthiaceae.

sumably being secondary to the higher numbers
in the tetraploid series x = 16, 14, 13, 12, 11.

EMBRYOLOGY

Although it was not possible to pursue the de-
tails of early endosperm development in Cam-
pynemanthe, the available fixed material of C.

LOWRY ET AL.—CAMPYNEMANTHE

575

TABLE 1. Chromosome numbers recorded in Cam-
pynemataceae and basic chromosome numbers for
genera of Melanthiaceae summarized from the /ndex
to Plant Chromosome Numbers (Goldblatt, 1985, et
praec.) and supplemented by data in Ambrose (1975),
Bodkin (1978), and Utech (1984, 1986).

Taxon (species Base Haploid
counted/total species) Number Numbers
Campynemataceae
Campynema (0/1) Uncounted
Campynemanthe (2/3)
C. neocaledonica' 11 11
C. viridiflora' 11 11
Melanthiaceae
Tofieldieae
Pleea (1/1) 15 15
Harperocallis 15 15
1/1
Tofieldia (9/20) 15 15, 14, 16,
30
Japanolirion (1/1) 12 12
Melanthieae
Veratrum (11/25) 8 8, 16, 40
48
Zigadenus (8/15) 16 16, 11
Schoenocaulon 8 8
/10)
Stenanthium (2/2) 8 8, 10
Melanthium 8
Amianthium 16 16
Narthecieae
Narthecium (2/5) 13 13, 26
Aletris (7/25) 13 13, 26
Metanarthecium 13 26
(1/2)
Nietreria 13
Ypsilandra (0/5) Uncounted
Helonias (1/1) 17 17
Heloniopsis (5/5) 17 17
Lophiola (1/1) 21 (or 7) 21
Chionographideae
Chionographis 12 12, 21, 22
(2/5)

Xerophylleae
Xerophyllum (2/2) 15 15

! Voucher data for original counts: C. neocaledonica,
New Caledonia, Plateau de Dogny, 900 m, Lowry 3945
(MO, NOU, P); C. viridiflora, New Caledonia, Mt. Mou,
1,150 m, Lowry 3857 (MO, NOU, P

neocaledonica (Lowry 3945), which comprised
nearly 100 seeds that had developed past the 64-
nucleate stage, indicates that the endosperm of
this species is nucleate and not helobial (Stenar,

576

* ax

* `; °

r 4 e
vo SP

Lea apa s Sem í

€ ahs sz -

«M.

NEL

(c
*

Q
e

S š ^
Mt at 4
TASS o* Wise

WU Sa
BLES Ye
FIGURE 2. Longitudinal section of a young seed of

Campynemanthe neocaledonica showing free endo-

sperm nuclei at the 64-nucleate pedi s, synergid; end,

endosperm nucleus. Scale = 50 yu

1949; Eunus, 1951) as in other Melanthiaceae in
which this embryological feature is known (in-
cluding Tofieldia, Narthecium, Heloniopsis,
Metanarthecium, Aletris, Zigadenus, Veratrum,
and Amianthium). In Amianthium muscaetoxi-
cum Walt. the embryo sac in postfertilization
stages forms a micropylar and chalazal endo-
sperm chamber characteristic of the helobial type;
in an earlier stage the micropylar chamber con-
tains 16 nuclei, and the chalazal chamber 8 nuclei
(Eunus, 1951, fig. 31). In Campynemanthe, how-
ever, there appears to be no such. chalazal cham-
ri

r, no
nuclei at the chalazal end. All of the endosperm
nuclei occur within a single cell (i.e., embryo sac),

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

although th bundant toward the cha-
lazal end (Fig. 2). The presence of nuclear en-
dosperm in C. neocaledonica suggests that this
character may be useful in distinguishing Cam-
pynemataceae from Melanthiaceae.

LITERATURE CITED

AMBROSE, J. D. 1975. Comparative Anatomy and
Morphology of the Melanthioideae (Liliaceae).
Ph.D. Dissertation. Cornell University, Ithaca,
New Yor

Bawa, K. S. &J. H. BEACH. 1981. Evolution of sexual
systems in flowering plants. Ann. Missouri Bot.
Gard. 68: 254-2

BopkiN, N. L. 1978. A Revision of North American

Melanthium L. (Liliaceae). Ph.D. Dissertation.
University of Maryland, College Park, Maryland.
DAHLGREN, R. & H T Cu IFFORD. 1982. The Mono-
cotyled p dy ic P
London.
A. M. Lu. 1985. Campynemanthe (Cam-

pynemataceae): morphology, microsporogenesis,
early ovule ontogeny and relationships. Nordic J.
Bot. 5: 321-330.

& P. - 1985. The Families of the
Monocotyledons: ructure, evolution and tax-
onomy. Springer- Verlag. Heidel

DUTT, B. S . 1 Hypoxidaceae. Bull. Indian
Natl. Sci. Acad. 41: 368-372.

UNUS, A. M. 1951. Co "e to the embryology
of the Liliaceae. V. Life of Amianthium

numbers, 1982-1983. Monogr. Syst. Bot. Mis-

sou E Bot. Gard. 13: 1-224.

Sy saei and relationships of the

d c Pacific family Campynemataceae (Lili-

ales). Bull. Mus. Natl. es Nat., Paris, ser. 4, sect.

, Adansonia 8: 2.

LLOYD, D. G. & C. J. "e 1986. The avoidance of
interference between the presentation of pollen and
stigmas in angiosperms. I. Dichogamy. New Zea-
land J. Bot. 24: 135-162.

STENAR, H. 1949. Zur Kenntnis der Embryologie und
der Raphiden-Zellen bei Bowiea volubilis Harvey
und anderen Liliazeen. Acta Horti Berg. 15: 45-

UTECH, F. H. 84. Floral vascular anatomy of Ja-
panolirion osense Nakai (Liliaceae) and its tribal
relationship. Ann. Carnegie Mus. 53: 447-461.

1986. Floral morphology and vascular anat-
omy of Amianthium muscaetoxicum (Walter) A.
Gray (Liliaceae-Veratreae) with notes on distri-
bution and taxonomy. Ann. Carnegie Mus. 55:

481-504.

A REVIEW OF THE NEW WORLD SPECIES OF
ORTHROSANTHUS SWEET (IRIDACEAE)!

JAMES E. HENRICH? AND PETER GOLDBLATT?

ABSTRACT

oe a genus of some 9 species of Iridaceae —Sisyrinchieae, occurs in Australia and South

This review, arising out of research for floristic treatments for Flora Mesoamericana

-— five American species: O. acorifolius and O. Oeerssapunpis are endemic in South America;

to Mesoamerica and Mexico. The taxonomic history of Orthrosanthus i in the New World is outlined
followed by a key, and the nomenclature, brief descriptions, and distribution ranges for each species

are provide

The genus Orthrosanthus comprises nine
species of Iridaceae, tribe Sisyrinchieae Baker. It
has an unusual disjunct distribution, occurring
in Australia and central Mexico to South Amer-
ica, a pattern shared in the family only with Lib-
ertia, although the latter does not extend into
Mesoamerica. In the New World the taxonomy
of Orthrosanthus has long been confused, with
several authors recognizing different numbers of

thus is usually regarded as closely allied with
Libertia and Sisyrinchium and is distinguished
from them by an oblong to cylindric, included,
and often sessile to subsessile ovary and capsule.
Sisyrinchium and Libertia have globose to sub-

spherical seeds without prominent angular ridges.

Preparing a treatment of Iridaceae for Flora
Mesoamericana and Flora de Nicaragua, we have
reviewed the literature dealing with Orthrosan-
thus and have examined the ample herbarium
material now available. Our conclusions regard-
ing the systematics of Orthrosanthus appear to
merit the review presented here, since they differ
extensively from currently accepted taxonomy,
as represented in most major herbaria. Also, in-
formation about Orthrosanthus is scattered in
the literature and there is no modern summary
of the systematics and geography of the genus in
the New World. Cooke (1986) treated Orthro-

santhus for Flora of Australia, where four species
are now admitted. We recognize five New World
species (Fig. 1): O. acorifolius (Kunth) Ravenna,
O. chimboracensis (Kunth) Baker, O. exsertus

short history of Orthrosanthus in the New World
is outlined below, followed by a diagnostic key
for the species and a review of their systematics
and nomenclature.

HISTORICAL REVIEW

Orthrosanthus was erected by Robert Sweet
(1827) for the Australian O. multiflorus Sweet.
The first New World species ultimately assigned
to Orthrosanthus were described by Kunth (1815)
as Moraea (a distantly related African genus, tribe
Irideae). Kunth described three species now rec-
ognized as Orthr th Moraea chimboracen-
sis, M. gladioloides, and M. acorifolia. Baker
(1876: 113) was the first to include New World
species in Orthrosanthus, recognizing O. chim-
boracensis (Kunth) Baker (with M. acorifolia as
a synonym and M. gladioloides as a variety) and
a second species O. spicatus (Baker) Baker from
South America. The latter is a short plant with
a winged flowering stem and congested, sessile
inflorescence units (rhipidia). The flowers have
a short perianth tube, and the globose capsules
are carried on slender pedicels above the sub-
tending bracts as in Sisyrinchium and Phaio-
phleps. Ravenna (1968) placed the species in
Phaiophleps as P. brasiliensis, but we doubt that
it belongs in this heterogeneous alliance. It should

! Supported by Grant DEB 81-19292 from the United States National Science Foundation.
A.

2 Missouri Botanical Garden, P.O. Box 299, St.

ouis, Missouri 63166,

o

U.S.
3 B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri

63166, U.S.A.

ANN. Missounmi Bor. GARD. 74: 577-582. 1987.

578

be referred back to Sisyrinchium, pending critical
study of its affinities. Baker’s (1892) later, more
detailed treatment recognized seven species in
Orthrosanthus, the same two in the New World
and five in Australia.

In contrast to Baker’s treatment, Klatt (1861)
first placed New World species of Orthrosanthus
in Sisyrinchium. He described the distinctive

central Andean O. occissapungus and a p
f O. chimboracensis, S. moritzianum.
Klatt (1882) placed New World Vedi
in Solenomelus Miers, recognizing Solenomelus
chimboracensis (including M. acorifolia and
Sisyrinchium moritzianum) and Solenomelus
aet ed (including Sisyrinchium occissapun-
, close of the nineteenth century Rus-
by dun ed O. nigrorhynchus, not real-
izing that Sisyrinchium occissapungum was an
earlier name for the plant, and two years later
Kuntze described the conspecific O. tunariensis.
Orthrosanthus occissapungus has white flowers,
characteristic long narrow capsules, and typically
single- to few-flowered rhipidia.

A review of Orthrosanthus by Steyermark
(1948) is consistent with Baker's earlier treat-
ment in recognizing only one American species
(at least he makes no reference to O. occissapun-
gus or its synonyms). Steyermark's contribution
is noteworthy for recognizing four varieties and
two forms distinguished by capsule size, degree

of exsertion of the capsule pe the SAQUE
eaf

h

bracts, flower color, inflorescenc
rona and ees distribution.

, Ravenna (1965) described Orthro-
mo ASN (= O. chimboracensis var.
centro-americanus Steyerm.) and made the com-
bination O. monadelphus subsp. exsertus (R.
Foster) Ravenna, Ravenna (1969) also Pise
O. O. chim-
boracensis s subsp. tunalieniis (Kuntze) Ravenna.
Without mentioning Steyermark's (1948) treat-
ment of the genus, in particular O. chimbora-
censis var. acorifolius, Ravenna (1977) trans-
ferred Moraea acorifolia to Orthrosanthus as O.
acorifolius (Kunth) Ravenna. In 1981 he indi-
cated that he regarded Orthrosanthus as com-
prising four species and one subspecies.

TAXONOMY

Orthrosanthus Sweet, Flora Australasica, t. 11.
l Baker, Handbk. Irid. 117-119. 1892;
Cooke, Fl. Australia 46: 10-13. 1986. TYPE:
O. multiflorus Sweet.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Elvetria Raf., Fl. Telluriana 4: 30. 1838. TYPE: E. mul-
tiflora (Sweet) Raf. (= O. multiflorus Sweet).

Evergreen tufted perennials with short, persis-
tent, creeping rhizomes. Leaves several, mostly
basal, ensiform and equitant, linear to linear-
lanceolate, coriaceous, without a midvein,
crowded at the apex of the rhizome. Flowering
stem erect, branched, the iones ascending,
either long or short, term

t

relatively short; spathes subequal. Flowers acti-
nomorphic, subsessile or shortly stalked, blue or
white; tepals free (united in a short tube in some
Australian species), subequal, spreading hori-
zontally from the base; filaments free or united
in the lower half, anthers erect; ovary included
in the spathes, + sessile, or exserted in O. exser-
tus, style short, dividing into 3 relatively long
branches extending between the stamens, the
branches stigmatic apically. Capsules ellipsoid to
cylindric, sometimes pubescent, included or
exserted; seeds angular to fusiform.

Species: 9; highland areas from Mexico to Pan-
ama, the Coastal Cordillera of Venezuela, and
through Andean South America to Bolivia and
northern Argentina (Fig. 1), and in Australia.

KEY TO THE NEW WORLD SPECIES

la. ead with pedicels is a 15(-25)

m long; capsule glabr
laf

2a.
capsules slender, Pin e mm long:
seeds elongate-angular, 2.5-3 mm E
0.5 mm ua (Peru, Bolivia, and nort
ern Arge . $. O. oceissapungus
Spathes ira an (2-)3 or more flow
capsules oblong to ellipsoid, 10-18 mm
long; seeds angular to irregularly globose,
1.2-2 mm long and wi
3a. Mature capsules borne well above
the subtending bracts; pedicels 15-
25 mm long (central Mexico: Mex-
ico, Puebla, Distrito Federal, Ta-
maulipas, Nuevo Leon, Michoacan)
. O. exsertus
. Mature capsules usually included;
pedicels 5-10 mm long (Costa Rica,
ode: Colombia, bows dor,
EEEE ; imboracensis
] a ide or with short p ne
m long; ca ipa sue aT DE
se

t3
g

Uu
og

c

m long; leaves 2-4 mm
wide (southern Mexico to northern Pan-
ama, except Belize and Nicaragua) .......
4. O. monadelphus
4b. Capsules 8-11 mm long; leaves 4-12 mm
wide (Venezuela) ...... 1. O. acorifolius

1987] HENRICH & GOLDBLATT—NEW WORLD ORTHROSANTHUS SWEET 579
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100 90
FiGurE 1. Distribution of the species of Orthrosanthus in the New World.

l. Orthrosanthus acorifolius (Kunth) Ravenna,
M

santhus chimboracensis var. acorifolius
(Kunth) Steyerm., Lloydia 11: 19. 1948.
Marica acorifolia (Kunth) Martens & Ga-
leotti, Bull. Acad. Brux. 10: 110. 1843 (mis-
applied, probably to O. monadelphus). TYPE:
Venezuela. Distrito Federal: in crepidinibus
montis Silla de Caracas, ca. 1,250 m, Bon-
pland & Humboldt s.n. (lectotype, P— Gen-
barium; the sheet is designated

more type material probably existed and may
be found.

iab erede chimboracensis Mi qiia f. albus

Steyerm., Lloydia 11: 19. YPE: Venezuela.
Trujillo La Quebrada oe by d line
Lara-Trujillo rmark

55339a (holotype, F).

Plants to 65 cm tall. Leaves to 50 cm long,
(0.4-)0.9-1.2 cm Il broadly linear, gradually
stem about as long as the

long, pubescent; seeds angular to somew
rounded, ca. 1-1.2 mm at the widest diameter.

Distribution. Orthrosanthus acorifolius is ap-
parently restricted to the Coastal Cordillera and
Andes of Venezuela and eastern Colombi

This species is usually very robust and has
comparatively broad leaves sometimes matched

580

in Orthrosanthus by those of O. chimboracensis.
The ovary and young capsules are heavily pu-
bescent, as in O. monadelphus, to which O. aco-
rifolius is probably most closely related.

2. Orthrosanthus chimboracensis (Kunth) Ba-
ker, Gard. Chron. n. ser. 6: 67-68. 1876.
Orthrosanthus chimboracensis var. typicus
Steyerm., Lloydia 11: 15-16. 1948. Moraea
chimboracensis Kunth, Nov. Gen. & Sp. PI.

: 322. 18 Tunguragua:
Chim-

Urcu, ca. 1,640 m, Bonpland & Humboldt
s.n. (lectotype, P—Herb. Bonpland, here
designated). More type material may be
found; the sheet we have designated as lec-
totype has the data “Chimborazo” only and
the number 3/88.

iini: a w ms Gen & Sp. Pl. 1: 322.
. Cajamarca: locis frigidis Pe-
m et Micuipampam, ca

Sisyrinchium moritzianum Klo si x Klatt, Linnaea
78. 1. TYPE: Venezuela (as Colombia).
erida: Paramo de la Culata, ae 1204 (iso-

type, K).

Plants (20—)60—115 cm tall. Leaves 15-70 cm
long, 8-12 mm wide, narrowly lanceolate to lin-

£e
=

blue. Capsules 14-18 mm long, glabrous, the
pedicels 5-10(-15) mm long; seeds angular, 1.2—
2 mm at the longest axis.

Distribution. | Orthrosanthus chimboracensis
ranges from Costa Rica and Venezuela south-
ward through the Andes from Colombia to Peru.
It is a montane species seldom occurring below
2,000 m.

Plants are typically less robust than Orthro-
santhus acorifolius and can be recognized by a
sparsely pubescent to glabrous ovary and com-
pletely glabrous capsule with a short pedicel 5-
l long

3. Orthrosanthus exsertus (R. Foster) Ravenna,
Wrightia 7: 10. 1981. Orthrosanthus chim-
boracensis var. exsertus R. Foster, Contr.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Gray Herb. 155: 49. 1945. TYPE: Mexico.
Distrito Federal: on the sides of ravines near
Eslava, Pringle 8827 (holotype, G; isotypes,
C, CAS, F, G, GH, MEXU, MO, NY, O,S,
TEX, UC, US).
Orthrosanthus chimboracensis var. exsertus f. albus
m., Lloydia 11: 17. 1948. TYPE: Mexico.
Tamaulipas: Santa Rita, Ranch Tamaulipas, 1,500
m, Runyon 875 (holotype, US; isotype, TEX)

Plants 40-70 cm tall. Leaves 40-55 cm long,
3-8 mm wide, narrowly acuminate. Flowerin
stem somewhat longer than the leaves; rhipidia
several-flowered. Flowers to 2.5 cm diam., blue,
or occasionally white; ovary exserted from the
spathes shortly after anthesis. Capsules 1-1.7 cm
long, glabrous, broadly acuminate apically, well
exserted from the spathes on pedicels 15-25 cm
long; seeds angular to somewhat rounded, 1.2-
1.5 mm at the widest diameter.

Distribution. Orthrosanthus exsertus is en-
demic to highland areas of southern and central
Mexico (Nuevo León, upon. Puebla, Mi-
choacán, Distrito Federal, Méxic

The smallest of the New World pam of Or-
throsanthus, O. exsertus is distinctive in having
comparatively narrow leaves and glabrous cap-
sules exserted from the spathes and bracts on
pedicels 15-25 mm long. Recognized only in 1981
as a distinct species, and first as a variety in 1948,
Orthrosanthus exsertus was recorded as early as
1829 or 1830 by Schiede and a few years later
by Liebmann. Their collections are among the
three syntypes cited by Klatt for Sisyrinchium
occissapungum, now O. occissapungus, and neo-
typified here by a collection from Peru.

4. Orthrosanthus monadelphus Ravenna, Bol.
Soc. Argentina Bot. 10: 317. 1965. TYPE:
Guatemala. Baja Verapaz: cumbre El Chol,
ca. 2,200-2,500 m, Ravenna 266 (holotype,
Herb. Ravenna, not seen; isotypes, F, HBG).

Orthrosanthus chimboracensis var. diis americanus
Steyerm all: 19. € 48. T

of rae unción Mita, Steyer-
mark 31913 fliclotvne. F).
Mice chimboracensis or intermedius Stey-
Lloydia 11: 19-20. 1948. TYPE: Costa Rica.
iei Volcán Irazu, Allen 674 (holotype, F).

Plants (15—)30—60 cm tall. Leaves 30-45 cm
long, 8-10 mm wide, linear. Flowering stem to
60 cm; rhipidia several-flowered and spaced
somewhat irregularly along the stem; spathe

1987]

margins narrowly hyaline. Flowers 1.4-3 cm
diam., blue. Capsules 10-13 mm long, usually
pubescent, sessile or nearly so, only the apices
exceeding the spathes; seeds angular to some-
what rounded, 1.2-1.5 mm at the widest diam-
eter.

Distribution. Orthrosanthus monadelphus is
common in highland areas of Mesoamerica. It is
recorded from southern Mexico to Guatemala
and El Salvador, and locally to the south in Costa
Rica and northern Panama.

This species is probably most closely allied to
the Venezuelan Orthrosanthus acorifolius, with
which it shares a similar, nearly sessile, and pu-
bescent ovary and capsule. The filaments are
united in the lower half, not much more so than
in the other American species. Although first rec-
ognized as a distinct subspecies in 1948, and as
a species in 1965, it is interesting to note that
the collection cited by Martens & Galeotti (1843)
as Marica acorifolia may be the first record for
Orthrosanthus monadelphus [Mexico: Oaxaca,
rochers gneissiques de Penoles, Misteca alta, ca.
2,150-2,300 m, Galeotti 5368 (?BR, not seen)].

5. Orthrosanthus occissapungus (as O. ocisa-
punga) (Ruiz ex Klatt) Diels, Engler & Prantl,
Nat. Pflanzenfam. ed. 2. 15a: 478. 1930.
Sisyrinchium occissapungum Ruiz ex Klatt,
Linnaea 31: 379. 1861. NEOTYPE: Peru. La
Libertad: along the Río Negro 3 km south
of Huamachuco, West 8113 (neotype, MO;
isoneotypes, GH, UC). The three cited syn-
types of S. occissapungum have not been
found and are presumed destroyed, hence
our designation of a neotype. For complete-
ness, the syntypes are cited here as follows:
Peru. Huanuco: ad Pillao etc., Ruiz ex Herb.
Lambertii (B, not seen); Mexico. Oaxaca:
Chinantla, Liebmann 310 (location un-

own) (? = O. exsertus); Mexico. Veracruz:
Jalapa, Schiede 1029 (location unknown) (? =
O. exsertus).

Orthrosanthus nigrorhynchus Rusby, Mem. Torrey Bot.
C 26. 1896. TYPE: Bolivia. Cochabamba:
near Cochabamba, Bang 1074 (lectotype, NY, here
designated as the best preserved and most com-
plete of three sheets at NY, the institution where
Rusby worked; isolectotypes, BM, F, G, GH, K,

MO, :

Orthrosanthus tunariensis Kuntze, Rev. Gen. Pl. 3:
309. 1898. TYPE: Bolivia. Cocha ba: im Tu-
narigebirge, 3,600 m (collecto and location of the
type not in the protologue and unknown to us).
Orthrosanthus EE siberaceasis subsp. tunariensis

HENRICH & GOLDBLATT — NEW WORLD ORTHROSANTHUS SWEET

581

(Kuntze) Ravenna, Revista Inst. Munic. Bot.
(Buenos Aires) 2: 30. 1969.
Orthrosanthus chimboracensis sensu Rusby, non
(Kunth) Baker, in Bull. Torrey Bot. Club 29: 224.
01.

Plants to 80 cm tall. Leaves 30-50 cm long,
-8 mm wide, narrowly linear, sharply acute.
Flowering stem to 80 cm; rhipidia typically sin-
gle-flowered. Flowers 2.6-3.4 cm diam., white.
Capsules (1 5-)22-30 mm long, glabrous, extend-
ing above the spathes on pedicels to 5 mm long;
seeds elongate-angular, 2.5-3 mm at the longest
axis.

Distribution. Orthrosanthus occissapungus is
restricted to northwestern Argentina, Bolivia, and
Peru, where it grows at elevations above 3,200 m.

The most unusual of the New World species
of Orthrosanthus, O. occissapungus can be rec-
ognized by its one- to few-flowered spathes, long
slender capsules, Fei elongate seeds. The flowers
are always white in O. occissapungus and only
occasionally so in de species in the New World.

As indicated in the nomenclature above, we
have been unable to locate the original type ma-
terial of Sisyrinchium occissapungum. The syn-
type from Peru, collected by Ruiz, had the manu-
script epithet '"*occissapungum" according to
Klatt. It was housed in the Berlin Herbarium and
must be presumed destroyed. It was almost cer-
tainly conspecific with O. nigrorhynchus de-
scribed in 1896. The two other specimens cited
in the protologue are from Mexico: Schiede 1029
(also presumed destroyed) from Jalapa in Ve-
racruz, and Liebmann 310 from Chinantla, Oa-
xaca. The latter specimen is cited as “Herb. mihi"
by Klatt, that is, in his own herbarium, which
was at one time at Berlin but is now at Stock-
holm. No Liebmann collection with this number
has, however, been located at either place. Du-
plicates of the Liebmann collection at Copen-
hagen, where Liebmann’s main collection is pre-
served, likewise do not include the missing type
number. Other Liebmann specimens at Copen-
hagen and at Paris collected at the same locality
and time are the Mexican Orthrosanthus exsertus
and represent the first collections of that species.

e are reg g pung
as a valid combination made by Diels (1930:
478) but recognize that it would not be accepted
according to the current Code of Botanical No-
menclature if made today. Diels did not cite a
basionym or indicate in any way that he was
making a combination in citing the species as O.
ocisapunga (sic) Ruiz. Nevertheless, there can be

582

no doubt that he was placing Sisyrinchium oc-
cissapungum Ruiz ex Klatt in Orthrosanthus as
the earliest name for the species known until this
time as either O. nigrorhynchus or O. tunariensis,
neither of which he mentioned in his enumera-
tion of the species of Orthrosanthus. The name
O. occissapungus was accepted by Macbride
(1936) and Vargas (1944) but was attributed by
both authors to Diels (as O. ocisapunga Ruiz ex
Diels). We do not believe that Diels intended to
describe the species, and therefore we cannot ac-
cept this treatment. We also note that it was not
Diels’s practice to cite the authors of species in
later combinations e.g., his listing of O. chim-
boracensis Bak. [instead of (Kunth) Baker], and
so there is no inconsistency in his not mentioning
the authors of the basionym for O. occissapun-
gus.

LITERATURE CITED
BAKER, J. G. 1876. Systema Iridacearum. J. Linn.
Soc., Bot. 16: 61-180.
1892. Handbook of the Irideae. George Bell

n.
1986. Iridaceae. In Flora of Australia

DIELs, L. 1930. Iridaceae. Jn A. Engler & K. Prantl
(editors), Die Natiirliche Pflanzenfamilien, ed. 2.
15a: 463-505.

Foster, R. C. 945. Studies in the Iridaceae. III.

Contr. ed Herb. 155: 3-54.

KLATT, F. W. 1861. Berichtigungen und Nachtráge
zu der Monographia Generis Sisyrinchium. Lin-
naea 31: 370-386.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

. Erganzungen und Berichtigungen zu
Baker's Systema Iridacearum. Abh. Naturf. Ges.
v

— Il

815. ys de nei s Bon-

Latine-Allemande, Paris. Gide Fils, Paris.
. Iridaceae. In Flora of Peru.
Mus. Nat. Hist. Bot. Ser. 13 (1, no. 3): 707-
717.
Martens, M. & H. GALEOTTI. 1843. Enumeratio syn-
optica plantarum phanerogamicarum ab Henrico
Bull

Acad. 2 Sci Belgique 10: 110-134.
RAVENNA, P. 1965. Notas sobre Iridaceae. II. Bol.
Soc. ae Bot. 10: 311-322.

l . Notas sobre Iridaceae. III. Bonplandia
2: 27 3-291.

. 1969. Notas sobre Iridaceae. IV. Revista Inst.
Munic. Bot. (Buenos Aires) 2: 25-38.
1977. Notas sobre Iridaceae. V. Mus. Nac.
Hist. Nat. Santiago Notic. Mens. 21: 7-9.
. 1981. On the presence of the genus Orthro-
(Irid )in the Argentine fl Wrighti

7: 10.

RusBv, H. H. 1896. On the collections of Mr. Miguel
E in Bolivia, Part III. Mem. Torrey Bot. Club
6: 1-130.

ee J. A. 1948. Orthrosanthus chimbora-
censis and its varieties (Iridaceae). Lloydia 11: 14—
20.

Sweet, R. 1827. Flora Australasica, t. 11. James
Ridgway, London.

VARGAS, C. C. 1944. Iridaceae Cuzcoensis. Revista
Univ. Cuzco 33(87): 167-177.

PATCH FORMATION AMONG ISRAELI CRUCIFERS:
HOW DO THEY GET AWAY WITH IT?!

MICHAEL AUERBACH? AND AVI SHMIDA?

ABSTRACT

ny m in be ee and desert floras of Israel frequently form large monospecific
r . This ted

Man
patches but do
theory of in
herbivore loads, a
defense against herbivores. In editerr.

Papilionaceae, Apiaceae). This is not true for desert
year to year.

allow Mediterranean patch-forming species to

of herbivory. In contrast, desert pa cies ee pins Se rely on pe ropa e phe ovre as wi
s. Patc

MEA Among . agis PAE patc
on-patch formers. In addition, patch for

A interactions which predicts that suc
nd that herbaceous plants iion rely on unprediethbilid in time and space as a
Isr nean crucifers fl of o
species-rich families with similar sciri da orms (Lami aceae, se Liliaceae, Solanaceae, Poaceae,

tch formers are "ale and have larger pet
mation is correlated

situation is unexpect from some recent
should attract high

ower earlier than members of other

ucifers

We hypothesize that their displaced phenologies id: possession of potent VIDA

uce monocultures while not sustaining high lev

E as
y be reinforced by plant-pollinator
tals than do

vels of floral ultraviolet

ormation m

high le

ier aed and patterning. These differences may result from perdes: iüerepecific but enhanced

intraspecific

competition for pollinators among pa

tch formers. These patterns are not iud among

desert species, although desert patch formers do initiate flowering before other crucifers.

The family Brassicaceae (Cruciferae) has fig-
ured prominently in the development of theory
concerning insect—plant interactions. Responses

& Feeny, 1977), as well as considerations of fam-
ily-wide attributes (Feeny, 1976, 1977), com-
prise much of the empirical and theoretical sup-
port for concepts such as ''associational
resistance" (Tahvanainen & Root, 1972) and
*plant apparency" (Feeny, 1976). Feeny (1977)
also cited family-wide attributes of crucifers to
support his contention that escape in time and
space and allelochemic diversity protect herba-
ceous plants from potential herbivores. Several
characteristics of crucifers make them particu-
larly amenable for studies of herbivore-plant in-
teractions. They are well represented in floras of
many regions of the world (Hedge, 1976), all
species thus far examined contain glucosinolates
(Kjaer, 1976), and many species are important
cultivated crops.

The family contains approximately 400 genera
and 3,000 species, most of which are annual herbs

(Vaughan et al., 1976). The Irano-Turanian re-
gion is home to about 900 crucifer species and
was probably the center of origin for at least the
Old World taxa (Hedge, 1976). Brassicaceae is a
dominant and conspicuous family in terms of

ticularly in late winter and early spring (Shmida
& Auerbach, 1983). Most of the 111 or more
species native to Israel are Irano-Turanian or are
descendants of Irano-Turanian stock (Zohary,
1973).

Crucifers frequently form enormous mono-

r
plant interactions which predicts that such re-
source concentration should attract high herbi-
vore loads, and that herbaceous plants should
rely on unpredictability in time and space as a
defense against herbivores. Here, we examine the
crucifer flora of Mediterranean and desert re-
gions of Israel in terms of plant morphology,
phenology, and propensity of some species to
occur naturally in large monocultures. In partic-

! We thank Nitsa Dagan for assistance with data analysis and Diana Lieberman for her critical review of this

manuscript. William Kunin and Micha

Boaz provided unpublish

ed data. Most analyses presented here were

conducted while one of us Sl was a Lady Davis Postdoctoral Fellow in the Department of Botany, Hebrew

Evers. The support and e
? Department of Biology, ir. of Nort

ANN. MIssourRI Bor. GARD. 74: 583—594. 1987.

y
akota, Grand For
3 Department of Botany, Hebrew University, Jerusalem, Israel.

Davis Trust is gratefully acknowledged.
s, North Dakota 58202, U.S.A.

584

ular, we address the question, “how do patch
formers avoid colonization by a high density and/
or diversity of phytophages’”?

ISRAELI CRUCIFERS

We obtained data on plant species from Flora
Palaestina (Zohary & Feinbrun-Dothan, 1966),
herbarium records of the Hebrew University De-

artment of Botany, Hebrew University, field
observations from January to June 1983, more
than 10 years of vegetation samples taken by
A.S., and phenological records from ECOPAS.
The last is a computer data base compiled by
members of the Department of Botany, Hebrew
University, and the Society for the Protection of
Nature in Israel. Each month phenological rec-
ords are submitted by a network of observers,
resulting in a comprehensive record of Israel’s

ora.

Tables 1 and 2 list species of native crucifers
growing in the Mediterranean (76 species) and
desert regions (37 species) of Israel (two species
are listed in both regions). For each species we
show growth form, floral characteristics, pubes-
cence, abundance in Israel, and a subjective mea-
sure of size of monospecific patches. Within both
assemblages we excluded extremely rare species
that we never encountered in the field (approx-
imately 20 species).

We must clarify one entry in Tables 1 and 2.
Our index of patch formation ranges from
“+++” (largest patches) to ““—” (no patches).
Although this categorization is subjective, we
classified species as patch formers only if they
formed large monospecific associations. A patch
in Our usage is not a clump of several individuals
of one species; rather, it is an extensive single-
species association. Patches range from approx-
imately ten meters to several kilometers on a
side. For instance, in spring 1983 (highest spring
rainfall in 30 years), we measured patches of Eru-
caria hispanica L., near Arad, Israel, and Sinapis
arvensis L., near Bet She'an, Israel, that extended

combined all patch formers for statistical anal-
yses.

Using these data, we compared phenological
attributes of the Brassicaceae with those of seven
other species-rich families with similar growth
forms in Israel (Table 3). Phenological compar-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

isons were based on peak flowering time (month
in which most individuals of a species produce
most of their flowers) and on range of flowering
time (range of months in which flowers have been
observed for each species). For each family we
constructed cumulative frequency distributions
of flowering range and peak and compared the

of association (Kolmogorov-Smirnov 2 sample
test, Conover, 1980). We also compared average
month of peak flowering between the Brassica-
ceae and other families with a parametric test
(Welch approximation of the t-test, Remington
& Schork, 1970), since these data were approx-
imately normally distributed. We limited these
comparisons to Mediterranean species because
desert species are constrained to flowering over
a short, variable period when rains occur. Com-
parisons were done on all species and the subset
of annual species for each family, except for Lil-
iaceae which only has one annual species.

e then compared morphological and phe-
nological attributes of patch- and non-patch-
forming crucifers. These analyses were conduct-
ed separately for Mediterranean and desert
species. We compared phenologies based on cu-
mulative frequency distributions of flowering
range and flowering peak and average month of
peak flowering. In addition, we compared the
average month of start of flowering using a para-
metric test (Welch approximation of t-test). Av-
erage plant height and petal size from ECOPAS
were compared using f-tests, whereas analyses of
associations of pubescence and ultraviolet pat-
terns with patch formation were based on two-
by-two contingency tables.

RESULTS
INTERFAMILY DIFFERENCES

Comparison of the phenology of Mediterra-
nean crucifers with members of other families
indicates that in general crucifers flower earlier
in the year, both in terms of flowering range and
flowering peak (Table 3). The single exception is
no significant difference between the peak flow-
ering time of the Brassicaceae and Liliaceae (Ta-
ble 3). The average duration of flowering of all
crucifers differs from that of all Lamiaceae, As-
teraceae, Solanaceae, and Poaceae (Table 3).
Among all families we examined, medians of
flowering range and peak occur earlier for annual
species than for perennials except for the Solana-
ceae, in which the situation is reversed. Peren-

585

AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION

1987]

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[Vor. 74

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AMOUMOUMOOUMOHOYL

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ANNALS OF THE MISSOURI BOTANICAL GARDEN
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586

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1987]

Continued.

TABLE 1.

Relative

Petal

Mean
Height
(cm)

Patchiness

Flower
Phenology

Pubes-
cence

Growth
Form

Species

++

LC

R. rostratus DC.

Rapistrum rugosum (L.) All.

AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION 587

++

LC

Rorippa amphibia (L.) Besser

Sinapis alba L.
S. arvensis L.

++

CC

RR
CC

> >

Sisymbrium damascenum Boiss. et Gaill.

S. officinale (L.) Scop.

arnn

>Z z z

Thlaspi arvense L.

Turritis laxa (Sibth. et Smith) Hayek

nials also have longer average durations of flow-
ering than annuals, except in the Solanaceae.

INTRAFAMILY COMPARISONS

Tables 1 and 2 show that most crucifer species
In Israel, as in other areas, are annuals. As men-
tioned above, the few perennial species in both
the Mediterranean and desert floras generally
flowerlaterandlongerthantheannuals,although
exceptions do exist (e.g., Fibigia clypeata L.). Ad-
ditionally, a distinct guild can be recognized

by small plants (5-15 cm), with small, generally
white or cream-colored flowers that appear early
in the year (mid January through February).
Within the two floral associations, we conducted
all statistical analyses on crucifers four times: all
species, annuals only, pygmy guild excluded, and
perennials and pygmies excluded

For the Mediterranean crucifers, patch-form-

ing S
species (Table
formers generally paralleled our rank of patch
formation (+++ > ++ > + > —,cases 3 and
4,++>+++ > + > —, cases | and 2, Table
4). In three of four comparisons, patch formers
also had significantly larger petals than did non-
patch formers (Table 4), with patch rank gen-
erally paralleling petal size (++ > +++ >
—, all cases). Few systematic differences
exist among phenologies; patch formers gener-
ally do not flower earlier or longer than non-
patch formers, although there are a few excep-
tions. There is no dominant flower color among
Mediterranean patch- forming species; oe
white, cream, and p comm
Pubescence is significantly associated with non-
patch-forming species in all analyses.
Unlike Mediterranean species, the desert patch-
forming species are neither taller nor have larger

durations of 4 generally do not differ be-
tween the grou

Patch sans in the desert, with the exception
of pygmy species, generally have purple flowers
(8 of 12 species). Once again, pubescence is sig-
nificantly ree aa with non-patch-forming
species (Table 4).

Horovitz & Cohen (1972) analyzed ultraviolet

[VoL. 74
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T
T
x
<

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588

*suongrA31qqe 10j | 9[QE.] JO PUIA IF ‘[OBIS] JO LIOY HSP ay} UT səroəds IIIN JO soINQUIIe [eoISojoIe pue [eordo[oudiojq ‘Z «ATAV_L

1987] AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION 589

characteristics of the flowers of 22 species of cru-
cifers native to Israel (17 Mediterranean, 5 des-

Š " ert). Among these, high ultraviolet reflectance
a H sÉ : H i p and/or patterning Is positively associated with
= + patch formation (G,,; = 12.3, P < 0.001). The
* same is true for the subset of Mediterranean

°, species (Gaa; = 6.59, P < 0.025), but not the

8B Š ou e desert ones (G,4; = 0.5, P > 0.05).

Oz ZlpOARonY

= DISCUSSION

E SE ——— WHY NOT FORM PATCHES?

x E Several lines of evidence suggest that patch

formation should increase the susceptibility of a
Š 5 plant species to insect attack. Many herbivores,
e Ç: Qm > > Z n. particularly those with narrow host ranges, are
more likely to find, to remain on, and to repro-
uce on hosts that are concentrated in space
(Root, 1973; Risch, 1981). This is one reason
pod ood (d why monocultures of herbaceous plants fre-
quently support greater densities of insect pests
than do polycultures (Pimentel, 1961; Root, 1973;
Feeny, 1976; Altieri et al., 1977; Risch, 1981).
The diversity and abundance of phytophagous
ll l gr + insects attacking a plant species in a given area
are also influenced by the botanical diversity of
the area (Strong et al., 1984). Members ofa mixed-
species plant assemblage often escape attack be-
CQ C QN 6 ae cause increased species diversity reduces the sus-
ceptibility of each plant species to discovery. This
phenomenon has been referred to as “‘associa-
tional resistance" (Tahvanainen & Root, 1972)
< < < < < pn. or “plant defense guilds” (Atsatt & O'Dowd,
1976). Considerations of alternative resource
availability for phytophagous insects as well as
their frequent use of visual or chemosensory cues
in host-plant location suggest that this form of
escape should increase in effectiveness as phy-
tophage dietary specialization increases (Root,
1973; Atsatt & O'Dowd, 1976).

Rhoades & Cates’s (1976) and Feeny’s (1976)
general theories of plant defensive chemistry ex-
pand the concept of plant escape from enemies
in ecological time into evolutionary arguments
concerning selection for different classes of al-
lelochemicals. Both postulate that as the occur-
rence of a plant species or tissue becomes more
predictable (apparent) in space and time there
should be greater selection for generalized, quan-
titative (dosage-dependent) defenses as opposed
to specialized, qualitative defenses (toxins).
Quantitative defenses are presumed to be more
metabolically costly but harder for a herbivore
to circumvent than qualitative ones. Thus, it is

Phenology
4
4
4
5
4
9

Pubescence

Mean
Height
(cm)
3
0
5
0
5
5

Growth
Form

Species

TABLE 2. Continued.
Reboudia pinnata (Viv.) O. Schulz
Savignya parviflora (Del.) Webb

Schimpera arabica Hochst. et Steud. ex Boiss.

Sisymbrium erysimoides Desf.

Zilla spinosa (L.) Prantl

590

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

BLE 3. Comparison of phenologies of - Brassicaceae with other families in the Mediterranean flora of
Feb. = 2, etc.

ae] Months were converted to integers (Jan. = 1,

or flowering range and peak analyses.

Inequality signs indicate earlier (<) and later ue in the growing season.

Compared with Brassicaceae

Flowering Flowering Flowering
Range Peak Duration Flowering Flowering
Number of (median (median (mean number Range Peak
Family Species month) month) months (s.d.)) (distribution) (distribution)
Brassicaceae 76 3.71 3.61 2.94 (1.47)
Annual 64 3.46 3.52 2.88 (1.20)
Perennial 12 5.50 4.75 -—
Lamiaceae 80 5.79 5.27 3.71 (1.49)* >+ >+
nnual 17 4.86 4.42 3.35 (1.41) >++ >++
Perennial 63 6.07 5.50 —

Asteraceae 169 5.74 4.88 3.43 (1.74)* >+ >+
Annual 105 5.17 4.54 3.25 (1.69) >++ >++
Perennial 64 6.63 5.83 — — —

Liliaceae 66 4.33 3.66 2.91 (1.34) >+ =
Perennial 65 4.33 3.67 — — —

Solanaceae 15 6.96 6.50 5.13 (2.64)* >+ >+

6 7.60 7.00 5.33 (0.82)** >++ >++
Perennial 9 6.50 6.17 — —

Poaceae 168 5.67 4.86 3.77 (1.95)* >+ >+
Annual 111 5.18 4.59 3.40 (1.61)** >++ >++
Perennial 57 6.65 5.68 — —

Papilionaceae 206 4.52 4.19 3.13 (1.10) >+ >+
Annual 173 4.33 4.08 2.92 (0.67) >++ >++
Perennial 33 5.70 5.21 —

Apiaceae 87 5.35 4.83 3.03 (1.24) >+ >+
Annua 48 4.93 4.55 2.77 (1.10) >++ >++
Perennial 39 5.79 5.39 —

* = * Significantly different from all, or annual (**) crucifer species (t-test, Welsh approximation, P < 0.05).
+ = Significantly different from all, or annual (+ +) crucifer species (Kolmogorov-Smirnov 2 sample test,

P « 0.05).

argued that a plant that is apparent by virtue of
its growth form, abundance, and longevity, such
as a common tree species, must invest a consid-
erable amount of energy into quantitative defen-
ses, whereas an annual herbaceous plant can in-
vest less energy in qualitative defenses, provided
it also remains unapparent in time and/or space.
By definition, unapparent plants should have
small patch sizes and low p density (Fox,
1981
Escape in time and space should be effective
against specialized herbivores in ecological time
and against nonspecialized herbivores over evo-
lutionary time. Specialized herbivores that have
evolved a means of circumventing an allelo-
chemic defense frequently respond to the allelo-
es for host location
and phagostimulation; at herbivores have, over
evolutionary time, turned an unapparent re-

source into an apparent one (see Courtney, 1985,
for various interpretations of apparency). Once
this occurs the plant benefits from anything that
reduces its apparency to adapted enemies, such
as having an irregular phenology or growing in
multispecies assemblages. Against nonspecial-
ized herbivores, escape in space and/or time re-
duces the frequency of encounters between po-
tential enemy and plant tissue, thereby decreasing
the likelihood of an herbivore evolving a detox-
ification mechanism.

Since most crucifers are short-lived herbs and
contain qualitative allelochemicals, one might
expect them to rely quite heavily on escape in
time and space from phytophagous insects. In
fact, Feeny ar regarded crucifers as typical
unapparent plants and much of his theory of
plant apparency is based on crucifers (see ref-
erences in Feeny, 1976). Most Israeli crucifers

1987] AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION

do rely on escape on one time scale by nature of
their annual growth form with concomitant short-
term availability each year of leaves. However,
the propensity of many species to form large
monotypic patches confers a high degree of spa-
tial predictability and resource concentration on
these species. In addition, location of patches of
Mediterranean patch-forming species frequently
does not change from year to year. This situation
would appear to be disadvantageous in light of
the theory discussed above.

HOW DO THEY GET AWAY WITH IT?

Mediterranean patch-forming crucifers in Is-
rael suffer little herbivore damage despite their
short-term spatial predictability. During a season
of sampling (1983), we never observed high den-
sities or diversities of phytophages on patch-
forming species. Neither high numbers of sup-
ported insect herbivores nor large amounts of
leaf area removal were detected with spot cen-
susing during ten years of vegetation sampling.

We suggest that Mediterranean patch-forming
crucifers are able to grow and persist in dense
monospecific associations owing to possession of
glucosinolates (and other allelochemicals in some
species), and because they flower earlier than oth-
er local species. Both displaced phenologies and
glucosinolates prevent seasonal tracking and use
of crucifers over evolutionary time by presently
non-crucifer-adapted herbivores. By growing
earlier than most other annual species, these cru-
cifers have reduced the number of chance en-

dietary shifts in phytophages. If patch formation
occurred later in the growing season, more en-
counters between pest and potential host would
ensue, and over evolutionary time one would
expect an increase in diversity of insect species
adapting to patch-forming crucifer species.
Many crucifers germinate, grow, and flower
before the annual peak of insect biomass. Indeed,
many of the patch-forming species flower before
the annual appearance of crucifer-adapted her-

mida, unpubl. data). Occasionally, these
adapted herbivores do appear on patch formers
during the late stages of flowering or seed set,

Groups

NP > P**
NP > P*
NP > P*

Pubescence

Start of
Flowering
P < NP*
P < NP**

NP
NP
NP

Flowering
Range
(distribution)
P=N

P=
P=
P=N
P=N
P=N

Flowering
Peak
(mean)

P=N
P > NP*
P
P = NP
P = NP
P=N
P=N

= NP

-forming (P) and non-patch-forming (NP) crucifer species in the (A) Medi-
P <

NP

P=N
= N
P =
(B) DESERT
P=N
P=N
P=N

(A) MEDITERRANEAN

(distribution)

Duration
P > NP*
P
P=N
NP
P=N
P=N
P=N

pygmy guild excluded; (4) perennials and pygmies excluded.
P s

Petal Size
P > NP**
P > NP***
N
P > NP*
P=N
P=N
P=N
P=N

Plant Height

P > NP***
P > NP***

P=N

TABLE 4. Comparisons of morphological and phenological attributes of patch
terranean and (B) desert floras of Israel. In reference to phenological traits, the inequality signs indicate earlier (<) and later (>) in the growing season.

are defined as: (1) = all species; (2) = annuals; (3)
* = Significant difference, P < 0.05; ** = P < 0.01; *** = P < 0.001.

Group

592

when herbivory has a minimal effect on the fit-
ness of n

Germination pane in the desert is far less
‘redictable than in Mediterranean regions be-
cause of var A eae a early-spring rains
(Sbmida et al., 1985). Thus, desert crucifers are
far more unpredictable in time than their Med-
iterranean counterparts. Year-to-year variation
in composition, location, and size of patches is
often tremendous. Despite these differences,
flowering is still well under way by the time most
herbivores, even crucifer specialists, arrive. In-
terestingly, pubescence, a characteristic fre-
quently interpreted as a possible herbivore de-
fense (Coley, 1983) as well as an adaptation that
reduces rates of evapotranspiration (Lieberman
& Lieberman, 1984), is more common among
non-patch- than patch-forming species in both
floristic regions.

e can only speculate why crucifer-adapted
herbivores have not undergone phenological dis-
placement to exploit early-season crucifers. Se-
m for Synenrony may li countered by phys-
early-season
cold temperatures. The exsul: variable phe-

aAlonoical
°

ony. As Feeny (1977) noted,
*Short growth season, shifting pattern of geo-
graphic distribution, and association with harsh
and somewhat unpredictable climatic conditions
are all characteristics which are likely to favor
escape by plants from their adapted enemies";
this has been echoed elsewhere (Janzen, 1970;
Rhoades & Cates, 1976; Feeny, 1976). In addi-
tion, selective pressure for phenological changes
among Mediterranean crucifer-feeding insects
may not be very great, since cruciferous hosts
are available later in the season. There also may
e phytochemical barriers that function with
phenology in much the same manner as we en-
visage for nonadapted herbivores. Many cruci-
fers contain diverse glucosinolates as well as oth-
er allelochemicals, and the performance of
crucifer feeders frequently varies greatly among
host plants (Root & Olson, 1969; Feeny, 1976;
Rodman & Chew, 1980).

HOW ARE PATCHES FORMED?

The ability to form patches clearly involves
more than early phenologies, since we found few
significant phenological differences between

editerranean patch- and non-patch-forming
crucifers (Table 4). In Mediterranean and desert

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

regions of Israel, late-winter rains initiate ger-
mination of seeds of crucifers and other annuals.
By early spring, patches of various crucifers dom-
inate the landscape of much of Israel (Shmida &
Auerbach, 1983). Bell & Muller (1973) described
similar patch formation by an introduced cru-
cifer, Brassica nigra, in some annual California
grasslands. There, B. nigra produces monospe-
cific patches throughout the grasslands despite
synchronous germination of all annual species
following late-autumn rains. Bell & Muller (1973)
found that patch formation by B. nigra ips
production of potent allelopathic chemicals.
do not know if allelopathy is important in nth
formation by Israeli crucifers. Growth rate ap-
pears to have a role in patch formation. Many
crucifers in Israel and elsewhere (Feeny, 1977)
mature and set seed extremely rapidly, a trait
associated with their occurrence in environmen-
tally harsh or disturbed sites. In addition, patch
formation may involve differential responses to
microsite variability in edaphic parameters and
grazing pressure.

WHY FORM PATCHES?

Patch formation may be reinforced by com-
petition for pollinators and increased pollination
ficiency. Most crucifers, including Israeli

pecies, are either facultative or obligate out-
crossers (Fryxell, 1957; Crisp, 1976; D. Zohary,
pers. comm.). Seasonally advanced flowering
among Mediterranean crucifers may reduce in-
terspecific competition for pollinators, provided
flowering does not occur before pollinators are
available. Patch formation among these species
may also increase pollination success over that
of solitary individuals if pollinators are limited.
Thus, patch formation could be reinforced by
pollination success.

While patch formation may concentrate re-
sources and produce attractive displays for pol-
linators (Handel, 1983; Rathcke, 1983), intra-
specific competition for pollinators among
members of a patch may increase if patch size
becomes sufficiently large. For example,
Rathcke's (1983) density-visitation model en-
visages a facilitation in pollination as patch size
increases up to the point where pollinators are

V O

advanced phenologies result in flowering before
the annual peak in pollinator availability

1987]

(Schemske et al., 1978). As intraspecific com-
petition intensifies, selection for enhanced at-
tractiveness may result in differences between
patch- and non-patch-forming species in polli-
nation-associated traits.

In our analyses, two morphological attributes
frequently associated with pollination, plant
height and petal size (Faegri & van der Pijl, 1979;
Waddington, 1979; Waser, 1983a), did vary be-
tween non-patch- and patch-forming species.
Among Mediterranean species, patch formers are
taller and they generally have larger petals than
do non-patch-forming species. Although we do
not know the intensity of intraspecific competi-
tion for pollinators in patches, increases in height
and petal size were also positively associated with
our measure of patch size. In addition, plant
height, petal size, and indices of patch size were
positively correlated with propensity for out-
crossing as measured by pollen/ovule ratios (Ku-
nin, unpubl. ms.). Thus, taller patch formers ap-
pear to have a greater tendency towards
outcrossing than do smaller patch formers, and
species that do not form patches have the lowest
tendency to outcross. That high levels of ultra-
violet reflectance and patterning is positively
correlated with patch formation found among
Mediterranean species also suggests that patch
formation may result in increased intraspecific
competition for pollinators.

Differences in height and petal size do not oc-
cur among desert species; however, desert patch-
forming species do begin flowering before non-
patch formers, a difference observed among
Mediterranean species only when the early-flow-
ering pygmy guild is excluded from analyses. Ad-
vanced flowering among the desert patch formers
may reduce interspecific competition for polli-
nators (Waser, 1983b) and possibly permit pol-
linator specialization. Some pollinators in other
deserts are capable of synchronizing with the
ephemeral phenologies of their hosts (Baker &
Hurd, 1968; Crawford, 1981). With the excep-
tion of the pygmy guild, a high proportion of
desert patch formers have purple flowers (67%
patch vs. 27% nonpatch), which suggests pos-
sible pollinator specialization or segregation be-
tween patch- and non-patch-forming species.

In summary, we suggest that growing in patches
aids in pollinator attraction, especially in con-
junction with atypical phenologies. Displaced
phenologies also may function to segregate pol-
linator and herbivore guilds temporally. If pol-
linator efficiency is high and seed dispersal low,

AUERBACH & SHMIDA—ISRAELI CRUCIFER PATCH FORMATION

393

patch formation may be self-perpetuating, be-
cause of concentration of seeds into localized seed
banks

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Israel.

FLAVONOID SYSTEMATICS OF SEVEN SECTIONS
OF LUDWIGIA (ONAGRACEAE)!

JOHN E. AvERETT,? PETER H. RAVEN,? AND ELSA ZARDINP

ABSTRACT

Da ita are presented for the flavonoids of 24 species vm seven sections of Ludwigia, all of which

sections reported suggests that these sections are advanced in this fe

al of eight flavonoids, three glycoflavones,

ature, as does a EUM reduction

of structural types and a reduction in the number of glycosidic substitutions in them

As part of a comprehensive study of the fla-
vonoids of Onagraceae, we are herein reporting
results from seven sections of Ludwigia. As ex-
plained in earlier papers (Averett et al., 1978,
1979), the objectives of the overall study on On-
agraceae are to provide an analysis of the fla-
vonoids at the generic level for the entire family
and to gain insight into the evolution of flavo-
noid compounds by correlating substitutional and
structural changes with a phylogeny based on
other systematic data. This is the first of several
papers in which we shall present the results of
flavonoid analyses of Ludwigia. The only pre-
vious report on flavonoids in Ludwigia is a brief
summary of data for the whole genus (including
that reported here) in Averett & Raven (1984).

udwigia is the only genus of the tribe Jus-
siaeeae and comprises approximately 82 species
found in wet habitats in both temperate and trop-
ical regions worldwide (Raven, 1963). Ludwigia
appears to represent a branch of the family dis-
tinct from all other members (Eyde, 1977, 1978,
1981; Raven & Tai, 1979). It is therefore of par-
ticular interest to consider the evolution of fea-
tures in this isolated evolutionary line.

This report deals with the species included in
the original broad circumscription by Munz
(1942; see also Raven, 1963) of sect. Myrtocar-
pus, a group of some 23 species centered in trop-
ical and subtropical South America that ap-

peared to be “phylogenetically central" in the
genus (Ravon, 1963). Based on his studies of the

and

, cyto logy
of this complex, Ramamoorthy (1979) divided
sect. Myrtocarpus into seven sections, com-
menting that the species had been grouped pri-
marily on the basis of shared primitive charac-
ters. Subsequent work has suggested that his sect.
Michelia is not distinct from sect. Myrtocarpus
sensu stricto, and that Ludwigia mexiae (Munz)
Hara is sufficiently distinct from other members
of sect. Pterocaulon that it is best treated as the
monotypic sect. Cinerascentes (Ramamoorthy &
Zardini, 1987). Thus we now recognize seven
sections in this group, delimited as follows: sect.

ocarpus with 20 species (14 examined
herein), including some with the most primitive
assemblages of characters in the genus; sect.
Pterocaulon with five species that comprise a well-

ve monotypic
Amazonia, Heterophylla, Tectiflora, Humboldt-
ia, and Cinerascentes—that are each specialized
relative to sect. Myrtocarpus (Ramamoorthy &
Zardini, 1987).

MATERIALS AND METHODS

Dried leaf materials from 24 species of seven
sections of Ludwigia were examined for flavo-
noids. Approximately 120 populations in total

! We gratefully acknowledge support from the U.S. National Science Foundation through individual grants

to Averett and Raven. Plant material was received from

. P. Ramamoorthy to whom we are especially indebted

for collecting in Brazil most : the species analyzed for this study. W. D. Stevens also collected material, and

his assistance is greatly appre
ment of Biolo
3 Missouri Botanical Ga , P.O. Box

ANN. MIssouRI Bor. GARD. 74: 595-599. 1987.

ate
ogy, aes of Missouri, St. Louis, Missouri 63121, U.S.A.
rden 299, St. Louis, Missouri 63166, U.S.A.

596

and as many as 25 populations of some of the
more variable species were sampled. The sam-
ples included most chromosomal races of the
species concerned. Voucher specimens are listed
in el.
The leaf material was extracted overnight in
85% methanol and the resulting extract was ex-
amined by two-dimensional paper chromatog-
raphy. Certain of the extracts were analyzed us-
ing TLC (polyamide and cellulose) as ~~ In
some cases, the flavonoids were crudely
rated on Sephadex LH 20 witha iese hana
system as described by Hiermann et al. (1978).
For structural elucidation, replicate chromato-
grams were run and the isolated compounds cut
from the paper for further purification and anal-
ysis. The quantity of leaf material varied ac-
cording to usage, but approximate amounts were
0.5-1.0 g for general screening, 5-10 g for rep-
licate chromatograms, and 20-30 g for column
hromatography. Identifications of the glyco-
sides, their aglycones, and sugars were made as
previously described (Averett et al., 1978, 1979)
and were compared with standard Rf values and
absorption maxima (Averett, 1977). In addition,
most of the aglycones and sugars were run, along
with authentic reference compounds, by circular
thin-layer chromatography as described by Ex-
ner et al. (1977). Base hydrolysis was employed
to determine acylation but the unction
was not determined.

RESULTS AND DISCUSSION

Eight flavonoids were found among the species
sampled (Table 2): orientin (1), isoorientin (2),
orientin-O-acylate (3), quercetin 3-O-rhamno-
side (4), quercetin 3-O-arabinoside (5), quercetin
3-O-glucoside (6), quercetin 3-O-diglucoside (7),
and quercetin 3-O-rutinoside (8). Compounds 1-
3 are glycoflavones and compounds 4-8 are fla-
vonol glycosides. All of the compounds are based
on structures having two hydroxyl substituents
in the B-ring.

Some infraspecific variation was found, es-
pecially in such variable species as Ludwigia pe-
ruviana, L. elegans, and L. laruotteana. Al-
though variation was present, no flavonoid unique
to any species was found in a single population.
We also detected differences in concentrations of
compounds between populations of some species,
but made no attempt to document these differ-
ences. Interspecific variation within the larger
sections is apparent but, like populational vari-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

ation, is largely confined to the presence or ab-
sence of certain compounds. The more interest-
ing variation is found in intersectional
comparisons. Because of this, samples are ar-
ranged by species and grouped in their respective
sections in Table 2

Compound 3, the acylated glycoflavone, is
found in only five species in three different sec-
tions, including the closely related sections Cine-
rascentes and Pterocaulon. Compounds 5 and 7
are the next most frequently absent, occurring in
only six and seven species, respectively, and typ-
ically in low concentration when present. Com-
pounds 1, 2, and 8 are the least variable and are
found in 18, 20, and 19 of the species, respec-
tively. Compound 2 is present in all species in
which glycoflavones occur, and compound 8 is
present in all species in which flavonols are found.
Compounds 4 and 6 occur in 13 and 14 of the
species sampled. Glycoflavones are found in 20
of the 24 species, flavonols in 19, both glycofla-
vones and flavonols in 15, only glycoflavones in
five, and only flavonols in four

All ofthe species have at least two compounds
and none has more than seven. The average
number of compounds per species is 5.37 for the
entire group. The comparison of numbers of
compounds at the sectional level is especially
interesting. Species of sect. Myrtocarpus, which
has relatively generalized features, and sect.

(Piero aalon. 3.0 (Humboldtia and Heterophyl-
la), and 2.0 (Amazonia and Tectiflora).
Flavonols are present throughout Onagraceae
and glycoflavones are present in all tribes except
Fuchsieae and Epilobieae (Averett & Raven,
1984). Glycoflavones are especially well repre-
sented in Circaea (Boufford et al., 1978; Averett
& Boufford, 1985). The presence of glycoflavones
is considered primitive relative to the presence
of other groups of flavonoids (Harborne, 1977),
and the distribution of these compounds within
Onagraceae does not contradict that contention.
Except for Ludwigia sericea, which has only
flavonols, all species examined of sect. Myrto-
carpus and the single species, L. mexiae, of sect.
Cinerascentes have both glycoflavones and fla-
onols. The remaining monotypic sections ex-
hibit either glycoflavones only—sections Heter-
ophylla and Amazonia—or flavonols only—
sections Humboldtia and Tectiflora. Collective-
ly, the species of sect. Pterocaulon exhibit both

ABLE 1. Voucher specimens of Ludwigia used for flavonoid analysis in this study. Specimens of all material
deposited at MO, unless otherwise indicated.

Ludwigia section Amazonia Ramamoorthy
Ludwigia densiflora (Micheli) Hara. BRAZIL. RONDONIA: Duarte 7329 (MO, RB).
Ludwigia section Cinerascentes Ramamoorthy & Zardini
Ludwigia mexiae (Munz) Hara. BRAZIL. PARA: Ramamoorthy 652.
Ludwigia section Heterophylla Ramamoorth
Ludwigia inclinata (L. f.) Gómez. MEXICO. OAXACA: Breedlove & Raven 13686. CosrA RICA. PUNTARENAS:
Stork & Horton 8912 (US). BRAZIL. AMAPA: Froes & Black 27732 (IAN).
Ludwigia section Humboldtia
Ludwigia sedoides (H. & B.) Hara. PANAMA. CANAL ZONE: D'Arcy 12350. BRAZIL. PARA: Archer 8411 (RSA).
Ludwigia section Myrtocarpus (Munz) Hara
Ludwigia albiflora Ramamoorthy. BRAZIL. GoIAS: Ramamoorthy 545. MINAS GERAIS: Ramamoorthy 427

MO, SP).

me d (Hassler) Hara. BRAZIL. MATO GROSSO Do suL: Ramamoorthy 610 (MO, SP), Ramamoor-
thy &

Ludwigia jou (Camb.) Hara. BRAZIL. Goras: Ramamoorthy 532, 560, 564. MINAS GERAIS: Ramamoorthy
403,4 4; Ramamoorthy & Vital 140; Ramamoorthy et al. 148, 153, 161, 168, 176, 178, 179, 181
(MO, SP), 184, 195, 299, 301, 308, 309, 319. RIO DE JANEIRO: Ramamoorthy et al. 291. SAO PAULO:
Ramamoorthy 379, 384, 395; Ramamoorthy & Vital 112; Ramamoorthy et al. 196

Ludwigia hassleriana (Chodat) Hassler. BRAZIL. MATO GROSSE DO SUL: Ramamoorthy 629, Ramamoorthy et

b

Ludwigia irwinii Ramamoorthy. BRAZIL. MINAS GERAIS: Ramamoorthy et al. 142. SAO PAULO: Ramamoorthy
80; Munz 15406 (NY, POM, US).

Ludwigia laruotteana (Camb.) Hara. BRAZIL. Goras: Ramamoorthy 419, 420. MINAS GERAIS: alga
101; Ramamoorthy & Vital 90; RE et al. 143, 147, 160, 172, 188, 311. SAO PAULO: Ram
moorthy 69.

Ludwigia martii (Micheli) Ramamoorthy. BRAZIL. MINAS GERAIS: G/aziou : ” (B, C, F, P, R).
Ludwigia myrtifolia (Camb.) Hara. BRAZIL. MINAS GERAIS: Ramamoorthy
Ludwigia nervosa (Poir.) Hara. M ZELAYA: maka 8275. onl BAHIA: ti eoa be et HR 328.
DISTRITO FEDERAL: Ramamoorthy 526. goras: Ramamoorthy 561; Ramamoorthy & Vital 544
rosso: Ramamoorthy 571. MATO GROSSO DO SUL: Ramamoorthy 605, 607. MINAS GERAIS: cede
et al. 170. sao PAULO: Ramamoorthy 393 (MO, SP); Ramamoorthy & Vital 78 (MO, SP).

Ludwigia peruviana (L.) Hara. BRAZIL. MINAS GERAIS: Ramamoorthy 366. PARANA: Ramamoorthy 207, 275,

281.

Ludwigia pseudo-narcissus (Chodat) Ramamoorthy. BRAZIL. PARANA: Ramamoorthy et al. 283.
Ludwigia rigida (Miq.) jvc SURINAM: Pulle 475. VENEZUELA. COJEDES: Pittier 11711 (B, US, VEN).
P sericea (Camb.) H RAZIL. MINAS GERAIS: Ramamoorthy 68; Ramamoorthy et al. 157, 158,
s PARANA: Ramamoorthy et al. 215, 216, 288, 289. SANTA CATARINA: Ramamoorthy et al. 240.
SAO PAULO: Ramamoorthy 44
Ludwigia pistas (Camb.) Hara. BRAZIL. BAHIA: Ramamoorthy et al. 336. DISTRITO FEDERAL: Ram
moorthy 513; Ramamoorthy et al. 349, 351. Goras: Ramamoorthy et al. 342, 344, 345, 506. MATO
Grosso: Ramamoorthy & Vital 579. MINAS GERAIS: Ramamoorthy 163, 164, 165, 405; Ramamoorthy et
al. 187.
Ludwigia section Pterocaulon Ramamoorth
Ludwigia decurrens Walt. NICARAGUA. ZELAYA: Stevens 4916. BRAZIL. MINAS GERAIS: Ramamoorthy et al.
303, 306. SANTA CATARINA: Ramamoorthy et al. 258, 259.
po erecta (L.) Hara. MEXICO. OAXACA: Breedlove & Raven 13669 (DS, MO). NICARAGUA. ZELAYA:
Stevens 8274. CUBA. ORIENTE: Ekman 6537 (S). COLOMBIA. HUILA: Smith 1204 (GH, UC, US).
hrai RHODESIA. NDANGA: Goodier 977.
Ludwigia filiformis (Micheli) Ramamoorthy. BRAZIL. GOIAS: Ramamoorthy & Vital 555. SAO PAULO: Rama-
moorthy

73:
pu longifolia (DC.) Hara. BRAZIL. MINAS GERAIS: Ramamoorthy & Vital 96; Ramamoorthy et al. 150.
A CATARINA: Ramamoorthy et al. 231, 233, 237.
a major (Micheli) u n BRAZIL. RIO GRANDE DO SUL: Ramamoorthy et al. 245.
Ludwigia section Tectiflora Ramamoorthy
i latifolia (Benth.) Hara. NICARAGUA. RIO SAN JUAN: Neill 3361. GUYANA. WEST DEMARARA: Ma-
e & Fanshave 22951 (NY, U, US). PERU. SAN MARTIN: Ferreyra 18506, Williams 7153 (F, US).

597

598

fA 13 A

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

f Ludwigia. + = flavonoid detected; dm flavonoid

TABLE2. Distribution
not detected. Key: 1 = orien

n, 2 = isoori rientin, 3 — orientin-O-acylate,

4 = quercetin-3-O-rhamnoside, 5 =

quercetin-3-O-arabinoside, P. — quercetin-3-O-glucoside, 7 — quercetin- -3- O-diglucoside, and 8 = pene nes
oside.

O-rutin
Glycoflavones Flavonols
l 2 3 4 5 6 7 8

Sect. Amazonia

L. densiflora + + 0 0 0 0 0 0
Sect. Cinerascentes

L. mexiae + + + 0 + 0 0 +
Sect. Heterophylla

L. inclinata + + + 0 0 0 0 0
Sect. Humboldtia

L. sedoides 0 0 0 + 0 + 0 +
Sect. Myrtocarpus

L. albiflora + + 0 + 0 + 0 +

L. bullata + + 0 + 0 + 0 +

L. elegans + + 0 + + + + +

L. hassleriana 0 + 0 + 0 0 + +

L. irwinii + + 0 + + + + +

L. laruotteana + + 0 + + E E ~

L. martii 0 + 0 0 0 0 + +

L. myrtifolia + + 0 + 0 + 0 +

L. nervosa + + 0 + 0 + 0 +

L. peruviana + + 0 t + + + +

L. pseudo-narcissus + + 0 0 0 0 0 +

L. rigida + + 0 0 0 0 0 +

L. sericea 0 0 0 + 0 + 0 +

L. tomentosa + + 0 + + + 0 4
Sect. Pterocaulon

L. decurrens + + 0 0 0 + + +

L. + + + 0 0 0 0 0

L. filiformis 0 0 0 + 0 + 0 +

L. longifolia + + + 0 0 0 0 0

L. major + + + 0 0 0 0 0
Sect. Tectiflora

L. latifolia 0 0 0 0 0 + 0 +

glycoflavones and flavonols, but only one species,
L. decurrens, has both classes of compounds. One
species, L. filiformis, has only flavonols, and the
remaining three species have only glycoflavones.
Thus, if sect. Pterocaulon is a monophyletic group
derived from sect. Myrtocarpus, as is indicated
from morphological studies, then within this sec-
tion of five species one has lost the ability to
produce glycoflavones and three to produce fla-
vonols; that, at sd would be the most parsi-
monious explanat

The five denied sections we are consid-

ering here do not appear to be more closely re-

lated to sect. Myrtocarpus, on the basis of their

overall characteristics, than they do to any other
dwigia

t
and there is no evidence of a direct relationship
between any two. The overall similarity of fla-
vonoids between sections ee and Hum-
boldtia and between sectio d Het-
erophylla could not, en. be taken as an
indication of relationship between those taxa.
Rather, the similarity of flavonoids between these

599

AVERETT ET AL.—FLAVONOID

1987]

groups seems to reflect parallel and independent
loss of particular classes of flavonoids and/or in-
dividual compounds, a trend that has character-
ized the evolution of the genus overall. A further
evaluation of their relationships, which must be
multidimensional, would need to take into ac-
count the remainder of the genus. It does appear,
in terms of admittedly largely plesiomorphic
characteristics, that sections Pterocaulon an
Cinerascentes are more clearly related to sect.
Myrtocarpus than are the others.

n summary, our analysis has revealed a pat-
tern of loss of individual flavonoids and groups
of flavonoids in the seven sections of Ludwigia
that we have considered in this paper. Further
resolution of the relationships of these species
must await more detailed study.

LITERATURE CITED

AVERETT, J. E. 1977. Absorption maxima and R
values as an aid to the identification of selected
flavonoids. Phytochem. Bull. 1 0-26.

FFORD. 1985. The flavonoids and

flavo noid ps of Circaea (Circaeeae, On-

agracea ux Sy y pa 3-373.

984. Flavonoids of Ona-
graceae. redu Missouri Bot. Gard. 71:
, B. KERR & . RAVEN. 1978. Flavon oids
of Onagraceae: ppm sect. Epilobium. Amer
J. Bot. 65: IRAE:
N & H. BECKER. 1979. Flavonoids
i oe Epilobieae. Amer. J. Bot. 66: 1151-
155.

TICS OF LUDWIGIA

BourroRD, D. E., P. H. RAVEN & J. E. AVERETT. 197
Glycoflavones i x cuis (Onagraceae). Biochem.

Syst. & Ecol. 6: 59-60.
EXNE E Lei KER. 1977. Circular
chroma a co hy-

o
lution in Ludwigia (Onagraceae). I. Androecium,
placentation, merism. Ann. Missouri Bot. Gard.

64: 644-655
. 1978. Re Mdh pi structures and evolution
in Ludwigia (Onagrace and seed. Ann

. II. Fruit

Missouri Bot. Gard. 65: 6—675.

1981. Reproductive structures and evolution

in Ludwigia (Onagraceae). III. Vasculature, nec-

taries, conclusions. Ann. Missouri Bot. Gard. 68:
379-412.

HARBORNE, J. B. 1977. Flavonoids and the evolution
of angiosperms. Biochem. Syst. Eco 2
HIERMANN, A., J. ExNER, H. BECKER & J. E. “AV VERETT.

1978. Gel filtration of flavonoids. Phytochem.

1942. Studies in Onagraceae XII. A
revision of the New World species of Jussiaea.
Darwiniana 4: 179-284.

1979. A sectional revision of

Ann. Missouri Bot. Gard. 66: 8

. M. ZARDINI. 1987. The systematics and

evolution of Ludwigia sect. Myrtocarpus sensu lato

Die prends Monogr. Syst. Bot. Missouri Bot.
ard. 19: 1-120.

RAVEN, P. H. TA The Old World species of Lud-
wigia (including Jussiaea), with a synopsis for the
genus (Onagraceae). Reinwardtia 6: 327-427.

& W. Observations of chromo-
somes in Ludwigia —ewaspupa Ann. Missouri
Bot. Gard. 66: 862-879.

A CHEMOTAXONOMIC CLASSIFICATION OF THE
SOLANACEAE!

PETER TETENYI’

ABSTRACT

Alkaloids and steroids in the Solanaceae are reported extensively in the baa nse By examining

the in deen pai leading to pape alkaloids, the pathways can be v
which th . Arrangem

c us compounds can be der

sualized as a spi
ment of the genera of Solanacea

their Padus contents in mesi em to pus igh — traditional classifications of the family,
w subfamilie

t
unalia must be allied with Jaborosa in tribe olas

Although solanaceous species are well known
to afford an array of alkaloids and steroids, the
family has not been arranged yet according to
chemical features. Indeed, it is difficult to find
reports of most features because, except for the
occurrence of calcium oxalate crystals, only their
absence is recorded. Thus Philipson (1977) con-
cluded that a lack of iridioids characterizes the
Solanaceae within the Unitegminae. Dahlgren
(1975, 1980) came to similar conclusions, adding
the deficiency of polyacetylenes as a character-
istic. Sporne (1980) believed that absence of leu-
coanthocyanins and ellagitanins are chemical
characteristics of the family and indicated that
whether or not the seeds contain endosperm may
or may not be significant.

In an investigation of the cytochrome-C and
plastocyanin amino acid sequence, Boulter et al.
(1979) placed the Solanaceae far from the As-
teraceae in their family tree, and although they
related it closely to the Scrophulariaceae, they

& Emberger (1960) that the Solanaceae and Cap-
rifoliaceae are closely related based on embry-
ological characters. Nevertheless, mature sola-
naceous plants are typical alkaloid-accumulators,
whereas mature caprifoliaceous plants have
phenol-glycosides (Hegnauer, 1973). In the same
publication Boulter et al. (1979) put the tomato
alongside the potato and separated tobacco and
the woody Solanum crispum Ruiz & Pavón, and
they held Capsicum frutescens L. to be signifi-

1c

! This paper was part of the Second I

s due to their
e must be a pei ed as separate
and exclusive uod synthesis. Acnistus and

cantly different from the preceding species based
on the amino acid complement.

The use of active principles found in different
Solanaceae to construct systematic schemes can
be accepted only when it can be demonstrated
that the biosynthetic routes leading to these
chemical structures are homologous (Tétényi,
1973). The valid chemical patterns are in the
various biosynthetic pathways and not in the

thetic pathways in alkaloid production in the So-
lanaceae in Table | and Figure 1. Numbers in
the following paragraphs correspond to those of
Table 1 and Figure 1.

The first evidence supporting this scheme lies
in the well-known, genetically determined chem-
ical differentiation in infraspecific chemotaxa of
Duboisia myoporoides R. Br. (Tétényi, 1970). The
characteristic active alkaloid ingredient, nicotine
(1), in one chemotaxon ofthis species is the result
of the synthesis of ornithine and tryptophan to
an alkaloid. Another chemotaxon of D. myopo-
roides is characterized by a splicing of the as-
partate pattern (lysine) and acetyl-CoA to the
alkaloid biosynthesis, and the main alkaloids are
then anabasine (A) and isopelletierine (B). In a
third infraspecific chemotaxon, an entirely dif-
ferent pattern occurs: an a-face nucleophilic at-
tack instead of the 8-face one of the N-methyl-
A'-pyrrolinium salt (Fig. 2; Leete, 1979) leads to
the pathways indicated by the solalkoid spiral
(Fig. 1), that is, the linking of ornithine and ace-
tyl-CoA to hygrine (2) and then the development

on the Biology and Systematics of the Solanaceae

presented at the Missouri Botanical Garden on 3-6 August t 1983.
? Research Institute for Medicinal Plants, P.O. Box 11, 2011 Budakalász, Hungary.

ANN. MISSOURI Bor. GARD. 74: 600—608. 1987.

1987]

TABLE l.

TÉTÉNYI—CHEMOTAXONOMIC CLASSIFICATION OF SOLANACEAE

Alkaloids of Solanaceae included in and arranged conforming to the solalkoid spiral.

Nicotine and its derivatives
ug (1)
HN;
p 11-5]
Tropane alkaloids
hygrine (2)
C,H,;N;O,
[496-49-1]
cuscohygrine (4)
C. H, NO
[454-14-8]
tropinone (5)
C,H,,NO
[532-24-1]
isovaleryloxytropane (6)
13H23 y.
[490-96-0]
tigloidine (7)
C.H; NO;
[495-83-0]
scopolamine (8)

Other alkaloids
fabianine (—)
O

anabasine (A)
C l oH ThE
[494-52-0]
hygroline (2)

aH;
[1617-83-0]

physoperuvine (5)

8 15
[60723-27-5]

tigloyloxytropane (7)

134421 3
[55727-41-8]
hyoscyamine (8)

iH; NO;
[101-31-5]

8-carboline (C)

isopelletierine (B)
C,H;,NO
[539-00-4]
withasomnine (3)
12 15 a
[10183-74-1]
belladonnine (4)
C,,H4;N; 4
[6696-63-5]
tropine (5)
C,H,;NO
[120-29-6]
senecioyloxytropane (6)
134421 O,
[77101-57-6]
valtropine (7)
physochlaine (8)
C,,H2;NO,
[54357-41-4]

capsaicine (D)

CHa, N n N.O 184427 3
[6871-51-8] [244-63-3] [54357-41-4]
betaine (E) Pictur (E)
C. Hi NO, C;H;; NO,
[107-43-7] [62- 49- 7]

Explanation: common name; symbols in parentheses refer to Figure 1
formula

[registry number]

of other simple and ester tropane alkaloids. Thus
pyruvate-leucine yields valeroidine (6) with iso-
valeric acid; aspartate-isoleucine yields tiglo-
idine (7) with tiglic acid; and om phe-
nylalanine and tropic acid,
becomes the characteristic alkaloid of the third
chemotaxon.

The genus Duboisia R. Br. provides further
evidence of this kind of chemical differentiation
separating the infraspecific taxa of D. myopo-
roides from one another. In D. hopwoodii F.
Muell., nicotine (1) and its derivatives are syn-
thesized by the 8-face attack of ornithine, where-
a

this alkaloid is accumulated through the con-
verse formation. Thus the scheme in Figure 1 is
true for Duboisia as a genus as well as for its
components.

Two other genera examined in subfamily An-
thocercidoideae (described on p. 607)— which

includes Duboisia —show the same diversity in

asine (A), cus e (4) opine (5), va-
leroidine (6), tigloidine (7), and scopolamine (8)
have been detected.

Alkaloid synthesis in subfamily Cestroideae
Schldl. also proceeds according to Figure 1, but

its derivatives, a somewhat different picture from
the Anthocercidoideae. Thus nicotine (1) is the
sole product in the genus Cestrum L., whereas
fabianine, a tetrahydroquinoline alkaloid arising
from a biosynthesis preceding the pyridine-nu-
cleotide cycle, is accumulated as well as nicotine
in Fabiana Ruiz & Pavón. The aspartate-lysine
path is switched in Streptosolen Miers, and an-
abasine is accumulated in addition to nicotine.
The alkaloid spectrum of Nicotiana L. and Sal-
piglossis Ruiz & Pavón by intervention of acetyl-
CoA, accumulates isopelletierine (B) in addition

602

SOLALKOID SPIRAL

P-hydroxypyruvate
serine

Pere oe

tryptophan
aspartate

ae
ornithin

phenylalanine

DOSE DN

(0 nicotine

(2) hygrine

(3) withasomnine

(4) cuscohygrine

(5) tropinone-physoperuvine
isovaleryl

(8) sonecioyl d

ED) tigloyl

(8) scopolamine-hyoscyamine

tropane

FIGURE 1.

from plants belongi ng t e Solan Am cids
alkaloid biosynthesis in “issuu are e clarified ir in rn

to the above substances. Nicotiana is even en-

kaloid of Vestia Willd., which contains a 8-car-
boline skeleton.

My scheme might not be tenable if the Ces-
troideae were not differentiated chemically also.
In fact, the alkaloid biogenesis of hygrine deriv-
atives is found in this subfamily too, although
only Brunfelsia L. accumulates just cuscohygrine
(4) and pyrrole-3-carboximidine, which was also
detected in Nierembergia Ruiz & Pavón, whereas

4 th z A in v. L: L

ANNALS OF THE MISSOURI BOTANICAL GARDEN

>

A

B isopelletierine
c

D

E

realt

[Vor. 74

pyruvate
leucine-valine

anabasine

B-carboline
capsaicine

choline-betaine

Solalkoid m Psp on ey ea Piet load patterns of most important alkaloids isolated
of

d d starting and combined points. Patterns

Ruiz & Pavón. However, the alkaloid spectrum
in Schizanthus is unique in the whole family with
its hygroline (occurring also in the Erythroxy-
laceae) and because the sencioyl and angeloyl
tropane esters (6) are formed from pyruvate-leu-
cine and ornithine. Formation of the alkaloid
valeroidine in Schizanthus by this biosynthetic
pathway is characteristic of the family Solana-
ceae as well as of Anthocercis Labill., Cyphanth-
era Miers, and Duboisia. However, taking into
account the two kinds of hygrine synthesis, the
predominance of nicotine in its various deriva-
tives (A, B, C), and the primary substances brun-
felsamine and fabianine, we can assert that the

1987]

TETENYI—CHEMOTAXONOMIC CLASSIFICATION OF SOLANACEAE

(n- (2R) - Hygrine

FIGURE 2.

chemical differentiation of the Cestroideae, in
having an alkaloid synthesis with several simi-
larities to that ofthe Erythroxylaceae, Rubiaceae,
Elaeagnaceae, and Malpighiaceae, significantly
surpasses that of the Anthocercidoideae.
Chemical differentiation in subfamily Sola-
noideae is of a different character than in the
Cestroideae, for here the hygrine derivatives are
always preponderant, and the differences are in
the ever-increasing complexity of the molecular
entities synthesized by the different genera as if
overcoming a barrier. Similar to the case of Brun-

(4)
Adans. starts from hygrine (2) and, like CypAo-
mandra Sendtner, reaches the formation of tro-
pinone (5). The alkaloid physoperuvine (5) of
Physalis L. is equivalent to this, although tiglo-
idine (7), an alkaloid of this genus, is an ester
but not with tropic acid. In Withania Pauquy
aspartate-isoleucine and ornithine are synthe-
sized to 3'-tigloyloxytropane without the pres-

(-)- (25)- Nicotine

Nucleophilic attack of N-methyl-A'-pyrrolinium salt from the a- and 6-face (Leete, 1979).

ence of any tropic ester, but the alkaloid spec-
trum of this genus is very broad; thus
withasomnine (3) is formed by coupling orni-
thine to phenylalanine. Isopelletierine (B) from
lysine and acetyl-CoA and even the alkaloid cho-
line (E) arise from the hydroxypyruvate synthesis
pathway. These are only the main types repre-
sented. In the genera Leucophysalis Rydb. (Phys-
aliastrum japonicum (Franchet & Savat.) Hon-
da) and Lycium L. of this subfamily, the unique
alkaloid detected so far is betaine (E), formed
from the serine cycle in the same way as choline.

synthetic routes from nicotine (1) to scopolamine
(8), while on the other hand, they also form and
accumulate the alkaloid valeroidine (6) like An-
thocercis and Duboisia. In the genus Solandra
Sw., valtropine (7), originating from the coupling
of isoleucine with ornithine, follows a path of
alkaloid biosynthesis homologous to that in Cy-
phanthera and Grammosolen Haegi. However,

604

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Differences of t um derivative alkaloids in subfamily Atropoideae. Data from

TABLE 2.
Romeike (1978) except where noted otherw

Genus
Man-
Hyoscy- Scopo- Whit- dra- Prze- Atro- Physo-
Alkaloids Atropa amus lia leya gora walskia | panthe | chlaina
Physochlaine +
Hyoscyamine + + + + + + + +
3'-tigloyloxytropane + + +
Belladonnine + +
Tropine + + + +
Tetramethylputrescine +

A-N-methylornithine
(Hedges & Herbert,
1981) +

a related feature in the genus Anthotroche Endl.
may be that hyoscyamine (8) predominates in
the spectrum. The hygrine (2) of Salpichroa Miers
also shows relationship to Duboisia, bu e
hyoscyamine (8) is the characteristic alkaloid,
similar to the content of tigloidine (7) in Acnistus
Schott and the small amount of scopolamine (8)
E Latua Phn, ds combination of pyruvate-va-

(D) supports
the curious alkaloid ere and the isola-
tion of Capsicum L. as stated by Boulter et al.
(1979)

The system shown in Figure | is also supported
by the alkaloid synthesis in subfamily Atropo-
ideae (described on p. 607). In contrast to the
Solanoideae, the simple alkaloid synthesis of the
Atropoideae is subordinate to that of tropane
esters. This is illustrated by the data in Table 2,
for initially nicotine formation is only inhibit-
ed—as exemplified by Atropa L. and Hyo-
scyamus L.—and the genera Scopolia Jacq. and
Whitleya Sweet (Anisodus Link) are the most
advanced cases in which a simple tropane base
could be detected. Occurrence of tigloyl ester (7)
can be verified in Mandragora L. only by the
presence of cuscohygrine (4) and a little scopol-
amine (8). Przewalskia Maxim. and Atropanthe
Pascher accumulate mainly hyoscyamine (8),
which is characteristic in the whole family and
occurs in each genus. Although the result of the
synthesis in Physochlaina G. Don is also pre-
dominantly hyoscyamine, its peculiar alkaloid,
physochlaine (8), is the tropane ester of
4 si beastacsticacid: nol "Piu oM"

=
=
Q
€

r

rivative.

The alkaloid-forming distinctions of solana-
ceous taxa at various levels have been arranged
according to Figure 1. The requirements were
satisfied by this scheme, which we have termed
the solalkoid spiral. One objection may be that
I have not mentioned the connection between
terpenoid and alkaloid biosynthesis. This is be-
cause the steroid synthesis that is characteristic
of the family is exclusive, alternative, and with-
out transition, in contrast to the process of al-
kaloid synthesis, which is gradual and proceeds

y inhibition

"This is poai by the omg * infraspecific
chemotaxa carried out with anolides on
Withania somnifera Dunal UE 1973) as well
as our own analysis of steroidal alkaloid taxa of
Solanum dulcamara L. (Tétényi et al., 1977). No
instance of a steroidal alkaloid in a withanolide-
containing species or the reverse is known, al-
though this may be due to deficiency in our
equipment or methods.

It seems that these two routes of steroid syn-
thesis — steroidal alkaloids versus lactones — rep-
resent a phylogenetic branching alternative with
cholesterol as its starting point. Oxidation leads
to withanolides compared with partial etherifi-
cation to neutral saponins, followed by cycliza-
tion to steroidal alkaloids.

Duboisia —subfamily Anthocercidoideae —
synthesizes only neutral sapogenins, and Cy-
phanthera, Anthotroche, Crenidium Haegi ur-
solic acid only, a further feature pointing to their
primordial endemism. Simple sapogenins were
detected in subfamily Cestroideae: in Combera
Sandw., Fabiana, Nierembergia, Browallia L.,

1987]

TETENYI—CHEMOTAXONOMIC CLASSIFICATION OF SOLANACEAE

605

FIGURE 3. Chemotaxonomic syste t fg
in four subfamilies containing 16 tribes and i Duro Dahlgren o Axes separate most importan
]

isopelletierine type [N-I], O = LE -carb
[H-T], © = capsaicine. Anthoc

3. Nicotianeae = Don. (8).
etieae A. T. Hunz. (1). Solanoideae.—7. Juannullo

Nicandreae Wettst. (1).—14. Lycieae A. T. Hun

des L.A
24 Salpislosolid|eae Benth. (7
oeae A. T. Hunz. (8).
neae-Phy idem (9). —10. Jaboroscae Miers (7).—11. Datureae Reichb. (2).—12
z. (3). Atropoideae.— 15. Discopodiineae Baehni (2).— 16. Atro-

kaloid [ALO = lactone [L]. aru ids (A): V = ae.

oline-betaine, A = hygrine-tropane type
5

hocerceae G. Don. (7). Cestroideae. E bid aaa Don

).—
).— 5. Schwenckieae A. T z. (3).—6. Parabouch-
—8. in Reh G (15).—9. Sola-
. Solandreae Miers (2). —13.

peae Reichb. (9). Numbers in ei mide numbers of genera

Salpiglossis, Vestia, and Streptosolen, as well as
in Cestrum and Nicotiana, but these two have
steroidal alkaloids.

Subfamily Solanoideae, apart from Exodeco-
nus Raf., shows a prevalence of steroidal alka-
loids when the genera Solanum, Cyphomandra,
Capsicum, and Lycopersicon Miller are consid-

pogenin-containing 7rechonaetes Miers, should
be evaluated as quite distinct. This differentiated
steroid synthesis divides subfamily Solanoideae.
The connection between alkaloid and steroid
biosynthesis is shown by the fact that Solanum,

arrested at cuscohygrine, includes species accu-
mulating steroidal alkaloids building in arginine
released from the ornithine cycle in their genesis.
The genera Physochlaina and Scopolia o
subfamily bd ones synthesize kei
yoscyam thus the
rss Is nearer to the subtribe of the Solanoi-
deae characterized by formation of the same ste-
roids.
If we base a chemical classification of the So-
lanaceae on the facts mentioned above, we can-
not accept the analysis of Evans (1979) or
Romeike (1978), who considered only alkaloid
properties in evaluating Wettstein’s century-old
system. I have developed in Figure 3 a chemo-
taxonomic classification of the Solanaceae. It

"-

606

TABLE 3. Characteristic alkaloids and steroids for the chemotaxonomic system of the Solanaceae (corre-

sponding to Fig. 3).

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Tribe Genus Alkaloid Steroid
1 Anthocercis oe, 10 scopolamine”? t
Anthotroche hyoscyam
Crenidium ee anabasine'* T
Cyphanthera scopolamine,'* nicotine!*
Duboisia SEROREN isopelletierine! sapogenin?
Grammosolen scopolamine!?
Symonanthus tigloyl wont
2 Cestrum nicotine? solasodin?
Vestia B-carboline'? diosgenin?
3 Combera — sapogenin?
abiana "en send sapogenin?
Nicotiana betaine,’ isopellet solasodin?
Nierembergia B- nea paces + EE E E sapogenin‘*
4 Browallia sapogenin?
Brunfelsia cu uscohyrine" + brunfelsamine!* —
Salpiglossis isopelletieri sapogenin?
Schizanth a LUN hygrolines' —
Streptosolen nabasine? sapogenin?
8 sicum capsaicine? solanidin?
Cyphomandra tropinone! solasodin!
Exodeconus — neotigogenin?
Jaltomata cuscohygrine? —
Lycopersicon — tomatin’
nu cuscohygrine (solamine)* solasodin?
9 Leucophysalis etaine? —
Margaranthus cuscohygrine* —
Physalis tigloidine,' physoperuvine' physalin?
Withania choline, isopelletierine, 3'-tigloyloxytropane! withanolide?
Witheringia physalin?

10 Acnistus — 19 withanolide!!
Dunalia — withanolide!!
Jaborosa — steroidlactone!!
Latua hyoscyamine! —

Salpichr hyoscyamine! —
Trechonaetes — sapogenin?

11 Brugmansia nicotine, scopolamine! —

Datura nicotine, scopolamine! withanolide?

12 Solandra hyoscyamine! —

13 Nicandra tropinone! withanolide?

14 Lycium betaine? withanolide?

16 ropa choline, '? Mio —
Atropanthe hyoscya —

yoscyamus choline? hyoscyamine’ sapogenin?
Mandragora hyoscyamine —
Physochlaina physochlaine! solanidin?
Przewalskia hyoscyamine!' —
Scopolia choline,'? hyoscyamine! solanidin?
Whitleya choline,'? hyoscyamine! —

T Ursolic acids present.'^

! Romeike,

1978; ? Gibbs, 1974; ? Hegnauer, 1973; * Lorenti et al., 1981; * Evans, 1979; * San Martin et al.,
1980; ? Paris & Moyse; 1971: 8 Evans & Somanabandhu, 1980; ° Antoun et al., 1981; '? Evans & Ramsey, 1981;
u D'Arcy, 1979; '? Gessner, 1977; ? Evans & Ramsey, 1983; '* El Imam & Evans, 1984; 15 Buschi & Pomilio,
1986; '^ Lloyd et al., 1985; 1 Faini et al., 1980.

1987]

considers the chemosyndrome of the Solanaceae,
has reference to the cross-sectional presentation
of Dahlgren (1975, 1980), and draws upon the
systematic data of D’Arcy (1979), Haegi (1979,
1981), and Hunziker (1979).

The surface dimensions of the four subfamilies
correspond to the number of included genera,
while the curves involving the subfamilies label
the affinity points of the homologous chemical
qualities. The two main trends of alkaloid syn-
thesis in the family— accumulation of nicotine-

steroidal alkaloid taxa. Tribes of the subfamilies
are shown by numbers and dotted lines. I have
divided the tribe Solaneae Reichb. into subtribes
Solaninae (Solanineae Dunal) and Physalinae
(Physalidineae Reichb.) on the basis of different
and exclusive types of steroid synthesis and con-
sidering the data of Baehni (1
Acnistus and Dunalia are placed in tribe Jabo-
roseae Miers in accordance with Baehni on the
basis of their steroidal lactone content. Latua
was similarly treated—its bent embryo agrees
with this placement and its alkaloid chemistry
precludes assigning it to subfamily Cestroideae.
e characteristic alkaloid or steroidal data for
the genera shown in Figure 3 are also shown in
able 3. I have not found reliable recent data on
the alkaloid or steroid active ingredient of four
tribes and 43 genera; however, I feel justified in
presenting this review, a novel chemotaxonomic
evaluation of facts known to others, and a clas-
sification system for the Solanaceae.

This new system consists of four subfamilies.
We must separate the Anthocercidoideae from
the Cestroideae, and the Atropoideae from the
Solanoideae because of differences in area of or-
igin, morphology, Bowers; ane enue oe

and are endemic to Australia. They differ mor-
phological from the Cestroideae by having long,
d corolla tubes,
grains. pm
chemosyndrome combines predominantly hy-
grine derivatives— characteri Such pude.
amine — with neutral sapogenins, while the Ces-
troideae have chiefly nicotine derivatives together
with simple and complex steroidal alkaloids, and
only one tropane ester alkaloid in common with
the Anthocercidoideae

The Atopadas; which have a cistmcuive al-
kaloi differ

HRALVIUNL pa J F

TÉTÉNYI—CHEMOTAXONOMIC CLASSIFICATION OF SOLANACEAE

607

also in having dispersed from their l cen-
ter. They are adapted to withstand cold seasons
in Eurasia and have become isolated on islands
mountains in Africa. The Atropoideae

having imb
noideae have valvate—sometimes twofold val-
vate—aestivation. The Atropoideae chemosyn-
drome combines highly derived tropane ester
alkaloids with steroidal alkaloids, while the So-
lanoideae possess one of these biosyntheses but
have the other only in inhibited form as simple
tropanes or as steroidal lactones. These evolu-
tionary patterns are sufficient to warrant recog-
nition of the taxa as subfamilies.

Subfamily ANTHOCERCIDOIDEAE Tété-
nyi, subfam. nov

Plantae frutescentes caulibus lignosis in Australia
habitantes. Aestivatio valvato-aperta. Co Mid

profundae didynama v
extrorsae longitudinaliter dehiscentes. die capsu-
lares aut baccatus. Embryo parum curvus. Plantae
praecipue alcaloidam *Scopolamin" ds eoi -sapo-
ninem continentes.

Woody shrubs inhabiting Australia. Corolla
with rolled, inflexed-valvate aestivation of the 5
lobes, these striated along main veins, varying
in length but never as long as the tube. Stamens
didynamous or equal, epipetalous low in corolla
tube, the anthers dehiscing extrorsely by longi-
tudinal slits. Fruit capsular or baccate. Embryo
only slightly curved. Plants containing predom-
inantly the alkaloid scopolamine and steroid sa-
ponins. Type: Anthocercis Labill.

Subfamily ATROPOIDEAE Tétényi, subfam.
nov.

Plantae ex orbe antiquo oriundae. Herbae locis cal-

regionis ‘montium excelsium vel insularum. Corolla

tus. Plantae praecipue alcaloidam “‘Hyoscyamin”
steroidalcaloidam continentes

Herbaceous plants of the Old World, occurring
exceptionally as endemic woody shrubs on
mountains or islands in warmer regions. Corolla
tubular or campanulate, the aestivation of lobes
imbricate. Stamens 5, equal. Fruit baccate or
capsular. Embryo curved circularly or in a spiral.

608

Plants containing pre onu nanu y the alkaloid
lalkaloids. Type: Atro-

pa b

LITERATURE CITED

ANTOUN, M. D., D. ABRAHAMSON, L. R. TvsoN, C. M

McLAUGHLIN, G. PECK & J.

. Potential antitumor agents md
Physalin B and 25, 26-epidihydrophysalin C from
Witheringia e Lloydia 44: 579—585.

BAEHNI, C. uverture du bouton chez les
fleurs de Solanées. eile 10: 399-492.

BOULTER, D., EACOCK, A. GUISE, J. T. GLEAVES &
G. ESTABROOK. 1979. piede ts between
partial amino acid sequences. Phytochemistry 18:

—608

BuscHi, C. A. & A. B. PoMiLio. 1986. Identificación
del compuesto letal de Nierembergia hippomani-
ca. II. Latinamerican Phytochemical Congress, La
Plata, Argentina. Abstract

CHADEFAUD, M. & EMBERGER. 1960. Traité de
Botanique Systématique, Volume 2. Masson, Paris.

DAHLGREN, R. A system of classification of the

apom Bot. Not. 128: 119-147

1980. A revised system of classification of

L^ e o wies J. Linn. Soc., Bot. 4.

v, W. G. 1979. Classification of Solanaceae.
. G. Hawkes, R. N. Lester & A. D. Skelding

META The Biology and Taxonomy of the So-

Academic Press, London.

W. C. Evans. 1984. Tropane
alkaloids of species of Anthocercis, Cyphanthera

a

and Crenidium. Pl. Med. 50: 86-87.
Evans, W. C 79. Tropane alkaloids of the Sola-
nac . G. Hawkes, R. N. Lester & A. D.

Skelding (editors), The Biology and Taxonomy of
the Solanaceae. Academic Press, London.

. A. RAMSEY. 1981. Tropane alkaloids
from Anthocercis and Anthotroche. Phytochem-

istry 20: 497-

Alkaloids of the Solanaceae
tribe Anthocercideae. Phytochemistry 22: 2219-
2225.

& A. SOMANABANDHU. 1980. Nitrogen-con-
taining non-steroidal secondary metabolites of
Solanum, Cyphomandra, Lycianthes and Mar-
garanthus. Phytochemistry 19: 2351-2356.

FAINI, F., R. TORR

carboxy-8-carbolin cid a
rss An Vestia ia. PI. Med. 38: mi 132.
GESSNER, O. 1977. Gift- und Arzneipflanzen von
Mitteleuropa. Carl Winter Universitäts Vlg., Hei-
delberg.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

GiBBs, R. D. 1974. Chemotaxonomy of Plants.
McGill-Queen's, Montreal & London
HAEGI, L. 1979. Australian genera of the Solanaceae.
J. s, R. N. Lester & A. D. Skelding
(editors), The Biology and Taxonomy of the So-
lanaceae. Academic Press, London.
A conspectus of Solanaceae tribe An-
thocercideae. Telopea 2: 173-
HEDGES, S. H. & . HERBERT. 1981. 6-N-meth-
ylornithine: a n natural constituent of Atropa bel-
ladonna. Phytochemistry 20: 2064-2065.
1973. Chemotaxonomie der Pflanzen,
asel.

ae: A. T.
a synoptic surv n J.
& A. D. Sk elding pens The | Biology and Tax-
onomy of the Solanaceae. Academic Press, Lon-

Se. uth n Solanaceae:
, R. N. Lester

LaviE, E. 1973. Applying chemistry to genetics in
certain Solanaceae. Pp. 181-188 in m n in
Botanical Classification. Nobel Symp. 25. Aca-
demic Press, London & Ne

LEETE, E. 1979. VAL and metabolism of the
tropane alkaloids. Pl. Med. 36: 97- Be

LLovp, H. A., H. M. FALEs, M. E. GOLDMAN, D. M
JERINA, T. PLOWMAN p R. E. ume 1985.
Brunfelsamine: a novel convulsant. Tetrahedron
Lett. 26: 2623-2624.

LonENTI, A. E., A. A. VITALE, C. A. Bus

ZALEZ, C. D. “pass si . M. IRRIBAREN

& A. B. PoMILIO. 1981. Antimicrobial activity
of some Argentine higher plants. Fitoterapia 52:
81-85.

Paris, R. R. & H. Moyse. 1971.
médicale. Masson, Paris.

PHILIPSON, W. B 77. Ovular morphology and the

CHI, M. D.

Précis de matiére

classification of the dicotyledons. Pl. Syst. Evol.
Suppl. 1:
ROMEIKE, A. 1978. Tropane alkaloids — occurrence

and systematic importance in angiosperms. Bot
Not. 131: 85 rm

SAN MARTIN, A., J. R A, O. V. GAMBARO & M
CASTILLO. i980. noe alkaloids fom Schi-
zanthus hookeri. Phytochemistry 19: 20 00

SPORNE, K. 19 re-investigation of character

ati
correlations among dicotyledonous plants. New
Phytol. 85: 419-449,
TÉTÉNYI, P. 19 Infraspecific Chemical Taxa of
Medicinal Plants. Chemical Publishing Co., New

de Homology of biosynthetic bacs the
base i chemotax l n Chem-
Istry in eo a Classification. Nobel Symp. 25.
agg tee iin London & New York.

H & N. Vo-Honc. 1977. Steroid
BE in ` dulcamara populations.
Herba. Hung. 16: 55-60.

FLORA OF THE VENEZUELAN GUAYANA — III!

JULIAN A. STEYERMARK2

ABSTRACT

Continued pote of various families of the flora of the Venezuelan Guayana have resulted in the
hu

xa: Stegolepis albiflora, S.

beri, S. humilis (Rapateaceae); Panopsis cuaensis,
i chae, G. bolivarensis, G
sancarlosiana sip na, Neea amaruayensis,

d nardi, N. pisite = ode acaba N. cedenensis, N. clarkii, N. davidsei, N. guaiquinimae,

cola, N. lie.

neri, N. mapourioides, N.

. marahuacae, N. parimensis, N. robusta,

" sebastianii, N. A redis N. pies (Nyctaginaceae); Brunellia neblinensis (Brunelliaceae):

Matayba ptariana subsp. guaiquinimae (Sapin
sutula, C.
Bonnetia bolivarensis, B. euryant

daceae); Catostemma cla

C. marahuacensis, C. rie tyla, C. sancarlosiana, Scleronema si spe (Bombacaceae);
. guaiquinimae, B. ptariensis, B. trist la

placea fruticosa var. chimantae (Theaceae). Daphnopsis guaiquinimae, D.

monocephala (Rubiaceae), a total of 50 spe
or the Venezuelan Gua piel zur of Proteaceae,

rkii, C. ebracteolata, C. hir-

variety.
uapira, Neea, Catostemma, Symplocos, and newly described taxa of Bonnet

RAPATEACEAE
STEGOLEPIS

Stegolepis albiflora Steyermark, sp. nov. TYPE
Venezuela. Bolívar: Meseta de Jaua; Cerro
Sarisarinama, northeastern part, 4?41'40"N,
64*13'20"W, 1,410 m, 10 Feb. 1974, Stey-
ermark et al. 108873 (holotype, VEN; iso-
type, NY).

erbae perennes usque 1.5 m; vaginis prope apicem
Peroni tenuiterque multinervatis eligulatis; pedun-
culis 7-18 axillaribus 4.5-7 dm longis 0.8-1.5 mm
diam.; capitulis globosis 1.8-2.5 cm diam., spiculis 6—
14 maturis lanceolatis acutis 7210 mm longis 2-3 mm
latis; petalis albidis.

Herbaceous perennials up to 1.5 m tall. Sheaths
eligulate, finely and many-nerved near the apex
or along one side, 20-25 cm long, 4-6 cm wide.
Leaf blades rich green both sides, 58-70 cm long,
3.5-5 cm wide, the nerves and midrib on lower
side somewhat more prominent than on upper
side. Peduncles numerous, 7-18, 45-70 cm long,
0.8-1.5 mm diam., many-sulcate. Heads glo-
bose, tan or tawny, 1.8-2.5 cm diam. Spikelets
spreading in various €— "sasawa acute,
7-10 mm long, 2-3 mm e. Bracteoles 14-15,
graduate, the lower outer ones ien be
ular, 222.5 x 2-2.5 mm, the middle ones obtuse,
the upper ovate-lanceolate, obtuse to subacute,
7 x 4 mm. Sepals lanceolate, subacute, 6—7 mm
long, 2.5-3 mm wide. Petals white, oblanceolate
or lanceolate, acute, recurved, 5.5-7 mm long,

1.5-2 mm wide toward the middle. Anthers lin-

ear, 3.5 x 0.4 mm. Ovary depressed-globose, |
m high. Style 1 mm long. Seeds ire barrel-

ee rounded at both ends, 1.5 x 1 mm

Paratypes. VENEZUELA. BOLÍVAR: Cerro Sarisari-
ñama, summit, W-central part, 4?45'N, 64?26'W, 1,922—
2,100 m, 22-27 Feb. 1967, Steyermark 97839 (VEN);
Meseta de Jaua, Cerro Jaua, summit, SW
forest,
64?34'10"W, 1,750-1,800 m, 22-28 Feb. 1974, Stey-
ermark, Carreño & NE n (VEN);

eseta de Jaua, o Jaua, summit, SW part,
4*47'22"N, 64?33' 35"W, 2 228-2,250 m, 2j Feb. 1974,
Steyermark, Carrefio & Brewer-Carias 109581 (VEN);
Meseta de Jaua, Cerro Jaua, summit, E-central part,
4°35'N, 64?15'W, 14 Feb. 1981, Steyermark, Brewer-
Carias & Liesner 124320 (VEN), 124321 (VEN); trail
to Sima menor, Cerro Sarisarifiama, Ravelo 17 (MY).

This species is remarkable for having white
petals and is the only white-flowered member of
the otherwise yellow-petaled genus Stegolepis.
Maguire (1976) identified all the from
the Meseta Jaua (Cerro Jaua and Cerro Sarisa-
rinama) as S. choripetala, a species originally de-
scribed from Cerro Sipapo and not known else-
where. The new taxon differs from S. choripetala
not only in the white petals, but also in the small-
er mature spikes, the greater number of more
slender, shorter peduncles, and in the finely
nerved sheath in the apical portion.

The collections cited were obtained from var-
ious parts of the huge Meseta de Jaua, indicating
that the species is well distributed over the whole

! Flora ofthe

V 1 y : e 4 under NI a 1¢ T 4 Fal
? Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.

ANN. MissouRi Bor. GARD. 74: 609-658. 1987.

«/(DCDiQ&1&^5nQo«
J

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Stegolepis iid — A. Habit
—G. Stamen. Based on holoty

FIGURE |.
natural position. — F. Pet

summit and is autochthonous to that table
mountain.

Stegolepis huberi Steyermark, sp. nov. TYPE:
ela. Bolívar: Kukenán tepui, 5?16'N,
60?48'W, 2,500 m, 28 Apr. 1984, Otto Hu-

ber 9467 (holotype, VEN; isotype, MO).

erbae perennes humiles 23-35 cm; vaginis 7-10
m latis omnino nervatis ligulatis, au-
m

latis; laminis coriaceis ligulato-lanceolatis acutis <
20 cm longis 1.8-2.5 cm latis, costa media haud m

nifesta subtus tenuiter aden puse nervis ne
soletis; pedunculis 17-25 cm longis v mm latis
infra capitu tulum 4-7 mm dilatatis; capitu lis ompressis
4—6-floris 2—3.5 cm latis; spiculis elliptico- dcos

. — B, C. Base A leaf with auricle of sheath. — D. Spikelet. — E. Flower,

sub fructu 15-18 mm longis; bracteolis oblongo-lan-
d vel lanceolatis acutis vel acuminatis 4-10 mm
longis 1.1-3 mm latis; petalis flabellato-rhomboideis
10 mm ns 7-8 mm latis.

Dwarf perennials 23-35 cm tall. Sheaths mem-
branous, subscarious, 7-10 cm long, 3-3.5 cm
wide, venose throughout, the auricles rounded
at apex, 10-15 mm long, 17-22 mm wide. Leaf
blades coriaceous, galate: -lanceolate, 15-20 cm
m wide, acute, finely nerved on

midrib obsolete on both sides. Peduncles com-
pressed, somewhat alate, 3—6-costate, 17-25 cm
long, 2-3.5 mm wide except 4-7 mm where di-
lated below inflorescence. Heads compressed, 4—

1987]

6-flowered, 2-3.5 cm wide, 1—1.7 cm high. Spike-
lets (in fruit) elliptic-lanceolate, 15-18 mm long,
5-seriate. Bracteoles 18-24, oblong-lanceolate or
lanceolate, acute to acuminate, lower ones 4-5
mm long, 1.1-1.2 mm wide, the others 7-10 mm
long, 2-3 mm wide. Sepals (in fruit) oblanceolate,
acute, 13 mm long, 5 mm wide. Petals flabellate-
rhomboid, 10 mm long, 7-8 mm w

This species is the shortest known in the genus.
All the plants seen (Huber, pers. comm.) in a
large colony were similarly small. The member
of the genus to which it shows closest affinity is
S. ptaritepuiensis Steyerm., from which it differs
in the shorter peduncles and leaves, membra-
nous, striate-nerved sheaths, smaller heads with
shorter spikelets, l lets, midri
not perceptible on either side, and a longer au-
ricle of the sheath.

Stegolepis humilis Steyermark, sp. nov. TYPE:
Venezuela. Bolivar: Camarcaibarai tepui,
SW-facing shoulder, 5°52'N, 62?1'W, 1,800-
1,825 m, 22-24 May 1986, Julian A. Stey-
ermark, R. Liesner & B. Holst 132006 (ho-
lotype, MO; isotype, VEN). Figure 1.

erbae perennes humiles 27-52 cm, vaginis mani-
feste nervatis 12 cm longis ligulatis, auriculis apice
rotundatis 5-7 mm longis; rpm nis linearibus apice

acutis 27-45 cm longis 1.3-2. i

subtus prominenti, nervis subt

subtiliter prominentibus; pedunculi s

lanceo-

lanceolatis acutis 15 mm longis 5-6 mm latis; petalis
late flabellati i bt to-rotundatis 12-13 mm

longis 18 mm latis.

Dwarf perennials 27-52 cm tall, the caudex
flattened, 11-12 cm long, 7-9 cm wide. Leaf
sheaths conduplicate, firmly membranous, con-
spicuously nerved, 12 cm long, 7 cm wide, lig-
ulate, the auricles rounded, these 5-7 mm long,

STEYERMARK— VENEZUELAN GUAYANA FLORA- III

611

12-13 mm wide. Leaf blades silvery green below,
linear, symmetrical to slightly subfalcate at the
acute apex, 27-45 cm long, 1.3-2.5 cm wide,
finely nerved above, prominently nerved below,
midrib prominent below, 1 mm wide. Peduncles
2-3, 5-costate with rounded ribs, 27-52 cm long,
1-1.5 mm diam. except below the inflorescence
where flattened and dilated to 4-5 mm wide.
mainly 1, sometimes 2, compressed.
Spikelets d fusiform, 15-18 mm long,
e. Bracteoles dark mahogany or
chestnut is indurated, 19-21, the lowest
suborbicular-ovate, broadly acute, 4.5-6 mm
long, 4-5 mm wide at base, the upper ones lan-
ceolate, acuminate, 12 mm long, 5 mm wide at
base. Sepals broadly lanceolate, indurated above
the middle, sharply slenderly acute, 15 mm long,
mm wide. Petals with a broadly rhombic-
flabellate blade, broadly subtruncate-rounded at
apex with a mucronate center, 12-13 mm long,
18 mm wide, unguiculate 11 mm. Anthers 10
mm long; filaments 9.5 mm long. Style subulate,
12 mm long.

Paratypes. "VENEZUELA. BOLÍVAR: Murisipán tepui

summit, 5°52

Steyermark & eee re (MO,

tepui, summit °52'N 2

: rd 1986, Liesner. y odds & Ha 21 075 (MO,
N).

"N

This taxon is related to Stegolepis terrama-
rensis Steyerm. from Cerro Marahuaca, from
which it differs in the solitary, or rarely two spikes,
acute leaf apex, fewer and more slender pedun-
cles, smaller size, shorter auricles of the nerved,
nonindurated ligulate sheath, and more con-
spicuous midrib and secondary nerves. It differs
from the other species with one to few spikelets
in size and details of spikelets, bracteoles, leaf
nervation, and peduncle

LITERATURE CITED

MaGuinE, B. 1976. Rapateaceae. In J. A. Steyermark
& C. Brewer-Carias, La Vegetación de la Cima del
Macizo de Jaua. Bol. Soc. Venez. Ci. Nat. 32(132-
133): 279.

PROTEACEAE

PANOPSIS

KEY TO THE SPECIES OF PANOPSIS

la. Leaves sessile or subsessile; reticulation coarsely areolate pod areoles 1-5 mm diam.; leaves crowded

on the stem, pseudoverticillate; fruit subglobose,

4-5 cm

P. sessilifolia

lb. Leaves petiolate; reticulation generally more minutely AR with areoles 0.5-1 mm diam.; leaves

612

lam

ANNALS OF THE MISSOURI BOTANICAL GARDEN

scattered on the stem, not pseudoverticillate; fruit fusiform or longer than broad or less than 4 cm in
di

[Vor. 74

2a. Stems, leaf blades, and petioles glabrous or d so
3a

Leaves abruptly acuminate-cuspidate

wh

pex, 8.5-11.5 cm long; reticulation minute and

ics on both sides; inflorescence m ^ cm long with axes 0.7-1 cm; below 150 m elev.,

Terr. Fed. Amazonas

P. cuaensis

3b. Leaves rounded at apex, 3.5-6 cm long; reticulation elevated and manifest on both sides;

N
c

bana, Edo. Bolív
. Young stems, portions of the leaf blades, and icr oles
l

cm with axes 2-3 cm long; plants at elev. of
P. ptariana

pubesce

4a. oo mainly 8-20 cm long, 3-7 cm wide; inflorescence 15-30 cm long; plants of d

elev

rubescens

4b. pati 2-15 cm long, 2.5-4 cm wide; inflorescence 5-8 cm long; plants of 1,150-2, cri
l

eiev.

5a. Trees 19—22 m tall; leaves 8-15 cm lon
5b. Small shrubs 1-1.5 m tall; leaves 2-9 cm lon

g P. tepuiana

6a. Flowers sessile to 2 mm pedicellate; style — EPA E of lower leaf surface

with larger areoles than on upper surface, subeleva
9 cm long; plants of the Sierra Parima, Terr. Fed. jean

a
c

. Flowers on pedicels 2.5-7 m

n upper surface; leaves ai
rimensis

mm long; style strigillose below the middle and iud

the base; reticulation subelevated on lower surface, scarcely evident or obscure on
upper se ly leaves 2-7 cm long; plants of sandstone table mountains of eastern
F:

Edo. Boliv

ornatinervia

Panopsis cuaensis Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:
Río Cuao, Río Orinoco, 125 m, 17 Jan. 1949.
Bassett Maguire & Louis Politi 28409 (ho-
lotype, NY).

Arbuscula, ramis glabris; foliis petiolatis, petiolis 5—
8 mm longis glabris, laminis discoloribus subtus mar-
ronino-brunneis oblongo-ellipticis apice acuminatis vel
subcuspidatis basi subacutis vel subobtusis 8.5-11.5
cm longis 3-4 cm latis ubique glabris, nervis lateralibus
vix bidon utroque latere 9, venulis tertiariis ubique
ICHU
visis; Vinftuclescentiae rhachidi terminali 15-17 cm longa
adpresso-pubescenti pilis pallidis instructa, axibus tri-
bus vel quattuor 7-10 mm longis; fructu fusiformi ex-
tremitatibus rotundatis 2.2-2.5 cm longo 1 cm lato
dense brunneo-velutino.

Small tree with glabrous branches. Petioles 5-
8 mm long; leaf blades discolored, dull olive green
above, maroon brown beneath, oblong- elliptic,

1 "s

bruptiy ac t apex, sub-
acute to subobtuse at base, 8. a 11.5 cm long, 3—
4c

e side, divaricate at
approximately 10-15°, ees and tertiary vena-
tion immersed, the tertiary venation finely and
minutely reticulate-subimpressed, the midrib el-
evated below, shallowly depressed above. Flow-
ers not seen. Fruiting rachis terminal, 15-17 cm
long, the 3-4 lateral axes 7-10 mm long, the
rachis and axes pale appressed-pubescent. Fruit
fusiform, rounded at both ends, 2.2-2.5 cm long,
1 cm wide, densely brown velutinous.

This species differs from Panopsis rubescens

(Pohl) Pittier in its completely glabrous leaves
which terminate abruptly in a shortly acuminate
or cuspidate apex, fewer and less distinct lateral
foliar nerves, glabrous stems, and smaller, short-
er fruits rounded at each end.

Panopsis parimensis Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:
Departamento Atabapo, helechales y for-

maciones secundarias en la i

25 km NNE de Parima “B,” cabeceras del

Río Ocamo, 3?3'N, 64*13'W, 1,150 m, 12

Jun. 1981, Otto Huber 6136 (holotype, VEN;

isotypes, MO, NY).

Fr .5 m, ramis novellis dense ferrugineo-to-
monies foliis Min ae petiolis 2-8 mm longis fer-
rugineo-tomentellis; foliorum laminis elliptico-obova-
tis vel elliptico-oblongis apice subacute obtusis basi
subacutis vel acutis 4.5—9 cm longis 2-4 cm latis supra

floribus sessilibus vel
sieur usque . 2 mm longis; perianthio 4-4. 5m
o,

tepali co

ong
instructis; stylo 2.3 mm longo glabro.

Shrub, 1.5 m tall; young leafy stems densely
ferruginous tomentose; mature branches dark
gray, glabrous. Petioles 2-8 mm long, ferrugi-
nous tomentose; leaf blades elliptic-obovate or
elliptic-oblong, subacute-obtuse at apex, sub-
acute to acute at base, 4.5-9 cm long, 2-4 cm

1987]

wide, finely strigose above with pale hairs, more
densely strigose along upper midrib, more dense-
ly appressed below, especially along the midrib,

with shorter ferruginous hairs; lateral nerves ca.

9 each side, not prominent but more manifest
than the tertiary venation; tertiary venation of
upper surface minutely and finely reticulate, sub-
elevated, impressed on lower surface. Inflores-
cences terminal, paniculate, densely ferruginous
tomentose, 4-8 cm long (including the peduncle),

5-7 cm wide, with 4 divaricately spreading
branches up to 4 cm long, and 1 mm diam. Pe-

duncle 8 mm long. Bracts subtending the branch-
es of the inflorescence subulate, 4 mm long,
densely ferruginous tomentose. Flowers irregu-
larly crowded on the axes, solitary or 2-3-fas-
ciculate, sessile to 2 mm pedicellate. Perianth 4—
4.5 mm long, the segments densely appressed

STEYERMARK — VENEZUELAN GUAYANA FLORA- III

613

pubescent without. Hypogynous disk slightly an-
gulate. Ovary ferruginous setose. Style 2-3 mm
long, glabrous.

This species differs from Panopsis ornatinervia
Steyerm. of eastern Venezuelan Guayana in the
upper leaf surface having a minute subelevated
reticulation and the lower surface a larger areo-
lation of impressed veinlets. In P. ornatinervia,
the upper surface has a pebbly rugulose, but not
reticulate, pattern, whereas the lower surface has
a subelevated and finer reticulation. Moreover,
in P. ornatinervia the leaves are rounded at the
apex, whereas those of P. parimensis are sub-
acutely obtuse and usually larger. Finally, the
flowers of P. parimensis have shorter pedicels
than those of P. ornatinervia and have a glabrous
style.

ROUPALA

KEY TO SPECIES OF ROUPALA

la. edi ciel less than 2 m tall; leaves 1.3-3 cm long, 0.7-2.5 cm wide; petioles 1-2 mm long; ovary
R.

glabro

minima

lb. Shrub ç or sues 3-20 m tall; leaves 3.5-17 cm long, (2-)2.5-10 cm wide; petioles 10-50 mm long; ovary

pubescent
2a. prem rounded or obtuse at apex

3

3a. vigi and rachis of inflorescence glabrous; leaves glabrous below
ioles 3-5 mm long; leaf blades 3.5-7.5 c
Hs ds 12-15(-30) mm long; leaf blades (5-)7-11(-17) cm long, acute at base ..

par pie

m long, obtuse at base R.
R. obtusa

3b. ee tomentose or furfuraceous-puberulent; rachis minutely ferruginous puberulent or

castane
5a Fern

the base, 2.5-7 cm

ves minutely puberulent or furfuraceous

us-furfuraceous or puberulent; le
th 7-9 mm lon ne ovary hirsutulous; leaf blades elliptic or ovate-oblong, obtuse at

R. sororopana

5b. Perianth 12-13 mm resa ovary shortly appressed-pubescent; lage blades ovate to sub-

orbicular-ovate, truncate or broadly rounded at base, 7.5-10 c

N
leg

. Leaves acute to acuminate at a

wide ............ R. chimantensis
6

to glabrescent toward apex

apex
6a. Stems, petioles, and lower surface of leaf blades glabrous or glabrescent; perianth strigillose
R.

montana

6b. itn ps and lower surface of leaf blades manifestly pubescent; perianth tomentellose

villou

78. Perianth densely pale brown and villous with spreading hairs; rachis of inflorescence

t ntose; fruit minutely tomentose
7b. Perianth tomentose; rachis of inflorescence yellow tomentose; fruit glabrous ...

Roupala paruensis sd sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:
Cerro Parú, be south to southeast to
edge descent to tributary of Cano Asisa, rocky
sabanita, open Río Ventuari, 2,000 m, 10
Feb. 1951, R. S. Cowan & John J. Wurdack
31378 (holotype, NY).

Arbor 7 m, ramis glabris; foliis petiolatis, petiolis 3-
a aminis ovato-oblongis apice ob-
datis -— obtusis 3.5-7.5 cm longis 2-
3.8(-4) cm glabris; nervis eval supra
haud manifestis vel Obsoletis subtus subimpressis vel

. griotii
R. suaveolens

subelevatis utroque latere 4—5; floribus
uy ctes rua > cm lo
2-4 mm y ees fructu o

c TI 2 lato glabro; sem

O aios ORE 1.2-1.4 cm longis 0.8-1 cm
latis

non vISIS; 1N-

Tree 7 m, with glabrous branches. Petioles 3—
5 mm long, glabrous; leaf blades alternate, co-
riaceous, ovate-oblong, obtuse or rounded at
apex, obtuse at base, ETP m long, 2-3.8(-4)
cm wide, glabrous both sides, narrowly subrev-
olute, lateral nerves 4—5 each side, obscure or

614

obsolete above, subimpressed or subelevated be-
low. Inflorescence 18 cm long, rachis glabrous;
pedicels 2-4 mm long, glabrous. Fruit obliquely
obovoid, 1.5-2.5 cm long, 1-1.2 cm wide, gla-
brous; seeds brown, ovate, rounded at each end,
1.2-1.4 cm long, 0.8-1 cm wide.

This species i dily distinguished from Rou-
pala obtusata Kl., its closest relative, by the
smaller, basally obtuse leaf blades with shorter
petioles.

NYCTAGINACEAE

The genera Guapira and Neea are represented
in the Venezuelan Guayana by a large number
of taxa. Schmidt (1872) treated the known species
of these genera at that time as they occurred in
Brazil, Guapira then included within the genus
Pisonia. Later, Heimerl (1896) published addi-
tions for an account of the West Indian taxa and
(1897, 1914, 1932) described various new species
from tropical America. Huber (1909) also de-
scribed several species of these genera from Am-
azonian Brazil. Standley (1931) took up the fam-
ily for northwestern South America, recognizing
the genus Torrubia of Vellozo with a dozen species
previously assigned to Pisonia. Lundell (1968)
transferred the taxa formerly assigned to Tor-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

rubia, as well as some others which had been
placed under Pisonia, to the genus Guapira Au-
blet (1775), an earlier legitimate name. Little
(1968) transferred d additional species from
Torrubia to Gua

Unfortunately, s has been no recent study
published for the taxa occurring in the Venezue-
lan Guayana. During the many years that have
elapsed since the publication of works by Schmidt,
Heimerl, and Standley, many unnamed or mis-
identified collections have accumulated in her-
baria. S large number of the taxa represented by
these separated, while oth-
ers appear to be closely related and differentiated
on characters relating to indument, peduncles,
leaf shape, size, venation, branching of inflores-
cence, and cauliflory. In some cases it is difficult
to be certain of the generic distinction where only
pistillate flowers are present. In general, how-
ever, Guapira and Neea may be separated using
staminate material. Yet Burger (1983) suggested
that the two genera may have to be united under
Guapira. Further studies will be necessary to judge
the merits of generic separation.

The present study of the taxa of the Venezue-
lan Guayana has resulted in the following 26 new
taxa, with keys provided for the species of Gua-
pira and Neea.

GUAPIRA

KEY TO THE SPECIES OF GUAPIRA OF THE VENEZUELAN GUAYANA

la. Leaves 1-4 cm long, 0.8-1.8 cm wide, rounded or manifestly obtuse at apex

G. oe ylla
do

lb. Leaves larger than 4 cm long and 1.8 cm wide, mainly acute to acuminate at apex, or, if rounde

obtuse, the leaves larger

2a. Leaves 25-30 cm long, 15 cm wide

2
G. sipapoana
3

2b. Leaves 5-15 cm long, 2-15 cm wide
3

a. Lower and/or upper surface of leaves, or lower midrib, pilosulous with lax, spreading, or

I divaricate
4a. Up

densely

per je n pide and shining; inflorescence subglobose or subhemispheric,
red and c ted G.

davidsei

+
c

cymosely or widely spreading

. Upper leaf s Miu Or a d upper midrib, pubescent, and not shining; inflorescence

Sa. Petiole 0.5 cm or less long; peduncle 1.5-2.5 cm long
5b. Petiole 1-2 cm long; peduncle (1—)3-9.5 cm lon

G. pubescens
6

6a. Peduncle 7-9.5 cm long; petiole densely pubescent with spreading hairs 0.2—0.5
mm long; young stems densely pubescent with spreading hairs 0.2-0.5 mm lon
G.m

marcano-bertii

6b. eae (1-)3.5-5 cm long; petiole and young stems with hairs less than 0.

usbyana

w
S

G.r
. Lower —Ó of en including midrib, glabrous or pubescent, but the indument not
ate

spreading or divaric

7a. Lower surface of leaves glabrous or essentially so, the midrib or nerves with scattered
ntu

microscopic tom

uncle "E petiole densely ferruginous tomentose
8b. Peduncle and petiole glabrous or sparsely

cle and/or axes of in reais ae to moderately puberulen

Oa. Leaves often broadest above the middle, conspicuously venose; Tu

9a. Pedun
l

G. amacurensis

pubescent

1987]

STEYERMARK — VENEZUELAN GUAYANA FLORA —III

615

nerves conspicuous, 9-11 each side, subelevated or impressed on both

sides, conspicuously anastomosing with t

he tertiary veinlets, ascending at

an angle of 45? or more; tertiary veinlets forming a prominent network

e G. fragrans

10b. Leaves often broadest near the middle, not venose, opaque; lateral nerves
Rr dar 5—6 each side, impressed, divarica seed spreading at an angle
G.

15-30°; tertiary veinlets obsolete or E cu

guianensis
11

9b. gian: and/or axes of inflorescence glabrous .... us

lla. Tertiary veinlets Me and finely pier ee on both sides of leaf

blades; eastern Edo. Boliva - G. bolivarensis
llb. Tertiary veinlets as i p d

Te mazonas . 12

12a. ` Leaves ss to es hee -lanceolate; staminate — cylin-
dric-tubular, 1.8-2 mm wide; stamens 8 glabriflora

12b. Leaves ovate or elliptic- jee staminate perianth ere rd
mm wide; stamens 10 G. neblinensis

7b. Lower surface of leaves, midrib, or nerves with a minute tomentum of nonspreading

S
13a. Tertiary venation on upper and lower leaf surface very conspicuous or elevated ...
G.

13b. Tertiary venation either not evident, inconspicuous, or not elevated
14a. Pri ncipal lateral nerves 8-10 e
f us to Du ions on biis branches generally cuspidate at

G.c

apex; dry frui gam

—
+
c

the sparsely rufous m

sa iesu NT
PNE EE 14

ach side; lower leaf surface usually with a

ong, 3-4 m uspidata

. Principal lateral nerves pi : each side; eae leaf surface glabrous except for
idrib a

nd sometimes sparsely tomentose or glabrescent

secondary nerves; leaves on fertile branches rounded, subtruncate, or abruptly

shortly acute at apex; dry fruit 8-9 mm long, 5-5.5 mm wi ide

Guapira amacurensis Steyermark, sp. nov. TYPE:
Venezuela. pare bosque pluvial, E of Rio
El Palmar, near limits of

1965, Luis Marcano-Berti 551 (holotype,
VEN; isotypes, MO, NY

5-25 m, ramulis dense-ferrugineo-tomen-
tosis; foliis ovatis, elliptico- ovatis vel lanceolato- ellip-

costa media subtus sparsim
ubique glabris; inflorescentiis dense ferrugi
tellis, pedunculis ferrugineo-tomentosis pilis 0.2-0.5

S
mentoso; staminibus 7, nd foemineo su
extus dense ferrugineo-tomentoso

Tree 15-25 m tall, the younger branches
densely ferruginous tomentose. Petioles 1-3.5 cm
long, densely ferruginous tomentose; leaf blades
turning black or dark brown, coriaceous, elliptic-
ovate, lance-elliptic, or ovate, acute, obtusely
acute to acuminate at the apex, acute to cuneate
at the generally inequilateral base, 6.5-14 cm
long, 3-6.3 cm wide, glabrous both sides except
the midrib on lower side sparsely rufous tomen-
tellous or glabrous; lateral nerves 6-11 each side,
obsolescent above, slightly more evident below.
Inflorescence of staminate plant 2-3 cm high, 4—
5 cm wide, ferruginous tomentose; peduncle 1.7—

DENS G. ayacuchae
4 cm long, 1-1.5 mm wide, not enlarged at junc-
tion with the lowest inflorescence axes, the fer-
ruginous hairs somewhat lax. Staminate flowers
sessile to 1 mm pedicellate; bracts 0.5-1 mm
long, densely ferruginous tomentose. Staminate
perianth infundibuliform, 7 mm long, 3 mm wide
bove; stamens 7, the filaments exserted 3-5 mm
beyond the orifice. Pistillate perianth tubular, up
to 7 mm long, densely ferruginous tomentellous
without.
Paratype. VENEZUELA. BOLÍVAR: Represa Guri, 55
km NE of Ciudad Piar, 7?35'N, 63?7'W, 200-300 m,
4-5 Apr. 1981, Liesner & Gonzalez 11174.

This species differs from Guapira bolivarensis,
described below, in the larger staminate perianth,
generally inequilateral leaf base, and the dark
brown to blackish leaves upon drying.

Guapira ayacuchae Steyermark, sp. nov. TYPE:

Steyermark & Otto Huber 113856 (holo-
type, VEN; isotype, MO)

Arbuscula 3-4 m, foliis late ovatis ramulorum
tilium. apice late rotundatis subtruncat tis vel ed SE

5- 15 cm | longis 4—9 cm latis, supra costa dies lan
lateralibusque ferrugineo- -pubescentibus aliter glabris
vel glabrescentibus subtus costa nervis lateralibusque

616

s ferrugineo-pubescentibus inter nervos magis
ilabresc en ntibus; nervis iiri adn. utroque latere 6-7;
rcte rrugineo-tomentellis, pe-

m longo

ext rrugineo-tomentello, perianthio
PUES tubuloso 2.5-3 mm longo extus ferrugineo-
tomentello; staminibus 5; fructibus in sicco ellipso-
ideo-oblongis 8-9 x 5-5.5 mm in vivo ovoideis 10 x
9 mm; pedicellis fructiferis 4-5 mm longis

Small tree 3-4 m tall. Petioles 1-2 cm long;
leaf blades membranous, broadly ovate, on the
fertile branches broadly rounded or subtruncate
to shortly and abruptly acute at apex, rounded,
truncate, or cuneate at base, 5-15 cm long, 4—9
cm wide, the upper surface glabrescent or re-
motely and minutely puberulent, the midrib and
lateral nerves rather densely rufous-ferruginous
tomentose, the lower surface more glabrescent,
here the midrib less densely rufous tomentose
and the secondary nerves only sparsely tomen-
tose to glabrescent; lateral nerves 6-7 each side,

Staminate inflorescence umbel-
lately 4-branched, the 4 primary axes 8 mm long;
peduncle 2 cm long, 0.9-1 mm wide, this and
axes of the inflorescence minutely densely ru-
fous-ferruginous tomentellose. Flowers on ped-
icels 1-1.8 mm long. Staminate perianth infun-
dibuliform, 3.5-4 mm long, 2-3 mm wide at
summit, moderately rufous-ferruginous without.
Stamens 5, filaments 6-7 mm long, exserted 2-
3 mm. Pistillate inflorescence umbellately
4-branched, the primary axes 10-12 mm long, 1
mm wide; peduncle 5 cm long, this and the axes
of the inflorescence closely rufous-ferruginous
tomentellose. Flowers on pedicels 1-2 mm long,
4-5 mm long in fruit. scene perianth tubular,
2.5-3 mm long, 1-1.5 m , ferruginous to-
mentellous without. Fruiting | primary axes 1-2.5
cm long, 1-1.5 mm wide, the secondary axes 8-
17 mm long. Anthocarp black, broadly ellipsoid-
oblong in dried state, ovoid in living state, sub-
sulcate, glabrous, 8-9 mm long, 5-5.5 mm broad
in dried state, when fresh 10 mm long, 9 mm
wide.

Paratypes. VENEZUELA. TERRITORIO FEDERAL
AMAZONAS: alrededores del aeropuerto Puerto Ayacu-
cho, 120 m, Morillo 3142 (VEN); Tobogán de la Selva,
5°22'N, 67?33'W, 150 m, 14 May 1980, Steyermark,
Davidse & Guanchez 122545 (MO, VEN); 6 km N of

; Estación de Piscicultura, erto Ayacucho
5°37'N, 67?36' , Huber 618, 662 (VEN); Ori-
popos, 7 km N of Puerto Ayacucho, Miller 1618 (MO,

VEN). COLOMBIA. DEPARTAMENTO VICHADA: Casuarito,

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

across from Puerto Ayacucho, 5?40'N, 67°40’W, 100
m, Gentry & Stein 46330 (MO, VEN).

This species has been confused with Guapira
cuspidata (Heim.) Lundell of northeastern Ven-
ezuela. It differs from that species in the larger,
glabrous fruit with longer pedicels, 5 instead of
7 stamens in the staminate flowers, generally
rounded to subtruncate or abruptly shortly acute
apex of the leaves on the fertile branches, fewer
and more distantly separated pairs of lateral
nerves on the leaf blades, glabrous or glabrescent
lower surface of the leaves between the latera
nerves, more sparsely puberulent upper leaf sur-
face with more conspicuously rufous puberulent
midrib and lateral nerves, and fewer ultimate
axes of the pistillate infructescences.

Guapira bolivarensis um. sp. nov. TYPE:

enezuela. Bolívar: Departamento Piar,

summit of Amaruay-tepui, south side, east-

ern half, 5°55’N, 62°13'W, 950-1,100 m, 11

May 1986, Ronald Liesner & Bruce Holst
20800 (holotype, MO; isotype, VEN).

6 m, ramulis sparsim puberulis pilis dnd
neo-glanduliferis munitis; foliis peat ova
ice acutis vel acuminatis basi a -] gis ia
8 cm latis utrinque glabris, nervis s Tateralibus utroque
i el ulum ad-

m bel glabro; perianthio mascu-

o 1.8-3.3 cm longo; peri-

Mert ‘foemineae peduncu
anthi 5 mm longo superne

1.1 mm lato extus glabro.

Tree 6 m tall, the young branches sparsely pu-
berulent with appressed-ascending, ferruginous-
glandular trichomes. Leaves alternate or oppo-
site. Petiole 0.7-2 cm long, glabrous to sparsely
puberulent; leaf blades elliptic to ovate, acute to
acuminate at apex, acute at base, minutely dark-
dotted beneath, 6-17 cm long, 3-8 cm wide, gla-
brous both sides; principal lateral nerves 8-10
each side, irregularly spaced, subhorizontal or
ascending at an angle of 10—25*; tertiary venation
finely reticulate both sides, the veinlets promi-
nulous. Staminate inflorescence somewhat
broader than long, 4-flowered, 2-2.5 cm high,
2.5-4 cm wide, subumbellate to broadly panic-
ulate with 3-8 axes, the lower axes larger, gla-
brate or sparsely puberulent near the apices; pe-
duncle 4-7.8 cm long, 1-1.3 mm broad, glabrous.
Staminate perianth narrowly infundibuliform, 4
mm long, 1.5 mm wide at summit, glabrous

1987]

without except for a few minute hairs at base.
Stamens 10, the filaments unequally exserted up
to 4 mm above orifice. Pistillate inflorescence
terminal, 3-5 x 3.5 cm, sparsely 3-5-flowered,
the 2-3 axes sparsely puberulent; peduncle in
anthesis 1.8-3.3 cm long, sparsely puberulent.
Pistillate perianth tubular-cylindric, 3-3.5 mm
long, 1.1 mm near orifice, glabrous without.
Fruiting axes 2.5-5 mm long. Young fruit nar-
rowly fusiform, 11-12 mm long, 3 mm wide at
middle.

Paratype. VENEZUELA. BOLÍVAR: Amaruay-tepui,
steep slopes on W side, 5*55'N, 62?15'W, 550-800 m,
20 May 1986, Liesner & Holst 20932 (MO, VEN).

This taxon resembles Guapira glabra (Hei-
merl) Steyerm., comb. nov. (Pisonia glabra Hei-
merl, Kew Bull. 1932: 220. 1932) in the glabrity
of the leaves and floral parts, prominently retic-
ulate tertiary venation on both leaf surfaces, and
subhorizontal to shallowly ascending lateral
nerves but differs in having 10 instead of 8 (rarely
9) stamens, short and narrower staminate peri-
anth, and shorter staminate inflorescence with
shorter axes.

Guapira davidsei Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Delta Ama-
mento Tucupita, 5-14 km ESE

1979, Gerrit Davidse & Angel González
16382 (holotype, VEN; isotype, MO)

Arbor 8 m, foliis oppositis. vel ternatis lanceolato-
vel oblongo 12cm lon-

C1 OUIOIIBU 1

presso-puberulis costa media pilis patentibus 0.1 mm

culo 1-2.5 cm longo dense ferrugineo-adpresso-pubes-
centi; floribus 10-35 sessilibus; perianthio masculo
p a 4—4.5 mm longo supra medium 2.5-

m lato extus adpresso-pubescenti pilis glanduli-
feris r mixtis munito.

Tree 8 m tall, the younger branches appressed
puberulent, the older ones sparsely so. Leaves
opposite or 3 at a node. Petioles 3-10 mm long,
densely appressed pale puberulent; leaf blades
lance-elliptic or oblong-elliptic, acute at apex,
acute at base, 6-12 cm long, 2-5 cm wide, gla-
brous and shining above, minutely subappressed
puberulent below with pale hairs 0.1 mm long,
the midrib below elevated, with minute, spread-
ing, slightly rigid hairs; lateral nerves 7-9 each

STEYERMARK — VENEZUELAN GUAYANA FLORA —III

617

side, inconspicuous, impressed. Staminate inflo-
rescence terminal, densely congested, globose or
subhemispheric, 8-14 mm high, - mm
broad, the primary axes suppressed. Peduncles
1-3, 1-2.5 cm long, with densely ferruginous,
subappressed hairs. Flowers 10—35, sessile. Bracts
lanceolate-oblong, subobtuse or rounded, 1 mm
long, densely t without. Sta-
minate perianth infundibuliform, 4—4.5 mm long,
2.5-2.7 mm above middle, densely ferruginous
appressed-pubescent without, mixed with glan-
dular Aun Stamens 7-8, exserted 2-2.2 mm
beyond ori

Guapira davidsei is well marked by the con-
gested globose inflorescence with dense ferrugi-
nous pubescence. From G. ferruginea (Klotzsch
ex Choisy) Lundell it is distinguished by having
much larger, acutely pointed leaves and numer-
ous glandular hairs intermixed with the pubes-
cence on the outer surface of the staminate peri-
anth.

Guapira glabriflora Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:
between San Carlos and El Solano, Depar-
tamento Río Negro, 11-17 Mar. 1979, Luis
Marcano-Berti & P. Salcedo 54-979 (holo-
type, MER).

rbuscula, ramulis glabris; petiolis 6-10 mm longis;

5 mm ‘longo (in sicco 2.2 mm ) apicem versus 1.8-2

berulo pilis glanduliferis atque apice dense papillato-
puberulo aliter glabro; staminibus 8

Small tree. Petioles 6-10 mm long, glabrous;
leaves opposite, the blades drying fuscous, ellip-
tic-lanceolate to oblanceolate, obtusely acute at
apex, cuneately acute at base, 6.5-12.5 cm long,

.5-4.7 cm wide, glabrous both sides, the lateral
nerves 6-9 each side, inconspicuous, impressed.
Staminate inflorescence 10-15-flowered, the pri-
mary axes 2-3, 3-15 mm long, bearing the flow-
ers directly or with short secondary axes up to 5
mm long bearing a few sessile flowers, the axes
minutely and sparsely ferruginous puberulent.
Peduncle 1.5-4.5 cm long, glabrous or glabres-
cent or with sparse ferruginous glandular or non-

618

glandular hairs. Bracts minute, ovate-deltoid,
subacute, 0.3-0.4 mm long, puberulent. Perianth
tubular, slightly and inconspicuously enlarged
above the middle, 4.5—5 mm long (2.2 mm dried),
1.8-2 mm wide at summit, 1 mm wide (0.6 mm
dried) in lower 3, nearly glabrous without except
for a few sparse, ferruginous papillate hairs in
basal 1 mm and densely papillate-puberulent
apically bordering and between lobes. Stamens
, the filaments exserted 4 mm beyond orifice.

This species is distinguished by its glabrous
leaves and nearly glabrous peduncle and stami-
nate perianth. It differs from Guapira neblinensis
Mag. & Steyerm. in the narrower oblanceolate
to elliptic-lanceolate leaves and narrower sta-
minate perianth.

Guapira marcano-bertii Steyermark, sp. nov.
TYPE: Venezuela. Territorio Federal Delta
Amacuro: east of Rio Grande, ENE of El
Palmar, near limits of Estado Bolívar, 26
May 1964, Luis Marcano-Berti 209 (holo-
type, VEN; isotype, MO).

LE Eg

Arb
Us amaka 0.2-0.5 mm es munitis; pe-
tiolis 1-2 cm

-8 incon
escencia oo ae A vejer
axibus principalibus 2-4, 12- gis 1.5

ia pe edunculo fructifero a 5cm s. 2-2.5
mm s pis e tomentoso pilis patentibus 0.2—0.4 mm
longis ne anthocarpio anguste ellipsoideo 9-10
mm longo 3.5—4 mm lato sparsim puberulo 10-costato.

Tree with young stems densely villosulous with
spreading ferruginous hairs 0.2-0.5 mm long.
e 1-2 cm long, densely ferruginous villo-
sulous with spreading hairs 0.2-0.5 mm long; leaf
blades membranous, broadly elliptic-ovate,
shortly and abruptly acute to short acuminate at
apex, obtuse to acute at base, 7-12.5 cm long,
3-7 cm wide, the midrib above with spreading
villous hairs 0.5-0.7 mm long, the upper surface
elsewhere with shorter and longer pale hairs 0.5-
0.7 mm long, the lower surface densely soft-vil-
losulous with crisp hairs 0.5 mm or more long,
the midrib with divaricate hairs up to 1 mm long.
Infructescence subumbellate to irregularly panic-
ulately branched, the 2-4 p l
mm long, 1.5 mm diam., dud ferruginous
villosulous with spreading hairs to 0.2 mm lon
Fruiting peduncle 7-9.5 cm long, 2-2.5 mm

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

diam., densely villosulous with spreading hairs
0.2-0.4 mm long. Fruiting bracts lanceolate, sub-
acute or obtuse, densely tomentose without.
Fruiting pedicels 1-3 mm long. Anthocarps nar-
rowly ellipsoid, 9-10 mm long, 3.5—4 mm wide,
sparsely puberulent, longitudinally 10-costate.

This taxon differs from Guapira pubescens
(HBK) Lundell in the much longer petioles and
the longer and more abundant pubescence on all
parts. From G. cuspidata (Heim.) Lundell it dif-
fers in the longer peduncles, longer fruits, and
the longer spreading pubescence throughout.

Guapira sancarlosiana Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas: 4.3
km NNE from San Carlos on Solano Road,
IVIC study site, 1°56'N, 67°3'W, 119 m, 8
Jan. 1981, H. L. Clark & Pedro Maquirino
7794 (holotype, MO). Figure 2.

Arbor 7-12 m, ramulis juvenilibus dense —
seque rufo- ferrugineo- tomentosis; foliorum lamini

subtus arcte rufo-tomentosis pilis dense adpressi is om-
nino obtectis ovato-ellipticis vel sublanceolato-ellip-
ticis apice breviter abrupteque acutis vel breviter acu-
minatis interdum rotundatis basi cuneatim acutis 7.5-
14.5 cm longis 4.5-7.5 cm latis; nervis lateralibus ut-
roque latere 9-11 subtus elevatis, venulis tertiariis su-
pra subtiliter conspicueque elevatis reticulatis subtus

mosissima, axibus primariis 0.8—3 cm re pedun-
culo terminali 0.2-3 cm longo 1.5-2 mm m. dense

4.8 mm longo sup m 3.8-4 mm lato extus

dense rufo-tomentoso; bs 5-6.

Tree 7-12 m tall, the young stems densely ap-
pressed rufous-ferruginous tomentose. Petiole
1.5-3.5 cm long, densely and minutely appressed
rufous tomentose; leaf blades coriaceous, glau-
cous above, ovate- to sublanceolate-elliptic,
shortly abruptly acute to shortly acuminate at
apex, sometimes rounded, cuneately acute and
often asymmetric at base, 7.5—14.5 cm long, 4.5-
7.5 cm wide, glabrous or moderately strigillose

ulate and elevated. Staminate inflorescence many
and closely flowered, much branched, 1.5-3.5 cm
high, 3-7 cm wide, the primary axes 0.8-3 cm

STEYERMARK

1987]

FIGURE 2. Guapira sancarlosiana.

long, paniculately branched with 5—6 short axes,

earing small clusters of scattered flowers along
the length of the axes. Flowers sessile. Perianth
shortly infundibuliform to subcampanulate, 4—
4.8 mm long, 3.8-4 mm above middle, densely
rufous tomentose without. Stamens 5—6, 3 ofthe
filaments exserted 2.2 mm beyond the perianth.

VENEZUELA. TERRITORIO FEDERA

AMAZONAS: same data as type, 6909 (NY), 7031 (NY),
Clark 7251 (MO); Clark & Maquirino 7776 (MO); be-
tween San Carlos and Solano, Marcano-Berti & Sal-

Es

—VENEZUELAN GUAYANA FLORA —III

s —A. Flowering hei staminate plant. —
rescence.—C. Staminate flower opened. Based on holotyp

619

B. Portion of staminate inflo-

mre 126-979 (MER); supra ostium fluminis Casi-
uiare, 1854, Spruce 3751 (NY).

This species is well characterized by the co-
riaceous leaves which are glaucous above and
bear densely appressed rufous puberulence be-
neath, by the upper and lower leaf surfaces with
elevated reticulate tertiary venation, and by the
densely branched staminate inflorescences bear-
ing numerous flowers with five stamens, of which
three are exserted. Spruce 3751 was labeled as

620

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FIGURE 3. Guapira sipapoana. — A. Fruiting branch.—B. Fruit.—C. Apical end of fruit. Based on holotype.

an unpublished new species doubtfully referred
to Neea (as “Neea? rubiginosa”).

Guapira sipapoana Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:
eserva Forestal Sipapo, left margin of Rio
Sipapo, Bloque 1, May 1971, Carlos Blanco
1158 (holotype, VEN). Figure 3.

Arbuscula, ramulis juvenilibus dense rufo-tomen-
tosis; foliis elliptico- ovatis apice acuminatis basi ro-
tundatis 25-29 cm longis
14.5-15 cm latis supra costa media puberula aliter gla-
bris subtus puberulis pilis erectis 0.1-0.2 mm longis
praeditis, venulis tertiariis reticulatis subtus promi-
ene reticulatis; infructescencia axillari, axibus pri-
mariis 1.5-2 cm longis; pedunculo 3.8 cm longo 3 mm
lato denos rufo-tomentoso; anthocarpio lineari-ellip-

1987] STEYERMARK— VENEZUELAN GUAYANA FLORA-~III 621

soideo 1.5-2 cm longo 4-5 mm lato rufo-tomentoso nently so with large areoles on lower surface.
conspicue 10-costato. Infructescence in the upper axil, the primary axes
1.5-2 cm long, the secondary axes subumbellate,
2.5-3 cm long, branched above into shorter axes
1.5-2 cm long. Anthocarp linear-ellipsoid, 1.5-
2 cm long, 4-5 mm wide, rufous tomentose, 10-
costate.

Small tree with young branches densely rufous
tomentose. Petiole 3.8 cm long, 3 mm wide,
densely rufous tomentose; leaf blades very large,
elliptic-ovate, shortly acuminate at apex, round-
ed or obtuse at the unequal base, 25-29 cm long,
14.5-15 cm wide, upper surface glabrous except This taxon is distinctive in having very large,
for the puberulous midrib; lower surface, in- prominently reticulate leaves with loose pubes-
cluding midrib, lateral nerves, and tertiary vein- cence on the lower surface and in having long,
lets, puberulous with erect, slender hairs 0.1-0.2 pubescent fruits.

mm long; tertiary veinlets reticulate, promi-

NEEA

KEY TO THE SPECIES OF NEEA

la. Leaves sessile or nearly so, usually obtuse at base, the petiole to 1 mm long; young stem and petiole
hirtellous with subspreading rufous-brown hairs; inflorescence both axillary and terminal on the stem

N. ignicola
. Leaves petiolate, generally acute to acuminate at base, the petiole 2-35 mm long; young stem and
petiole glabrous or pubescent with appressed hairs; inflorescence either cauliflorous on the old stem

=

2a. Inflorescence cauliflorous on the old stem .... 3
3a. Leaves oblanceolate-elliptic or je elliptic, 8-12.5(-15) cm long, 2.5-4.5(-5.5)cm ~ UN

3b. Leaves mainly obovate, oblong, or oblong-ovate, 10-30 cm long, 5-16 cm wide .......
4a. Principal secondary leaf nerves 6-8 each side, slightly elevated on lower surface
N. e 5

4b. Principal secondary leaf nerves 8-12 each side, conspicuously elevated on lower surfac

Sa. Lower leaf surface and nerves completely glabrous; fruiting peduncle 1-3.5 cm long;
flowers sessile; inflorescence dichotomous or with short axes on an elongated rachis

wid avidsei
5b. Lower leaf surface glabrous, but midrib and secondary nerves with minute spreading
s; flowering peduncle 0.5-0.6 cm long; flowers with pedicels 2-2.5 mm long;
inflorescence much branched divaricately N. liesneri
2b. Inflorescence Wigs ducas the stem or its branches
Leaves broadly rounded at the apex, obovate or elliptic-obovate N. obovata
6b. Leaves mainly acute to acuminate at the apex, of other shapes than 7

abov
7a. Main secondary nerves of leaf blades 15-25 each side, subhorizontal: or ascending at an
angle less than 20°, relatively close together, 3-4 mm ovalifolia
7b. Main secondary nerves of leaf blades generally amis ee 12 each side (fainter inter-
mediate nerves may be present), ascending at an angle usually greater than el or if m
4

es 12 nerves or at a smaller angle, then the nerves more than 4 mm apart „uu
8a. Peduncle 6-11 cm lon 9
9a. Stem, peduncle, and axes of inflorescence densely ferruginous tomentose, peri-
anth 3 x 1.5 mm, densely ferruginous tomentose N. bernardii
9b. Stem glabrous, peduncle and axes of i nflorescence minutely year Te or PE
rescent; perianth 8-9 x 3 mm, glabro grandis
8b. Peduncle 1-5 cm 10
10a. apa and axes of inflorescence glabrous throughout or essentially so ......... 11
—— perianth slightly or moderately ferruginous pubescent toward
N. clarkii
11b. Staminate perianth glabrous i p ly put

l
12a. Staminate perianth 6-6.5 x 3.5-4 mm; inflorescence paniculately
and irregularly Rory oe and a with elongated ra-
chis and axes, 6-15 cm long, 5-10 cm wide; peduncle 3-5 mm in
diameter; principal se d = nerves 19-12 each side; tertiary vein-
lets conspicuously ie and subelevated on upper leaf ay tose
robusta

622

ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

12b. Staminate perianth 2 x 0.8 mm; inflorescence umbellately or tri-

mously branched, 1-2 cm long (or high), 2.5-4 cm wide;
0

diameter; principal ges nerves 5-

obsolete er ace
r leaf surface lustrous; lower leaf decies with subele-
i rl

vated, subreticulate tertiary venation more or less manifest;
leaves ira or rarely rounded at apex; Jima (pistillate)

1-1

n diameter; leaves ovate to subovate ... N. tepuiensis

13b. Upper kar eres dull; lower leaf surface with obsolescent
ertiary venation; leaves obtusely acute to acute at apex; pe-
duncle ai 0.8-1 mm in diameter; leaves prua
late

N. subglabrata

10b. Peduncle and axes of inflorescence i to densely ferruginous or puber-

T% sometimes with glandular trichome

. Junction of lowest axes of inflorescence with summit of peduncle not

ug enlarged, 0.8-2 mm w
15a. Peduncle usually with pai ired ferruginous bra
ri

cts
ry axes of inflorescence 4-7, umbellate; pistillate peri-

anth ferruginous tom E main lateral nerves of leaf blades
sharply ascending at 50—60? N. bracteosa
16b. Primary axes of inflorescence 3, not umbellate; staminate
perianth glabrous except for papillate exterior of lobes; main
lateral nerves of leaf blades shallowly ascending at 15-25°.

15b. ea ebracteat

N. amaruayensis
17

anth a or glabrescent; bracts mainly pM ex-

ou “for the puberulent margins marahuacae

N
°

. Perianth variously pubescent; bracts ferruginous dorsally and
on margins

18a. Ultimate axes of inflorescence racemose with alter-

18b.

nately arranged flowers; upper and lower Sei of the
leaf blade with minut e, ferruginous indumen
9a. Lower leaf

th Z . IM

indume eol alc,
hioen e much- branched; periant nth and inflo-
rescence axes densely ferruginous tomentose with
some glandular ie petal to 1.2 cm long ....

N. parimensis
19b. Lower leaf surface glabrous or glabrescent; peri-

sparsely branched; perianth and inflorescence axes
rather sparsely ferruginous tomentose without
ong ........... N. tristis
Ultimate axes of inflorescence cymose or corymbiform,
or with fasciculately arranged flowers; upper midrib of
leaf blades glabrous, the lower midrib glabrous or the
basal part sparsely puberulent 20
20a. Petiole and upper part of stem with pale, minute,
spreading papillalike trichomes; perianth with

edu

florescence densely ferruginous ba sat

fruit 3.5-5 mm sh N. clarkii
21b. Tertiary venation grossly reticulate, subel-

evated andm smile on both sides of leaves;

axes and peduncle of inflorescence aie rsely

to moderately ge scent; antho-

carp 6-7 mm wi .N.gi guaiquinimae

14b. edes of lowest axes of inflorescences with summit of peduncle le
wide

, 2.5-4 mm

1987]

STEYERMARK— VENEZUELAN GUAYANA FLORA-III

623

22a. Perianth 3-4.5 mm long, 1.5 mm wide; petiole and young stem
N.

sparsely puberulent
Perianth 6-8 mm mg 2-3.5 mm wide; petiole and young stem

22b.

og

. sebastianii

glabrous or glabresc 23
23a. Tertiary ene manifest, elevated on lower surface, im-
pressed on upper surface; flowers in small clusters at or near

the end of the ultimate axes N. neblinensis
23b. Tertiary venation obsolete or scarcely manifest; e if
2 mainly scattered along the length of the ultimate

m 24
oe Pistillate perianth glabrous without; leaves drying
blackish; principal secondary nerves 9-12 on each side

N. D NT

24b. Staminate perianth with minute, sparse to modera
erruginous indument; leaves drying deri “sasa

principal secondary nerves 6—8 on each s

"N. mapourioides

Neea amaruayensis Steyermark, sp. nov. TYPE:
Venezuela. Bolívar: Amaruay-tepui, west
side, steep slopes, 5?55'N, 62°15'W 0-
740 m, 2 May 1986, Ronald Liesner & Bruce
Holst 20514 (holotype, MO; isotype, VEN).

Frutex vel arbor 2.5-5 m; foliis prs dini piee vel
lanceolato- sey apice breviter obtuse s 6.5-
15 cm longis (2.5-)3.5-7.5 cm latis ne ruis
nervis Enea principalibus utroque 8-10 ad an-
gulum 15-25* adscendentibus, venulis tertiariis reti-
lati infl
iin masculina 12- 21- flora 0. 9-1. 4 sha longa 1.3-2 cm
8 mm longis 0.5 mm
eis sparsim minute ferrugineo- ubera inaequaliter
insertis; pedu nculo l- 3 cm Jonga sparsim m vel modice
ferrugineo-p munito, bibrac-
teato, bracteis oppositis oblongo- ovatis vel oblongis-
1 mm dense ferrugineo-puberulis 4 mm
spositis; perianthio masculino cylindri-
i i 3-3.5 mm i medio 1.5 mm
lato extus glabra; staminibus 8 inclusis

Shrub or tree 2.5-5 m tall. Leaves mainly op-
posite; petiole 7-20 mm long, glabrous; leaf blades
elliptic-ovate, oblong- or lanceolate-elliptic,
abruptly shortly and ens subobliquely and ob-
tusely acute, 6.5-15 cm long, (2.5-)3.5-7.5 cm
wide, with dark dots beneath, glabrous both sides;
principal lateral nerves 8-10 each side with ad-
ditional shorter intermediate ones, anastomos-
ing 2-5 mm from margin; tertiary venation re-
ticulate, more conspicuous on upper than lower
side. Staminate inflorescence relatively small,
0.9-1.4 cm long, 1.3-2 cm wide, 12-21-flowered,
the axes unequally inserted, the 3 main ones 5-
8 mm long, 0.5 mm wide, sparsely minutely fer-
ruginous puberulent, each axis terminating in 3
shorter axes 0.7-2 mm long, these each sup-
porting 2-3 sessile flowers. Peduncle 1—3 cm long,
sparsely to moderately ferruginous puberulent,
bibracteate with a pair of oblong-ovate or ob-

bs obtuse, densely ferruginous bracts 1-2 mm
ong and 1 mm wide situated 4 mm above base.
Staminate perianth subinfundibuliform-cylin-
dric, 3-3.5 mm long, 1.5 mm wide at middle, 1—
1.2 mm wide at summit, glabrous without. Sta-
mens 8, included; filaments 0.5-1.5 mm long;
anthers suborbicular, 0.6-7 mm long, 0.5-0.6

=

Paratype. VENEZUELA. BOLÍVAR: Amaruay-tepul,
southwest slope, 750-900 m, 24 Apr. 1986, Holst &
Liesner 2702 (MO, VEN).

This taxon possesses paired bracts on the pe-
duncle, as in Neea bracteata Steyerm., but differs
in the perianth of the staminate flower being
completely glabrous except for the papillate mar-
gins and exterior surface of the shallow perianth
lobes. It differs further in the longer petioles, the
unequally inserted lower nonumbellate axes of
the staminate inflorescences, and the shallowly
ascending lateral nerves of the leaves at an angle
of 15-25? instead of the sharply ascending ones
of N. bracteata arising at 50—60?

Neea bernardii Steyermark, sp. nov. TYPE: Ven-
ezuela. Bolívar: de Santa Elena de Uairen al
sur-este, 800-900 m, 23 Apr. 1957, L. Ber-
nardi 6746 (holotype, MER; isotypes, MO,
NY

Arbuscula vel arbor, m dense ferrugineo-tomen
tosis; foliis in sicco brunneis oppositis late ovatis d
liptico-ovatis vel oblanceolato- ellipticis apice ke y
acutis vel breviter
11.5 cm longis (2.7- )3-6. 5 cm latis subtus costa ui ER ia

tere 6-10
ee i
cula subcorymbos ae 5c ta
chotome ramosa den zou tua rruginea; axibus pri-
mariis duobus, (0. 421-3 cm longis 1.5 mm diam. dense

624

adpresso-ferrugineo-t tosi d l -)8-10
cm longo 1.5-2 mm diam. dense. adpresso- EU.
tomentoso apice haud manifeste ee perianthio
masculino late cylindrico 3-4 mm o 1.5 mm lato
extus dense ferrugineo tomentoso; UE 8.

Small or large tree to 40 m tall, with densely
ferruginous
posite; petiole (0.8—)1.5-3 cm long, appressed-
ferruginous tomentose; leaf blades drying dull
brown to rufous-brown, broadly ovate, elliptic-
ovate, or oblanceolate-elliptic, obtusely acute to
shortly acuminate at apex, cuneately acute at base,
5.5-11.5 cm long, (2.7-)3-6.5 cm wide, glabrous
on both sides except strigose beneath on midrib;
lateral nerves 6-10 each side, impressed and in-
conspicuous above, faintly manifest below; ter-
tiary venation scarcely manifest above, incon-
spicuously finely reticulate beneath. Staminate
inflorescence terminal, subcorymbose, 1.5-5 cm
high, 2.5-10 cm broad, dichotomously me
with 2 primary axes (0.4—)1—3 cm long, 1.5 m
diam., these branched into 3 Seana sec-
ondary axes (3-)12-16 mm long, each with 2
branches bearing 3-7 clusters of sessile or sub-
sessile flowers, the axes with densely appressed
ferruginous tomentum. Peduncle (2.5-)8-10 cm
long, 1.5-2 mm diam., densely appressed fer-
ruginous tomentose, its summit 2-3 mm broad,
not enlarged at junction with lowest areas. Bracts
ovate, acute, | mm long, densely ferruginous to-

entose. Perianth broadly cylindric or tubular-
subinfundibuliform, 3—4 mm long, 1.5 mm wide,
densely ferruginous tomentose. Stamens 8, oc-
cupying 7^ of the length of the tube. Fruit oblong,
14 mm long, 7 mm wide, glabrous.

iei i VENEZUELA. BOLÍVAR: El dare in
margin of Río Grande, 300 m, 20 Feb. 1959, Ber-
sees 7156 (VEN); 40 km SE of Tumeremo, nea ath
Botanamo, 100 m, 7 May 1960, Little 17597 (US,
VEN)

Neea bernardii is distinguished by the greatly
elongated peduncle.

Neea bracteosa Steyermark, sp. nov. TYPE: Ven-
ezuela. Bolivar: Represa Guri, 55 km NE of
Ciudad Piar, 7?35'N, 63°7'W, 250-300 m,
4-5 Apr. 1981, R. Liesner & A. González
11277 (holotype, VEN; isotype, MO).

r8m, ferrugineo-pubescen-

tibus; petiolis 3-10 m m longis; foliorum laminis op-
positis. vel ad nodos superiores ternatis coriaceis ellip-

basi cuneatim acutis 6- 9 cm longis 2-3.5 cm latis,
nervis lateralibus utroque latere 8-10 inconspicuis; in-
florescentia foeminea umbellatim ramosa 1.5-2 cm alta

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

2-3.5 cm lata, axibus Aia ee 4-7 tenuissimis E
11 mm longis 0.5 mm latis. Pedunculo tenui 1.5-
m longo modice ferrugineo- hirtello pilis laxis E

g floribus (immaturis)
congestis; perianthio (immaturo) modice vel sparsim
ferrugineo-adpresso tomentoso.

Tree 8 m with young branchlets ferruginous
pubescent. Leaves opposite or sometimes ternate
at the upper nodes; petiole 3-10 mm long, gla-
brous or moderately to sparsely ferruginous pu-
bescent at the base; leaf blades coriaceous, dull
brown upon drying, elliptic or lance-elliptic, acute
to obtusely acute at apex, cuneately acute at the
mainly equilateral base, 6-9 cm long, 2-3.5 cm
wide; lateral nerves 8-10, fine and slightly evi-
dent below, scarcely evident above, arising at an
angle of 50-60°. Pistillate inflorescence 1.5-2 cm
high, 2-3.5 cm broad, with 4-7 umbellately
ies very ne axes 6-11 mm long, 0.5
mm wide, moderately fe t with
lax bi hairs 0.1-0.2 mm long; sec-
ondary axes 2-5, also umbellate, 3-5 mm long,
bearing several clustered flowers at the apices.
Peduncle slender, 1.5-3.2 cm long, | mm diam.,
moderately ferruginous with lax subspreading
hairs 0.1—0.2 mm long, bibracteate, not enlarged
at junction of summit of peduncle with lowest

ally present between '4-'^ the length of the pe-
duncle. Flowers (immature) congested, 85-100.
Perianth sparsely to moderately ferruginous pu-
bescent

This taxon may be differentiated by the bi-
bracteate, very slender peduncles with slender
umbellate axes.
Neea b ] l St , Sp. nov. TYPE:
Venez uela. Territorio Federal Amazonas:

etween Paso El Diablo and Cano de Cu-
lebra,25-30 km SE of Puerto Ayacucho, 100
m, 12 May 1980, Julian A. Steyermark, Ger-
rit Davidse & Francisco Guanchez 122366
(holotype, pistillate plant, VEN; isotype,
MO). Figure 4.

AT 6-20 m, ramis glabris, foliis coriaceis opacis
obovatis late oblanceolatis vel elliptico-oblongis apice
acutis vel

incrassatos 1-5 inflorescentiis pedunculatis praedita;
pedunculis brevibus 1-1.5 cm longis minute puberulis,

1987]

omnibus ramosis cum 3-4 axibus divaricatis subfas-
ciculatis vel subumbellatis 9-20 mm longis minute
adpressoque puberulis; abend foemineo infundi-
buliformi 3. 5 mm longo superne - mm lato extus

t .7 mm

longo 1. 5 mm lato extus glabro; staminibus 7 inclusis.

Tree 6-20 m tall, with glabrous branchlets.
Petioles 1-2.5 cm long, glabrous; leaf blades co-
riaceous, opaque, obovate, broadly oblanceolate,
or elliptic-oblong, abruptly and obtusely acute to
rarely rounded at apex, cuneately acute to obtuse
at base, 10-22 cm long, 5-12.5 cm wide; lateral
nerves 6-8 each side, slightly elevated below,
obsolescent above; tertiary veinlets obscure
above, slightly evident below, forming large ar-
eoles. Pistillate inflorescence cauliflorous, 1.5-2
cm high, 2-4 cm wide, with 1-5 short, many-
flowered, pedunculate inflorescences arising from
the old wood at the usually thickened nodes. Pe-
duncles 1-1.5 cm long, minutely puberulent, each
branched into 3-4 widely divaricate, subfascic-
ulate or subumbellate primary axes 9-20 mm
long, these branched above into short, alternate
secondary axes 2-7 mm long bearing the flowers,
minutely appressed pale tomentellose. Pistillate
perianth infundibuliform, 3.5 mm long, 2
wide, externally minutely puberulent; pistil 3 mm
long (ovary 1 mm long; style 1 mm long with
prominently penicillate stigmas 1 mm long); ster-
ile filaments 0.5-1. long. Staminate flowers
in sessile alternately arranged clusters along the
short axes of the inflorescence. Staminate peri-
anth tubular, somewhat narrowed at base, 3.7
mm long, 1.5 mm wide, glabrous. Stamens 7,
included within perianth; filaments 0.5-2 mm
long.

Paratype. | VENEZUELA.

at edge of bana (lower Amaz inga),

67*3'W, 119 m, H. Clark 6605 panna plant, MO).
This taxon has been confused with Neea flo-

ribunda Poeppig & Endl. but differs in the shorter

peduncles with more closely crowded, smaller

flowers of a different form and in the glabrous

staminate perianth.

Neea cedenensis Steyermark, sp. nov. TYPE: Ven-
ezuela. Bolivar: Departamento Cedeno:
bosque de galeria del Caño Chaviripa del
drenaje del Escudo Guayanes, carretera Cai-
cara—El Burro, 16 Apr. 1984, B. Stergios, D.
Tephorn, L. Nico & C. Gilbert 8611 (holo-
type, MO; isotype, PORT)

Frutex, ramulis modice puberulis trichomatibus

STEYERMARK — VENEZUELAN GUAYANA FLORA-III

TERRITORIO FEDERAL

625

brevibus patentibus instructis; laminarum foliis lan-
ceolatis vel lanceolato-ellipticis apice anguste subob-
tusis basi acutis 8.5-11.5 cm longis 2.2-3.5 cm latis
utrinque glabris costa media inferiore prope basem tri

umbellata ca. 27-flora inaequaliter ramosa, axibus
quinque dense ferrugineo-tomentellis; pedunculo 12
m ferrugineo-puberulo; perianthio mas-

m longo dens
culino WE Reed 5 Cs 3-4 mm longo
superne 1-1.2 mm lato indumento ferrugineo pilis ver-
miformibus munito; staminibus 8.

Shrub having stems moderately yis
with pale, short, spreading trichom .05-0.1

mm long. Petiole with minute, IE short
papillalike trichomes; leaf blades mainly oppo-
site, lanceolate or lance-elliptic, narrowed to a
subobtuse apex, acute at base, 8.5-11.5 cm long,
2.2-3.5 cm wide, 3-3'^ times longer than broad,
glabrous on both sides except for scattered pale-
erruginous trichomes at base of lower midrib,
the upper surface punctate; lateral nerves about
10 and inconspicuous both sides; tertiary vena-
tion inconspicuously reticulate. Staminate inflo-
rescence terminal, 0.8 cm long, 2 cm wide, ca.
27-flowered, subumbellate with 5 main axes un-
equally branched 1-2.5 mm above the base,
densely ferruginous puberulent, each axis un-
equally divided into 4 branches 1.5-2 mm long,
these 1-2-flowered. Flowers sessile. Bracts sub-
tending flowers lanceolate, acute, 0.8-1 mm long,
unequal, densely ferruginous puberulent. Sta-
minate perianth cylindric-subinfundibuliform,
3-4 mm long, 1-1.2 mm wide at summit, abun-
dantly streaked with appressed, ferruginous, ver-
miform indument without. Stamens 8; filaments
1.5-2 mm long; anthers suborbicular, 0.6 mm
long, 0.8 mm wide.

This taxon is characterized by short, spreading
trichomes on stems and petioles, vermiform fer-
ruginous indument on the staminate perianth,
small inflorescences, and inconspicuous tertiary
venation.

Neea clarkii Steyermark, sp. nov. TYPE: Vene-

m, Howard L. Clark 6980 (holotype, NY).

tex 3 m, ramulis glabris; petiolis 8-15 mm longis
glabris; foliorum laminis coriaceis lanceolato- pace dex
vel oblanceolato- ellipticis apice acutis vel acuminatis
—15) cm longis 25-45

illari trichotome ramosa, axibus dense r ar hasa to-

626

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

RE 4. Neea brevipedunculata. — A. Flowering branch, “seo: plant.

view. .—C. Pistillate perianth,
mens.

mentosis; pedunculo ferrugineo- tomentoso ? apice haud
dilatato; p

longo supra medium 1.5 mm lato extus minute spar-
simque adpresso-puberulo pilis pallidis munito; sta-
minibus 8, anthocarpio ellipsoideo 8.5-10 mm longo,
3.5-5 mm lato glabro

Shrub 3 m tall with grayish-white, glabrous
branches. Petioles 8-15 mm long, glabrous; leaf
blades coriaceous, fuscous, lanceolate-elliptic or

apical view showing thicken
—E. Portion of staminate inflorescence. Based on ho

— B. Pistillate perianth, exterior

ed perianth lobes. — D. Pistil with rudiments of sta-
lot UD.

oblanceolate-elliptic, acute to shortly acuminate
at apex, cuneately acute at the equilateral base,
8-12.5(-15) cm long, 2.5-5.5 cm wide; glabrous
both sides, densely dark punctate below; lateral
nerves 8-11 each side, finely impressed or ob-
solescent above, faint or finely elevated below;
tertiary venation inconspicuous. Staminate in-
florescence terminal or axillary (immature?), tri-
chotomously branched, 10 mm high, 1

STEYERMARK

1987]

— VENEZUELAN GUAYANA FLORA-III

627

wide, shortly pedunculate; 3 primary axes 2 mm
long (immature), ferruginous tomentose, termi-
nating in groups of 3 or more sessile flowers.
Bracts ovate-deltoid, subacute, 0.3-0.4 mm long,
densely ferruginous tomentose. Peduncle 4-15
(-30) mm long, 1-1.5 mm wide, ferruginous to-
mentose, not enlarged at junction of peduncle
with lowest axes of inflorescence. Staminate peri-
anth subinfundibuliform, 4 mm long, 1.5 mm
wide above the middle, minutely sparsely pu-
bescent with pale hairs without. Stamens 8, in-
cluded, unequal; filaments 1.3-2.2 mm long. Fruit
ellipsoid, 8.5210 mm long, 3.5-5 mm wide, gla-
brous.

Paratypes. "VENEZUELA. TERRITORIO FEDERAL
AMAZONAS: San Carlos, 21 Mar.-17 Apr. 1981, Delas-
cio, Christensen & Broome 9496 (VEN); 4 km NE of
San Carlos, Liesner 6422 (MO, VEN); prope San Car-
los and Río Negro, 1853, Spruce 3766 (NY); San Carlos
de Río Negro, 16 Feb. 1981, Otto Huber 6067 (VEN).

This species is related to Neea guaiquinimae,
from which it differs in the absence of glandu-
larity on the peduncles and axes of the inflores-
cence, the lack of tertiary reticulation on the
leaves, and in a denser indument on the peduncle
and inflorescence axes.

Huber's specimen matches the photographs of
two unpublished names by Heimerl of a Spruce
3766 collection from **prope San Carlos, ad Rio
Negro." The photograph from the Vienna her-
barium has a note by Heimerl pertaining to a
pistillate plant, *Insbesonders durch die Kráftig
berandeten Blatter u. wahrscheinlich auch durch
Cauliflori ausgezeichnet." This photo shows a
cauliflorous, dense, short inflorescence only 10
mm high and 8 mm broad. The other photo from
the Berlin herbarium does not show any inflo-
rescence. The specimen of Huber 6067 also shows
a small inflorescence 10 mm high and 8-12 mm
broad. Of the three small inflorescences present,
one appears at the apex ofa leafless short branch,
whereas the other two originate 5 mm below the
apex of a leafless stem.

The lower leaf surface of the Huber 6067 col-
lection is densely punctate, but the other collec-
tions assigned to Neea clarkii do not show punc-
tation.

Neea davidsei Steyermark, sp. nov. TYPE: Ven-
ezuela. Territorio Federal Amacuro: Depar-
tamento Tucupita, mountain area, ca. 13 km
by road ESE of town of Sierra Imata, 8?23'N,
62°23'W, 400 m, 4-6 Apr. 1979, Gerrit Da-

vidse & A. González 16558 (holotype, VEN;
isotype, VEN)
bor 8 m, ramulis glabris; petiolis 1-2 cm longis

Ar
glabris; foliorum laminis oppositis late obovatis vel
vel acutis

lo ong
1AW DUIUI ull

pedunculata,
pu es pilis adpressis hivis munitis, primariis 3 cm
longis superne ue 5mm longis pss lo
vetere fructifero 1-3.5 cm es 3m .; bracteis
ovatis subacutis 0.5 mm longis extus sparsim adpresso
puberulis; fl 1.2 cm longo
0. 8-1 cm lato haud costato.

Tree 8 m, the branches glabrous. Leaves op-
posite; petiole 1-2 cm long, glabrous; leaf blades
broadly obovate or broadly oblong-elliptic, nar-
rowed to a subacutely obtuse or acute apex, cu-
neately acute at base, 11-24(-30) cm long, 6.5-

1(-16) cm wide, wholly glabrous; lateral nerves
8-10 each side, ascending at an angle of 35—45*,
indistinctly anastomosing 10-15 mm from mar-
gins, elevated and somewhat prominent below,
obsolescent or faintly impressed above; tertiary
veinlets inconspicuous, grossly reticulate below,
obsolescent above. Pistillate inflorescence axil-
lary, cauliflorous on the old wood, pedunculate
or epedunculate. Flowers sessile. Old fruiting pe-
duncle 1-3.5 cm long, 3 mm diam. Primary axis
on old inflorescence 3 cm long, branched in up-
per !^ with alternate short axes 5 mm long, all
axes minutely and sparsely puberulent with pale
appressed hairs. Anthocarp 1.2 cm long, 0.8-1
cm wide, not ribbed.

Paratypes. VENEZUELA. BOLIVAR: between km 11
and 18.5, south of El Dorado, 215 m, 23 July 1960,
Stevermark 86585 (NY, VEN). TERRITORIO FEDERAL
TA AMACURO: Orocoima, Rio Toro, Bernardi 7573,
pe 3734 (NY).

The axillary, cauliflorous infructescence re-
lates this species to Neea liesneri of Cerro de La
Neblina, but the former differs in the longer,
stouter peduncle, sessile flowers, and completely
glabrous lower leaf surface.

The fruiting paratype has the leaves broadly
oblong-elliptic and acute, whereas in the holo-
type the leaf blades are broadly obovate nar-
rowed to a subacutely obtuse apex.

Neea guaiquinimae Steyermark, sp. nov. TYPE:
Venezuela. Bolívar: Cerro Guaiquinima,
summit, NE sector, 6?00'N, 63?28'W, 1,650
m, 9 Apr. 1979, Julian A. Steyermark, G.

628

C. K. & E. Dunsterville 117971 (holotype,
VEN; isotype, MO).

Arbuscula vel frutex 2-3 m, ramis glabris; petiolis
5-20 mm longis. minute eine vel glabris; fo-
liorum a basi
subacutis vel acutis 5-12. 5c cm longis 2-6 cm latis supra
glabris subtus glanduloso-punctatis, nervis lateralibus
utroque latere venulis tertiariis ubique manifeste gros-
seque reticulatis; inflorescentia foeminea trichotome
ramosa | cm alta 1.8 cm lata, axibus principalibus
quattuor 4-5 mm longis 0.8 mm diam., modice glan-
jene ae sub anthesi 2.5 cm longo | mm
dia b fructu 5-7 cm longo 1.5-2 mm lato, sparsim
glandulifero: “s 9 ea tubuloso 4 mm longo 1.2 mm
lato extus dense glandulifero. Anthocarpio oblongo-
ellipsoideo ina ovato-oblongo 12-13 mm longo 6-7
mm lato obtuse 10-costato glabro.

vv UU lU SK ALULIS

Small tree or shrub 2—3 m tall, the branchlets
glabrous. Petioles 5-20 mm long, glanduliferous
or the glands deciduous; leaf blades coriaceous,
elliptic-ovate, obtusely acute at apex, subacute
to cuneately acute at base, 5-12.5 cm long, 2-6
cm wide, glabrous above, densely dark gland-
dotted below, slightly revolute on margins; lat-
eral nerves 6-10 each side, slightly impressed or
obsolescent above, slightly elevated below; ter-
tiary venation non both sides subelevated, grossly
reticulate. P
branched, 1 cm high, 1.8 cm broad, with 4 pri-
mary axes 4-5 mm long, 0.8 mm diam., mod-
erately glanduliferous, the primary axes shortly
branched into 4 shorter, moderately glandulifer-
ous secondary axes 1-2 mm long terminating in
shortly pedicellate extensions, each bearing 2-3
sessile flowers. Peduncle in anthesis 2.5 cm long,
| mm diam., sparsely glanduliferous in fruit, 5—
7 cm long, 1.5-2 mm wide and glabrous, pendent
in fruit, the junction of peduncle with lowest axes
not conspicuously enlarged. Bracts narrowly lan-
ceolate, 1 mm long, densely glanduliferous. Pis-
tillate perianth tubular, 4 mm long, 1.2 mm wide,
densely glandular without; ovary ellipsoid, 1.3
mm long, sessile; style 1 mm long; stigmas 0.4
mm long, fimbrillate. Anthocarp oblong-ellip-
soid to ovate-oblong, 12-13 mm long, 6-7 mm
wide, obtusely 10-costate.

Paratype. | VENEZUELA. BOLIVAR: Cerro Guaiquini-
ma, W sector near forested border, 5?45'N, 63°43' W,
1,540 m, Steyermark, Berry, G. C. K. & E. Dunsterville
117499 (MO, VEN).

This species is distinguished by the sparsely to
moderately glandular pubescence of the peduncle
and inflorescence axes and by the manifestly sub-
elevated, grossly reticulate tertiary venation on
both leaf surfaces.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Neea huachamacarae Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:
Río Cunucunuma, base of trail to Cerro
Huachamacari, 3?39'N, 65?41'W, 220 m, 7
Mar. 1985, Ronald Liesner 18403 (holotype
VEN; isotype, MO).
r 10 m, ramis glabris; petiolis 1-2.5 cm longis
Mes "foliorum laminis oppositis oblanceolatis vel
oblongo-ellipticis apice acutis obtusis vel rotundatis
basi cuneatim acutis 6-15.5 cm longis 3-5.5 cm latis
glabris; nervis lateralibus utroque latere 9-12 incon-
spicuis; inflorescentia foeminea terminali, axibus dense
ferrugineo-adpresso-pubescentibus, pedunculo 1-2.5
cm longo 2 mm lato autem i

durato-tubuloso 7.5-8 mm longo 3-3.5 mm lato extus
glabro

Tree 10 m tall with glabrous branches. Leaves
opposite; petiole 1-2.5 cm long, glabrous; leaf
blades opaque, drying blackish, oblanceolate or
oblong-elliptic, shortly acute, obtuse, or rounded
at apex, cuneately acute at base, 6-15.5 cm long,
3-5.5 cm wide; lateral nerves 9-12 each side, not
elevated, i
spicuous. Pistillate e terminal, the
usually 4 primary axes 5-13 mm long, branched
into 10-15 secondary axes, the solitary flowers
or groups of flowers borne along the length of the
axes, not congested in groups at the ends of the
axes, the axes densely ferruginous appressed-pu-
bescent. Peduncle 1-2.5 cm long, 2 mm wide
except at the junction of the lowest axes of the
inflorescence with the top of the peduncle where
enlarged to 3-4 mm wide. Perianth subpandu-
rate-tubular, 7.5-8 mm long, 3-3.5 mm wide,
glabrous without. Ovary 2-3 mm long; style 5-
6 mm long, exserted 2 mm; sterile stamens 6
with filaments 3 mm long

CI tiar y venation incon-

This species is characterized by having the
flowers inserted along the length of the axes of
the inflorescence rather than in congested groups
at the ends of pedicellate branches. In this respect
it resembles Neea mapourioides but differs in the
glabrous pistillate perianth and the greater num-
ber of principal secondary nerves of the leaf
blades.

Neea ignicola Steyermark, sp. nov. TYPE: Ven-
ezuela. Territorio Federal Amazonas: 11 km
N of Puerto Ayacucho toward El Burro,
5?44'N, 67°30'W, 90 m, 26 Jan. 1978, Otto
Huber & Julio Cerda 1456 (holotype, VEN).

Frutex 2-3 m, ramulis dense hirtellis pilis patentibus

1987]

rufo- brunneis munitis; ed vix dici usque | mm
eis munito; foliorum la-

lateralibus utroque later
RÀ eii reticulatis; inforeseentia mascula terminali
axillarique 5-13 mm alta 7-10 mm la
axibus dun ae alternatim

axibus lanceolatis 1-1.5 mm longis; pedunculo

m longo 0.4-0.5 mm lato hirtello pilis rufo-
brunneis patentibus munito; perianthio masculo (im-
maturo) tubuloso extus sparsim hirtello pilis ferrugi-
neis munito; staminibus 6-7

Shrub 2-3 m tall; branches slender, 1-1.5 mm
diam., densely reddish-brown hirtellous with
spreading hairs 0.5 mm long. Petiole scarcely
developed, up to 1 mm long, hirtellous with lax,
subspreading hairs; leaf blades subsessile, mem-
branous, ovate-lanceolate, usually narrowed to
an acuminate apex, obtuse or rarely rounded at
base, 3.5-6 cm long, 1.3-2.5 cm wide, glabrous
both sides; principal lateral nerves 10 or some-
times more on both sides, elevated above, less
conspicuous beneath; tertiary venation reticulate
both sides, less so beneath. Staminate inflores-
cence axillary and terminal, 5-13 mm high, 7-
10 mm broad, densely hirtellous with rufous-
brown spreading hairs; the 4 primary axes alter-
nately and racemosely branched with flowers
solitary along the length of the axes, but with
terminal flowers in 2s or 3s, the secondary axes
scarcely or not developed. Peduncle filiform, 8—
14 mm long, 0.4-0.5 mm diam., hirtellous with
rufous-brown spreading hairs, not enlarged at
apex where joining the lowest axes of the inflo-
rescence. Bracts subtending base of primary axes
lanceolate, subacute, 1-1.5 mm long. Staminate
perianth (immature) tubular, sparsely hirtellous
with ferruginous hairs. Stamens 6-7

This taxon is readily differentiated by its sessile
or subsessile leaves and spreading pubescence.

Neea liesneri Steyermark, sp. nov. TYPE: Vene-
zuela. Territorio Federal Amazonas: De-
partamento Río Negro: Cerro de La Neblina,
0.4 km W of base camp on Río Mawari-
numa, 0?50'N, 66°10’W, 140 m, 9 Mar. 1984,
Liesner 16502 (holotype, VEN; isotype,
MO). Figure 5.

5 m; petiolis 1-2 cm longis; foliorum laminis
late obovatis vel lato oblongo-ovatis 2d subacutis
obtusis vel rotundatis basi acutis majo asymme-
tricis 16-30 cm longis 9-16 cm latis Men vila sub-

STEYERMARK — VENEZUELAN GUAYANA FLORA —III

629
tus costa media nervis lateralibusque elevatis promi-
tantibus 12-27 m

pressis munitis; peduncu dense pu-
berulo pilis adpressis munito; floribus pedicellatis, pe-
dicellis 2-2.5 mm longis dense puberulis pilis adpressis
obtectis; perianthio urceolato 6 mm longo basi 4 mm
lato apicem versus 1.5-2 mm lato extus dense puberulo
pilis adpressis obtecto; staminibus 7-9; anthocarpio
oblongo-ellipsoideo 20 mm longo 8 mm lato minute
pubescenti pilis subferrugineis adpressis obtecto.

Tree 5 m tall. Petioles 1—2 cm long; leaf blades
broadly obovate or broadly oblong-ovate, sub-
acute to obtuse or rounded at apex, acute at base,
the larger leaves conspicuously asymmetrical, the
smaller leaves nearly symmetrical, 16—30 cm long,
9-16 cm wide, glabrous above, the midrib and
lateral nerves beneath prominent and elevated
with minute spreading pubescence, elsewhere
glabrous; lateral nerves 9-12 each side, ascending
at an angle of 35-45?, anastomosing 5-13 mm
from the margin, regularly 12-27 mm equidis-
tant or irregularly spaced, conspicuously elevat-
ed below, inconspicuously impressed above; in-
termediate and tertiary venation below
inconspicuously grossly reticulate, inconspic-
uous above. Staminate inflorescences cauliflo-
rous on the old wood, broader than high, 2.5 cm
long, 5 cm broad, much branched, many-flow-
ered, the lateral primary axes divaricate, 1-2 cm
long, all parts densely puberulent with pale-ful-
vous, appressed hairs. Peduncle 5-6 mm long.
Staminate flowers on pedicels 2-2.5 mm long
and covered with a pale-fulvous, appressed pu-
berulence. Staminate perianth urceolate, 6 mm
long, 4 mm wide at base, 1.5-2 mm wide near
summit, densely puberulent with pale-fulvous,
appressed hairs. Stamens 7-9, the anthers 1.3-2
mm Ros apiculate or exapiculate, the shorter
filaments 1-2 mm long, the longer ones 3-4 mm
grad Pstillod 4—5 mm long. Anthocarp oblong-

elli , 20 mm long, 8 mm wide, minutely
Benin with appressed, subferruginous hairs.

Paratype. BRAZIL. AMAZONAS: Between Maloca and
Rio md south of Cerro de La Neblina, 50-100
m, Nilo T. Silva & Umbelino Brazáo 60782 (MO, NY).

This remarkably distinct species is character-
ized by the cauliflorous, many-branched, cymose
inflorescence developed on the old wood; large,
obovate, conspicuously nerved leaves; and large,
pedicellate, urceolate staminate flowers with
strongly developed asymmetrical bases.

630

FIG Neea liesneri.—A. Portion d ees branch of staminate plant.
Pistillodium with stamens. Based on holoty

Neea mapourioides Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:
.5-1.5 km east of San Carlos, 1?55'N,
67°5'W, 120 m, 1 Dec. 1977, Ronald Liesner
4096 (holotype, VEN; isotype, MO)

ex 3 m, ramulis glabris; petiolis 5-15 mm longis
glabris; foliorum laminis oppositis elliptico- Medio
3.5—

7.5 cm longis 4—7 cm latis ubique glabris; nervis la-
cate principalibus utroque latere 6-8; inflorescen-
tia mascula terminali 2.5-4.5 cm alta 3.5-7 cm lata,

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

— B. Staminate flower. — C.

axibus late divaricati il 4, dense fer-
rugineo o-tomentosis, axium ultimorum fa alter-
-2

o 1.5-2.5 cm lon

angustato; staminibus 7 ea TR

Shrub 3 m tall; branches glabrous. Leaves op-
posite; petiole 5-15 mm long, glabrous; leaf blades

1987]

coriaceous, opaque, fuscous, elliptic-obovate,
abruptly acuminate at apex, cuneately acuminate
at base, 5.5-17.5 cm long, 4-7 cm wide; lateral
nerves 6-8 each side, anastomosing 4-6 mm from
margin, impressed and slightly manifest below,
but essentially obsolescent; tertiary venation ob-
solete above, scarcely impressed beneath. Sta-
minate inflorescence terminal, 2.5-4.5 cm high,
3.5-7 cm wide, the axes widely spica dense-
ly ferruginous tomentose, the 3-4 primary axes
1.5-3.5 cm long with the diea part bearing
flowers, the ultimate branches bearing sessile
flower clusters scattered alternately ita the
length; peduncle 1.5-2.5 cm long, 1.5-2 mm wide,
but at the summit with the lowest axes s of the
inflorescence enlarged to 2.5 mm wide, sparsely
to moderately minutely brown puberulent. Bracts
of base of primary axes 1-1.5 mm long, ferru-
ginous tomentose, those subtending the flowers
ovate-deltoid, subacute, 0.5-0.8 mm long, fer-
ruginous tomentose. Staminate perianth narrow-
ly ellipsoid, 6.5-7 mm long, 2.8-3 mm wide,
narrower at summit, mie to moderately mi-
nutely ferruginous puberulent. Stamens 7, in-
cluded within the lower '4—'/ of the perianth tube,
unequal; anthers 1.5 x 0.5-1 mm; filaments 1—
3 mm long. Stylopodium 4 mm long.

This species possesses alternately arranged
flowers on the axes of the inflorescence as in Neea
huachamacarae but differs in the fewer lateral
nerves, the fuscous-brown instead of blackish
leaves when dried, and in having pubescence on
the staminate perianth, as opposed to the gla-
brous pistillate perianth in N. huachamacarae.

Neea marahuacae Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:
Cerro Marahuaca, forested slopes 1-2 km N
of Sima Camp, 3?43'N, 65?31'W, 1,100 m,
8—9 Mar. 1985, Ronald Liesner 18428 (ho-
lotype, VEN; isotype, MO).

x2m, ramulis glabris; petiolis t= 20 mm longis;

ferrugineo aarin erigi tenui 1.5-2 cm longo
.5-0.7 m sim puberulo pilis adpressis
munito; oot ok ae tubuloso 1.5 mm longo
mm lato extus glabro vel glabrescenti; staminibus
5-6 inaequalibus inclusis.

Shrub 2 m. Young stems glabrous. Petioles 7—
20 mm long; leaf blades opaque, elliptic-oblan-

STEYERMARK— VENEZUELAN GUAYANA FLORA- III

631

ceolate, acuminate at apex, acute to acuminate
at base, 7.5-10.5 cm long, 2.5-4.5 cm wide, gla-
brous; lateral nerves 6-8 each side, inconspic-
uous; tertiary venation obsolete or inconspic-
uous. Peduncle slender, 1.5-2 cm long, 0.5-0.7
mm diam., simple or forked near base, sparsely
appressed-pubescent with reddish-brown pubes-
cence, not enlarged at junction of summit with
lowest axes of inflorescence. Staminate inflores-
cence terminal, subumbellately 5-branched, 5-8
mm high, 10-15 mm wide, sparsely rufous-brown
appressed-puberulent; primary axes 2-3 mm long;
secondary axes 0.5-1 mm long, each terminating
in branched congested clusters, each of these
bearing 4—5 flowers. Bracts at base of flower clus-
ters ligulate-lanceolate, acute, 1.2 mm long, 0.3-

.4 mm wide, glabrous except for the minutely
puberulous margins near the apex. Staminate
perianth tubular, 1.5 mm long, 0.7 mm wide,
glabrous or glabrescent without. Stamens 5-6,
unequal, included; anthers suborbicular, 0.1 mm
long (immature).

Paratype. VENEZUELA. TERRITORIO FEDERAL
AMAZONAS: Cerro Marahuaca, Sima Camp, S-Central
portion of forested slopes along east branch of Cano
Negro, 3*43'N, 65°31'W, 1,140 m, 21-22, 24 Feb. 1985,
Steyermark & Holst 130545 (MO).

This montane species of Cerro Marahuaca dif-
fers from related species in the glabrous or gla-
brescent staminate perianth and in the mainly
glabrous bracts, except for the apical margins.

The paratype differs from the type collection
in having ovate-elliptic to elliptic-obovate leaves,
which are larger, indistinctly nerved, and sub-
acutely obtuse at the apex.

Neea parimensis Steyermark, sp. nov. TYPE: Ven-
ezuela. Territorio Federal Amazonas: Sierra
arima, Simarawochi, Río Matacuni,
one 64°36’W, 6-7 km west of Venezue-
iom ngu 795-830 m, 18-23
; an A. Steyermark 106982

dudas E isotype, NY).

Arbor 10 m, ramulis dense adpresso- -pubescentibus;

rugineo-tomentosis cum pilis glandulosis interspersis;
pedunculo usque 12 mm longis vel nullo dense fer-
rugineo-tomentoso cum pilis glandulosis interspersis;

632

bracteis cp pn quattuor cupulam facientibus a
ovatis ongis extus dense ferrugineo-tomen-
tosis; rans masculo urceolato 4 mm longo > 5
mm lato extus dense ferrugineo cum pilis denials
interspersis; staminibus 7 inclusis, filamentis 1.2 mm
longis.

Tree 10 m tall with densely appressed-pubes-
cent stems. Petioles 5-8 mm long, densely ap-
pressed-pubescent; leaf blades membranous, ob-
lanceolate, acute at apex, narrowed to an acute
base, 4-7 cm long, 1.5-2.5 cm wide, the upper
surface glabrous, the lower surface densely ap-
pressed-puberulent with minute scalelike hairs;
lateral nerves about 8 each side, scarcely evident.
Staminate inflorescence terminal, 4 cm high, 5
cm broad, much branched, the flowers alter-
nately arranged on ina numerous axes, the 4-8
principal axes unbranched in the basal 8-13 mm,
then forked into alternately or fasciculately dis-
posed secondary axes, densely ferruginous to-
mentose mixed with glandular hairs. Peduncle
none or up to 12 mm long, densely ferruginous
tomentose with intermixed glandular hairs. Bracts
at base of flowers 4, forming a cupule, broa
ovate, subacute, 0.7 mm long, densely ferrugi-
nous tomentose without. Staminate perianth ur-
ceolate, 4 mm long, 2.5 mm wide at the middle,
constricted below the summit, densely ferrugi-
nous tomentose without, mixed with short glan-
dular hairs. Stamens 7, included within the lower
half of the perianth tube; filaments 1.2 mm long.

This species is characterized by having a
broadly urceolate staminate perianth, much-
branched inflorescence, lower leaf surface with a
dense appressed indument, and a short peduncle.
Glandular hairs are scattered on the perianth,
peduncle, and axes of the inflorescence.

Neea robusta Steyermark, Sp. nov. TYPE: Vene-

0 m, 14-28 Feb.
Otto Huber 1693 (holotype, VEN).

1978,

Arbor 3 m, Mona: glabris; petiolis 1-3.5 cm longis
glabris; foliorum laminis obovatis elliptico- a
elliptico- oblongis mus aiu ui QAI apice pleri
que brevit
basi cuneatim acutis 8-24 cm

e glabri

5.
4.5-9.5 cm latis

cm longa 5-10 cm lata

oppositis vel alternis, axis parte apicali minute ad-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74
presso- -ferrugineo puberula axibus aliter glabris, flori-

extus glabro; staminibus 9-10 inclusis; pistillodio 3
mm longo; infructescentia 6-15 cm longo 7-15 cm lato;
pedunculo fructifero 1-6 cm longo 3-5 mm lato ad
apicem dilatato; anthocarpio in sicco 1.2 x 0.5 cm in
vivo 2 x 1 cm.

Tree 3 m tall, the branches glabrous. Petioles
1-3.5 cm long, glabrous; leaf blades coriaceous,
obovate, elliptic-lanceolate, elliptic-oblong, or
oblong-elliptic, mainly shortly and obtusely acute
or obtuse at apex, sometimes rounded, cuneately
acute at the base, 8-24 cm long, 4.5—9.5 cm wide,
glabrous both sides; lateral nerves 9-12 each side,
slightly elevated on upper surface, less conspic-
uous and impressed on lower surface, anasto-
mosing 5-10 mm from the margin; tertiary ve-
nation subelevated and conspicuously reticulate
on upper surface, less conspicuously reticulate
on lower surface. Staminate inflorescence elon-
gated, subpyramidal, paniculately branched, 6-
15 cm long, 5-10 cm wide, the upper axis longer
than the lateral ones, 6-9 cm long, 6 mm diam.,
the 4-8 lateral axes divaricately spreading, 0.3-
2 cm long, 2-3 mm diam., the upper ones gen-
erally shorter and alternate, the lower axes long-
er, opposite or alternate, glabrous except mi-
nutely appressed ferruginous puberulent apically.
Peduncles in anthesis 1-1.2 cm long, in fruiting
stage 1-6 cm long, 3-5 mm diam. Flowers sessile
and scattered but more congested apically. Bracts
subtending flowers deltoid, obtuse, 0.2 mm lon
Staminate perianth carnose, suburceolate, nar-
rowed and contracted below summit, rounded at
base, 6-6.5 mm long, 3.5-4 mm wide, narrower
at summit. Stamens 9-10, included, unequal; an-
thers broadly rhomboid; filaments 2-4.5 mm
long. Pistillode 3 mm long. Infructescence 6-15
cm long, 7-15 cm wide. Anthocarp 1.2 x 0.5 cm
in dried state, 2 x 1 cm fresh.

Paratypes. | VENEZUELA. TERRITORIO FEDERAL
AMAZONAS: Cano Yagua at Cucurital de Yagua, 3?36'N,
UEM, 120 m, Davidse et al. 17362 (MO, Mp

año Yagua, Chipital, 3°29'N, 66°41'W, 120 m,
vide et al. 17320 (MO, VEN); Cucurital de eon

ño Caname, 3?40'N, 67°22'W, 100 m, Davidse et al.
16880 (MO, VEN); 1-2 km SE and E of San Carlos de
Yos Lo 1°56’N, 67°3'W, 120 m, Liesner 6868 (MO,
N); m NE of San Carlos on Solano Road,
Mm un (MO, NY, VEN); 4.3 km NE of San Carlos
on Solano Road, Clark 7038, 7164 (NY); S and SW
of San Carlos de Río Negro, Liesner 6732 (MO, VEN);

Huber Pn al. 374 (VEN); NE and E base of Cerro

1987]

Cucurito, 120 m, Huber & Tillett 2973 (VEN); 12-15
km NE of San Carlos de Rio Negro, road to Solano,
100 m, Morillo et al. 4170 (VEN). BRAZIL. AMAZONAS:
Serra de Neblina, between Maloca and Rio Cauaburi,
50-100 m, Silva & Brazáo 60765 (MO, NY)

This species is characterized by the large, ur-
ceolate, glabrous staminate perianth, the irreg-
ularly paniculately branched, large inflorescence
and general glabrity of parts. The more numerous
secondary nerves, thick peduncle, and conspic-
uously reticulate tertiary venation of the upper
leaf surface distinguish it from related species of
the Venezuelan Guayana.

Neea sebastianii Steyermark, sp. nov. TYPE: Ven-
ezuela. Territorio Federal Amazonas: high
rebalsa (seasonally flooded forest), Isla Se-
bastian, Rio Casiquiare above Chapezon,
between Boca and Solano, 1?58'N, 67?3'W,
120 m, 31 Jan. 1980, Ronald Liesner &
Howard Clark 8959 (holotype, VEN; iso-
type, )

rbor 8 m, ramulis juvenilibus dense miniteque ad-
MSS -puberulis pilis ferrugineis antes petiolis 3-
foliorum laminis ova-
to- ellipticis apice obtuse acutis vel ndis basi cu-
neatim acutis 7.5-14 cm longis 3.5-6.5 cm latis supra
glabris subtus praesertim costa media nervisque mi-
nute puberulis aliter pagina inferiore minute puncti-
culatis atque glandulis sessilibus obtectis; nervis late-
ralibus utroque latere 10-12; inflorescentia mascula 1—
2.5 cm alta 2-2.5 cm lata, axibus 4—5 dense breviter
ferrugineo-tomentosis 3-5 mm pedunculo 1. m
2.5 cm longo 1-2 mm m 2.5-4 m

maturo) dense ferrugineo-tomentoso; staminibus 6-8
inclusis.

Tree 8 m, the younger stems with a dense,
minutely appressed ferruginous puberulence.
Petiole 3-15 mm long, sparsely puberulent; leaf
blades subcoriaceous, ovate-elliptic, obtusely
acute, or rarely obtuse or rounded at apex, cu-
neately acute at the slightly asymmetric base,
7.5-14 cm long, 3.5—6.5 cm wide, glabrous above,
below minutely puberulent, especially the midrib
and nerves, with erect hairs and also with sparse
sessile glands on the leaf surface; lateral nerves
10-12 each side, impressed below. Staminate in-
florescence terminal, 1-2.5 cm long, 2-2.5 cm
wide with 4—5 axes 3-5 mm long, densely fer-
ruginous tomentellose. Bracts subtending flowers
ovate, 0.5-1 mm long, densely ferruginous to-
mentose. Peduncle 1.5-2.5 cm long, 1- m
diam., enlarged to 2.5-4 mm wide at the junction
above with the lowest primary axes. Perianth

STEYERMARK — VENEZUELAN GUAYANA FLORA-III

633

suburceolate, 3-4.5 mm long, 1.5 mm wide (im-
mature), densely ferruginous tomentose. Sta-
mens 6-8, included; anthers rhomboid, 0.8-1 mm
long

This species is related to the other newly de-
scribed species, Neea neblinensis Maguire &
Steyerm., N. huachamacarae Steyerm., and N.
mapourioides Steyerm., but has smaller peri-
anths and has sparsely puberulent petioles and
young stems.

Neea iia . sp. nov
Venezuela. Bolivar: between Betania iid
ae = de ed 40 km west of Santa
Elena, 4?35'N, 61?28'W, 900 m, 15 Dec.
1978, Julian A. Steyermark et al. 117615
(holotype, VEN; isotype, NY

or 15 m, ramulis juvenilibus superne sparsim
puberulis pilis laxis 0.1 mm praeditis; petiolis 3-15
mm longis glabris vel sparse puberulis; foliorum la-
minis oe apice acutis vel obtuse acutis basi
cuneatim angustatis 4—9.5 cm longis 2.5-4 cm latis
ubique ga bris, nervis lateralibus utroque latere 6-7
eben venulis terti
la

we
` oo

ferentibus; pedunculo ten m
lato apice haud dilatato; E sub floribus lanceo-
latis 0.8-1.2 mm longis extus parce puberulo-ciliatis;
perianthio mascula infundibuliformi 3-3.5 mm longo
superne 1-1.6 mm lato basi 0.3 mm lato extus inferne
pilis laxis paucis praeditis atque loborum ee
papillatis aliter glabris; staminibus 6 inclusis

Tree 15 m, the young branches near tip sparse-
ly laxly pubescent, elsewhere glabrous. Petioles
3-15 mm long, glabrous or with a few sparse, lax
hairs; leaf blades opaque, drying dull brown, ob-
anceolate, acute or abruptly obtusely acute at
apex, cuneately narrowed at the equal to slightly
asymmetric base, 4—9.5 cm long, 2.5-4 cm wide,
glabrous both sides; lateral nerves 6-7 each side,
m hina the margin, the ter-
uous. Staminate inflo-
rescence subhemisphe
cm wide, with 4—6 umbellately disposed primary
axes 5-17 mm long, glabrous or sparsely laxly
pubescent; secondary axes 2-5, filiform, umbel-
lately disposed, 3-5 mm long, glabrous or laxly
sparsely pubescent, these branched into pedicel-
late axes 0.5-2 mm long, each bearing 1-3 flow-
ers subtended by lanceolate, acute bracts 0.8-1.2
mm long, sparsely ciliate-puberulent. Peduncle
slender, 2—3.8 cm long, 0.8-1 mm wide, gla-

634

brous, not manifestly enlarged at its junction, 1.5
mm wide where united with the lowest primary
axes of the inflorescence. Staminate perianth in-
fundibuliform, 3-3.5 mm long, 1-1.6 mm wide
at summit, 0.3 mm wide at base, glabrous except
for a few sparse hairs near base and papillate-
margined lobes. Stamens 6, included; anthers
broadly rhomboid, 0.6 x 0.3 mm; filaments 1.5-
2 mm long.

Paratype. VENEZUELA. BOLIVAR: Río Aponguao 2,
151-152 km S of El _ sA et al. 10524 (NY,
VEN); Uei-tepui, between Luepa and Cerro Venamo,
1,100-1,300 m, EDEN 318 (NY. VEN).

Neea tepuiensis Steyermark, sp. nov. TYPE: Ven-
ezuela. Bolivar: Chimantá Massif, Torono
tepui, summit, south-facing forested slopes
above valley of South Cano, 1,955-2, m,
23 Feb. 1955, Julian A. Steyermark & John
Wurdack 1104 (holotype, VEN; isotypes, F,
MO, NY).

rbor 4-5 m praeter gemmas terminales ferrugineas
omnino glaberrima; petiolis 2-10 mm longis; foliorum

ovatis vel obovato-ellipticis vel foliis parvis interdum
suborbicularibus apice anguste obtusis vel rotundatis
basi praeter folia parva rotundata vulgo subacutis vel
anguste acutis (2.7-4.3)7-9.5 cm longis (1.5-3)3.5—5.5
cm latis, nervis lateralibus principalibus utroque latere
5-6; inflorescentia foeminea 1 cm alta s cm iura im-
longis
s 2-3 mm sae gerentibus
cula 2 mm iris 0.8 mm la

pice haud manifeste dilatato; anthocarpio anguste el-
idee oblongo 9 mm longo 4 mm lato glabro.

Tree 4-5 m tall, glabrous throughout, only the
terminal bud glandular-ferruginous pubescent.
Leaves opposite or subopposite; petiole 2-10 mm
long; leaf blades shining above, elliptic-ovate to
obovate-elliptic, or some of the smaller leaves
suborbicular, obtuse to rounded at apex, mainly
subacute to acutely narrowed at base, or some
of the smaller leaves rounded at base, the small
leaves 2.7-4.3 cm long and 1.5-4.5 cm wide, but
leaves mostly 7-9.5 cm long and 3.5-5.5 cm wide,
finely impressed-nerved above, finely and slight-
ly elevated-nerved below, secondary nerves 5-6
each side, faintly anastomosing 5-13 mm from
margin, ascending to an angle of 15—20°; tertiary
venation slightly evident and reticulate beneath.
Pistillate inflorescence 1 cm high, 1.7 cm wide
(immature?) with 4 primary axes 6-7 mm long,

.6 mm wide, each branched into 4 smaller sec-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

ondary axes 2-3 mm long ending in 3 axes bear-
ing the flowers. Flowers sessile. Pistillate peri-
anth 6 mm long, 2 mm wide. Staminate perianth
tubular, glabrous except for the papillate-puber-
ulent npes at the summit, 2 mm long, 0.8 mm
wide. B th lanceolate, sub-
acute, 0.8-1 mm long, | glandular-pubescent. In-
fructescence terminal, bearing 3-4 main axes 7-
11 mm long and 1 mm wide, these suns
into 4 secondary, subumbellate axes 4-7 m
long, these eventually terminating in 4 idle or
slightly branched pedicels 1.5-2 mm long bear-
ing the fruits. Anthocarp narrowly ellipsoid-ob-
long, 9 mm long, 4 mm wide, glabrous, slightly
striate.

Paratype. VENEZUELA. BOLIVAR: Chimanta Massif,
Agparaman tepui, forested middle slopes near Río Ti-
rica, 1,365 m, 5 Mar. 1955, Steyermark & Wurdack
1255 (F, NY, VEN).

This species attains the highest altitude on the
sandstone tabular mountains of the Venezuelan
Guayana, where it grows at an altitude of 1,365-
2,090 m. It is characterized by the glabrity of all
parts and is distinguished from N. robusta n.
erm. byi its smaller flowers and
by the obsolete venation of the tertiary veinlets.
From N. subglabrata Steyerm. it differs in leaf
shape, greater diameter of the peduncle, usually
subobtuse to rounded leaf apices, lustrous upper
surface of leaves, and subelevated, subreticulate
tertiary venation of the lower leaf surface.

LITERATURE CITED

BURGER, W. 1983

. Nyctaginaceae. Jn Flora Costa-
ricensis. Fieldiana, Bot. N.S. 13: 190-191,
196

193-

HEIMERL, A. 1896. Additamenta ad cognitionem flo-
e Indiae occidentalis. III. Bot. Jahrb. Syst. 21:
615-636.
897. Beitráge zur Systematik der Nyctagi-
naceen. Jahresber. Staats-Oberreal. Wien XXIII.
—39.

1914. a In R. Pilger a

Pl antae Uleanae novae vel minus cognitae. No-
tizbl. Kónigl. Bot. "Ga rt. Berin 6: 126-

1932. Nyctaginaceae. Jn N. Y. Sandwith (ed-
itor), Contr. to the Flora of Tropical Am. XII. Kew
Bull. 1932: 220-221.

Huser, H. 1909. Materiaes para a Flora Amazonica.
VII. Plantae Duckeanae austro-guyanenses. Bol.
Belém Mus. Paran. Emilio Goeldi 5: 347-351.

LITTLE, E. L. 1968. Transfers to Guapira from Tor-
rubia (Nyctaginaceae). Phytologia 17: 367-368.

LUNDELL, C. L. 19 tudies of tropical American
plants—V. Wrightia 4: 72-84.

SCHMIDT, J. A. Nyctagineae. /n Martius, Flora
Brasiliensis 14(2): 351-369.

STEYERMARK

1987]

— VENEZUELAN GUAYANA FLORA~III

635

STANDLEY, P. C. 1931.
western South America. Fie
Bot. Ser. 11: 73-90.

The Nyctaginaceae of North-
ld Mus. Nat. Hist

BRUNELLIACEAE
BRUNELLIA
Brunellia nebli is St k & Cuatrecasas,
nov. TYPE: Venezuela-Bexzil border.

za de La Neblina, en bosque alto, 1,500
m, 29 Apr. 1964, J. Ewel 209 (holotype,
NY).

Arbor, kes sad a imparipinnatis 6-jugatis; fo-
liolorum laminis ha ud discoloribus chartaceis ellipti-

nervis ipe eas 16-17 utroque latere distantibus 8-
; foliolis inferioribus 1-2 m

lobato 3.5-4.2
triangulari-lanceolatis acutis 1.5-1.8 mm longis 1-1.2
mm latis; folliculis subrotundatis 3 mm longis 2.5 mm
latis dense hirtellis basi pilis hispidis munitis; enl
nibus obconico-subglobosis subcompressis 3 mm 1

is 2.5 mm latis.

Tree with opposite imparipinnate leaves. Leaf
rachis minutely puberulent. Leaflets in 6 pairs,
not discolored, chartaceous, elliptic-oblong, acute
at apex, asymmetrically acute to subobtuse at
base, 15-18.5 cm long, 4.5-6 cm wide, crenate-
serrate, the upper leaflets sessile, the lower ones
1-2 mm, shortly petiolate; lower surface gla-
brous, except minutely puberulent with short
spreading hairs on midrib, lateral nerves, and
tertiary veinlets; upper surface glabrous. Infruc-
tescence densely tomentellose, the branches 1 mm
diam. Calyx 4—5-lobed, 3.5-4.2 mm in fruit,
densely hirtellous, the lobes triangular-lanceo-
late, acute, 1.5-1.8 mm long, 1-1.2 mm wide.
Follicles subrotund, 3 mm long, 2.5 mm wide,
densely hirtellous with additional hispid hairs at

ase. Seeds shining, obconic-subglobose, nar-
rowed toward one end, 3 mm long, 2.5 mm wide.

This taxon differs from Brunellia comocladi-
folia Humb. & Bonpl. and especially the subsp.
ptariana (Steyerm.) Cuatrec. in the smaller fol-
licles, shorter calyx with slightly narrower calyx
lobes, larger seeds, and the glabrous lower leaf
surfaces, except for the minutely tomentellose
nerves. From B. hygrothermica Cuatrec. it may
be distinguished by the concolorous leaves with
glabrous lower leaf surface, more numerous pairs

of lateral nerves, more depressed and less con-
spicuous serrations, and shorter calyx. From B.
gentryi Cuatrec. it is differentiated by the slightly
larger calyx diameter, larger calyx lobes, longer
and numerous hispid hairs of the follicles, and
concolorous leaves not silvery green beneath.

SAPINDACEAE
MATAYBA

Matayba ptariana Steyermark subsp. guaiqui-
nimae Steyermark, subsp. nov. TYPE: Ven-
ezuela. Bolívar: Departamento Heres, Me-
seta de Guaiquinima, sector NE de la meseta,
cerca de la cumbre, 5?58'N, 63?29'W, 1,400
m, 27 Mar. 1985, Otto Huber 10388 (ho-
lotype, MO; isotypes, NY, VEN)

A subsp. ptariana foliolis minoribus apice rotun-
datis, venulis tertiariis haud elevatis, calycis lobis sub-
iai predic dense — pubescentibus, petalo-

mis interioribus longitudinem petali
nnii en recedit.

Shrub, 2-3 m tall. Leaves bifoliate, glabrous,
the leaflets oblong, rounded at the apex, subob-
tuse or subacute at the base, 2-4 cm long, 1-2.2
cm wide. Inflorescence axillary, 4.5 cm long, the
flowering part 2.5 cm long, 6-7 mm wide, sub-
racemose, 23-flowered. Peduncle 2 cm long, 1
mm wide, solitary, minutely appressed-pubes-
cent; bracts ovate-oblong, subacute, 0.7-1 mm
long, 0.5 mm wide. Pedicels 2 mm long, densely
pubescent with ascending, appressed, pale hairs.
Calyx lobes 5, broadly suborbicular-ovate
rounded to obtuse, appressed ne eee withbut.
1x 1 mm, sericeous within. Petals pilose-ciliate,
subalbiciine broadly rounded above, gee de id
short-unguiculate at base, 1.3 mm long,
broad; 2 inner segments oblong, rounded, b mm
long, 0.8 mm wide, densely pilose both sides.
Stamens 8, exserted; anthers suborbicular, 0.8
mm long; filaments 1.7-1.8 mm long, pilose be-
low, glabrous above the middle; disk cupulate,
glabrous, 0.5 mm high.

At first, this collection appeared to be distinct
from Matayba ptariana. However, a second col-
lection (Huber 10339), assigned to subsp. pta-
riana, in fruit from a lower altitude at 480 m
(“Rio Trueno, 35 km west of caserio of Chig-
uao”), although showing the rounded leaf apices
together with inconspicuous impressed veinlets
on the lower surface of the new subspecies, has
much larger leaflets and longer petioles similar

636

to typical M. ptariana subsp. ptariana. Until ad-
ditional material is forthcoming, it seems best to
retain a subspecific rank for the Huber collection
of the higher altitude.

BOMBACACEAE

NOTES ON CATOSTEMMA AND SCLERONEMA
(BOMBACACEAE)

In the course of preparation of the genus Ca-
tostemma for the Flora of the Venezuelan Gua-
yana, it was found that all the Venezuelan ma-
terial had been identified either as C. commune
Sandw. or had remained unidentified. An ex-
amination of available material in the herbaria
of F, K, MO, NY, US, and VEN indicated that
more than one species of Catostemma was rep-
resented in Venezuela and revealed further the
presence of the genus Scleronema. The present
study includes an attempt to delimit the genus
Catostemma in Venezuela, together with a de-
scription of a new species in Scleronema.

Publications by Sandwith (1931, 1948), Ducke
(1937), and Paula (1969) have greatly enhanced
our knowledge of both genera with the result that
the characters differentiating Catostemma from
Scleronema can now be more clearly defined,
involving chiefly stamen differences. In Sclero-
nema the filaments are dilated and thickened
apically with the small, strongly sessile anther
appressed across the summit of the filament,
whereas in Catostemma the uniformly slender
filaments bear a larger incumbent anther folded
inward over the summit. Additional differences

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

are the longer and more numerous filaments and
longer style branches in Catostemma, also from
available fruiting material, the elongated, ellip-
soid, oblong, or obovoid, tardily dehiscent fruit
of Catostemma as opposed to the globose or
ee ds indehiscent fruit of Scleronema.

Sandwith (1931) was able to differentiate two
species of Catostemma from Guyana, C. com-
mune Sandw. and C. fragrans Benth., on the ba-
sis of vegetative characters found in the seedling
leaves and in the venation, especially ofthe lower
leaf surface. In the present study vegetative dif-
ferences have been noted and employed to dis-
tinguish several taxa. Unfortunately, leaves of
seedling plants are still unknown from most of
the taxa recognized. In addition to the type of
tertiary venation of the lower leaf surface, useful
characters have been found in the glabrity or
stellate tomentum of the buds and petioles.
Moreover, the absence or presence of stellate to-
mentum and its distribution on the style and
style branches, and the presence of bracteoles and
their position on the pedicels serve to delimit
various taxa within the complex. Much of the
material in herbaria is sterile or lacks flowers,
and this has made delimitation of the taxa dif-

cult.

The author wishes to acknowledge to the cu-
rators of F, K, MO, NY, US, and VEN his great
appreciation for the loan of herbarium material.
Based on the material examined, a description
is provided for the taxa of Catostemma, together
with a citation of specimens, followed by a key
to the species.

CATOSTEMMA

KEY TO THE SPECIES OF CATOSTEMMA

la. Adult leaves compound

lb. Adult leaves simple

C. digitatum
2

2a. Petioles and young stems hirsutulous with spreading to ascending stiffish hairs 0.2-0.4 mm long

2b. ape and young stems glabrous or with minute or closely appressed tomentum
ioles and New stems stellate- tomentose

C. hirsutulum
3

4

a. Petioles

tion

E di wer ‘leaf surface irregular or scarcely et Pe leaf surface glabrous; ejon
nt from pedicels; en nerves of leaf blades 5-6 each side; style and style branches

C. clarkii

ox ste late pubesce
. Petioles and y
venatio a oflo ower leaf su

+
c

rfacem

oung stems n nope with larger, less crowded tomentum; tertiary
r less regular, subparallel and subhorizontal; lower

leaf surface minutely stellate- SU to glabrous; bracteoles Arai on pedicels; lateral
af blades

nerves of le 8-12 each

side; style glabrous except at bas

. C. fragrans
5

3b. dg pa young stems glabro
Sa.

veins of lower leaf ipd see in more or less subparallel lines obliquely connecting
C. m

"a dM nerves

arahuacense

1987] STEYERMARK— VENEZUELAN GUAYANA FLORA —III 637

5b. Tertiary m Ws lower leaf surface irregularly reticulate and anastomosing, not regularly
obliquely par
6a. Style stellate- n throughout; di branches stellate-pubescent; us scales
wit ssed, stellate pubes pubistylum
. Style Mes ee at ol or lowest ^; stylar |
or the innermost ones pilos
7a. Bracteoles absent fent een calyx lobes densely strigose within; innermost
bud scales pilose apically ebracteolatum
7b. Bracteoles present, either on the pedicels or immediately subtending the base
of the calyx; calyx lobes glabrous within or partly strigose near apex; bud scales
glabrous 8
8a. itl mediately subtending the calyx or on the uppermost 1-
sel: leaves without revolute ier kane 9
the scales more or less con-
tiguous or overlapping with numerous, relatively elongated rays; leaf
base mainly acute to cuneate, more rarely obtuse „u... C. commune
9b. Pedicels with moderately stellate ironies | the scales minute, scat-
dd. and more separated with fewer and shorter rays; leaf 2 mainly
unded, or sometimes subcordate sclerophyllum
Í Ma at least some of them, toward or below the middle 2 T pedicel;
leaves usually with revolute margins
10a. Pedicels 4-10 cm long; principal lateral nerves of the leaves 8-12 each
ies leaf blades up to 30 cm long and 11.5 cm wide; petals 17-23
m long; tree 10-45 m tall C. altsonii
10b. Pedicels 2.5-3 cm long; principal lateral nerves of the pis "uad each
side; leaf blades up to 15 cm long and 4.5 cm wide; petals 15 mm
long; shrub 2-4 m tall a sancarlosiana

1 lat hiv
o

n
=

oo
c

Catostemma altsonii Sandw., Kew Bull. 1928:
66. 1928. TYPE: Guyana. Macreba Falls,
Kurupung River, Sept. 1925, Altson 391
(holotype, K). Figure 6A, a.

Tree, 10-45 m tall, the branchlets glabrous;
bud scales glabrous. Seedling leaves 3-foliate.
ture leaves with petioles 0.8—6 cm

rounded and mucronate at apex, narrowed to a
cuneate or obtuse base, 8—30 cm long, 3-11.5 cm
wide, glabrous, the margins revolute; primary
lateral nerves 8-12 each side, impressed above,
slightly elevated below, at irregular angles and
not uniformly ascending, branching or anasto-
imo ass reaching the margins, 7-25 mm
apart; tertiary venation of the lower leaf surface
So reticulate, usually prominulous, usu-
ally impressed above on sterile branchlets. Ped-
icels in anthesis 4—10 cm long, densely stellate-
3

e. acute, situated alter-
nately and mend boxer below and above the
middle. Calyx densely stellate-tomentose with-
out, 13-15 mm long, the tube 3-5 mm long, the
lobes ovate-oblong, obtuse, 10 mm long, gla-
brous within. Petals obovate-oblong, rounded,

17-23 mm long, 7-8 mm wide. Stamens ca. 35-
40; filaments 10-14 mm long; anthers 1-1.3 mm
long. Style 10 mm long, glabrous except in low-
ermost 4-6 mm, there stellulate; style branches
2-3.5 mm long, glabrous. Mature fruit ellipsoid,
15-20 cm long, 3-5 cm diam.; immature fruit
obovoid-oblong, ca. 5 cm long, 2 cm diam., mi-
nutely ferruginous furfuraceous, fruiting pedicsl
l

Distribution. Wallaba forest at 50-125 m el-
evation; known only from Guyana

Additional specimens examined. GUYANA: Macre-
ba Falls, Kurupung River, Mazaruni tributary, A/tson
391 (K, ie Essequibo-Mazaruni Divide, about 25
mi. south of Bartica, Forest Dept. Record No. 2316
(Field ios D323) (K); Membaru creek, u uppe er Mazaruni

Bartica-Potaro road, Forest Dept. Record
No. F1520 DM 4256) (K); 1'2 mi. Bartica-Potaro
road, Forest D t. Rec ord No. 6906( Field No. CAP149)

Forest Dept. Record No. 2315 (Field No. D

Potaro River below Tukeit, Maguire & Fanshawe 23486
(NY); Bartica-Potaro road, 24 mi. south of Bartica at
Forest Service's “24 mile Camp," 50-125 m, Mori et
al. 8902 (NY); Membaru Creek, Upper Mazaruni Riv-
er, Pinkus 238 (F, NY, US); Bartica-Potaro road, For-
est Dept. No. 107 (Field No. F 1520, Record No. 4256)
(NY).

638

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FiGURE 6.—A. Catostemma altsonii, leaf. a. Detail of lower leaf surface n irregularly reticulate tertiary

paqa —B. i e es fragrans, leaf. b. Deta
rtiary venatio

siut iun lattiat ascent Based on holoty

Two other species of Catostemma are known
from Guyana. The present taxon may be distin-
guished from C. fragrans Benth. by the glabrous

uds, petioles, and young stems, by the irregu-
larly reticulate tertiary venation on the lower leaf
surface, and by the irregular branching of the
main lateral nerves. From C une it can be

e es
longer pedicels, longer petals and calyx lobes, and
proportionately longer leaf blades

Catostemma clarkii Steyermark, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas:

il of lower
tostemma haha isis leaf. c

leaf surface showing subparallel, subhorizontal
gael of eae leaf surface showing subparallel,

Mari’s bana (low Amazon caatinga), 10.8
km NE of San Carlos on road to Solano,
1°56’N, 67°03'W, 119 m, 19 Aug. 1981,
Howard L. Clark 8126 (holotype, MO).

Arbuscula vel € gemmis ramulis juvenili-
busque minute confertimque cano-stellato-tomentel-
lis; petiolis minute cano- alate foliorum laminis sub-

di inute stellat laminarum

tus praete
juvenilium glabris anguste obovatis, nervis lateralibus

stellatis bracteolis desunt; calyce extus dense stellato-
omentello intus lobis glabris; petalis 20-22 mm longis;
ae inm stellato-tomentello ramis jure mm longis
leviter stellatis

STEYERMARK

1987]

— VENEZUELAN GUAYANA FLORA —III

639

Shrub or small tree to 12 m; young stems
densely and minutely gray stellate-tomentellose;
older branches glabrous. Petioles 4-25 mm long,
minutely and densely gray stellate-tomentellose
on young leaves, minutely gray stellate to gla-
brescent on mature leaves; leaf blades narrowly
obovate, rounded or retuse at apex, mucronulate
in the sinus, narrowed to a rounded or subcor-
date base; mature leaf blades rigid-coriaceous,
4.5-10.5 cm long, 2-4.5 cm wide, glabrous
throughout except for the Ciel stellate basal
portion of the lower midrib; young leaves mi-
nutely gray stellate- utei tm on the lower
midrib; main lateral nerves 5-6 each side, scarce-
ly manifest, lightly impressed on the upper side,
branching t g the margins, 8-15 mm
distant; tertiary venation beneath inconspic-
uously anastomosing. Pedicels ebracteolate, ca.
3.2 cm long, densely and minutely gray stellate-
tomentellose. Petals ligulate-oblong to subspat-
ulate, 20-22 mm long, 6-9 mm wide. Calyx
densely stellate without, the tube campanulate,

m long, the lobes ovate, obtuse, 11 mm long,
i ae ar Stamens numerous, about 40;
filaments 12—13 mm long; anthers 0.8-1 mm long.
Ovary conic, 7 mm long, 3 mm diam., tapering
to an elongate beak 2.7 mm long. Style 9 mm
long, sparsely stellate-pubescent from base to
apex, the branches 4-5 mm long, lightly stellate-
pubescent. Fruit ellipsoid, 6-6.5 cm long, 4.5 cm
diam., fruiting pedicel 4.5 cm long.

r 1

VENEZUELA

atype. lity as type, | Jan.
Ti Clark 6919 (MO).

This species is characterized by the minute,
pale gray, closely stellate tomentum of the veg-
etative buds, young petioles, and lower midrib
of the early leaves, by the sparsely stellate pu-
bescence on style and branches, and by the
scarcely manifest tertiary venation of the lower
leaf surface. The stellate tomentum of this species
is the most minute of the genus.

Catostemma commune Sandw., Kew Bull. 1931:
1. 1931; Oliver in Hook. Icon. Pl. 1986.
1891, as to flowers and leaf, but not to fruit.
TYPE: Guyana. Moraballi Creek, Essequibo
River, Aug. 1929, Sandwith 435 (holotype,

K)

Tree to 45 m. Young branches and stems and
bud scales glabrous. Seedling leaves usually
3-foliate (rarely 2- or 4—5-foliate), the leaflets ob-
lanceolate, obliquely acuminate at the apex, the
acumen triangular-lanceolate, 1-1.8 cm long, 3-

6 mm wide above the middle, obtuse, but the
midrib excurrent, cuneately attenuate at base ex-
cept for the rounded lower side of the outer leaf-
lets, 9-32 cm long, 3-8 cm wide, glabrous both
sides. Mature leaves with petioles 1—9 cm long,
glabrous. Mature leaf blades stiffly chartaceous
or subcoriaceous, elliptic o or obovate, pounce’ or
retuse at apex, rrent costa,
cuneate to rarely obtuse at base, 4-19 cm long,
2-9 cm wide, glabrous on both sides; main lateral
nerves 8-12 each side, impressed above, slightly
elevated below, ascending at an angle of 40-50°,
branching and anastomosing just before reaching
the margins, 8-15 mm distant; tertiary venation
faintly impressed-reticulate above, irregularly re-
ticulate with slightly elevated veins below. Ped-
icels 0.5-2 or 3 cm long, densely stellate-tomen-
tose, bracteolate. Bracteoles paired, approximate,
overlapping, alternate, squamiform or rarely
conspicuous, situated in the upper part of the
pedicel or immediately subtending the calyx,
ovate or suborbicular, obtuse to subacute, 1.5-

8(-6) mm long, 1-1.5(-4.5) mm wide. Calyx
densely stellate-tomentose without, the tube
campanulate, 4 mm long, 3-4 mm wide; lobes
ovate, 7 mm long, 6 mm wide. Petals d pad
spatulate, obtuse, 12 mm long, 4-5 mm w
Stamens numerous, 40-50; filaments 8-10 mm
long; anthers 0.8-0.9 mm long. Style 13 mm long,
stellate-tomentose in lower !^5, elsewhere gla-
brous; style branches 0.75-1.5 mm long. Fruit
oblong-ellipsoid, 7-10 cm long, 3-4 cm diam.,
densely ferruginous tomentellous.

Common names. Baromalli, common ba-
romalli (Guyana), baramán (Venezuela).

Distribution. In mixed tall forest of green-
heart, mora, and morakubea of Guyana, prin-
cipally in the Essequibo and Cuyuni river basins,
and in evergreen mixed forests, often with Eper-
ua, in Estado Bolívar and Territorio Federal Del-
ta Amacuro of eastern Venezuelan Guayana, at
altitudes of 80-850 m

Additional erue examined. WEST INDIES: cul-

tivated in Old Botanic Gardens, St. Vincent, April 1891,
H. Powell (K). GUYANA. Moraballi Creek, Essequibo

No. D402) (K); Upper Mazaruni Kit Leng 416 (seed-
ling leaves, NY); Matt Ridge, ma River,
Northwest Territory, 80 m, Cowan 39361 (F, NY),

39361-A (fruit, NY); P
Forest Dept. Record No. 7712 (K
Tutin 338 (K, US). VENEZUELA: TER

d Cuyuni River,
ORIO RAL
DELTA AMACURO: este de Rio Grande E on de

640

El Palmar cerca de xp "hen del Estado Eae Nov.
1965, Carlos Blanco 495 (MO, US, VEN); La Paloma,
Río Cuyubini, T 100-200 m, pppn rk
87583 (F, MO, NY, US, VEN); El Morro, Atabuina,
Caño Arature, base of Sierra Imataca, 750 m, Bernardi
7529 (K, NY, VEN); same locality and date, Buza 329
(K, NY, VEN); near Río Grande, east of Upata, border
between Estado Bolívar and Territorio Federal Delta
Amacuro, 8?14'N, 61°4'W, 300 m, de Bruijn 1631 (F,

€ I US, VEN), 1677 (US, VEN), 1623 (F,
M ; same locality, Breteler 3841
Dee tie leaves, NY, US), 4958 (NY, US), 4972 (NY);
ENE de El Palmar, cerca de los limites del Estado
Bolívar, Zabala 96 (VEN), 163 (VEN). BOLÍVAR: base
of Cerro Pauji, Quebrada 94, km 94 south of El Do-
rado, 250 m, Steyermark 867 11 (NY, US, VEN), 86712
(VEN), 86715 (F, US, VEN); La Isabel a Rio Grande,
El Palmar, Conejos 36 (MO, VEN); Cerro La Reforma,
above junction of Río Reforma with Río Toro (Río
Grande), Sierra Imataca, 200-250 m, Steyermark 88113
La Lira trail, km 27 S of El Dorado,

o, ESE of Villa Lola, Altiplanicie de dr.
315m, Foi rk 86049 (seedling leaves, y
Río Chirca, Bernardi 898 (3-foliolate leaves, Se east
of El Palmar, Marcano-Berti 319 (VEN); Río ree
between mouth of Rio Aparurén and Uriman, 40

Steyermark 76077 (F, VEN); región de las eens
del Rio Hacha, 450-850 m, Bernardi 2897 (NY); Rio
Caura, arriba del Salto Para, 2-3 km arriba del cam-
pamento “Las Pavas,” 250—300 m, J. Steyermark, G.

. K. & E. Dunsterville 112934 (MO, VEN); same lo-
cality, Morillo & Liesner 8944 (MO, VEN); 30 km S
of El Dorado, 140 m, Bernardi 2097 (NY, VEN); Río
Venamo, afluente del Ikabaru, Cardona 1706 (VEN);
Río Curutü, upper Paragua, 550 m, Cardona 2484
(VEN); Salto Ichun, Río Ichun, tributary of Rio Par-
agua, 4?46'N, 63?*18'W, 500 m, Steyermark 90371
(VEN); El Abismo, Río Samay, 4?27'N, 61?34'W, 550-
600 m, Holst & Liesner 2422 (MO, VEN).

In Sandwith's excellent discussion of the ba-
romallis (Catostemma) of Guyana (1931), a
number of characters were noted by which C.
commune Sandw., or common baromalli, could
be distinguished from C. fragrans Benth., or sand
baromalli, the latter being the type species of the
genus based on a collection by Robert Schom-
burgk 280 in 1837 from Berbice, Guyana. One
of the differentiating | characters mentioned by

C. commune was the
" bracteoles. Sandwith

grans (plate 1986 of Hook. Icon. Pl., 1891). Bracts
conforming to these dimensions are evidenced
by a specimen at K which had been sent from
the Old Botanic Garden of St. Vincent, collected
by H. Powell in April 1891, where it had been
cultivated for many years. This specimen was
cited by Sandwith as a paratype. However, the

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

actual type specimen at K (Sandwith 116) does
not have either large or conspicuous bracts. Thus,
the only flowering material available to Sandwith
was his own collection (the type) and that of
Powell from a cultivated specimen

Three subsequently collected specimens from
the wild state show small squamiform bracts only,
which do not exceed 2.8 mm in length and 1.5
mm in width. These bracts or bracteoles subtend
the very base of the bud, as in Tutin 338 and
Forest Dept. Record No. 7712 (both at K), or
occur on the uppermost 1-6 mm of the pedicel
below the base of the calyx, as in Forest Dept.
Record No. 2398, Davis 402 (K). All these spec-
imens were annotated by Sandwith as C. com-
mune Sandw. Similarly, flowering collections of
this species from adjacent eastern Venezuela show
only squamiform bracts va on the upper-
most 1-6 mm of the pe

Thus, the description s de bracts in C. com-
mune should be emended as follows: “bracteis
squamiformibus late ovatis vel suborbicularibus
obtusis vel subacutis 1.5-2.8(5—6) mm longis 1—
1.5(-4.5) mm latis binis vel arcte alternantibus."

Catostemma digitatum Shepherd & Alverson,
Brittonia 33: 587, fig. 1. 1981. TYPE: Colom-
bia. Antioquia: confluence of Quebrada La
Tirana and Río Anori, 3 km upriver (SW)
from Planta Waaa ca. 28 km SW of
Zaragota, 7°13'N, 75°3'W, 500 m, 9 Apr.

1977, W. S. Piu S. White & E. J. D.
Shepherd 397 (holotype, COL).

Tree to 30 m; young branches or stems gla-
brous. Buds sericeous-tomentose. Mature leaves
3-5-palmately compound. Petioles 6-21 cm long,
glabrous. Leaf blades chartaceous, lanceolate,
long-caudate in seedling leaves, subacute or ob-
tuse and mucronulate in mature leaflets, acute at
base, mainly glabrous; principal lateral nerves
10-16 each side, arcuately ascending near the
margins; tertiary venation below irregularly laxly
reticulate, scarcely prominulous. Flowers un-
known. Fruit obovoid, 9-12 cm long, 5.5-6 cm
diam.; seed 5-7 cm long, 2-4 cm wide.

Distribution. Known only from the depart-
ments of Antioquia and Santander, Colombia.

Additional specimens examined. COLOMBIA.
ANTIOQUIA: confluence of Quebrada La Tirana and Río
Anori, 28 km SW of Zaragota, 7?13'N, 75°3'W, 500
m, Alverson, White & Shepherd 397 (holotype, COL;
isotypes, HUA, MO, NY, WIS); slope facing Río An-
ori, near Planta Providencia, Denslow 1404 (seedling,
COL, HUA, MO, NY, WIS); confluence of Quebrada

1987]

La Tirana and Rio Anori, 3 km upriver (SW) from
Planta Providencia, ca. 28 km SW of Zaragota, 7?13'N,
75°3'W, 500 m, 9 Apr. 1977, W. S. Alverson, S. White,
& E. J. D. Shepherd 397 DE COL). SANTANDER:
Magdalena Valley, Cam E of Carare, 300 m,
Gentry & Renteria 20065 er MO, UIS), 20106
(COL, MO, UIS).

This is the only species thus far known in the
genus with the mature leaves compound.

Catostemma ebracteolatum oe sp. nov.
TYPE: Venezuela. Territorio Federal Ama-
zonas: Cerro Sipapo aa ga water course
above Intermediate Camp, 2 Feb. 1949,
Bassett Maguire & Louis Politi 28741 (ho-
lotype, MO; isotype, NY)

E on 15-30 m, ramulis juvenilibus glabris; gem-
e squamis intimis acuminatis pilosis extimis cau-
datis petiolis 3- -35 mm longis glabris; foliorum laminis
asi
acutis vel cuneatis majoribus 6-10 cm longis 2.5-6.5
cm latis ubiq ue glabris, nervis lateralibus utroque latere
7-9 supra impressis subtus leviter elevatis, venatione
tertiaria subtus leviter irregulariterque reticulatis; ped-
icellis 1.5-4 cm longis pilis dimorphis munitis, pilis
brunneis minoribus t pallidis lon-
gioribus patulis plus minusve seriali bus alternantibus
so, tubo 4

ete] lato-to

longis tabes dense strigosis; petalis ligulato- oblongis 15
mm longis; filamentis ad 9 mm longis; stylo parte in-
feriore » ce 2 stellato-tomentoso ceterum glabro,
ramis ad 4 mm longis glabris.

Tree 15-30 m tall, the young stems glabrous;
innermost bud scales acuminate, pilosulous, the
others glabrous, the outermost caudate. Seedling
leaves trifoliolate. Petioles 3-35 mm long, gla-
brous; leaf blades coriaceous, rich green above,
paler below, obovate, rounded and retuse at apex,
mucronulate, narrowed to an acute or cuneate
base, 6-10 cm long, 2.5-6.5 cm wide, glabrous;
lateral nerves 7-9 each side, spreading at first
from an angle of 10—20°, then ascending, branch-
ing before reaching the margins, impressed above,
lightly and narrowly elevated on the lower side;
tertiary venation irregularly and inconspicuously
reticulate below. Pedicels costate, especially to-
ward apex, 1.5-4 cm long, ebracteolate, with
2-tiered pubescence, the shorter brownish, ap-
pressed, stellate-tomentose hairs between paler
longer hairs + prominently spreading from pseu-
dovertical rows. Calyx in pre-anthesis 12.5 mm
long, split above the side from one portion,
densely stellate without, 4 mm long, 3-5 mm
wide above; lobes longer than the tube, 7-8 mm
long, suborbicular-ovate, obtuse, densely stri-
gose within. Petals ligulate-oblong, rounded, 15

STEYERMARK — VENEZUELAN GUAYANA FLORA—III

641

mm long, 5-6 mm wide. Stamens 25-30; fila-
ments up to 9 mm long; anthers 0.8-0.9 mm
long. Style pubescent for 2.5-4.5 mm from the
base up to !^-'^ of the length, glabrous above;
style branches up to 4 mm long, glabrous. Fruit
orange, rounded at apex, manifestly narrowed to
the base, obtusely 3-costate, 7 cm long, 4 cm

iam., minutely SONOMA fruiting pedicel 4
cm long, 5 cm dia

Paratypes. "VENEZUELA. BOLÍVAR: Sierra Ichün, cer-
canías del Salto María Espuma (Salto Ichün), base of
la Sierra Ichün, tributary of Río Paragua, 4?46'N,
68?18'W, 500 m, = a 1961, Steyermark 90371 (F,
NY, US, VEN). T ORIO FEDERAL AMAZONAS: 4.3
km NNE of San nue on Solano road, 1?56'N, 67°3' W,

100-300 m, Steyermark, Davidse & Guanchez 122136

(MO, VEN), 172386 (MO, VEN), 122402 (MO, VEN);

Sierra Parima, headwaters of Rio Siapa and Rio Pa-

dauiri, 1°22’N, 64°38’W, 1,260 m, Cardona 1507 (US,
VEN); Yavita, Lizot 1972-4 (US, VEN).

This taxon differs from other members of the
genus in its two-tiered pub on the pedicel,
with longer, pale, spreading hairs in more or less
separate vertical rows or lines rather prominently
differentiated from the more abundant, smaller,
brownish, more appressed stellate tomentum in
between. In the other species of the genus the
stellate tomentum is more uniformly appressed,
pale to dark brown, and uninterrupted by longer
hairs.

Catostemma fragrans Benth. in Hook. London
Bot. 2: 365. 1843; Baker in Hook. Icon.
Pl. 1793. 1888; Oliver in Hook. Icon. Pl.
1986. 1891, as to fruit. TYPE: Guyana: Ber-
bice, 1837, Rob. Schomburgk 280 (holotype,
K). Figure 6B, b.
um € Sagot ex vires Bull. Mus.
Hist. Nat. (Paris) 25: 387. 19. TYPE: French
nndis Acarouany, Sagot s.n. (holotype, P).

Tree (4-)1 5-30 m tall. Young branches or stem
stellate-tomentose. Buds stellate-tomentose.
Seedling leaves simple, 12-38 cm long, 3-7 cm
wide, obovate-oblong or obovate-elliptic, nar-
rowly elongated, cuspidate at apex, the cuspidate
portion linear-subulate, this 1-5 cm long, 0.7-
2.5 mm wide, narrowed to a subobtuse to sub-
rotundate base. Mature leaves with petioles 0.5—
5 cm long, stellate-tomentose; mature leaf blades
coriaceous or subcoriaceous, obovate-oblong,
obovate-elliptic, or oblong-elliptic, rounded or
retuse, sometimes cuspidate at apex, rounded at
base, the larger ones 17-22 cm long, 3-8 cm

642

wide, glabrous above, glabrous to sparsely stel-
late-puberulent below, sparsely to moderately
stellate-pubescent on the lower midrib and main
lateral nerves; lateral nerves commonly 8-12 each
side, impressed above, elevated below, + regu-
larly ascending at an angle of 45-60° and sub-
parallel nearly to the margins, 5-20 mm distant;
tertiary venation inconspicuous above, below
conspicuous and elevated with the veins sub-
parallel and subhorizontal. Pedicels 1.5-4 cm
long, densely stellate-tomentose, bracteolate.
Bracteoles 2 or 3, situated near the middle or in
the lower or upper part of the pedicels, some-
times only 2-3 mm below the base of calyx, paired
or usually proximate alternately, ovate, obtuse,
1-2.5 mm long, 1 mm wide. Calyx densely stel-
peti cna without, the tube 3-4 mm long,

m wide, the lobes broadly ovate or ovate-
pais obtuse, 9 mm lon 7 wide, gla-
brous within. Petals spathulate- abus obtuse,
13-15 mm long, 4-7 mm wide. Stamens about
35-40; filaments 7-8 mm long; anthers 1-1.2
mm long. Style 11-15 mm long, glabrous except
stellulate at the very base, the style branches 1.5-
2 mm long, glabrous. Fruit oblong-ellipsoid, 4—
8 cm long, 3-3.5 cm diam., densely and minutely
ferruginous tomentellous; fruiting pedicel 2 cm
long.

Common names. Sand baromalli, baromalli
or wallaba; baromalli (Guyana); kajoewaballi
barmani (Suriname).

Distribution. Sandy soils of wallaba forests
at low elevations in Guyana, Suriname, and
French Guiana.

Gegen specimens examined. GUYANA: near
Bartica, Moraballi Creek, Essequibo River, near sea
ede Sandwith 485 (K, NY); Coverden, Persaud 134
(F, NY); basin of Essequibo River, near mouth of Ono-
ro Creek, l cans , A. C. Smith 2739 (F, MO, NY, US);
a Riv . No rthwest pops bso 59*50'W,
,NY,U ica—Potaro road,

Creek, Rewa River, 35 mi. SSE of mouth, Forest Dept.
Record No. 2087 (Field No. D96) (K); lower Demarara

River, Demarara River, Hohenkerk 704-C (K). SURI-
NAME: km 6, Wijneweg, “B.B.S. 174" (K, MO); Wa-
yombo, Wood Herbarium 352 (K, NY); Maratakka
River, Snake Creek, Maas 10799 (K, NY, U); Cor-
antynes B.B.S. 901 (K); Tafelberg, Maguire
Foengoe Island, Nat. Park Raleigh Falls, Roberts 14764
(NY); Christianburg, Demarara River, Anderson 271
(K); Wilhelmina Gebergte, 2 km below affluence of
Oost River, 225 m, Maguire et al. 54098 (MO, NY).

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FRENCH GUIANA: south of Crique Gregoire, Sinnamary
River, Oldeman 1600 (NY); Sagot & Melinon (photo

of Guenetia macrosperma, F, MO); Godebert, Wach-
enheim s.n. (K) (as Guenetia macrosperma).

This species is the most widely distributed of
the species of Catostemma, being found 1n all
the Guianas, where it is common, especially in
the wallaba forests of Guyana. It differs from the
common taller forest species C. commune Sandw.
in it stellate-pubescent bud scales, petioles, and
young stems; simple, long-cuspidate seedling
leaves; completely different venation ofthe lower
surface of both seedlings and mature leaves; style
glabrous throughout, except at very base; and
somewhat shorter, usually alternately Henn
bracteoles on the pedicels.

Catostemma hirsutulum Steyermark, sp. nov.
TYPE: Venezuela. Bolívar: Chimantá Massif,
rich rain forest on northwestern slopes of
Abácapa-tepui, vicinity of camp | along Río
Abácapa, 420 m, 30-31 Mar. 1953, Julian
A. Steyermark 74781 (holotype, MO; iso-
types, F, MO, VEN).

r 20-25 m; gemmis petiolis ramulis juvenilib-
pat

visis; fructu obovoideo-oblongo 8-9 cm longo 4 cm
lato.

Tree 20-25 m tall. Vegetative buds, petioles,
and young stems densely hirsute with spreading-
ascending hairs 0.2-0.4 mm long. Petioles 3-9
mm long (on young shoots 35 mm); leaf blades
crowded, becoming subverticillate at the ends of
branches, 3-5 cm long, 0.9-3 cm wide, coria-
ceous, shining and deep green above, paler green
below, oblong-obovate, rounded and minutely
mucronulate at the retuse apex, narrowed to a
subacute to subobtuse base, on sterile shoots to
10-11 x 3-6 cm; lateral nerves 5-7 each side,
ascending at an angle of 50-60°, branching before
reaching the margins, the principal lateral nerves
faintly impressed above, slightly elevated below,
2-6 mm distant or on sterile shoots 10-12 mm;
tertiary venation beneath faintly to moderately
prominent with an irregularly reticulate pattern.
ies pendent, i eas obovoid-oblong, 8-
m long, 4 c iam., minutely stellulate-ve-
Saa

`D

STEYERMARK—

VENEZUELAN GUAYANA FLORA —III

643

1987]

Paratypes. VENEZUELA. BOLÍVAR: Quebrada Los
Brasileros, 4.5 km al SW de Icabarü, 4?20'N, 61?48'W,

0 m, 16 Dec. 1978, Weed Carreño Espinosa
& Dunsterville 117778 (MO, VEN). TERRITORIO FE-
DERAL AMAZONAS: Cerro Neblina, south of Base Camp,
Río Mawarinuma, 0*50'N, 66°11’W, 150—350 m, Gen-
try & Stein 47115 (MO).

This taxon differs from other species of the
genus in having the pubescence of the buds, pet-
ioles, and young stems of elongate, spreading-
ascending stiffish, hirsutulous hairs rather than
the densely appressed, short-stellate tomentum
or glabrity of the other members of the genus.
The paratypes differ from the type collection in
having larger leaves widest near, instead of above,

e middle, but possesses identical elongated,
hirsute pubescence.

rr arl

Catostemma ,Sp.nov
TYPE: Venezuela. Territorio Federal ini.
zonas: Departamento Atabapo, Cerro Ma-
rahuaca, “Sima Camp,” south-central por-
tion of forested slopes along eastern branch
of Cafio Negro, 3°43’N, 65°31'W, 1,140 m,
28 Feb.-1 Mar. 1985, Julian A. Steyermark
& Bruce Holst 130878 (holotype, MO). Fig-
ure 6C, c.

Arbor 20-30 m, ramulis Bonis is glabris; gemmis
glabris; petiolis 0.7-3.2 cm longis glabris; foliorum
laminis elliptico-ovatis vl fe ellipticis ad me-
dium latioribus apice rotundatis obtusis vel subacutis
interdum mucronulatis basi obtusis vel subrotundatis
8-14 cm longis 3.5-8 cm latis utrinque glabris, nervis
lateralibus utroque latere 9-12 plus minusve fere ad

venulis tertiariis subtus tenuibus nec elevatis plus mi-
nusve subparalle lis atque subhorizontalibus; floribus
non visis; fructu maturo ellipsoideo 2-3- obtuse costato
vel subangulat 2-sper-
mo fide Holst & Liesner) 8-9 cm longo 4-4. 5 cm diam.
(fructu immaturo magis elongato angustiorique stria-
tulo indumento minute olivaceo-tomentello munito).

Tree 20-30 m tall, the branches glabrous. Bud
scales glabrous. Petiole 0.7-3.2 cm long, gla-
brous; leaf blades chartaceous or subcoriaceous,
brittle, elliptic-ovate, oblong- or lanceolate-ellip-
tic, wiss: at the middle, pounced, boss or sub-

ute at pex, obtuse
Or T cba at the base, 8-14 cm long; 3.5-8
cm wide, glabrous both sides, the midrib im-
pressed above, subelevated below; lateral nerves
slender, 9-12 each side, impressed above, uni-
formly ascending at an angle of 45-50°, subpar-
allel and unbranched nearly to the margins, 3-
15 mm distant; tertiary venation subimpressed
above, faintly subimpressed below and rather in-

conspicuous, not elevated, forming a + subpar-
allel and subhorizontal pattern of veinlets. Ma-
ture fruit ellipsoid, 2-3-obtusely costate or
subangled, unilocular, 1-(2-testa Holst & Lies-
ner)-seeded, 8-9 cm long, 4—4.5 cm diam. (im-
mature fruit longer and narrower after shrink-
ing), striatulate with minute olivaceous indument.

Paratype. VENEZUELA. TERRITORIO FEDE
AMAZONAS: Cerro Marahuaca, same locality as type, as
Feb. 1985, Steyermark & Holst 130709 (MO).

The venation of the lower surface in this species
resembles that found in Catostemma fragrans
Benth. of Guyana, but C. marahuacenses differs
in having glabrous bud scales and less prominent
tertiary venation on the lower leaf surface. The
tertiary veinlets resemble those found in some
species of Scleronema, such as S. neblinense, but
C. marahuacenses differs in its elongate ellipsoid,
instead of globose, fruit.

Catostemma pubistylum Steyermark, sp. nov.
TYPE: Venezuela. Territorio Federal Ama-
zonas: IVIC main study site, 4.3 km NE of
San Carlos de Rio Negro, 1°56'N, 67°03' W,
119 m, 3 Aug. 1978, Howard L. Clark & P.
Maquirino 6742 (holotype, NY).

Arbor 4—5.5(-8) m; gemmis minute adpressoque
stellato- 5. ramulis juvenilibus glabris; petiolis
(5-)12-15 mm longis glabris; foliorum laminis obov-
ato- pane apice rotundatis retusisque basi rotun-
datis vel leviter subcordatis majoribus 9-16 em longis

roque e latere 5
irregulariter reticulatis; pedicellis 2-2.5 cm longis mi-
que adpresso-stellato- tomentoso ebrac-

rte

apicali strigosa ceteru ongo om
nino stellato-tomentello, ramis 4 mm longis stellatis.

Tree, 4-5.5(-8) m tall; buds minutely ap-
pressed stellate-tomentose; young branchlets gla-
brous. Petioles (5—)12—15 mm long, glabrous; leaf
blades thick-coriaceous with revolute margins,
shining above, oblong-obovate, rounded, retuse
and mucronulate at apex, narrowed to a rounded
or slightly subcordate base, the larger ones 9-16
cm long and 4.5—6.8 cm wide, glabrous both sides;
lateral nerves 5—6 each side, ascending at an angle
of 30-45*, prominently elevated below, 10-25
mm distant on the larger leaves, branching before

slightly prominulous veinlets. Pedice
long, minutely stellulate-tomentose, ebracteo-
late. Calyx densely stellate-tomentose without,

644

the tube 4 mm long, 4-5 mm above, the lobes
ovate, obtuse, or rounded, 6 mm long, 3.5-4 mm
wide, glabrous within except for the strigose api-
cal portion. Petals ligulate-oblong, rounded at
apex, 16 mm long, 6.5 mm wide. Stamens 45-
55; filaments up to 9 mm long; anthers 0.5-1
mm long. Ovary suborbicular-ovoid, 3 mm long,
3 mm broad at base. Style 6 mm long, stellate-
tomentellose throughout; style branches 4 mm
long, stellate-pubescent; fruit unknown.

bibi (Ad VENEZUELA. TERRITORIO FEDERAL
: same locality as type collection, Clark &
l. 8115 (MO).

This species Is characterized by a combination
of completely stellate style and stylar branches;
minutely appressed-stellate bud scales; glabrous
petioles, stems, and leaf blades, the latter strongly
revolute; and strigose apical inner portion of the
calyx lobes.

Catost losi Stey k, sp. nov.
TYPE: Venezuela. Territorio Federal Ama-
zonas: Mari's bana (low Amazon caatinga),
10.8 km NE of San Carlos on Solano Road,
1°56'N, 67*03'W, 119 m, 16 Aug. 1981, H.
L. & K. Clark 8117 (holotype, MO).

Frutex 2-4 m, ramulis praecipue ad vel partim
microscopico stellato- -pu ube rulis; g eci
glabris vel a exudato m

petiolis 2- 5; mm longis plerumque glabris vel partim
microscopico stellato- puberulis; foliorum laminis ma-

disnositit calyce extus dense stellato-tomentoso, tubo
3 mm longo 4 mm lato, lobis late ovato-oblongis ob-
tuse acutis 10-12 mm longis 5-6 mm latis intus glabris;
petalis 15 mm longis 6-8 mm latis; stylo 9-11 mm
longo glabro, ramis ^ mm longis glabris vel pilis stel-
latis minutis munit

Shrub 2-4 m tall, the branches mainly gla-
brous or partly stellulate-puberulent with micro-
scopic indument. Buds mainly glabrous or some-
times furnished with a microscopic pale exudate.
Petioles 2215 mm long, mainly glabrous or partly
stellate-puberulent with microscopic indument;
mature leaves thick, rigid, coriaceous, strongly

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

revolute, obovate, rounded and retuse at the usu-
ally mucronulate apex, 4.5-15 cm long, 2-4.5
cm wide, glabrous both sides; main lateral nerves
4—6 each side, scarcely evident and impressed or
subsulcate above, usually elevated and conspic-
uous below, 5-20 mm distant; tertiary venation
above with a minutely cellular impressed retic-
ulum surrounded by a slightly more impressed
network, reticulate below with mainly finely im-
pressed veinlets, some of these obliquely trans-
verse between the main lateral nerves. Flowers
few, (1-)2-7-fasciculate; pedicels 2.5-3 cm long,
densely stellate-tomentose, bracteolate, the 2
bracteoles alternately situated at or below the
middle, deltoid-ovate, obtuse, 0.8-1 mm long.
Calyx densely stellate-tomentose without, 3 mm
ong, 4 mm wide above, the lobes broadly ovate-
oblong, obtusely acute, 10-12 mm long, 5-6 mm
wide, glabrous within. Petals ligulate-oblong or
ligulate-spatulate, rounded, 15 mm long, 6-8 mm
wide. Stamens 35-40; filaments 9-12 mm long;
anthers 0.8-0.9 mm long. Style 9-11 mm long,
glabrous except stellate in basal 1.5 mm, glabrous
or partly minutely stellulate, with style branches
2 mm long.

This species differs from Catostemma altsonii
in its shrubby habit and low stature and in its
thick coriaceous and strongly revolute leaf blades
with only 4-6 main lateral nerves on each side.

—

Catostemma sclerophyllum Ducke, Trop. Woods
50: 39. 1937. TYPE: Brazil. Amazonas: near
Manaos, 20 May 1936, A. Ducke (holotype,
RB no. 29040).

Medium-sized tree. Bud scales and branches
glabrous. Petioles 2-5 cm long, glabrous; leaf
blades congested at the apex of the branches,
oblong-elliptic, rounded and retuse at apex, sub-
cordate or rounded at the base, 6-10 cm long,
4—7.5 cm wide, glabrous both sides, the main
lateral nerves 6-10 each side, branching before
reaching the margins; tertiary venation irregu-
larly reticulate between the main secondary lat-
eral nerves, less conspicuously elevated below
than the main lateral nerves. Pedicels 2.5-5 cm
long, moderately stellate-tomentose, bracteolate;
bracteoles alternate, immediately subtending the
calyx, 1-1.5 mm long, | mm wide, broadly ovate,
obtuse. Calyx stellate-tomentose without, 10-12
mm long, the tube 4-5 mm long, the lobes 5—6
mm long, glabrous within. Petals obovate-ob-
long, 12-15 mm long. Stamens 30-35; filaments
7-9 mm long; anthers 0.5-0.7 mm long. Style 6-
7 mm long, glabrous except stellate-tomentose

1987]

in the basal 2 mm portion; style branches 2-2.5
mm long, glabrous. Fruit ellipsoid, 6-7 cm long.

Distribution. Amazonian Brazil.

re gundihciy examined. BRAZIL. AMAZONAS:
Ducke 20-5-1936, Herb. Jard. Bot. Rio
de Janeiro M (holotype, RB; beri MO); Man-
A do Aleixo, Ducke 490 (F, US); Manáos,
minis oni Ducke 749 (F, US);
1469 (F); same locality,
Steward et al. (NY). Seen also near Santa Izabel on Rio
Negro by Ducke.

This species resembles C. commune but differs
in having the bracteoles on the pedicels imme-
diately subtending the calyx and alternately ar-
ranged; longer, less stellate-tomentose pedicels;
somewhat longer petals; and more rigid and
thicker leaves, subcordate to rounded at the base.
From C. fragrans it may be distinguished by hav-
ing glabrous buds and young stems and by having
completely different venation of the lower leaf
surface.

SCLERONEMA

The genus Sc/ has hitherto been known
only from Brazil and Guyana, the most recently
described species being S. guianense Sandw.
(1948). The following species is newly described
as the first one known from Venezuela.

Scleronema nebli St k, sp. nov. TYPE:
Brazil. Amazonas: Serra de Neblina, vicin-
ity of Base Camp, Cano Tucano, Rio Cau-
aburi, 100 m, 15 Nov. 1965, Bassett Ma-
guire, Julian A. Steyermark & Celia K.
Maguire 60181 (holotype, MO; isotype,
flowers, NY).

Arbor 20-40 m, ramulis juvenilibus stellato-tomen-
tellis; gemmis dense stellato-tomentosis; petiolis 1-4
cm longis stellato-tomentosis; foliorum laminis plan-
tarum incipientium simplicibus, elliptico- Mrd
subito poe cuspide 1.5-3 cm longo, 14-24

cm longis, m latis utrinque see iliorum
laminis maturis late ° oblongis vel ae apice ro-

l4 xps
3.5-7.5 cm latis subtus praeter costam mediam nervis

lateralibusque sparsim stellatis sabre. 1 ne
libus EE latere 7- I: subtus prominenti
vatis plus m argines uniformiter ad-
scendentibus subparallelisque; venulis tertiariis subtus

talibus; „pedicellis (1.3-)1. 8- 2.5 cm longis stellato-to-
s; calyce extus dense stellato-tomentoso lobis
tribus intus moderatim pubescentibus pilis hirsutulo-
strigosis praeditis; staminibus 14-16, filamentis apice
dilatatis 7-8 mm longis, antheris sessilibus 0.2-0.3 mm

longis; stylo 13.5 mm longo, parte basali 4 mm longa

STEYERMARK — VENEZUELAN GUAYANA FLORA-—III

645

stellata, ceterum glabro; fructu globoso 7 x 7 cm sub-
ruguloso subglabrescente.

Tree 20-40 m tall, the young branches stellate-
tomentose. Buds densely stellate-tomentose.
Seedling leaves simple, elliptic-oblong, abruptly
long cuspidate, 14—24 cm long, 5.5-7.5 cm wide,
the cuspid part 1.5-3 cm long, glabrous both
sides. Petioles 1-4 cm long, stellate-tomentose;
mature leaf blades chartaceous, oblong or ob-
ovate, rounded and sometimes mucronate at
apex, rounded to subacute at base, 5.5-14 cm
long, 3.5-7.5 cm wide, glabrous above (a few
stellate hairs sometimes on midrib), the lower
surface glabrous; lateral nerves 7-13 each side,
impressed above, prominently elevated below,
5-15 mm distant, + uniformly ascending at an
angle of 45-55?, + subparallel and unbranched
nearly to the margins; tertiary venation promi-
nent and elevated below with the veins + reg-
ularly subparallel and subhorizontal. Pedicels
(1.3 —2.5 cm long, ebracteolate, densely stel-
late-tomentose with shorter and longer hairs in-
termixed, the shorter brownish hairs predomi-
nating. Calyx minutely densely stellate-tomentose
without, tube campanulate, 4 mm long, 4 mm
broad at summit, the 3 lobes 4.5 mm long, 4 mm
wide, moderately hirsute-strigose within with
hairs 1 mm long. Petals lance-oblong, obtuse, 13
mm long, 4 mm wide. Stamens 14-16; filaments
7-8 mm long; anthers 0.2-0.3 mm long. Style
13.5 mm long, stellate-tomentose in the basal 4
mm. Fruit globose, 7 x 7 cm, faintly rugulose,
subglabrescent.

Paratype. M TERRITORIO FEDERAL

á Stein 47148 (fruiting material with one flower pres-
ent, MO), 47152 (seedling leaves, MO); same locality,
11 Mar. 1984, Liesner 16539 (MO).

This taxon possesses small sessile anthers ter-
minating the relatively few dilated ñlaments and
indehiscent, globose fruits. The morphologies of
the anthers and the dilated filaments are typical
of Scleronema, and globose fruits are known from
other species of the genus, in contrast to the more
elongated ellipsoid or oblong fruits of Catostem-
ma. Others features of S. neblinense shared by
the known species of the genus are the relatively
few stamens, the relatively few flowers of each
axillary fascicle, and the more or less character-
istic tertiary venation and subparallel nen
nerves. While the calyx lobes of Catostemma are
regularly three, those of Scleronema vary from
three to five. The three calyx lobes of S. nebli-

646

nense are also found in S. micranthum Ducke,
to which it is most closely related. The former
differs in the rounded apex of the perfectly glo-
bose fruit and in the greater number of lateral
nerves of the leaf blades.

LITERATURE CITED

BAKER, J. G. 1888. Catostemma fragrans Benth.
Hooker's Icon. Pl., pl. 1793.

BENTHAM, G. 1843. Contributions toward a flora of
South America —enumeration of plants collected
by Mr. Schomburgk, in British Guiana. London

5.

0. Plantes bir beg ou peu connues
e la region amazonienne. Arch. Jard. Bot. Rio de
Janeiro 5: 163-165.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

—. 1937. New forest trees of the Brazilian Am-
n. Trop. Woods 50: 37-39.

one D. 1891. Catostemma fragrans. Hooker’s
Icon. Pl., pl. 1

PAULA, J. E. 1969. Estudios sobre Bombacaceae — I.
Ci. e Cultura 21: 19.

SANDWITH, l N British Guiana.

Bull. Misc. Inform. 1928, no. 9: 365-379.

1931. Contributions to the flora of Tropical
America. IV and V. The baromallis of British
Guiana. Kew Bull. 1931: 46-54.

1948. Contributions to the flora of Tropical
America. XLVIII. Kew Bull. 1948: 304—306.
SHEPHERD, J. D. & W. S. ALvERSON. 1981. A new
Catostemma (Bombacaceae) from Colombia.

Brittonia 33: 588-590.

THEACEAE

BONNETIA

KEY TO RECENTLY DESCRIBED TAXA OF BONNETIA

la. Anthers broader than long or as broad as long, subreniform; ovary incompletely trilocular

lb. Anthers longer than broad, oblong or suborbicular; ovary completely trilocular
2

Style undivided

. euryanthera
2

3

Lateral nerves elevated on upper surface, impressed on lower surface; leaf blades 2-4 x 1.5-
5

cm

. tepuiensis ve peia

blades 1-2 x

3b. Lateral nerves ewe sia on upper surface, mainly not evident on lower surfac
4c

N
c

4a. Style shallowly 3-lobed

i Ps divided nearly or all x length into 3 branches, or merely shallowly 3- lobed at the

tepuiensis isi minor
eps Emo

quinimae

4b. vid deeply parted into 3 branches, divided nearly all the way to the base or at least V "0 from

top
P Petals white or p
6a. Uppe

oui send or 3-parted to
a. Petals 9-9.5 x

ink
r leaf ms e sepals subacute or obtusely acute; styles divided ?^ of
ba

4-6 mm; leaves 1.2-2.7 x 0.5-0.8 cm; sepals 9-9. 1 mm lon

B. chimantensis

7b. Petals 21-22 x 14-16 mm at the summit; leaves (2—)2.5—5 x 0.8-1. 7 cm; sepals

11-12.5 mm long

B. boli varensis

6b. Upper leaf Puce with impressed n above; sepals obtuse or ee at apex;

styles 3-parted '4 distance from the

5b. Petals yellow

B. toroniensis
8

a. Peduncle elongate, 3—6.5 cm long, often surpassing the leaves, ebracteolate
cm mu 2.7-3 cm wide; lateral nerves o

9a. Petals 3.5-4.2 c
leaves scarcely elev

sa surfaces of th
tristyla subsp. tristyla

ted
9b. Petals smaller; ues nerves of both leaf surfaces prominently elevated

tristyla subsp. nervosa

8b. digi lacking or at most 1.6 cm long, usually concealed by and much short
than

e leaves, verticillate-bracte

=
oO

10a. pend linear-oblanceolate, > 5-7 mm wide; petals 8 mm long; — es 8-
16 mm

long

B. huberiana

10b.

mig broadly lanceolate, obovate, or oblong-lanceolate, 4-15 mm wide (leaves

n vegetative shoots often wider); petals 9-16 mm long; peduncles 3-16 mm

ong
11a. Leaves lanceolate, acute at the apex, only Ec» ipli ug at ag base,
width for most of len

nearly the same w

th, 3.5-4 aed

impressed-nerved beneath, not pale punctate there, ihe Ph aum
B

ifest; sepals 12-13 m

Leaves oblanceolate or obovate, obtuse, rounded t

m long ptariensis

tthea

conspicuously narrowed : the base, broadest above the oet 1-3. S x

Is]

tomata

yen
manifest; sepals 9-10 mm ines

e wurdackii

1987]

In the first installment of the Flora of the Ven-
ezuelan Guayana (Steyermark, 1984), four new
taxa of Bonnetia were described. Continued ex-
ploration of the summit floras of the Guayana
Highland has yielded the following five new bon-
netias.

Bonnetia bolivarensis Steyermark, sp. nov. TYPE:
Venezuela. Bolivar; Ptari-tepui, cumbre,
5°47'N, 61°47'W, 2,400 m, 19 Nov. 1984,
Otto Huber 9818 (holotype, VEN; isotype,
MO)

Frutex 1 m; foliorum laminis dense rosulatis sub-
sessilibus ber Vier dr apice obtuse acutis basi
obtusis (2-)2.5-5 .7 cm; sepalis 11-12.5 x
5 mm; petalis 2122 x 14-16 mm; lag tribus 3-3.5
mm longis fere usque ad basem divisis

Shrub 1 m tall. Leaves crowded at summit of
branches, oblong-lanceolate, subsessile, nar-
rowed to an obtusely acute apex, obtuse at base,
(2-)2.5-5 x 0.8-1.7 cm, faintly impressed-nerved
both sides (the midrib slightly elevated below)
or the lateral nerves not evident. Flowers soli-
tary, sessile or subsessile; bracts immediately
subtending flower oblong-lanceolate, acute to
obtuse, 11-12 x 4.5 mm, dorsally carinate, se-
tulose marginally with dark setae 1 mm long.
Sepals lance-oblong, subacute, 11-12.5 x 4.5
mm, obtusely dorsally keeled basally and api-
cally; petals white, subcuneately obovate, sub-
truncate apically with unequally rounded sides,
narrowed to the base, 21-22 mm long, 14-16
mm wide at summit, 4 mm wide at base. Sta-
mens numerous, multiseriate; filaments 5 mm or
less long; anthers .6 mm. Pistil 9 mm
long, the styles 3, 3-3.5 mm long, divided about
25 of their length.

The larger flowers and larger subacute leaves
differentiate this taxon from B. chimantensis
Steyerm., B. tepuiensis Kobuski & Steyerm., and
B. toronoensis Steyerm. In its deeply 3-parted
style it further differs from B. tepuiensis.

Bonnetia euryanthera Steyermark, sp. nov. TYPE:
Venezuela. Bolivar: meseta de Jaua, cumbre,
sección oriental-central, afloramientos de
piedra arenisca en sitios expuestos con ve-
getación herbácea y arbustos achaparrados,
4°35'N, 64°15’W, 2,020 m, 14 Feb. 1981,
Julian A. Steyermark, Charles Brewer-Ca-
rias & Ron Liesner 124311 (holotype, NY;
isotypes, MO, VEN). Figure 7

Subfrutex 1 m, ramulis parum ramosis; cicatricibus

STEYERMARK— VENEZUELAN GUAYANA FLORA III

647
valde confertis; foliis alternis ad apicem confertis ob-

m latis, nervis lateralibus vix manifestis; inflorescen-
tia terminali conferta dense cymosa, 12-15 flora, ax-

S
datis 3-4.5 mm longis 1.5-2 mm latis marginibus ver-

vel subsessilibus; sepalis 5, inaequalibus exterioribus
ovalibus vel suboblongis rotundatis 5-6 mm longis 3-
is mm latis, interioribus majoribus ovatis obtusis 6-7

m longis 4 mm latis dorsalibus carinatis; petalis 5
suborbiculari, obovatis apice rotundato-subtruncatis
10-11 mm longis apicem versus 6-8 mm latis; fila-
mentis numerosis 2-seriatis; antheris subreniformibus
latioribus quam longioribus 0.5 mm altis 0.6-0.7 mm
s " thecatis; ovario a mad imperfecte 3- locu-

ietali placentis ;Sty-
lis trifidis, ramis 1- ‘I. 25 mm lonis

Dwarf shrub 1 m tall, sparsely branched with
the leaf scars closely crowded, about 2 mm dis-
tant. Leaves alternate, closely crowded near the
summit, coriaceous, oblong to obovate-oblong,
rounded or obtuse at apex, sessile or subsessile,
3-4.5 cm long, 1-2 cm wide; lateral nerves
scarcely evident on either side. Inflorescence ter-
minal, compact, densely cymosely subpanicu-
late, 2 cm high, cm wide with 2-3 short axes
3-5 mm long, 1.5 mm wide, 12-15-flowered, on
a short peduncle 2 mm long, 1.5 mm wide. Brac-
teoles ligulate-oblong, rounded, 3-4.5 mm long
1.5-2 mm wide, with minutely setulose margins
especially manifest in lower half. Flowers sessile,
crowded. Sepals 5, unequal, the outer oval or
oblong, rounded, 5-6 mm long, 3-4 mm wide,
the inner larger, ovate, obtuse, 6-7 mm long, 4
mm wide, dorsally carinate. Petals 5, pink, rose
and white, suborbicular-obovate, subtruncate-
rounded at apex, 10-11 mm long, 6-8 mm wide
near apex, 2 mm wide at base. Stamens numer-
ous; filaments filiform, 3-5 mm long, 2-seriate,
attached at base to a thickened tissue; anthers
4-celled, subreniform, broader than long, 0.5 mm
high, 0.6-0.7 mm wide, the lop d Sery l or
subelliptic. Pistil 5.5 mm long; o
long, l-celled (incompletely 3- elled), the he.
centae parietal and conspicuously intruding to-
ward center; ovules numerous. Style 1 mm long,
trifid, the style branches 1-1.25 mm long; stig-
mas terminal.

Paratype. VENEZUELA. Same locality and date as
type, E Brewer-Carias & Liesner 124322
(NY, VEN).

This species is remarkably distinct in the shape
of the anthers, which are broader than long or

648

20mm

G

0.7mm

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

4.5cm

5.5mm

lace euryanthera. — A. Fertile branch in bud stage. — B. Portion of iioi. branch enlarged. —

. Flower, lateral view. gus Petal.
tu dinal aces oe howing placentae and o

—E. Anther, poe

view. — F. Anther, dorsal view.—G. Pistil.—H. Longi-

Cross eis through ovary showing single

stil s :
locule occupied by sea parietal placentae with ss ie pw on holoty

as broad as long and subreniform to suborbic-
ular. Other distinctive features are found in the
incompletely trilocular ovary with parietal pla-
centae extending toward and almost fusing at the
center, their ovules crowded and touching the
ends of the placenta dissepiment, in this respect
simulating the drawing of the ovary of Bonnetia
celiae Maguire (Maguire, 1972, fig. 22k), which
is quite different from the completely trilocular
ovary of B. neblinae Maguire (Maguire, 1972,
fig. 23g). Several dissections of the ovary verify
observations which were also confirmed by bo-
tanical colleagues at the Missouri Botanical Gar-
den. A longitudinal section of the ovary shows
that the placental partitions are not united. They

nearly come together, but a slight space develops

angustifolia Maguire (A, b-c)

nifolium (B, b—c), Bonnetia neblinensis Maguire
(D, b=), and B. steyermarkii Kobuski (F, b-c).
In this respect they differ from the linear-oblong
anthers of Bonnetia jauaensis Maguire (Maguire,
1976) collected on another part of the summit
of Cerro Jaua of the Meseta de Jaua.

Cronquist (1981) stated that the gynoecium of
all four subfamilies of the Theaceae has axile
placentation, **(2-2)3-5(-10) carpels," and forms
a compound ovary with equal ovule and carpel

1987]

number, except for Piquetia, which has the car-
pels united only at the base. In his treatment of
the Bonnetiaceae, Maguire (1972) stated that the
ovary is **5-3-locular" and that the placentation
is *axial or a permutation thereof." In our species,
it would be more accurate to state that the ovary
is incompletely trilocular, or actually unilocular
with three parietal placentae extending inward
but not fusing or unite

Bonnetia guaiquinimae Steyerm., sp. nov. TYPE:
Venezuela. Bolívar: Cerro Guaiquinima,
cumbre, sector SE, 5?40'N, 63?26'W, 1,250
m, 26 May 1978, Julian A. Steyermark, Paul
Berry, G. C. K. & E. Dunsterville 117421
(holotype, MO).

Frutex 1.5 m; foliorum laminis subpetiolatis oblon-

vatis integerrimis; sepalis minute mucronatis 10-12 x
m —20 mm; filamentis 3.5-7 mm
locis stylo subulato apice leviter 3-lobato.

Shrub 1.5 m tall. Petiole 1-2 mm long; leaves
coriaceous, entire, oblong-lanceolate to oblan-
ceolate, acute at apex, gradually narrowed to a
subacute or subobtuse base, 5-6.5 x 1.3-1.6 cm,
enervate below, the midrib subimpressed below,
the lateral nerves elevated above. Sepals coria-
ceous, suborbicular-obovate, rounded at a short-
ly cuspidate apex, 10-12 mm long, 6-8 mm wide
above the middle, 3-4 mm wide at base. Petals
white, obovate, narrowed to a niin mid
base, 20 mm long, 15-20 mm wide at the sum-
mit, 2-3 mm wide at base. Filaments Mi une
3.5-7 mm long; anthers 1.5-1.8 x 0.7 mm. Pistil
9 mm long; style united, merely 3-lobed at apex.

This taxon is characterized by the shallowly
3-lobed style and the entire, oblong-lanceolate,
acute leaves, which are enervate beneath. It may
be distinguished from B. chimantensis Steyerm.
by the larger petals, shallowly trilobed style, and
larger leaves enervate beneath. It differs from B.
toronoensis Steyerm. in the larger sepals and pet-
als, and larger, entire leaves enervate below, while
it is differentiated from B. tepuiensis and subsp.
minor Steyerm. by the larger, minutely mucro-
nate sepals, longer filaments, and completely en-
tire leaves.

Bonnetia ptariensis Steyerm., sp. nov. TYPE:
Venezuela. Bolívar: Ptari-tepui, cumbre,
5?45'N, 61?45'W, 2,360-2,420 m, 23 Feb.
1978, Julian Steyermark, V. Carrefio E., R.

STEYERMARK — VENEZUELAN GUAYANA FLORA III

649

McDiarmid & C. Brewer-Carias 115645
(holotype, MO).

Frutex 2.5 m; foliis sessilibus lanceolatis apice acutis
majoribus 3.5-4.5 x 1-1.5 cm minute obscureque ser-
rulatis; floribus solitariis minute ae pedunculatis,
pedunculis teretibus vel subteretibus 4—6 cm longis;
nes lanceolatis vel suboblanceolatis acutis 12-13 x

m; petalis luteis obovatis RE rotundatis 16-
18 x T mm; antheris 0.8 x 0.7 mm; stylis tribus 6
mm longis fere usque ad basim divisis.

Subshrubs 2.5 m tall. Leaves coriaceous, lan-
ceolate, acute at apex, slightly narrowed to the
base, the larger ones 3.5-4.5 x 1-1.5 cm and
obsoletely pinnately nerved, the midrib slightly
elevated below, microscopically and obscurely
serrulate. Flowers solitary on (sub-)terete pedun-
cle 4-6 mm long; bracts immediately subtending
flower narrowly oblanceolate, acute, 12-13 x 3-
4 mm. Sepals subconvex, lanceolate or sublan-
ceolate, acute, 12-13 x 3-5 mm, 10-striate, mi-
nutely ciliolate, Mt ud carinate; Vine yellow,
obovate, rounded above, 16-18 mm long (pre-
anthesis), 12 mm wien near apex, - mm wide at
base; filaments 1.5—3.5 mm long (pre-anthesis);
anthers 0.8 x 0.7 mm long. Pistil 10 mm long;
styles 3, 6 mm long, free nearly to the base.

This species differs from the yellow-flowered
B. wurdackii Maguire in the larger, lanceolate,
and acute leaves, longer sepals, larger petals (even
in bud), longer filaments, and longer style
branches. Furthermore, the leaves do not show
the pale-punctate stomata, which are clearly vis-
ible in B. wurdackii. It is easily distinguished
from B. tristyla by the short pedicels, smaller
floral parts, and smaller leaves, and is well dif-
ferentiated from B. huberiana by the larger lan-
ceolate leaves, shorter pedicels, and larger floral
parts.

Bonnetia tristyla Gleason subsp. nervosa "Don
ermark subsp. nov. TYPE: Venezuela. Ter-
ritorio Federal Amazonas: Cerro X.
2 Jan. 1951, Maguire, Cowan & Wurdack
30632 (holotype, VEN; isotype, NY).
subsp. B. tristyla foliis supra costa media necnon

nervis lateralibus nonnullis (6-8) conspicue manifes-

teque elevatis, nervis lateralibus supra inaequaliter
prominentibus; petalis minoribus 25-38 mm longis re-
cedit.

Petiole absent or 1-2 mm long. Leaves subob-
long, oblong-oblanceolate, or oblong-obovate,
obtuse to rounded at apex, narrowed to a sub-
obtuse or subacute base, 4-8 cm long, 1.5-3.5
cm wide, the midrib on upper surface and 6-8

650

pairs of lateral nerves conspicuous and elevated,
the other intermediate pairs of lateral nerves less
conspicuously and lightly impressed, nerves on
lower surface subequal and lightly impressed. Pe-
duncle 3-4.5(-6.5) cm long. Sepals 12-20 mm
long, the outer 12-15 mm long, the inner ones
15-20 mm long. Petals 25-38 x 15-22 mm.

Paratypes. VENEZ ELA. TERRITORIO FEDERA

acana, summit, 3?45'N, 66?45'W, 1,000-2,000 m,
ena & Bunting 103103; Yapacana, 1,200 m,
Maguire et al. 30665; Cerro Avispa, Río Siapa, sum-
mit, 1?30'N, 65*51'W, 1,510 m, Cardona 3098; Neb-
lina, summit, Canon Grande slopes E of summit camp
“1,200-2,200” m, Maguire et al. 42179; Neblina, Cañ-
on Grande SSW of summit camp, 1,050-1,100 m, Ma-
guire et al. 42498; Yapacana, Maguire et al. 30632;
Neblina, Cañon Grande, slopes E of summit camp,
1,200-1,300 m, co al. 42235; vicinity of Cerro
Vi nilla, 30 km Ocamo, caños affluent to Río

Huber 6168; altiplanicie de
arenisca, E side of Río Siapa or Matapire, slightly above
leaving the Macizo of Aracumuni, 1?36'N, 65?41'W,
600 m, Huber 6006

This taxon differs from typical Bonnetia tri-
styla in having smaller petals, prominently ele-
vated upper midrib, and unequally prominent
lateral nerves on the upper surface of the leaf,
with 6-8 conspicuously elevated pairs alternat-
ing with fine, lightly impressed alternate nerves.
There is also a tendency for the base of the leaf
blade in subsp. tristyla to be cuneately narrowed,
whereas in subsp. nervosa, the base of the leaf
blade is usually slightly obtusely curved or
rounded above its junction with the petiole.

LITERATURE CITED

CRONQUIST, A. 1981. An Integrated System of the
C sini w Flowering Plants. Columbia Univ.

Press, New
ce B. 1972. The as of the Guayana High-
m. New York Bot. Gard.

D, b-c; F, b-«; 142,
Theaceae. In J. A. Steyermark & C.
Brewer- Carias (editors), La Vegetación de la Cima
del Macizo de Jaua. Bol. Soc. Venez. Ci. Nat.

1984. Flora of the Venezuelan
LI. Ann. Missouri Bot. Gard. 71: 326-

NOTES ON LAPLACEA

The last revision of the genus Laplacea, by
Kobuski (1950), recognized nine species. Two of

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

these, L. acutifolia (Wawra) Kobuski and L. ob-
ovata (Wawra) Kobuski, were considered by Wa-
wra (1886) as varieties of Laplacea semiserrata,
or of Wikstroemia fruticosa by Blake (1918).

Kobuski noted that great variation existed in
size of the flowers and fruits, even within a single
species, and that the keys to the species had been
“based almost entirely on leaf characters and pu-
bescence."

The species of Laplacea can be divided into
two groups, those with asymmetrical leaves and
those with symmetrical leaves, both of which
occur in the Venezuelan Guayana, the common
asymmetrical type represented by L. fruticosa
(Schrader) Kobuski, the symmetrical type by L.
pubescens Planchon & Linden. Variations occur
in each species (Fig. 8B, b, 8C, c). Specimens
from the Territorio Federal Amazonas (Maroa,
Duida, Huachamacari, Yutaje, and Cerro de La
Neblina), as well as a specimen from Cerro Sa-
risarihama of the state of Bolivar (Cardona 380)
(Fig. 8A, a) may be referred to typical L. fruti-
cosa. However, two specimens from the summit
of the Chimanta Massif (Steyermark & Wurdack
879 and 1126) (Fig. 8D, d) from the state of
Bolivar, differ conspicuously from other Gua-
yanan material, as well as from specimens out-
side the Venezuelan Rind conforming to L.
fruticosa. TI
with larger leaf sc scars; thicker, larger leaves, en-
ervate beneath, broader at the subsessile base as
to appear nonpetiolate; rather conspicuous hairy
tufts along the leaf margins in the sinuses of the
marginal teeth; longer, stouter pedicels; and larg-
er sepals and petals. The isolation of the Chi-
mantá specimens from the upper slopes of this
sandstone mountain, well known for its endemic
flora, may help explain the evolutionary trend
here noted toward the segregation of morpho-
logical characters at variance from typical L. fru-
ticosa. A specific or subspecific category for this
population might be envisioned. However, in
view of the degree of variation shown by other
taxa of this species, varietal rank is here pro-
posed.

br anches

Laplacea fruticosa (Schrader) Kobuski var. chi-
mantae Steyermark, var. nov. TYPE: Vene-
zuela. Bolivar: Chimantá Massif, central
section, along west branch of headwaters of
Rio Tirica above Upper Falls, 2,090 m, 17
Feb. 1955, Julian A. Steyermark & John J.
Wurdack 879 (holotype, VEN; isotypes, F,
NY). Figure 8D, d.

VENEZUELAN GUAYANA FLORA—III

1987] STEYERMARK—
URE 8.

of a sinus (1).—B. Leaf of Maguire et al. 37153

Steyermark et al. 120086. c. portion of margin with detail of a sinus (3).

A-C. Laplacea fruticosa var. fruticosa.—A. Leaf of Cardona 380. a. portion of margin Mas detail
3. b. portion of margin with detail of a sinus (2). — C. Leaf of

— D. Laplacea fruticosa var. chimantae,

12
leaf of Steyermark & Wurdack 879. d. portion of margin with detail of a sinus showing tufts of hairs (4).

A var. fruticosa a sessilibus vel subsessilibus ap-
etiolum nihil angustatis basi 5—

i marginalibus p o ab ids
abus urs I. robustis 2—3 c s 2.5-3
crassis, sepalis petalisque majoribus .

Tree 5—20 m tall, the leafy branches 5-10 mm
thick, the leaf scars conspicuous, orbisilat, 2.5-
4 mm wide. Leaves sessile to abruptly
contracted into a petiolar portion 1 -21 mm long;
leaf blades thick-subcoriaceous, asymmetric, ob-
lanceolate-oblong, rounded at the slightly emar-
ginate apex, gradually narrowed to a broad base,
9.5-12.5 cm long, 3—4.8 cm wide, 5-7 mm wide
at base, the midrib bordered on one side by foliar
tissue 3-4 mm wide, on other side 2-3 mm wide,
the lower surface enervate, mainly glabrous but
with scattered, pale, minute, appressed, simple
hairs over a pustulate surface, inconspicuously
appressed-subserrulate from about % distance
from base to apex with dark acicular teeth 1 mm
long, these bearing at their base a tuft of silky
hairs 1 mm long (seen best from lower margin);

cocc1l

upper surface enervate, glabrous. Peduncle 2-3
cm long, 2.5-3 mm thick, glabrous. Flower bud
just before anthesis 2-2.5 cm long, 1.5-2.5 cm
wide. Sepals broadly suborbicular, the two out-
ermost 20 mm long, 15 mm wide, the others 17—
25 mm long, 13-22 mm wide, minutely gray
sericeous most of the length without, glabrous
within. Petals broadly oblong-obovate, broa
rounded and emarginate at apex, mm long,
15-22 mm wide, minutely gray sericeous without
in the central 5-9 mm portion, glabrous on the
marginal 4-5 mm, glabrous within. Filaments 5-
6 mm long, glabrous; anthers up (e oblong,
broadly rounded at the m long,
1.5 mm wide. Ovary barrel-shaped, 8 mm long,
5 mm wide, densely sericeous.

Paratype. VENEZUELA. BOLIVAR: Chimantá Massif,
Agparaman tepui, southeast-facing forested slopes be-
low escarpment, 1,880-1,955 m, 26 Feb. 1955, Stey-
ermark & Wurdack 1126 (F, NY, VEN).

The broad, nonpetiolate leaf bases of var. chi-

652

mantae differ markedly from the generally nar-
rowed, subpetiolar ones of typical Laplacea fru-
ticosa, in which the leaf base is conspicuously
narrowed toward the base into a subpetiolar por-
tion 5-8 mm long and 1-2 mm wide with the
midrib bordered on either side by a very narrow
strip of tissue only 0.5-1 mm wide on each side.
In L. fruticosa var. pulcherrima (Melchior) Ko-
buski, the leaf base is broader than in typical L.
fruticosa, but the apex is gradually obtusely nar-
rowed and is not broadly rounded as in var. chi-
mantae. The tufts of hairs at the base of the
marginal teeth are absent from most specimens
examined but are present to a lesser degree in
Dusen 15451 from Brazil and Maguire et al.
37259 from Cerro de La Neblina in Venezuela.
The enervate lower leaf surface may also occur
in other specimens of L. fruticosa, but more fre-
quently the lower surface shows venation.

The second species of Laplacea in the Gua-
yana Highland, L. pubescens Planchon & Linden
ex Triana & Planchon, is represented by two col-
lections, Maguire, Wurdack & Bunting 37272
from the summit of Cerro de La Neblina, and
Maguire, Cowan & Wurdack 30285 from the
summit of Cerro Huachamacari, both from Ter-
ritorio Federal Amazonas. Elsewhere, the species
ranges in the Andes from Venezuela south to
Peru and Bolivia. On Cerro de La Neblina it
varies slightly from typical L. pubescens of the
Andes, whereas on Cerro Huachamacari it has
developed more reduced leaves, and in both
Guayanan collections the lower surface of the
leaves is enervate, as contrasted with the more
evident venation shown in most ofthe specimens
of L. pubescens. Laplacea fruticosa, the more
common species, also occurs on Cerro Huacha-
macari, but at a lower altitude of 1,100 m of
forested talus slopes, as well as on Cerro de La
Neblina.

Laplacea pubescens Planchon & Linden ex Triana
Planchon

: aguire, R.
Cowan & J. Wurdack 30285 putida NY).

var. pubescens foliorum laminis minoribus rece-

.5-4.5(-9) cm E 2(-3) cm latis oe

atis; pedicellis 6-8 m iis i sepalis 5-11 m
pues is; petalis cuneiformi- obovat 6-21 mm longi
supra medium 14-15 mm latis.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Leaf blades symmetrical, oblanceolate, nar-
rowly obtuse at the apex, strongly narrowed at
the base, 2.5-4.5(-9) cm long, 1-2(-3) cm wide,
closely crenulate above the middle, enervate be-
neath. Pedicels 6-8 mm long. Sepals 5-11 mm
long. Petals cuneiform-obovate, 16-21 mm long,
14-15 mm wide above the middle.

Paratype. "VENEZUELA. TERRITORIO FEDERAL
AMAZONAS: Cerro de La Neblina, savanna edge, 3 km
N of Cumbre Camp, 1,800 m, 12 Jan. 1953, Maguire,
Wurdack & Bunting 37272 (NY).

Both collections have the leaf surface enervate
beneath. However, the paratype (Maguire et al.
37272) has the larger leaves typical of L. pubes-
cens var. pubescens. In both collections the in-
dument of the lower surface is sparse compared
with specimens from the Andean portion of the
range of the species.

LITERATURE CITED
BLAKE, S. F. 1918. New isting VL i Ed in
Venezuela and Curacao by Messrs. Curran
Haman. Contr. Gray Herb. n.s. 53: 39.
KoBuski, C. E. 1950. Studies in the Theaceae, XX.

Mese S Plantae

olu mbianae, Kew Bull. Misc. Sarma te 1926:

43-44.
Wawra, H. 86. Ternstroemiaceae. /n C. F. Mar-
tius, Flora Brasiliensis 12(1): 289.

THYMELEACEAE
DAPHNOPSIS

Daphnopsis guaiquinimae ipsun. sp. nov.
TYPE: Venezuela. Bolív : Cerro Guaiqui-
nima, INE la extensión
al del cerro en el sector noreste, la parte
pedregosa, abierta y plana con vegetación
baja, 6°0'N, 63?28'W, 1,650 m, 9 Apr. 1979,
Julian A. Steyermark, G. C. K. & E. Duns-
terville 117977 (holotype, VEN; isotype,
MO). Figure 9.

x 1.5 m; ramulis juvenilibus sericeo-strigosis;
laa laminis elliptico- oblongis vel lance A eg iod
lipticis apice p
tusis vel subacutis 3-6 cm longis 1-3 cm we supra
glabris subtus minute strigillosis costa media subtus
dense strigillosis venulis tertiariis pagina superiori ir-
regulariter contortis; pedunculis filiformibus 10-20 mm

oblongis subobtusis utrinque sericeo-hirtellis, orificio
glabro; staminibus 8, 2-seriatis; squamis pa 7-
8 subulatis glabris fere liberis 1.7-2 mm lon

1987]

Shrub 1.5 m tall; juvenile stems 2-3 mm diam.,
sericeous-strigose. Petiole 2-3 mm long, strigil-
lose, leaf blades subcoriaceous, gray-white be-
low, elliptic-oblong or lance-elliptic, mostly ob-
tuse to rounded at apex, rarely subacute, obtuse
to subacute at base, 3-6 cm long, 1-3 cm wide,
glabrous above, minutely strigillose below,
slightly more densely on the lower midrib; prin-
cipal lateral nerves 7-10 each side; tertiary ve-
nation above irregularly contorted and twisted,
below more regularly reticulate. Inflorescence
terminal or axillary, the peduncles filiform, 10—
20 mm long, 0.5 mm wide, sericeous-strigillose.
Calyx subcylindric-obovoid, 7 mm long, 2 mm
wide at and above middle, 1 mm wide at base,
antrorsely sericeous without, densely antrorsely
sericeous within in the lower 4 mm; lobes 4,
unequal, the outer ligulate-oblong, obtuse, 1.2-
1.5 x 0.5 mm, theinner 1 x 0.08 mm, sericeous-
hirtellous on both surfaces, the orifice glabrous.
Stamens 8 in 2 series; anthers sessile, suborbic-
ular-oblong, 0.6 mm long, the upper 4 opposite
the calyx lobes, the lower 4 alternate with the
calyx lobes; style terminal, 0.3 mm long; stigma
clavate, 0.3 mm long. Hypogynous scales 7-8,
subulate, glabrous, 1.7-2 mm long, cleft nearly
or all the way to the base, free or nearly so.

The type and only specimen of this species was
indicated by Nevling (1967) as a new species of
Daphnopsis related to D. longipedunculata Gilg
ex Domke, described originally from Mount Ro-
raima. she chief differences possessed by the idi

number Oo f1

(8 vs. 4 as in thé Roraima specimen), digne.

1-2 cm long contrasted with 2.5-9 cm on the
Roraima specimen, and the shorter aie Domke
(1935) stated that the umbels in D. longipedun-
culata were on axillary peduncles but added
(“atque terminali?"), apparently suspecting that
they may also be terminal, as is the case in the
Guaiquinima specimens. He also stated that sta-
minate as well as pistillate flowers occur. This
indicates that the specimen Ule 8739 from Ro-
raima studied by Domke was monoecious. In
the Guaiquinima specimens, there is also an in-
dication of monoecism, since the anthers are well
developed and suborbicular-oblong on some
flowers but poorly developed and linear-oblong
on others, even on the same plant.

The hypogynous scales in the Guaiquinima
plant are regularly disposed as eight filiform ap-
pendages. Although they appear to be free, as in
Funifera utilis Leandro, they are connected at
their very base by a slight membrane on the disk,

STEYERMARK— VENEZUELAN GUAYANA FLORA —III

653

which appears to be adnate to the actual base of
the calyx tube, thus showing the relationship with
Daphnopsis. Conversely, they could be inter-
preted as completely free as in the genus Funi-
fera. Nevling (1967) at first interpreted the style
to be lateral (eccentric) in Daphnopsis longipe-
dunculata, but later (pers. comm., 1986) con-
firmed my observation that the style is terminal.

Daphnopsis nevlingiana Steyermark, sp. nov.
TYPE: Venezuela. Bolivar: Cerro Sarisari-
ñama, Meseta de Jaua, summit, 4?41'40"N,
64?13'20"W, 1,400 m, 16-18 Feb. 1974, Ju-
lian A. Steyermark, V. Carreño E. & Charles
Brewer-Carias 109199 (holotype, VEN; iso-
type, ).

A ical longipedunculata Gilg ex Domke fo-
liorum paginis superio oribus venatione tertiaria irre-
gulariter contorta
calycis tubo tys practer infra medium sparse pier

po apice | Gain ti tube seu eie
cedit.

Slender tree 3 m tall with bark difficult to tear,
the young branches strigose. Petiole 3-4 mm long,
moderately strigose. Leaf blades elliptic-oblong
or ovate-oblong, rounded at apex, acute at base,

7.5 cm long, 2-4.5 cm wide; upper surface
mainly glabrous except strigose along the de-
pressed midrib; lower surface uniformly short-
strigose with the lower midrib more densely stri-
gose with longer hairs; lateral nerves 7-9 each
side, faint above, slender and slightly manifest
below; tertiary veins of upper surface irregularly
contorted, loosely and inconspicuously reticu-
late. Pistillate calyx dull red with whitish tips,
cylindric, 5 mm long, 4-lobed; lobes unequal,
puberulous in upper half, glabrous below, the
larger ones 1 mm long, 0.8 mm wide, pubescent
over a larger area with longer hairs toward the
base; the smaller lobes 0.7 mm long, 0.7 mm
wide, ovate-oblong; calyx tube glabrous within
except for small sericeous patches below the mid-
dle. Hypogynous scales 4, linear-lanceolate, 1.5—
2 mm long, 0.2 mm wide in lower part, dull
yellow, dull reddish near the tip. Pistil 4 mm
long; ovary ellipsoid, 2.4 mm long, densely se-
riceous; style 1.5 mm long, glabrous; stigma
subglobose, 0.5 mm long.

This species differs from Daphnopsis longi-
pedunculata Gilg ex Domke by having peculiarly
contorted, irregular tertiary venation on the up-
per leaf surface; the calyx reddish white with white
tips and with glabrous interior except for sparse

654

00 0.

a

UN OL /

7mm

FiGurE 9. Daphnopsis guaiquinimae.—
scales. — D. Peduncle with flowers. — E. Gro

pubescence below the middle; and hypogynous
scales thickened and dull yellow with reddish
near the obtuse tips.

SCHOENOBIBLUS

Schoenobiblus amazonicus Steyermark, sp. nov.
TYPE: Venezuela. Territorio Federal Ama-
zonas: 25 km S of Puerto Ayacucho, 5?30'N,
67?35'W, 5 Aug. 1967, Wessels- Boer 1953
(holotype, MER; isotype, MY). Figure 10.

Arbor 6 m; foliorum laminis oblanceolato-obovatis
apice acuminatis base acutis 36 cm longis s cm
latis subtus praesertim costa media necnon nerv
teralibus puberulis; petiolis subtus dense strigillosis

calycis tubo lineari 17-18 m
extus sericeo intus supra e retrorse sparsimque
strigoso ceterum glabro, lobis quattuor linearibus apice

ANNALS OF THE MISSOURI BOTANICAL GARDEN

— A. Flo a branch.—B.
oup of flower

[VoL. 74

Flower, interior view.—C. Hypogynous

rotundatis 6 mm longis 1.5 mm latis extus sericeo-

i š
lanuginosis intus villosis; stylo filiformi 18 mm longo

Tree 6 m tall; buds fusiform, subobtuse, 6.5
mm long, 2.5 mm wide. Petiole 6-9 mm long,
2.5-3 mm wide, strigillose beneath; leaf blades
oblanceolate-obovate, acutely acuminate at apex,
acute at base, 36 cm long, 12.5-14 cm wide,
glabrous above, moderately strigillose on midrib
and lateral nerves below, these sparsely puber-
ulous on leaf surface, with the midrib elevated,

r
ceous axes, one of them solitary, the other forking
into 2-3 shorter secondary axes 2-4 cm lon

Flowers 10-20-umbellate; pedicels 10-18 mm

1987]

0. popa E amazonicus. — A. Flowering branch. — B. Flower.

FIGURE 1
E Sent

long. Calyx tube linear, 17-18 mm long, 0.6-0.8
mm wide, short-sericeous without, retrorsely
sparsely strigose within in the upper half, gla-
brous in the lower half; lobes 4, spreading, rev-
olute at margins and at apex, linear, rounded
at apex, 6 mm long, 1.5 mm wide, sericeous-
lanuginose without, villosulous within. Stamens
4; anthers suborbicular or broadly oblong, 1.1
mm long, mm b

exserted, erect-ascending, 5-5.5 mm 1
brous. Style filiform, 18 mm long, glabrous, at-
taining the orifice; stigma ellipsoid, 1.2 mm long.

STEYERMARK — VENEZUELAN GUAYANA FLORA —III

655

AN

F DOM EP APP EE
Dr ER eee See nga Sart ere

—C. Stamen, dorsal view.—D.

The new taxon differs from Schoenobiblus
daphnoides Martius of Brazil in the much broad-
er leaves and less-branched inflorescence, while
from other Venezuelan material identified as S.
daphnoides it is differentiated by the longer, more
slender calyx tube, narrower calyx lobes with
shorter, gray sericeous pubescence, and much
longer peduncle

LITERATURE CITED

, W. 1935. Neue Arten und Varietäten der
Nun en Daphnopsis Mart. et Zucc. und Fu-

656 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 74

dala (Leandro ex) C. A. Mey. aus Mittel- und
merika. Notizbl. Bot. Gart. Berlin-Dahlem
12: 723-1 24.
MEISNER, C. F. Thymeleaceae. Jn C. F. Mar-
tius, Flora Brasiliensis 5(1): 65. t. 26 (1).

NEVLING, L. 1959. Revision ofthe genus Daphnopsis.
p 352.

67. n J. A. Steyermark &
B. i A (editors), eme of the Chimantá
Massif. II. Mem. New York Bot. Gard. 17: 454.
SYMPLOCACEAE
SYMPLOCOS

KEY TO THE SPECIES OF SYMPLOCOS OF THE VENEZUELAN GUAYANA

la. Ovary 4-celled; pubescence of Sd or iiis xk at least on the upper part of the

, petioles

and lower leaf surface, and/or on the low ulei
lb. Ovary 3- or 5-celled; sre eal absent or of abba or sericeous hairs 2
2a. Calyx lobes acute to acuminate at apex; style pubescent
3a. het E shorter than the petiole, sessile or nearly so, 2-3- hielo calyx lobes 1.8-2
m long; corolla 7 mm long; leaves minutely and distantly crenulate U... . acananensis
3b. ne equaling or longer than the petiole, shortly pedunculate, 5-8- flowered: calyx
.5-3 mm long; corolla 8-9 mm long; leaves lo to subentire 0. ibaa

2b. Calyx dies obtuse to rounded at apex; style glabrous
4a. Ova celled

5a. los glabrous beneath, acuminate at apex, acute at base; petiole glabrous throughout
UM
5b. Leaves sparsely pubescent beneath on the nerves, obtuse at apex, rounded or ue
base; ae strigillose adaxially 5S. V rid NS
4b. Ovary 3-cell 6
6a. Calyx is without, except for the ciliate lobes; filaments not connate, free or nearly
so to th

bes;

e below; leaves with at least the lower midrib appressed-pubescent;
petioles bey iren. Or seri
7a. Calyx lobes longer than the calyx tube; leaves rounded to obtuse at apex, rounded
or obtuse at cm lon . Jauensis

. Calyx lobes shorter than or equaling the calyx tube; leaves acuminate at apex, acute
at base, 7-10 cm long
8a. Fruit densely strigillose, 5 mm long; stamens 25-40; lower leaf n sparsely
appressed-pubescent to glabrescent; bracts suborbicular, rounded or obtuse; co-
rolla 3 mm lon $ yapacanensis

. Fruit glabrate, 6-8 mm long; stamens 20-25; lower leaf surface glabrous; bracts
acute; corolla 4-4.5 mm long S. neblinae

oo
c

Symplocos acananensis Steyermark, sp. nov.
TYPE: Venezuela. Bolivar: gallery forest bor-
dering savanna, vicinity of Guadequen, Rio
Acanan, affluent of Rio Carrao, west of Cer-
ro Las Hermanas, 5?56'N, 62°17'W, 470 m,
1-3, 20 May 1986, Julian A. Steyermark, R.
Liesner & B. Holst 131868 (holotype, VEN;
isotype, MO)

superne libris inferne in tubum 3.5 mm longum coa-
lescentibus glabris 1.5-3.5 mm longis; stylo 5 mm lon-
o in dimidio inferiore piloso.

Shrub 1.5 m tall with mainly glabrous branch-
es. Petioles 1.2-5 mm long, glabrous; leaf blades
chartaceous, elliptic-oblong, abruptly short-acu-
minate at apex, obtuse t

F 1.5 m, ramis plerumque glabris; foliis pe-
tiolatis, "petiolis 1.2-5 mm lo a í

tiis axillaribus sessilibus vel fere 2-3 floris; bracteis sub

lycis tubo cupulato 1.2-1.5 mm longo glabro, lobis 5
late lanceolatis acuminatis 2 mm longis 0.8-0.9 mm
latis praeter margines ciliatos glabris; corolla 7 mm
longo, tubo 2.5 mm longo, lobis 5 oblongis apice ro
tundatis 4.5 mm yobis staminibus ca. 20, ments

sides, appearing subentire but the margins
with microscopically obtuse serrulations 0. 1—0.2
mm long, 2-4 of these dispersed in 1 cm of mar-
gin; lateral nerves 4-5 each side, faintly anas-
tomosing with the tertiary veinlets 4-8 mm from
the margin, obsolete above, slightly manifest be-
low; tertiary veinlets slightly reticulate below,
scarcely or not manifest above. Inflorescence ax-
illary, 2-3-flowered, sessile or nearly so, shorter
than the petioles; flowers fasciculate, sessile.
Bracts subtending flowers lanceolate, subacute
1 x ] mm, pubescent without, strongly ciliate,

STEYERMARK

1987]

—VENEZUELAN GUAYANA FLORA —III

657

with caducous black glands. Corolla 7 mm long,
the tube 2.5 mm long, 1.5 mm wide; lobes 5,
oblong, rounded at summit, 4.5 mm long, 2.1
mm wide. Calyx tube cupulate, 1.2-1.5 mm long,
1.5-2 mm wide above, glabrous; sa lobes 5,
broadly lanceolate, acuminate, 2 mm long, 0.8-
0.9 mm wide, glabrous except for the ciliate mar-
gins. Corolla 7 mm long, the tube 2.5 mm long,
the 5 lobes oblong, rounded at the apex, 4.5 mm
long. Stamens about 20, ca. 4-seriate; filaments
free above, forming a tube 3.5 mm long, the
longer filaments 1.5-3.5 mm long, the shorter
ones | mm long, all glabrous; anthers broadly
oblong, 0.2 mm long. Style 5 mm long, pilose in
the lower half. Fruit not seen.

This species is closely related to the group of
Symplocos schomburgkii Klotzsch ex Schomb.,
S. guianensis (Aublet) Giirke, and S. pilosiuscula
Brand. It is distinguished from S. guianensis by
the glabrous branches, from S. schomburgkii by
the sessile or nearly sessile inflorescence which
is fewer-flowered and shorter than the petiole,
the shorter corolla and calyx lobes, and minutely
crenulate leaf margins. The new species differs
from S. pilosiuscula in the acute to acuminate
calyx lobes.

RUBIACEAE

Chomelia glabricalyx Steyermark, sp. nov. TYPE:
Venezuela. Bolivar: primary forest and river
edge, Rio Caura, 5-10 km S of Las Pavas
(Salto Para), 6°12'N, 64?28'W, 240 m, May

— Gilberto Morillo 6811 (holotype,
N).

x 1 m, ramis glabris spinosis; foliorum laminis
ovato- vel lanceolato-ellipticis apice acutis vel sub-

rum atque int
glabris; inflorescentia. 6- flora, pedunculo filiformi 14-
E Poir m longo glabr o;

4. s ° 1

axils of the lateral nerves and sometimes with
few sparse ciliate hairs at the base; lateral nerves
5-6 each side, scarcely evident above, slightly
impressed below. Stipules subulate, projecting
0.5 mm long. Inflorescence 6-flowered; peduncle
filiform, 14-20 mm long, glabrous; flowers not
involucrate at base, sessile, or in fruit 0.5-1 mm
pedicellate. Calyx and hypanthium glabrous; hy-
panthium tubular, 1.5 mm long, | mm wide,
glabrous; calyx E unequal, ligulate- oblong,
obtuse, 0.5-1 m wide, gla
brous. Corolla bei, 4 mm long, 1.2 mm
wide; tube 2.5-3 mm long, glabrous without; lobes

oblong, obtuse, 1.5 mm long, glabrous below,
sparsely iis mas upward. Fruit narrow-cylin-
dric, 10 m .5 mm wide, 0.5-1 mm long
eae e rq

This taxon is related to the recently described
Chomelia delascioi Steyerm. and C. stergiosii
Steyerm e present taxon differs from C. de-
lascioi in the larger, barbellate leaves, shorter
corollas, and more numerously flowered inflo-
rescence, and from C. stergoisii in the shorter
corolla and calyx lobes and the completely gla-
brous hypanthium and calyx lobes.

ae huberi Steyermark, sp. nov. TYPE:
Venezuela. Bolívar: Distrito Roscio, Serra-
nia es Caco, 25 km NW of San Ignacio de
Yuruani, 2.5 km ESE of Wanaru, 5?12'N,
61?15'W, 1,150-1,200 m, 1 Mar. 1984, Otto
Huber 9123 (holotype, VEN; isotype, MO).

eta acel

Herba radicans, caulibus fepenusus 0. ie mm at
s 1.2 mm lon-

gis sparsim | strigillosis; foliis petiolatis, ie sare
mm longis dense strigosis; foliorum laminis ellipticis
anguste oblongo-ovatis vel elliptico-ovatis apice $ed
acutis vel rsen basi acutis vel obtusis 2-2.8 c
longis 1-1.5 cm latis utrinque breviter strigillosis, ner-
vis gi d utroque latere 5-7; inflorescentia 1 -flor
dunculo sub anthesi 1.5 mm longo sub fructu 4.5
mm longo dense strigilloso; hypanthio dense strigoso;

calycis ] ceolatis acutis 1.8—2.5 mm

5-1
it wa thioque glabro, semen 1.5 mm longo 1 mm
lato, lobis inaequalibus ligulato-oblongis obtusis 0.5-
1 mm longis glabris; corolla 4 mm longa tubo extus
glabro, lobis subtus glabris apicem versus sparsim stri-
gillosis; fructu anguste cylindrico 10 mm longo 3.5 mm
lato 0.5-1 mm pedicellato.

rub ] m tall; branches glabrous, spinose.

o
elliptic, acute to subacuminate at apex, cuneately
acute at base, 4.5—9.5 cm long, 1.5-4 cm wide,
glabrous both sides except for barbellate lower

longis 0.4-0.5 mm latis ieee sparsim strigillosis ad

gillosa; bacca (immatura) in statu vivo 1-1.5 cm diam.,
in sicco 5 mm longo 4.5 mm lato modice strigilloso.

Creeping herb with moderately strigillose stems
0.8-1 mm diam.; stipules linear-setaceous, 1.2
mm long, sparsely strigillose. Petioles ofthe larg-
er leaves 10-13 mm long, of the smaller ones 3-
5 mm long, densely strigose; leaf blades elliptic,
narrowly oblong-ovate, or elliptic-ovate, sub-
acute to subobtuse at apex, acute to obtuse at

658

eidec more

base, moderat
abundantly on nerves d Beneath: 2-2.8 cm long,
1-1.5 cm wide; lateral nerves 5-7 each side, faint
on lower side. Flowers solitary, the peduncle 1.5
mm long in anthesis, 4-5 mm long in fruit, densely
strigillose. Calyx 3.5 mm long, the tube subglo-
bose, 1.3 x 1.3 mm, densely strigose, the lobes
unequal, lanceolate, acute, 1.8—2.5 mm long, 0.4—
0.5 mm wide, sparsely strigillose on both sides,
at base within setose with erect hairs, each sinus
at the base furnished with 1 squamella. Corolla
white, infundibuliform, 4.8-5 mm long, sparsely
strigillose without, the tube 2.8-3 mm long, the
lobes broadly lanceolate, 2 mm lon 8 mm

wide. Stamens not exserted, attached Uh distance
of the tube length; anthers oblong, obtuse, 1.1
mm long. Style 3 mm long, glabrous. Immature
fruit urceolate-globose, 5 mm long, 4.5 mm wide
(dried), 1-1.5 cm diam. (living).

The present taxon approaches the endemic Ja-
maican species, Coccocypselum pseudotontanea
Griseb. Both possess white corollas and small
leaf blades, but the Venezuelan species differs in
having shorter stipules, shorter calyx lobes and
corolla, shorter and 1-flowered peduncles, one
squamella at the base of each side of the inner
part of the calyx lobe, larger fruits with appressed
pubescence, and the freely rooting habit.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Perama dichotoma Poepp. & Endl. var. mono-
cephala Steyerm., var. nov. TYPE: Venezue-
la. Territorio Federal Amazonas: Departa-
mento Atabapo, Cerro Huachamacari, ladera
sur-este 175-180 m, 12-14 Oct. 1984, Fran-
cisco Delascio 12358; hierba pilosa; hojas
arrosadas verde oscura por el haz, verde pal-
ido por el envés; pedünculo floral erecto,
violaceo-morado en su base; flores blancas
con el apice de los pétalos lila) (holotype,
VEN; isotype, MO).

A P. dichotoma var. dichotoma et var. scaposa c
libus simplicibus ipei pa recedit; capitulis so so-
litariis depressis 2-4 mm altis 5-6 mm latis.

Herb, 4-5 cm tall. Leaves rosulate, ovate or
elliptic, 7-10 mm long, 3-6 mm wide, densely
pilose above, pilose on main nerves below. Heads
solitary, ca. 10-flowered, depressed-hemispher-
ic, 2-4 mm high, 5-6 mm wide. Corolla laven-
der.

Perama dichotoma has the inflorescence usu-
ally consisting of several to numerous heads on
filiform forking branches. The new variety has
only a solitary head of flowers terminating a sim-
ple peduncle.

NOTES
NEW COMBINATIONS IN CENTRAL AMERICAN ARACEAE

Work in recent years with the Araceae of Cen-
tral America necessitates new combinations in-
volving species in the genera Monstera and
Philodendron which were erroneously published
as belonging to Rhodospatha and Syngonium re-
spectively.

Monstera costaricensis (Engl. & Krause) Croat
rayum, comb. nov. Rhodospatha cos-
taricensis Engl. & Krause, Das Pflanzenr. 4.
23B (Heft 37): 95. 1908. TYPE: Costa Rica.
Limon: Ferme de Boston, Atlantic wa-
tershed, 30 m elev. 10?01'N, 83?15'30"W,
Tonduz 14628 (holotype, B).

We had concluded that this distinctive species
of Monstera, occurring in primary forest at Finca
La Selva, was undescribed and had provisionally
assigned it the name “Monstera undulata.” It
now turns out that this species was previously
described by Engler & Krause (1908) in the genus
Rhodospatha; the transfer to Monstera is here-
with effected. Monstera costaricensis 1s distin-
guished from other species by its strongly un-
dulate petiole sheaths, tuberculate petiole bases
and pistils shaped like bowling pins. The lamina
may be either entire (as on the holotype) or per-
forate. Monstera costaricensis is now known in
the Atlantic lowlands of Costa Rica from La Sel-
va to Punta Mona. It probably occurs in Nica-
ragua and Panama as well.

Monstera dissecta (Schott) Croat & Grayum,
comb. nov. Tornelia dissecta Schott, Oes-
terr. Bot. Z. 8: 179. 1858. TYPE: Costa Rica.
Cartago: Volcán de Turrialba, Wendland 500
(holotype, GOET).

Although the name Monstera dilacerata is well
known and firmly entrenched in both taxonomic
and horticultural aroid literature, it has been ap-
plied in a highly indiscriminate manner and, in-
deed, may be impossible to confidently assign to
any real biological entity. As described by Mad-
ison (1977), “the type of M. dilacerata is an in-
ferior specimen consisting of a few leaves from
an immature cultivated plant of uncertain geo-
graphic origin." Such a specimen, even if it could
be located, would be virtually impossible to iden-
tify with any biological species in a genus as phe-

ANN. MissouRi Bor. GARD. 74: 659—660. 1987.

notypically plastic as Monstera; but the type of
Monstera dilacerata is apparently not extant,
having been destroyed in the Berlin herbarium
during World War II, and we know it only from
photos.

Madison (1977), determined to salvage this
well-known name by any means, adopted the
concept of Engler & Krause (1908). Although the
latter authors applied the name M. dilacerata
rather consistently (i.e., to the species here treat-
ed as M. dissecta), their concept would appear
to be completely irrelevant; what matters is the
interpretation of the type material. It could be
argued that Engler & Krause might have seen
living (perhaps even fertile) material from the
original collection, but they give no such indi-
cation and the point is moot.

Madison (1977) himself made matters signif-
icantly worse by applying the name Monstera
dilacerata with reckless abandon. Based on his
specimen citations, herbarium annotations, and
the use of his keys, Madison has employed the
name M. dilacerata for no fewer than four dis-
tinctly different species in Costa Rica alone (Croat
& Grayum, unpubl. data). Given the expansive
geographic distribution of Monstera dilacerata
sensu Madison (from Guatemala to Amazonian
Brazil), the application of the latter name to any
one of the four Costa Rican species would have
to be entirely arbitrary. Indeed, the type might
just as well have come from Brazil, and may
represent a different species altogether.

The only alternative seems to be the relegation
of the much-abused name Monstera dilacerata
Schott to the limbo of nomen dubium status,
where it will probably lie forever unless someone
can come up with a way (epidermal anatomy?)
to unequivocally interpret the type specimen, in
the unlikely event that it ever turns up. Three of
the four Costa Rican species in the Monstera
dilacerata complex have apparently never been
described (they will be described elsewhere). The
fourth clearly corresponds to Tornelia dissecta
Schott, the type of which is extant at GOET and
has been studied by the present authors (Madi-
son was apparently unaware of its existence). The
specimen is a good one and agrees in all details
with a species we know well from throughout
Costa Rica (but mostly from the Atlantic slope)

660

at elevations of ca. 100-1,800 m. Contemporary
collections from the general vicinity of the type
locality include: Grayum & Sleeper 3304 (CR,
MO); Grayum et al. 3490 (CR, MO); and Gra-
yum & Hammel 5739 (MO). The appropriate
new combination in Monstera is made above.

Philodendron rothschuhianum (Engl. & Krause)
Grayum, comb. nov. Syngonium
rothschuhianum Engl. & Krause, Das Pflan-
zenr. 4. 23E (Heft 71): 124. 1920. TYPE: Nic-
aragua. Matagalpa: Matagalpa, 1,000 m,
Rothschuh 229 (holotype, B).

This species has long been excluded from Syn-
gonium, perhaps having been first so treated by
Birdsey (1955) in an unpublished thesis. Croat
(1981) erroneously placed the species with P. an-
isotomum Schott. There is no longer any doubt
that it is a distinct species, distinguished from
the latter by having the posterior lobes more

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

broadly confluent with the anterior lobe and by
details of the inflorescence. The species ranges
from Nicaragua to western Panama (Veraguas)
from sea level to 1,000 m

LITERATURE CITED

BinpsEv, M. 1955. The Morphology and Taxonomy
of the Genus Syngonium (Araceae). Ph.D. Dis-
sertation. University of California, Berkeley, Cal-
ifornia.

Croat, T. B. 1981. A revision of Syngonium (Ara-
ceae). Ann. Missouri Bot. Gar : 649.

ENGLER, A. & K. KRAUSE. 1908. Ara ceae— Monst ster-

oideae. Pflanzenreich IV. 23B (Heft 37): 4-139.

MADISON, M. 1977. A revision of Monstera (Ara-

ceae). Contr. Gray Herb. Harv. Univ. 207: 3-100.

— Thomas B. Croat, Missouri Botanical Garden,
P.O. Box 299, St. Louis, Missouri 63166, U.S.A.;
and Michael H. Grayum, Missouri Botanical
Garden, P.O. Box 299, St. Louis, Missouri 63166,
U.S.A.

TWO CUBAN NOVELTIES IN XYRIS

Among the X: de kindly loaned by the curators
of F, NY, MO, US for a study of North
American Sechs were materials of two
species of undescribed Yyris, both from the Isle
of Pines. From descriptions of the white sand
savannas of that island, once so frequently vis-
ited by such collectors as N. L. Britton, E. P
Killip, and P. Wilson, it is easily seen that it 1s
ideal Xyris habitat, having nearly all of the Xyris
species endemic to Cuba as well as most of those
found both in southern Florida and along the
Mosquito Coast and Belize. It is seen that west-
ern Cuba is perhaps the most significant floristic
connector between the Florida peninsula and
esoamerica for not only Xyridaceae but for
Eriocaulaceae and many genera of sedges as well.
Thus these two novelties are presented as but a
small part of what promises to be a considerable
updating as the new Flora de Cuba progresses
and as Cuban-based botanists continue the work.

Xyris curassavica Kral & Urquiola, sp. nov. TYPE:
Cuba. Isle of Pines: vicinity of Los Indios,
13 Feb. 1916, N. L. Britton, E. G. Britton &
P. Wilson 14219 (holotype, NY; isotypes,
CM, F, MO, US). Figure 1

Planta perennis, densicespitosa, laxa, 1.5-4 dm alta,
caulibus contractis, radicibus gracilibus fibrosis. Folia
bud erecta vel leviter expansa, torta, (5-)10-20(-25)

vaginae integrae, laminis 4—5-plo breviores,

PE a ë Te a vel roseolae, marginibus in laminas
paume m ad apicem pii m (us-
lam-

inae wei RE anguste dete pallide luteo-
virides, 0.5-2 mm latae, ad apicem subulatae, raro
paucitrichomatiferae, marginibus integris leviter in-
crassatis, raro distante papillosis, paginis glabris, lon-
gitudine valde 1—3-nervatis. Vaginae scaporum prox-
imale bras glandaceae, folia principalia superantia,

pallide
3, fertilibus E bractea

Siri bpt ovatae,
4.5-5 mm longae, anguste rotundatae arginem
tenues, integri, tum erosae. Sevals teas libera, sub-
aequilatera, elliptica, curvata, 4—4.5 ga; [v
rinalis et firm 1 apice abrido-papil-
losa. Laminae petalorum late obovatae, luteolae, ca
.5 mm longae, apice anguste rotundatae, erosae. An
therae ee ca. 1.5 m pon vald
bifidae et sagittatae, filamentis ca. 0.5 m ngis.

ANN. ait Bor. GARD. 74: DEUS 1987.

Staminodia bibrachiata, brachiis apice sparsim peni-
cillatis. Capsula SOMME ca. 3m onga; placenta
ma "n nalis. Sem elc rtocylindra-

aie nga, succinea, , trans lucida, apicu-
ha dongiudine e valde et irregulariter anastomosocos-

Densely cespitose, soft-based perennial 1.5—4
dm high, the stems contracted, the roots slender-
fibrous. Leaves mostly erect to slightly spreading,
twisted, (5-)10-20(-25) cm long; sheaths '4-!5
as long as blades, entire, a lustrous tan or red-
brown, tapering gradually from the clasping base,
multicostate and carinate to the blade, there with
a scarious, rounded ligule to 2 mm long; blades
strongly flattened, narrowly linear, pale yellow-
green, 0.5- m wide, apically subulate and
sometimes with a apa coarse trichomes, the edges
entire and slightly thickened, sometimes re-
motely papillate, the surfaces longitudinally
strongly 1—3-nerved, often papillate or rugulose-
papillate proximally. Scape sheaths lustrous tan
proximally, open and short-bladed distally,
shorter than principal leaves. Scapes linear-fili-
form, flexuous, twisted, distally subterete or oval
in cross section, 1—3(—4)-costate, ca. 0.5 mm thick,
the costas smooth or distantly scabro-papillose.
Spikes broadly ovoid to subglobose, 5-7(-10) mm
long, the bracts few, loosely spirally imbricate,
lustrous pale red-brown with distinct or indis-
tinct ovate pale green (when young) dorsal areas,
the backs convex, toward apex low-carinate; ster-
ile bracts 2-3, smaller than the fertile bracts and

the broad keel firm, a
base to acute apex. Petal blades broadly obovate,
yellow, ca. 4.5 mm long, the narrowly rounde

apex erose. Anthers lance-oblong, ca. 1.5 mm
long, deeply bifid and sagittate, on stout fila-
ments ca. 0.5 mm long. Staminodia bibra-
chiate, the flat, narrow branches sparsely pen-
llac, Capsule ellipsoid, ca. 3 mm long, the
placentation marginal with placentas extending

short-cylindric, 0.5-0.7 mm long, apicula e
red-brown, translucent, longitudinally distinctly
but irregularly anastomosing-ribbe

Frequent in sand savannas, Isle of Pines. Cuba.

FIGURE l. Xyris curassavica (Alain & Killip 2194).—a. Habit sketch. — b. Leaf apex. —c. Sector of mid-blade. —
d. Leaf blade-sheath junction.—e. Apex of leaf sheath.—f. Leaf base.—g. Spike.—h. Fertile bract.—i. Lateral
sepal.—j. Petal blade and stamen.—k. Staminode.—1. Stylar apex.—m. Capsule, one valve removed, showing
placentation.—n. Seed.

662

1987]

iy yum

FIGURE 2. Xyris paleacea (Leon & Victorin 17823).—a. Habit sketch.—b. Lea
f. Spike. —g. Fertile bract.

mid-blade.—d. Leaf blade-sheath junction. —e. Leaf. —
valve with placenta. —j. Seed.

es aie specimens examined. CUBA. ISLE OF
PINES ios, white sand sabanas, 27 Dec. 1951,
Alain & Crus 2194 (US); Los Indios, pir rate 4 Feb.

22 Feb. 1953, Killip 42873 (CAS, F, GH, NY, U, US).

NOTES

f apex.—c. Sector of leaf at
—h. Cen sepal.—i. Capsule

This species most resembles X. bicarinata
Griseb., another Cuban endemic, in general hab-
it and somewhat in leaf but has thinner, paler
leaf bases; flatter leaf blades; more slender scapes;
and broader spikes with thinner fertile bracts less

664

tending to spread and recurve. The aspect is less
robust and smoother.

Xyris paleacea Kral & Urquiola, sp. nov. TYPE:
Cuba. ixi del Río: Laguna de Sta. Maria,
W of Sn. Luis, May 1940, H. Leon & M.
Victorin 1 mE (hope US). Figure 2.

Planta solitaria vel parum LR annua vel pe-
rennis,usque ad 3 dm alta, caulibus contractis, radi-
ld gracilibus fibrosis. Folia leviter ie expan-

, 4-6 cm longa; vaginae in tegrae, laminis 2-3-plo
dug carinatae, lampro-ferrugineae, marginibus

ientibus, aut eligulatis; lam-
inae compressae, mu ies eae 0.5-1.5 mm latae, ad
apicem anguste acutae, leviter incrassatae, marginibus
pallidis incrassatis, papillatis vel mein Twana

s. Vaginae sca-
porum brunneolae, nitidae, klani inio folia prin-
cipalia superantes, brevilaminae. Scapi filiformes, ap-

icem versus leviter compressi, ca. 0.5 mm lati, distincte
bicostales, costis pallidis glabris aut papillosis. Spicae

rtilibus breviores; bracteae fer-

m longae, apicem versus

anguste c piste, valde carinatae. Sepala

hteralia libera, ' Subaequilatera, pallide fusca, valde cur-
-5

cinea, apiculata, translucida, longitudine valde et ir-
regulariter anastomoso-costata

Solitary or in small tufts, annual or short-lived
perennial to 3 dm high, the stems contracted, the
roots filiform-fibrous. Leaves flabellately spread-
ing-ascending, 4—6 cm long; sheaths entire, ^-^
as long as the blades, carinate, lustrous red-brown,
narrowing gradually from the dilated, clasping
base to the blade, there either with a short, scar-
ious, rounded ligule or merging with blade base;
blades green-brown or maroon, flattened, slightly
twisted, 0.5-1.5 mm wide, the apex incurved,

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

narrowly acute, slightly thickened, papillate or
tuberculate-edged, the margins pale, incrassate,
smooth or papillose, the surfaces sm on-
gitudinally striate-nerved. Scape sheaths shorter
than principal leaves, at base tubular, lustrous
brown, multicostate, opening distally, short-
bladed. Scapes filiform, distally slightly com-
pressed, ca. m wide, distinctly bicostate,
the costas pale, smooth or papillate. Spikes lance-
ovoid, ca. 1 cm long, acute, of a few, loosely
spirally imbricate, red-brown, lacerate bracts with
pale green, papillose, subapical, lanceolate dorsal
areas; sterile bracts usually 2, oblong, strongly
carinate, 3.5-4 mm long; fertile bracts lance-
ovate, 5-6 mm long, toward the narrow, sub-
cucullate tip strongly carinate, the borders thin,
becoming strongly lacerate. Lateral sepals 4.5-5
mm long, free, subequilateral, strongly curved,
acute, the thick, narrow keel scabrid from near
base to tip. Flowers and stamens not seen. Cap-
sule ellipsoid, ca. 3 mm long, the placentation
marginal, the placentas extending from base to
near tip. Seeds ellipsoid, ca. 0.6 mm long, amber,
apiculate, longitudinally strongly but irregularly
anastomosing-ribbed.

Endemic to pine savanna, Pinar del Río, Cuba.

This species, thus far known only from the type
specimen, appears allied to X. brevifolia Michaux
and X. flabelliformis Chapman and is distin-
guished from them by a combination of (often)
longer, pale-incrassate-bordered leaves; overall
narrower spike outline with narrower, lacerate-
bordered fertile bracts; and slightly longer and
differently sculptured seeds.

— Robert Kral, Department of General Biology,
Vanderbilt x 1705, Station B,

do Urquiola, Sta. final, Edif. #52, Apto. C-4, Rpto.
Hermanos Cruz, Pinar del Río, Cuba.

A NEW “VIORNA” CLEMATIS FROM NORTHERN ALABAMA

In 1983, while examining specimens from the
Huntsville, Alabama, area donated to VDB by
the collector, Mr. James D. Morefield, I was par-
ticularly interested in a distinctive ““Viorna” that
he had collected previously from the lower west-
ern slopes of Round Top Mountain (summit el-
evation ca. 1,600 feet). This mountain is a dis-
tinctive westward lobe of Huntsville Mountain
and is south-southwest of Monte Sano Mountain
in the same trend, the whole massif a part of the
Cumberland Plateau. The Viornae subsection of
Clematis is distinguished primarily by the thick-
ened, bevelled-edged sepals which in the live
flower form (usually) an urceolate design, the
flowers themselves being arranged singly or in
few-flowered cymes in leaf axils with the primary
peduncle short to elongated but always having
leafy bracts. The subsection is confined to North
America and centers in the southeastern United
States; it is notable for its narrow endemics.

I visited the locality and, thanks to the accu-
racy of Morefield’s label information, was able
to locate the population quickly and to find an
abundance of plants in flower and early fruit.
Further exploration on the same mountain trend
resulted in discovery of yet another thriving pop-
ulation of this new Clematis. Named in honor
of its perceptive first collector, it is described as
follows:

Clematis morefieldii Kral, sp. nov. TYPE: United
States. Alabama: Madison Co., SE Hunts-
ville, along upslope side of eastward un-
paved extension of Deborah Avenue, 0.65
mi. SSW of Round Top Mtn., limerocky face
of slope, clay soil, in Cotinus-Quercus. Vines

—5 m, sprawling on boulders or shrubs and
forest reproduction; calyx greenish with rose
tints, 17 June 1983, R. Kral 70176 (fruiting
material from same locality, 27 June 1986,
R. Kral with J. R. Carter 73540) (holotype,
MO; isotypes, ALU, AUA, BM, CM, DOV,
DUR, EKY, F, FLAS, FSU, GA, GH, ILL,
ISC, K, LAF, MICH, MISSA, MO, NCU,
NY, OS, PAC, RSA, SMU, TENN, TEX,
UC, US, VDB, VSC, WAT, WILLI). Figures
L. 2

Planta perennis, scandens, usque ad 5 m E cir-

rhos efferens. er valde ides subteretes, 2-3 mm

crassi, valde cos obrunnei, parce vel jc albo-

villosi vel pilosi de araneosi. Foliola principalia re-

ANN. Missouni Bor. GARD. 74: 665-669. 1987.

ota, expansa, imparipinnata, usque ad 2 dm longa,
rhachidi breviter pilosa vel villosa, flexuosa; foliola

mucronata. tenuia, integra vel bi- vel-triloba, elon
pilosis, 4-15 mm longis; pagina er L^ a fla-
vovirens; pagina T sericea aut pilos
illares, solitares aut pau in cyma d

v
expansi mm var. ad — bibracteolis. Se-

oy

um, acuminatum, compre r ma
gine Tedania pons 30:35 mm longo, tmb
moso.

Perennial scandent vine to 5 m long. Stems
flexuous, copiously villous and/or arachnoid with
white hairs. Principal leaves imparipinnate, to 2
dm long, spreading, the rachis base shorter than
the lowest leaflets, the rachis axis flexuous, pi-
losulous or villous; leaflets paired, 9-11, spread-
ing or erect, reduced distally on rachis, the upper
1-3 forming tendrils, the lowest broadly to nar-
rowly ovate, 5-10 cm long, acute to acuminate,
mucronulate, thin, entire to 2- or 3-lobate, on
pilose petiolules 4-15 mm long; upper surface
smooth, yellow-green; lower surface sericeous or
pilose. Flowers axillary, solitary or (more often)
1—3(—5) in sessile cymes, the peduncles at anthe-
sis densely white-villous, erect or spreading, 15-
25 mm long, with 2 bracteoles at base. Sepals
oblong-lanceolate, 20-25 mm long, erect, the tips
acuminate, with narrow white borders, slightly
spreading to short-reflexed, the backs pink or
pale green-and-red, albosericeous, the edges thick,
white-tomentulose, the inner surface smooth,
longitudinally inconspicuously few-nerved. Sta-
mens linear, 12-20 mm long, the filaments flat-
tened, pilose from middle to apex, the anthers
including apiculus 3—3.5 mm long, pilose. Fruit

omboidal-ovate, 7-9 mm long, acumi-
nate, compressed, marginally thickened, seri-
ceous, the style 30-35 mm long, with a brown,
plumose coma.

Additional specimens examined. UNITED STATES.
ALABAMA: Madison Co., SE Huntsville, along upslope

ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 74

666

vine.—c. Flow-

FIGURE |.

Clematis morefieldii. —a. Habit sketch, lower node.—b. Sketch of node from mid

ering node, largest leaves removed. —d. Sector of mid-stem. (Drawn from Kral 70176.)

1987] NOTES 667

ane
Ws

M
SS

FiGuRE 2. Clematis morefieldii. —a. Upper flowering node.—b. Flower.—c. Dorsal (left) and ventral (right)
sides of sepal.—d. Three stamens. —e. Anther, enlarged. —f. Carpel.—g. Fruit with long persistent style. (Drawn
from Kral 73540.)

668

side of eastward dirt extension of Deborah Avenue,
0.65 mi. SSW of Round T tn., margin of mixed
woods on rocky limestone slope, locally common, elev
920 ft., 31 May 1982, J. D. Morefield 629 [JDM (More-
field Herbarium), VDB]; vine of loam pockets in ju-
niper-Cotinus-mixed A rather dry area, up-
per end of Drake Avenue, W face of mountian at E
side of Huntsville; calyx pinkish, 17 June 1983, R. Kral
70216 (VDB, and to be distributed)

This viorna, so far found only in the limestone
uplands around Huntsville, Alabama, is closely
related to the variable Clematis viorna L. Dr.
Carl S. Keener (1975: 45), an pee on w
genus, considers such variants
Small, C. flaccida Small, and C. p an
Erickson as part of that species, an opinion sup-
ported by the biosystematic study of the complex
by Dr. W. M. Dennis (1976). However, Keener
suggested (loc. cit.), “Nevertheless, critical pop-
ulation studies of C. viorna, especially in central
Tennessee and adjacent Kentucky, would be in-
structive and might reveal more precise taxo-
nomically definable topogamodemes."

It seems that this species, nested well inside
an area of much of the diversity in C. viorna, is
indeed distinctive. Of particular interest is the
character combination of villous and arachnoid
tomentum on the shoot, velvety lower leaflet sur-
face, and stouter, usually shorter, peduncles which
bear bracts only at the very base (this of partic-
ular significance as a character state in the vior-
nas). The inflorescence is curious,the lower flow-
ering nodes often producing sessile dichasia in
the axils, or the bracteoles of these dichasia sup-
porting more flower buds in their axils. The up-
per nodes are unifoliar and frequently display a
wandlike length of progressively reduced leaves
with the flowers paired in the axils, while the
ultimate and penultimate nodes are often single-
flowered; thus the o ct is one ofa narrow
raceme with well-spaced nodes.

The plants are found in consistent habitat,
namely the limestone measures that outcrop be-
low the sandstone caprock of the Huntsville
Mountain chain ofthe Cumberland Plateau which
borders the Highland Rim on the east side of
Huntsville. The vines root in a basic clay-loam
amongst boulders of massive limestone, often
sprawling over the rock itself. The forest type is
an open to dense mixture of Juniperus with hard-
woods typical of basic substrate, the more dom-
inant being Carya carolinae-septentrionalis, C.
ovata, Quercus shumardii, Q. muehlenbergii, Q.
alba, Q. stellata, Ulmus, Celtis, Acer saccharum,
and Fraxinus americana. However, the most no-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

table indicator is the Smoketree, Cotinus obova-
tus. The shrub layer is marked particularly by
Rhus aromatica, Symphoricarpos, Hypericum
Jrondosum, Forestiera ligustrina, and Viburnum
rufidulum. Associated and prevalent herbaceous
markers include Anemone virginiana, Delphin-
ium carolini m, Euphorbia corollata (var.),

aptera, Thaspium pinnatifidum, Spigelia
ARE Scutellaria ovata, Pycnanthemum
incanum (complex), Blephilia hirsuta, Dasysto-
ma macrophylla, Hedyotis purpurea, and many
composites — particularly Aster, Solidago, and the
handsome white-liguled Polymnia canadensis
var. ligulata. Most notable here are, however,
Silphium brachiatum and Solidago auriculata,
these always in close association with Cotinus
and the new Clematis. The only other Clematis
so far found in the same area is C. virginiana,
which is part of a different section. Plants of this
entire assemblage grow well on the clays and
cherts derived from surrounding limestones and
are markedly and abruptly different from the as-
sociations of the overlying shales and sandstones
of this region.

Further field exploration of the same geologic
and floristic system is being conducted with an
eye toward occurrence of the Cotinus associa-
tion, this now known to extend north into Frank-
lin and Marion counties in Tennessee and south
into Morgan County in Alabama. Where this tree
occurs there is likely habitat for Clematis more-
fieldii.

Mr. J. D. Morefield, a careful and perceptive
student of botany, is gratefully acknowledged.
During the few years of his adult residence in
Huntsville he developed an excellent personal
herbarium which adds much to our information
about the flora of northern Alabama, his Clem-
atis being but one of many interesting discov-
eries. Today he is a graduate resident at the Ran-
cho Santa Ana Botanic Garden and is involved
in revisional and floristic studies centering in the
Basin and Range physiography. The Clematis is
therefore named in his honor and as a reminder
that he is missed back east.

LITERATURE CITED
PEDE W.M. 1976. A Biosystematic Study of Clem-
s, Sect. Viorna, Subsection Viornae. Ph.D. Dis-
wnatio on. The University of Tennessee, Knoxville,

ssee
E R. O. 1943. Taxonomy of Clematis sec-
tion Viorna. Ann. Missouri Bot. Gard. 30: 1-60.

1987] NOTES 669

KEENER, C. S. 1975. Studies in the Ranunculaceae of — Robert Kral, Department of Biology, Vander-
the southeastern United States III. Clematis L. pilt University, Nashville, Tennessee 37235
Sida 6: 33-47. , : i

SMALL, J. K. 1933. Viorna Reichenb. Pp. 526-528 in U
Manual of the Southeastern Flora. Chapel Hill

ition.

TWO NEW SPECIES OF CORDIA (BORAGINACEAE)
FROM CENTRAL AMERICA

Increased collecting efforts in Central America
associated with various floristic projects have
provided a wealth of new material. A large num-
ber of new species have been reported, and many
others, previously poorly known, are now rep-
resented by a sufficient number of collections to
be understood more adequately. Since 1970, ef-
forts in southern Central America have rendered
earlier works incomplete in terms of taxa in-
cluded and distributional data. Recent studies of
Cordia in Mexico and Central America (Miller,
1985), and preparations of treatments for floras
of Nicaragua (Miller, in press) and Panama (Mil-
ler, in press) have uncovered two new species

rom Central America.

Cordia liesneri James S. Miller, sp. nov. TYPE:

L. Liesner 1976 (holotype, MO 2664908;
isotypes, AAU, US). Figure 1.

Frutex vel arbor parva 3-6(-8) m alta, ramunculis
glabris. Folia persistentia, petiolis 6-1 1(-18) mm lon-
gis; laminae anguste elliptico-ovate, (11—)14.5-30(-40)
cm longae, (3.8—)6.5-9.5(-13.5) cm latae, glabrae, apice
longi-acuminatis, A rotundatis ad obtusis. Inflores-
centiae axillaris, (2.5—)6—8.5 cm latis. Flores sessiles,
bisexualis; calyx tubulari-campanulatus, 5-6.8 mm
longus, 3-lobatus; corolla alba, tubularis, 8.4-11 mm
longa, 5-lobata, lobis oblongis, reflexis; stamina 5, filis

6-10 mm longis, puberulis, antheris oblongis,

raliter ovoideo, 9-18 mm longo, 9-12 mm lato, ros-

trato ad apiceum

Shrub or small, slender-trunked tree 3-6(-8)
m tall, the twigs glabrous. Leaves persistent, co-
riaceous; petioles 6-11(-18) mm long, dl
canaliculate adaxially, glabro

—9.5(-13.5) cm wide, the apex
long-acuminate, the acumen (2.3-)2.8-3.5(-4.6)
cm long, the base rounded to obtuse or rarely
acute, the margin entire, the adaxial surface gla-
brous or rarely minutely strigillose, the abaxial
surface glabrous or minutely strigillose. Inflores-
cence axillary or sometimes internodal, pendu-
lous, cymose, (2.5-)6—8.5 cm broad, the branches
shortly brown-canescent. Flowers sessile, bisex-
ual, monomorphic; calyx tubular-campanulate,
5-6.8 mm long, 3-4.7 mm wide at the mouth,

ANN. Missouni Bor. GARD. 74: 670—673. 1987.

lacking ribs, sparsely and minutely brown-strig-
illose, 3-lobed, the lobes often somewhat un-
equal, deltate to ovate, 0.6-1.1 mm long; corolla
white, tubular with reflexed lobes, 8.4211 mm
long, 5-merous, the lobes oblong, 2.4—4 mm long,
1.8-2.5 mm wide, the tube 4.4-7.2 mm long;
stamens 5, the filaments 7.6-10 mm long, the
upper 2.1—3.2 mm free, pubescent on the lower
free portion and at the point of insertion, the
anthers oblong, 1.3-2 mm long; ovary ovoid to
broadly ovoid, 1.3-2 mm long, 1-1.3 mm broad;
disc crateriform, 0.6-0.7 mm tall, 1-1.1 mm
broad, glabrous; style 5.4-9 mm long, the stylar
branches 2-3(-6) mm long, the stigma lobes cla-
vate to nearly discoid. Fruits borne in the cup-
shaped calyx, red or orange at maturity, drupa-
ceous, the stone inequilaterally ovoid, rostrate at
the apex, 9-18 mm long, 9-12 mm broad, the
surface with low ridges, the endocarp bony.

Distribution. Cordia liesneri is known only
from the Golfo Dulce region of Puntarenas, Cos-
ta Rica, where it occurs in wet forests below 200
m in elevation.

Among the Central American members of sect.
Myxa (Endl.) DC., Cordia liesneri is distinctive
in its oblong-ovate leaves longer than those of
related species, axillary inflorescences, and bright
red fruits containing large and prominently ros-
trate stones. This species is perhaps most closely
related to Cordia lucidula I. M. Johnston with
which it shares a similar habit of growth, rela-
tively large, glabrous leaves, a 3-lobed calyx, and
red drupacous fruits. However, it differs from C.
lucidula by bui axillary inflorescences and
rostrate endoc

Collections of C oua liesneri have existed since
Skutch and Allen collected in the Golfo Dulce
region in 1947 and 1951 respectively. However,
they have been identified incorrectly as Cordia
protracta I. M. Johnston, which differs by its
terminal inflorescences and pentamerous calyx,
and as Cordia eriostigma Pittier, which differs
in having terminal inflorescences and a campan-
ulate corolla. Allen (1956) briefly described C.
liesneri under the name C. protracta and stated
that flowering occurs in December and that fruits
mature in January

Additional specimens examined. |. CosrA RICA.
PUNTARENAS: region between Esquinas and Pas Sur

1987] NOTES 671

FicurE 1. Cordia liesneri.—A. Fruiting branch, after Liesner 1976 (MO).—B. Open corolla, after Burger &
Liesner 7224 (NY).

de Osa, elev. 30 m, Allen 5772 [DS, F (2), GH, US]; ner 7224 (CR, F, NY); slopes adjacent to airport, Rin-
region between Equinas and Palmar Sur de Osa, elev. cón de Osa, Liesner 1858 (AAU, CR, MO, US); Golfo
75 m, Allen 5827 (DS, P); about 5 km W of Rincón Dulce and Rio Terraba, elev. 30 m, Skutch 5303 (F,
de Osa, Osa Peninsula, elev. 50-200 m, Burger & Lies- | MICH).

FIGURE 2. Cordia cardenasiana.
Pip intact striate calyx. All after Contreras 6835 (holotype).

Cordia cardenasiana James S. Miller, sp. nov.

TYPE: Guatemala. Petén: Cardenas, on rocky

hill, 24 Mar. 1967, Elias Contreras 6835

(holotype, LL 279752; isotypes, DS, F, LL,
US). Figure 2.

Arbor ad 15 m, ramunculis glabris. Folia deciduis,

laminis ellipticis, 4.4—1 1.4 cm longis, 2.4—4.5 cm latis
glabris. Inflorescentia cymo-paniculatis. Flores hetero-

`

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

—A. Flowering branch.—B. Open flower.—C. Apex of a flowering branch

styli; calyx tubularis, 5—6 mm longus; corolla tubularis
4 ; m

mm longum; discus annularis. Fructus non visi.

Tree 15 m tall, the twigs glabrous. Leaves de-

ciduous; petioles 8-30 mm long, Pep en
ipe glabrous; blades elliptic, 4.4-11.4 c
long, 2.4-4.5 cm wide, the apex acute to acu-

1987]

minate, the base acute, the margin entire, the
adaxial surface glabrous but evenly papillose, the
abaxial surface glabrous. Flowers on short ped-
icels to 1.5 mm long, distylous, borne with the
leaves; calyx tubular, 5-6 mm long, 3-3.8 mm
wide at the mouth, unequally lobed and tearing
upon dehiscence or dehiscing circumscissily,
striate or faintly costate, inconspicuously short-
puberulent; corolla white, tubular with some-
what spreading lobes, 14.2-16.8 mm long,
5-merous, the lobes oblong, 5.2-6.2 mm long,
4.8-6 mm wide, the tube 4.7-5 mm long; sta-
mens 5, the filaments 9.8-11.5 mm long, t
upper 7-7.6 mm free, glabrous, the anthers ob-
long, 2.3-3 mm long; ovary obloid, 1.2-1.4 mm
long, 1.3-2.5 mm broad; disc annular, 1.4 mm
tall, 1.3 mm broad, glabrous; style 6.8-7 mm
long, the stylar branches 1.4-1.6 mm long, the
stigma lobes clavate. Fruits unknown.

Distribution. Cordia cardenasiana is known
only from the type collection made in Petén,
Guatemala.

Cordia cardenasiana is a small tree that must
be quite attractive in flower. Like related species,
it presumably produces all of its flowers in a short
period during the dry sesaon. It is assigned pro-
visionally to sect. Gerascanthus (P. Br.) G. Don,
but a definite sectional placement cannot be made
until fruits are known. Members of sect. Ger-
ascanthus have ellipsoid fruits with a fibrous wall,
and the base of the style remains attached. Mem-
bers of sect. Rhabdocalyx A. DC. have fruits with
a bony endocarp; some species are otherwise in-
distinguishable from sect. Gerascanthus. Al-
though Cordia is diverse in pollen morphology,
the two sections share pollen grains of the same
type (Nowicke & Ridgway, 1973), and this char-
acter cannot zagi the sectional placement of
C. cardenasia

Cordia ae en is probably most closely
related to C. gerascanthus L., with which it shares
glabrous leaves and a terminal, cymose-panic-
ulate inflorescence. Although most populations
of C. gerascanthus have pubescent staminal fil-
aments, populations with glabrous filaments are
known from southeastern Mexico (Johnston,
1950; Miller, 1985); filaments of C. cardenasiana

NOTES

673

are glabrous. Among the Mexican and Central
American species of Cordia, C. cardenasiana is
distinctive in having parallel-sided corolla lobes,
a character that it shares only with C. alliodora
(Ruiz & Pavón) Oken, C. gerascanthus, and C
globulifera 1. M. Johnston. A key is provided
below, and a key to the remaining species of the
section from this region is in Miller (1986).

la. n with ant domatia; indument of stellate
hai C. alliodora
Ib. Pant lacking ant domatia; indument of sim-
ple hairs.
2a. Inflorescence cymose-paniculate.
Calyx striate or faintly costate, gla-
brous; corolla less than 18 mm long
RSV RAYS OAE en rune C. tals Mita
3b. Calyx costate, pubescent; corolla mo
th m lon C. Md
2b. Inflorescence pat AER nearly umbel-
Vet x C. globulifera

I thank the curators of the following herbaria
for lending collections: AAU, CR, DS, F, GH,
LL, MICH, MO, US. I also thank my wife, Leslie
Miller, for support and the illustration of Cordia
liesneri, John Myers for the illustration of Cordia
cardenasiana, and G. Schatz for helpful com-
ments on the manuscript.

LITERATURE CITED

ALLEN, P. H. 1956. The Rain Forests of Golfo Dulce.
Univ. Florida Press, Gainesville, Florida

JoHNsTON, I. M. 1950. Studies in the Boraginaceae
XIX. Cordia section Gerascanthus in Mexico an

. Arnold Arbor. 31: 179-187.

MILLER, J. S. _ Systematics of the Genus Cordia
(Boraginaceae) in Mexico and Central America.

h.D. Dissertation. St. “Loui is Univ., St. Louis,

Missouri

1986. Cordia macvaughii, a new species of

Boraginaceae from western Mexico. Syst. Bot. 11:

179-187

Central America

— T Boraginaceae. In W. D. Stevens (editor), Flora
de ein dr (in press).
A revised treatment of Boraginaceae for Pan-
. Ann. - Miles uri Bot. Gard. (in press).
NowICKE, J. W. & J. E. RipGwAv. 1973. Pollen stud-
ies in the ur Cordia (Boraginaceae). Amer. J.
Bot. 60: 584-5

—James S. Miller, Missouri Botanical Garden,
P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

SARACHA SPINOSA—A NEW COMBINATION IN
PERUVIAN SOLANACEAE

The name Saracha R. & P. was long misap-
plied to the group of herbaceous neotropical
plants now correctly known as Jaltomata Schldl.
(Gentry, 1973). With reestablishment of the name
Jaltomata for the herbaceous plants, the need for
reinstatement of Saracha to the group of shrubs
of upland South America that had long been er-
roneously known as Poecilochroma Miers be-
came evident. The redefinition of Ja/tomata was
followed by D'Arcy (1973: 626, 1986: 28), Gen-
try & Standley (1974: 42), Hunziker (1979: 53),
and Nee (1986: 76) and the reinstatement of
Saracha by Hunziker (1979: 56). Both genera
are in need of revision, and not all combinations
have been made to recognize the restored generic
names. However, determination of a collection
made in the course of preparing a flora for Huas-
carán National Park in the uplands of the Cor-
dillera Blanca in northcentral Peru (Smith, in
prep.) requires the following new combination:

Saracha spinosa (Dammer) D'Arcy & D. N.
Smith, comb. nov. Poecilochroma spinosa
Dammer, Bot. Jahrb. Syst. 37: 637. 1906.
TYPE: Peru. Weberbauer 2907 (B, not seen,
destroyed; photo, MO).

Copiously armed shrub with long, trailing or
weeping branches; twigs glabrate, sometimes with
a simple Pp, paia ed many short

d lea

A

Bray-
ish, acicular spines 3-25; mm long. Leaves entire;
elliptical, obtuse or rounded at each end, mostly
12-20 mm long, coriaceous, glabrous or with
minute simple trichomes, shiny above, the mar-
gins subrevolute; petioles 2-5 mm long. Inflo-
rescences consisting of solitary terminal or ax-
illary flowers; peduncles wanting; pedicels slender
but broadening upwards, 10-15 mm long. Flow-
ers pendant, the calyx cupular, ribbed, dentate,
sometimes irregular, 6-8 mm long, sometimes
splitting; corolla apically plicate in bud, narrowly
campanulate, ca. 20-25 mm long, ca. 15 mm
wide at the mouth, the apex truncate with 5 del-
toid teeth 2 mm long; yellow and evenly mi-
nutely puberulent outside, inside basally uni-
formly dark violet, this reduced upwards to a
field of lobes and spots and pure yellow at the
apex; filaments ca. 15 mm long, inserted at the
base of corolla tube, glabrous and unappendaged,

ANN. Missouri Bor. GARD. 74: 674—675. 1987.

the anthers oblong, apiculate, basally cordate, 4—
5 mm long, yellow, included; ovary obconical, 4
mm long, the basal third enveloped in nectary,
glabrous, the style glabrous, exserted beyond the
anthers and nearly equalling the corolla. Fruit
(not preserved) a leathery berry about the size of
a gooseberry, perhaps 7 mm across.

Saracha spinosa is a weak, clambering shrub
that assumes a down-curving, weeping habit when
it lacks support. The branches can reach several
meters in length. The corollas are not so broadly
campanulate as in other members of the genus.
The plicate folds in the corolla in bud were not
found by Hunziker (1979: 58) i in the species he

discussed. Saracha

distinct from
other members of the genus in n having formidable
slender spines which appar erive from short
leafy shoots in the manner of Dunalia spinosa,
Iochroma horrida, and some other related So-
lanaceae. The spines on the specimen at hand
are copious, ascending, 8-25 mm long, and
needlelike. Fruit was not available for dissection
to ascertain the details referred to by Hunziker
(1979: 58) in his notes on three other species of
Saracha.

In his discussion under Poecilochroma in the
Flora of Peru, Macbride (1962: 58) suggested that
this species is similar to P. lindeniana Miers and
to P. lobbiana Miers, but both of these are un-
armed and have more open campanulate corol-
las, as is to be seen in the type illustration and
photos examined of these two species

The collection made in this study bears a re-
markable similarity to a specimen of Dunalia
collected by López & Sagástegui which we ten-
tatively refer to D. spinosa (Meyen) Dammer.
The leaves of the López & Sagástegui collection
are small and elliptical like those of our collection
of Saracha and unlike the larger oblanceolate
leaves of other specimens of D. spinosa, includ-
ing the type collection (Meyer, BM, destroyed,
photo MO). The flowers, too, are superficially
similar but narrower, and of course the filaments
have the basal appendages characteristic of the
genus Dunalia (Hunziker, 1959-60: 212). Had
we not examined the interior of the flowers, we
would have thought these the same species, not
different genera.

The great similarity between Saracha spinosa

1987]

and the neighboring Dunalia within adjacent parts
of Peru argues a need for unusual care in deter-
mining specimens of these plants and caution in
accepting past determinations. Under his treat-
ment of Saracha (Poecilochroma) spinosa Mac-
bride cited the following four specimens, which
appear to be at least geographically consistent:
Dept. La Libertad: Prov. Bolivar, Ferreyra 1254;
Prov. Santiago de Chuco, toward Angasmarca,
West 8162. Dept. Ancash: Prov. Huari, 3,600
m, Weberbauer 7014. Dept. Ayacucho: Prov.
Huamanga, above Quinua, Weberbauer 5542.

Saracha spinosa is apparently restricted to in-
terandean valleys of the western and central
chains of the Peruvian Andes. It has an extended
range of about 700 km, within which, at least
according to Weberbauer (1945: 420), the species
is sometimes locally common. Collection sites
known to us show that it occurs in shrubland
ranging from about 78°05’ W to 74*08'W and from
about 7?22'S to 8?05'S. The species has an ap-
parent elevation range from 3,300 to 3,700 m.
The type locality, which was imprecise, is near
or within Huascarán National Park.

The Park is in the Ancash Department of cen-
tral Peru, 300 km (air distance) north of Lima.
The reserve occupies nearly all the Cordillera
Blanca, which is the world's highest tropical
mountain range. The cordillera is located be-
tween 8?50'S and 10?00'S latitude and between
77°05'W and 77?49"W longitude with a north-
south length of 158 km and an area of 340,000
ha (131 sq. mi.). The elevation range is from
3,240 m to the 6,770 m summit of Nevado Huas-
carán Sur, the bulk of the Park above 3,500 m.
The outcrop is a mixture of igneous and sedi-
mentary rocks. The cordillera was extensively
glaciated and still has many glaciers and icefields.
Its topography is complex and supports a mosaic
of vegetation types. The most diverse and den-
sest shrub communities are found in the valleys
reaching lower elevations (3,500—3,800 m), where
the microclimate is warmer and moister. Al-
though valleys with these conditions are found
on both sides of the cordillera, the greatest num-
ber are on the eastern side.

In over a year of field collecting throughout

NOTES

675

the entire park, Saracha spinosa was located only
once, in Quebrada Rurichinchay near the valley
bottom in a community dominated by Miconia
salicifolia and with Alnus acuminata, Myrica pu-
bescens, Vallea stipularis, and Weinmannia aff.
laxiflora

SPECIMENS EXAMINED

Saracha spinosa. PERU. DEPT. ANCASH: Prov. Huari,
Huascarán National Park, Quebrada Rurichinchay be-
tween boundary and Quebrada Pachachaca. 3,600-
3,700 m, D. N. Smith 1 2475 (CPUN, HUT, MO, USM,
dupla). DEPT. AYACUCHO: Prov. Huamanga, road from
La Quinua to Abra Apacheta de Tambo, 12,000 ft.,
Plowman & Davis 4651

Dunalia aff. spinosa. PERU. DEPT. LA LIBERTAD: Prov.
Bolivar, Laguna de Los Ichus, al pié de rocas, 3,600
m, López & Sagástequi 3241 (MO).

This study was supported by National Science
Foundation Grant BSR-8305425

LITERATURE CITED

D’Arcy, W. G. 1973 [1974]. Solanaceae. In Flora
of Panama. Ann. Missouri Bot. Gard. 60: 573-
780

1986. The genera of Solanaceae and their
t ypes. Solanaceae Newsletter 2(4): 10-3
GENTRY, J. L Restoration of the genus Jal-
tomata (Solanaceae), Phytologia 27: 286-288.
TANDLEY. 1974. Solanaceae. Jn Flora
ofG Field Mus. Bot. 24(10-1&2): 1-151.
HUNZIKER, A. J. 1959-60 [1960]. Studios Lai So-
lanaceae. II. Sinopsis taxonómica del género Du-
nalia H.B.K. Bol. Acad. Nac. Ci. Córdoba (Ar-
gentina) 41: 211-244
— oo. American Solanaceae: a syn-
optic s . Pp. 49-85 in J. G. Hawkes et al.
(editors) The Biology and ERU of the So-
cademic Press, Lon
Mich n 1962. Solanaceae. Jn Flora of Peru.
eld Mus. Bot. 13(5B-1): 3-269.
NEE, M. 1986. Solanaceae I. Jn Flora de Veracruz,
Fasc. 49. Inst. Nac. Invest. Kcu Bióticas, Xa-
lapa, Veracruz, Mexico.
WEBERBAUER, A. 1945. El digo PME de los An-
des Peruanos. Min. Agric.,

— William G. D'Arcy, Missouri Botanical Gar-
den, P.O. Box 299, St. Louis, Missouri 63166,
U.S.A.; and David N. Smith, Missouri Botanical
Garden and Department of Botany, Iowa State
University, Ames, Iowa 50011, U.S.A.

YUTAJEA, ANOTHER NEW GENUS OF RUBIACEAE
FROM THE GUAYANA HIGHLAND

Of the genera of Rubiaceae endemic to the
region of the Guayana Highland, none had pre-
viously been described from the Serrania de Yu-
taje in the Territorio Federal Amazonas of Ven-
ezuela. A recent expedition to this region collected
material of a rubiaceous tree, here described as
a new genus. It is fitting to name the genus for
Yutaje, a sandstone mountainous area where nu-
merous endemic species are already known.

Yutajea Steyermark, gen. nov. TYPE: Y. /iesneri
Steyermark. Tribe Isertieae.

Arbor. Stipulae interpetiolares persistentes late lan-
Inflorescentia thyrsiformi-paniculata

alyx eases tandem in k
S

5111 Adv

sape DEN ° E ovulis numerosis.

Yutajea liesneri Steyermark, sp. nov. TYPE: Ven-
ezuela. Territorio Federal Amazonas: Dep-
to. Atures, 5-8 km NW of Yutaje settle-
ment, along stream flowing south from east
side of unnamed peak, 3 km west of Río
Coro Coro, west of Serranía de Yutaje,
05?40'N, 66°9’'W, 700-1,000 m, 10 Mar.
1987, Ronald Liesner & Bruce Holst 21826
(holotype, MO; isotype, VEN).

_ Arbor € 6- metralis li ib d

r

4 mm latis; foliis oblanceolatis vel pera ipii
apice acumina rra iie cutis 13.5-20 cm longis 5.5-8.5
cm latis e. thyrsiformi- sein late
ovoidea mul crag saratapas , axibus p
2-16 modice vel dense hirtellis in verticillos

dinatos ema in dichasia composita desinsatibus. ca-
lyce hypanthioque 4-6 mm longo subadpresso-pubes-
centi; calyce ante anthesin truncato vel paullo undulato
sub anthesi in lobos 3-4 irregulariter rumpenti, lobis
suborbicularibus vel suborbiculari-ovatis apice rotun-
datis, marginibus minute ciliolatis; pec rosea late
cylindrica vel subinfundibuliformi 8-9 mm longa 4-7
mm lata, tubo extus glabro intus seis partem ba-

silarem 1.5 mm glabrum adpresso-pubescenti atque
prope orificium barbato pilis longioribus instructo; lobis
6 paullo inaequalibus suborbiculari-ovatis vel obova-
to-oblongis apice obtusis vel rotundatis ubique glabris
marginibus minute papillatis; staminibus 6, antheris
linearibus

ANN. MISSGURI Bor. GARD. 74: 676—678. 1987.

Tree 6 m tall, the twigs appressed-pubescent
distally. Stipules broadly lanceolate, acute, 8 mm
long, 4 mm wide, strigose. vid oblanceolate
to lance-elliptic pex, acute at base,
13.5-20 cm long, 5.5-8.5 cm wide, the up
surface glabrous, the lower surface pilosulous on
midrib and lateral nerves, sparsely pubescent on
some tertiary veins, glabrous on surface between
the tertiary veins; lateral nerves 12-17 each side,
anastomosing near margin, elevated below; mid-
rib sulcate above and elevated below; tertiary
veinlets inconspicuous above, finely reticulate and
impressed below. Inflorescence thyrsiform-pan-
iculate, broadly ovoid, 35-50-flowered, 6-11 cm
long excluding the peduncle, 4.5-7 cm wide, the
main axes 12-16 in 3-5 verticils ending in often
compound dichasia, moderately to densely hir-
tellous; lowest axes 12-25 mm long, the others
7-15 mm long. Bracts subtending axes ovate,
acute, 2.5-4 mm long, m wide, minutely
hirtellous. Peduncle terminal, 8-12 cm long, 2.5-
3 mm diam., moderately pubescent. Calyx and
hypanthium 4-6 mm long, subappressed-pubes-
cent; hypanthium obconic, 3-5 x 3-5 mm. Calyx
truncate or slightly undulate before anthesis,
splitting irregularly during anthesis into 3-4
thickened, suborbicular or suborbicular-ovate
obes, these rounded at apex, 3-3.5 mm long, 4-
8 mm wide, minutely ciliolate at the margins.
Corolla pink or roseate, fleshy-thickened, broad-
ly cylindric or subinfundibuliform, 8-9 mm long,
4—7 mm wide, the tube 5 mm long, 4-7 mm wide,
glabrous without, within densely appressed-pu-
bescent except for the glabrous basal 1.5 mm,
and with longer barbate pubescence at the orifice;
lobes 6, slightly unequal, suborbicular-ovate or
obovate-oblong, obtuse or rounded at apex, 4-5
mm long, 2.5-3 mm wide, glabrous both sides,

thers linear, 3.5-5 mm long, 0.8-1.
the thecae transversely rugulose, terminating in
an oblong, rounded connective 0.5-1 mm long,
0.5 mm wide; filaments ligulate-linear, thick-
ened, 1.5 mm long, 0.7 mm wide, inserted 1.5-
2.5 mm above the base ofthe corolla tube, broad-
er than the base of the thecae. Style 4-6 mm
long, papillate-verrucose; stigmas 4—6, ligulate,
0.5 mm long. Disk annular, the margin undulate,
1 mm long, 4 mm diam. Ovary 6-celled, ovules
ca. 8 in each cell.

1987] NOTES 677

LECT

FiGURE 1. Yutajea liesneri.—A. Habit.—B. Portion of inflorescence.—C. Calyx and hypanthium before
anthesis. — D. Calyx and hypanthium during anthesis. — E. Stamen (lateral view). — F. Stamen (ventral view). —
G. Corolla, interior view. — H. Transverse section through ovary, semidiagrammatic. —I. Corolla, subinfundi-
buliform type.—J. Disk, from above, in depression at base of calyx tube. — K. Floral bract, exterior view.—
Detail of lower leaf surface. — M. Style and stigmas. — N. Flower with broadly cylindrical corolla type with slightly
unequal lobes.

Yutajea is a member of the tribe Isertieae as stamens inserted near the base of the corolla tube
circumscribed by Kirkbride (1979) and may be and in having sub-basifixed anthers. The slightly
placed next to its closest related genus, /sertia. ^ unequal corolla lobes and the uniformly pubes-
From I/sertia it differs especially in having the cent interior of the corolla tube are additional

678

characers at variance with /sertia. The corolla of
Yutajea is short-cylindric or subinfundibuliform
with imbricate lobes. Boom (1984) described the
corolla tube in /sertia as “cylindrical, short or
elongate" and the lobes as **valvate or imbricate
in bud.” In her abstract of characters delimiting
the tribe Isertieae, Kirkbride (1979) character-
ized the aestivation of the corolla as **valvate"
but later (p. 315) gave the aestivation as “‘valvate
or valvate-imbricate." As Boom indicated, Zs-
ertia may have either valvate or imbricate aes-
tivation. In /sertia the stamens are inserted near
the mouth of the corolla tube, the anthers are
dorsifixed, and the orifice of the corolla is usually
villous or barbate, except in J. scorpioides Boom,
while the remainder of the interior of the corolla
tube is glabrous in all the species with the ex-
ception of I. longifolia (Hoffsg. ex Roemer &
Schultes) Schumann.

Yutajea adds another endemic genus of the

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Rubiaceae to the list of twelve previously de-
scribed from the Guayana Highland (Steyer-
mark, 6).

The collections cited in this paper were made
under Grant No. 3264-86 of The National Geo-
graphic Society.

LITERATURE CITED

Boom, B 84. A revision of /sertia (Isertieae: Ru-
biaceae). Brittonia 36: 425-454.

KIRKBRIDE, M. CRISTINA GARCÍA. 1979. Review of
} š 1T : Danhi \ B : :

rittonia 31:

STEYERMARK, J. 1986. Holstianthus, a new genus of
Rubiaceae from the Guayana Highland. Ann. Mis-
souri Bot. Gard. 73: 495-497.

—Julian A. Steyermark, Missouri Botanical Gar-
den, P.O. Box 299, St. Louis, Missouri 63166,
U.S.A.

BOOK REVIEW

Duke, James A. 1986. Isthmian ee
ical Dictionary, Third edition. Pawan
mar Scientific Publishers, Jodhpur odis
ISBN 81-85046-35-2. Price $60

The Isthmian Ethnobotanical Dictionary, con-
cerned with useful plants of Panama, is 205 pages
in length, each page measuring approximately
81^ x 11 inches. It is in its third edition, the first
appearing in 1972, the second in 1978. The latest
contains 325 black-and-white line drawings of
flowering plants and a few ferns

The author explains that the Dictionary “°
contains herbal folklore in tropical plants which
complements the voluminous folklore on tem-
perate plants." In his introduction he indexes
“some frequent ailments and the more common
herbs used to treat them." Actually, many of the
plants listed are trees and shrubs. The diseases
range from abscesses to yellow fever. For ex-
ample, under *'indigestion" he lists by common

betical and composite fashion, he lists the com-
mon names ofthe plants without discussing them;
secondly, the diseases themselves (in full capi-
tals), including a brief layman's definition of the
isease and the genera of plants used therapeu
tically; lastly, the binomial name (in full Mida
and a discussion of how the plants are used by
z natives. While emphasis is placed on medic-
inal plants, the discussion may include such uses
of the plants as food, in construction, in the fash-
ioning of fishhooks, etc. Occasionally, the treat-
ments extend extra Panama, e.g., under bamboo
there is an elaborate discussion of about 1,500
words, a goodly segment of Hsing bou to the
uses of the grass in India and Ben
y the use of abbreviations in uma
names are used in

5
Panama, “Darien Spanish (D)"; “English yao
etc.

In the eyes of a botanist, apart from the wealth
of ethnobotanical data, the three most striking
features of the Dictionary are: 1) the complete-
ness of the list of ethnobotanical plants; 2) the
accuracy in the spelling of binomial names and
the authorities; and 3) how easily one familiar

ANN. MissouRni Bor. GARD. 74: 679-680. 1987.

with the vegetation of Panama can recognize the
individual species from the line drawings.

here are several items which I find disturbing,
the prime being a lack of discussion in the In-
troduction concerning the various ethnic groups
in Panama, plus the lack of a map of their lo-
cations in the Republic. The author sometimes
leaves the impression that the Kuna Indians are
the only aboriginals in Panama worth consid-
ering ethnobotanically. Why not more consid-
eration of the Chocó Indians, granted that they
are not as sedentary in Panama as are the Kunas?
After all, Duke tells us (p. 26): “I am called ‘Bo-
rojo' among the Darién Chocó because, for a
while I was distributing ‘borojo’ seedlings like
Johnny Appleseed." The Guyami Indians of
western Panama are hardly given a nod. The
obvious answer is that Duke spent a considerable
portion of his early scientific life among the Ku-
as.

From the viewpoint of a print job, the book
leaves something to be desired, considering that
the price of the book is $60.00 (U.S.). The paper
is of questionable quality, occasionally the ink is
smudged; all too often one encounters a blank
space in a sentence, a word or words having been
deleted; occasionally the alignment of words in
a column or sentence is poor.

An important point: books whose titles con-
tain the word “dictionary” are usually segregated
on library shelves under “for reference only."
Such a restriction may limit the uses of the book
and have an impact on its popularity.

I suggest that in the next edition a small su-
perscript number be appended after each bino-
mial to indicate the plant family; this could be
checked against a list of numbered families in an
appendix.

In the introduction, Duke in a cursory, but

=

its vegetation. His concern for the future disap-
pearance of the magnificent jungles of Darién
strikes a familiar note; his defense of folklore
medicine is convincing: . I could spend a
month with the herbals and dic medical texts
and come up with hundreds of examples of folk
medicine that have been vindicated or justified
by subsequent scientific research." As far as Pan-
ama's herb lore is concerned, he has already done

680

the basic spadework in admirable fashion. One
must not forget that James Duke collected more
than 6,000 “numbers” (herbarium specimens,
each number usually in duplicate or triplicate)
in the Republic of Panama (cf. Dwyer, in the

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Botany and Natural History of Panama, Mis-
souri Botanical Garden, 1985), a memorial to

his assiduity and a base for his ethnobotany.—
John

. Dwyer, Missouri Botanical Garden, P.O.
Box 299, St. Louis, Missouri 63166, U.S.A

ERRATUM

The incorrect author citation was used in the intro-
duction to the paper titled “Notes on the Breeding
Systems of Sacoila lanceolata (Aublet) Garay (Orchi-
daceae),” by Paul M. Catling [74(1): 58-68. 1987]. In-
stead of “var. paludicola Luer” it should be “var. pal-
udicola (Luer) eroi Wunderlin, et Hansen,
Phytologia 56: 308.

Volume 74, No. 2, pp. 183-462 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN was published on 29
September 1987.

THE GENERA OF EUPATORIEAE
(ASTERACEAE)

R. M. King & H. Robinson

Smithsonian Institution

Starting in 1966 the authors have provided a series of partial revisions of the
Eupatorieae, publishing over 200 papers on the group. The Genera provides a
synthesis of their contributions to our understanding of this species-rich group:
one in every 150 species of flowering plants belongs to the Eupatorieae.

This new work treats all 180 genera of the tribe with a description, full-page
illustration, discussion, and listing of recognized species and their distributions.
Keys to all the subtribes and genera and regional keys for portions of tropical
America are included. A nomenclator of the over 6,000 specific and infraspecific
names that have been placed in the tribe at one time or another indicates their
present disposition.

Published as Monographs in Systematic Botany from the
Missouri Botanical Garden, volume 22, 581 pages, 81⁄2 by 11 inches,
color frontispiece, hardbound. October 1987.

Price $70.00

For U.S. shipments: add $1.50 for one copy and $.50 for each additional copy.
Orders should be pre-paid; a $1.00 fee will be added to orders requiring invoices.
No shipments are made until payment is received. Mail order with your check or
money order, payable to Missouri Botanical Garden to:

Department Eleven

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P.O. Box 299

St. Louis, MO 63166-0299, U.S.A.

Please send copy(ies) of The Genera of Eupatorieae to:
O Payment enclosed.
Name : :
— ———— TEL Send invoice
Address | ec Sg ($1.00 will be added to total)

iT 743)

CONTENTS

A Revision of Panicum Subgenus Poulin Section Rudgeana (Poaceae: Paniceae)
nando O. Zuloaga

Arundoclaytonia, a New Genus of the Steyermarkochloeae (Poaceae: Arundinoideae) from
Brazil Gerrit Davidse & R. P. Ellis

Siphocampylus oscitans (Campanulaceae: Lobelioideae), a New Name for Burmeistera
weberbaueri from Peru Bruce A. Stein

Synopsis of the Genus antera (Campanulaceae: nd in Peru
Stein

New Species of Possiflora Subgenus sovr from Ecuador
J. E. Lawesson

E The Genus Attalea C Palenie) i in Panama

X4 :
Ç Novelties In i

Fer-

Bruce A.

L. B. Holm-Nielsen &

eror C. de Nevers

LY HE

(Loranthaceae and Viscaceae) Job Kuip
E A Revision of Pisi (Sapindaceae)
O. Téllez V. & B. G. Schubert .... z

: Systematics of the Southern African Genus cea (Iridaceae Iridoideae)
Goldblatt . dd.

Peter Goldblatt wars od

ç (Iridaceae) i in the Southwestern Cape, South Africa ae

x A. H. Gentry & J. Steyermark -n -— |
cte Una Nueva Especie del Genero Dioscorea (Dioscoreaceae) del Estado de Queretaro, Mexico -

Peter `
es on the ME id Tenn; of Watsonie Giboni (IV. pyramidata, W. -ardernei) k

, Cytology, and. Embryology of Campynemanthe (Liliales: ae AL
"Porter P. pe I, r Goldblatt * Hiroshi [PC icd ; a

Annals
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. Missouri
— Botanical

US

Volume 74, Number 4
Winter 1987

Annals of the
Missouri Botanical Garden

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Missouri Botanical Garden

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Volume 74
Number 4
1987

Annals
of the
Missouri
Dotanical

Garden

NZ

CONTRIBUTIONS TO A SYMPOSIUM ON THE
EVOLUTION OF THE MODERN FLORA OF THE
NORTHERN ROCKY MOUNTAINS:
INTRODUCTORY REMARKS

The following nine papers represent contri-
butions from a symposium presented on June
11, 1985, at the University of Montana. The
ymposium was sponsored jointly by the Paleo-
botanical Section and the Pacific Section of the
Botanical Society of America and was held dur-
ing the annual meetings of the Pacific Division
of the American Association for the Advance-
ment of Science.

The main purpose of the symposium was to
bring together current information about the

life and earth sciences, and they have constructed
their written reports accordingly.

A frequent problem for nonpaleobotanists in-
terested in learning about the past vegetation in
a given region is that it is difficult to locate this
information. Reports about past vegetation are
by no means rare, but they are scattered, and we
have lacked good syntheses. The following pa-
pers attempt to solve that problem for the north-
ern Rocky Mountain region. Each is a succinct
yet authoritative summary of a particular body
of information authored by leading specialists in

ANN. MISSOURI Bor. GARD. 74: 681-682. 1987.

that field. A number ofauthors have gone beyond
presenting reviews and have included informa-
tion and ideas in their treatments that have not
been published elsewhere. Thus, each report is
intended to be a definitive work on a particular
duration of geologic time.

Readers will note what may appear to be two
major inconsistencies from paper to paper. One
has to do with geologic time and the application
of epoch names, while the other is the use of the
terms “flora” and “vegetation.”

Paleobotanists generally agree on the absolute
ages of the various plant-bearing strata, but they
may disagree on the absolute age of the boundary
between epochs. Thus, the same deposit may be
regarded as latest Eocene by one worker and ear-
liest Oligocene by another. Where this might cause
confusion in this series of papers, authors have
been asked to provide the absolute age as well
as the epoch.

Workers dealing with modern plants and with
those from the not too distant past use the term
"flora" to indicate a list of taxa present and
“vegetation” to include all aspects of those plants.
Historically, fossil “floras” were little more than
lists of species identified from a deposit. Later,
such lists expanded to include taxonomic de-
scriptions, comparisons, and, more recently, in-

682

formation about frequency of occurrence of re-

ins and environment of deposition.
Interpretations of community structure, mean
annual temperature, mean annual range of tem-
perature, seasonality, and amount and seasonal
distribution of rainfall are often included in mod-
ern treatments as well. ile these are clearly
aspects of “vegetation,” the term “flora” has per-
sisted and is often used instead. Thus, those
working with ancient plants may not iudi
between these two terms and may use "flora
when “vegetation” is appropriate.

A second purpose of the symposium was to
honor four workers who pioneered the study of
the past vegetation of the northern Rocky Moun-
tain region. These are: Herman F. Becker, Ro-
land W. Brown, Edu Dorf, and Harry D.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

MacGinitie. Their contributions have been sum-
marized by Jack Wolfe, and the summary is in-
cluded as part of this publication. Drs. Becker
and MacGinitie passed away after the sympo-
sium was held and their bibliographies are in-
cluded here. The accomplishments of these sci-
entists provided a solid foundation on which
modern work rests. Even more importantly, they
provided leadership and encouragement to the
current generation of paleobotanists, and the pa-
pers published as a part of this symposium owe
much to their efforts.

— Charles N. Miller, Jr., Department of Botany,
University of Montana, Missoula, Montana
59812, U.S.A.

DEDICATION

This symposium is dedicated to four col-
leagues, all unfortunately deceased, who have
contributed significantly to the paleobotany of
the northern Rocky Mountains: Herman F.
Becker, Roland W. Brown, Erling Dorf, and Har-
ry D. MacGinitie. The work of all four men has
provided a foundation of collections and knowl-
edge on which this symposium is partially based.
Two ofthese valued colleagues were, in fact, alive
at the time (June 1985) the symposium was held.
We had intended to honor Herman Becker dur-
ing the course of the meetings at which the sym-
posium was held, but he was at that time in the
throes of his terminal illness and was unable to
attend. Also regrettable is the death in January
1987 of Harry MacGinitie. Early in his paleo-
botanical career, Becker solicited and received
much advice from Brown, Dorf, and MacGinitie,
colleagues whom Becker greatly respected; thus,
dedication of this symposium to all four men is
particularly appropriate.

As Dilcher (1987) notes, Herman F. Becker
(1907-1985) entered paleobotany late in life, and
thus his paleobotanical career, mostly on the staff
of the New York Botanical Garden, covered less
than two decades. In that time, however, Becker
monographed several latest Eocene (Oligocene
in some chronologies) floras from southwestern
Montana and discussed in shorter contributions
some enigmatic plant fossils. The Montana floras
contain early records of many microthermal taxa,
and Becker's extensive collections of these floras
have been a major basis for expanding knowl-
edge of the development of microthermal vege-
tation and the evolution of microthermal taxa.

Roland W. Brown (1893-1961) was for 30 years
the Mesozoic and Cenozoic paleobotanist of the
U.S. Geological Survey (see Mamay, 1963) and
was particularly concerned with the Cretaceous
and Tertiary floras of the Rocky Mountains and
adjacent High Plains. His contributions included
numerous short, largely taxonomic papers;
Brown's sharp eye for details kept his contem-
poraries “on their toes" in regard to determi-
nations. However, Brown's major contribution
was the monograph “Paleocene floras of the
Rocky Mountains and Great Plains," which rep-
resents the accumulation of a massive amount

opment of the flora and vegetation in this region,

ANN. Missouni Bor. GARD. 74: 683. 1987.

or the general distribution of Paleocene vegeta-
tion mad on Brown’s monograph.

For many years, Erling Dorf (1905-1985) was
a highly respected professor in Geology at
Princeton cea His paleobotanical con-
tributions "ass from studies of Devonian plants
from Beart Butte, Wyoming, to studies of
the 1. iin ci floras of California to
taphonomic observations following the eruption
of Paricutin in Mexico. Dorf was also concerned
with general patterns of climatic change as evi-
denced by land floras. Of particular relevance to
this symposium, however, is Dorf’s fieldwork on
rocks and their floras near the Cretaceous—Ter-
tiary boundary in the northern Rocky Mountain
region and particularly his monographic treat-
ment of the latest Cretaceous Lance and Medi-
cine Bow floras; this monograph remains the most
recent and significant treatment of latest Creta-
ceous plants in the northern Rockies.

Harry D. MacGinitie (1896-1987) was a source
of encouragement and advice to all concerned
with the Tertiary floras of western North Amer-
ica (Wolfe, 1987). His published contributions
span 42 years (1933-1974) and include several
major monographs of both Paleogene and Neo-
gene floras, including those of the Rocky Moun-
tains. MacGinitie's systematic work was always
of high quality. The areas of historical plant ge-
osraphy, paledecolce on analyses of fossil plant

ange were, however,
of major ‘concern to him, and embedded in his
floral monographs are thoughtful and extensive
discussions of the development of the vegetation
and flora of western North America. Many now
generally accepted concepts regarding the floris-
tic and vegetational history of western North
America during the Tertiary have their origins
in MacGinitie’s work.

LITERATURE CITED

DircHER, D. L. 1987 [1988]. Memorial to Herman
F. Becker (1907-1985). Ann. Missouri Bot. Gard.
he P gta
MAY, S. H. 1963. Memorial to Roland W. Brown
ee: isc Geol. Soc. Am. Bull. 74: 79-

Wo tre, J. A. 1987 [1988]. Memorial to Harry
MacGinitie (1896-1987). Ann. Missouri Bot. he
74: 684—688.

— Jack A. ie Paleontology and Stratigraphy
Branch, MS-919, U.S. Geological Survey, Fed-
eral Center, nr Colorado 80225, U.S.A

Harry D. MACGINITIE

MEMORIAL TO HARRY D.

Back in the early 1950s, a bus trip from Port-
land, Oregon, to Boston would have been a long
one, especially for a kid just entering college. But
not for this freshman, because I had along Harry
D. MacGinitie's recently published “Fossil Plants
of the Florissant Beds, Colorado." The copy is
now well worn, the binding tattered. Why a bud-
ding paleobotanist could and should have read
this book is part of the story that follows.

In December 1952, I visited Berkeley—then
the Mecca of Tertiary paleobotany —and met with
R. W. Chaney, the widely accepted leader of this
field. Chaney assigned Marie Pabst the task of
entertaining this possible novitiate. Among the
many questions asked of Pabst were: who pro-
duced good work, who was a model? Unhesitat-
ingly, Pabst reached for the bookshelf and pulled
out MacGinitie’s (1941) “A Middle Eocene Flora
from the Central Sierra Nevada." “This is the
best work that has been done in western Tertiary
paleobotany," she declared. “This is the work to
emulate.”

Mac MacGinitie was a paleobotanist’s paleo-
botanist to Pabst, as well as to his other col-
leagues. He considered all evidence that bore on
a scientific problem, and his conclusions and hy-
potheses were written logically and clearly, es-
chewing jargon whenever possible. He consid-
ered all reasonable alternatives and stated basic
assumptions. If he changed his mind on a subject,
he clearly stated this and the evidence that re-
sulted in the change. His knowledge of leaf ar-
chitecture of the angiosperms was immense, and

ANN. MISSOURI Bor. GARD. 74: 684—688. 1987.

MACGINITIE (1896-1987)

the great bulk of his determinations of fossils are
still valid. Most importantly, Mac shared his
knowledge continually by discussions with his
colleagues and would even spend his time in the
herbarium helping with identifications. In both
personal contact and his publications, Mac has
had a profound me in Tertiary paleobotany
in North Amer

By modern * oubli or perish" standards,
Mac's publications were few. Each, however,
contains a few to several essays that could have
readily been extracted as separate papers. And,
each 5 UN new approaches to Ter-
tiary paleobot

Mac's first uo NA in 1933 was the first
comprehensive account of an upland Miocene
flora from the Columbia Plateaus. In this paper,
Mac's discussion of the climatic implications of
the flora is the first to attempt to assign numbers
to climatic parameters and clearly reveals his
interest in, above all, paleoclimate. I recall one
discussion with Mac in which he stated that he
entered paleobotany largely because he thought
that fossil plants were the best indicators of pa-
leoclimates. He was well aware ofthe significance
of altitude, and his assignment of about 2,500
feet to the Trout Creek has not been significantly
altered by newer (and supposedly more reliable)
techniques.

The Weaverville flora, published in 1937, was
Mac's Ph.D. thesis, which was completed under
R. W. Chaney in 1935. This year also saw another
major event in Mac's life; Beatrice MacGinitie

1987] WOLFE—

(nee Hess), who became his wife on February 2,
1935, was a constant source of encouragement
and emotional support to Mac for the rest of his
life.

The Weaverville flora reveals Mac’s increasing
involvement with both the detailed climatic im-
plications of fossil-plant assemblages and the
possible causes of climatic change. Further, the
detailed discussion of the geologic occurrence of
the fossils and of basinal geology placed the
Weaverville flora in a taphonomic context. Mac
attributed the Weaverville flora to the early Oli-
gocene, recognizing that the flora contained ele-
ments that indicated an age younger than the
Goshen flora.

The monograph of the Eocene Chalk Bluffs
flora, published in 1941, shows a maturation of
many concepts. As in his earlier works, Mac con-
sidered the paleoecological significance of the
fossil taxa in a depositional framework. He pre-
sented solid evidence for the co-occurrence of
taxa whose modern counterparts have markedly
different climatic tolerances and suggested that
at least some tolerances have changed through
time. Mac documented that the neotropical ele-
ment in the Chalk Bluffs was today largely found
in the Mexican uplands and that the flora also
contained a large paleotropical element. This
monograph also contains the first significant dis-
cussion of the plant biogeography of the North
American Eocene. Mac laid the cornerstones for
future discussions of the historical biogeography
of the western North American Tertiary.

The taxonomy in the Chalk Bluffs paper has
largely withstood the test of time. Mac, using
solid leaf-architectural criteria, demonstrated that
the many leaves of “Aralia,” a major element in
western American Paleogene floras, represented
a platanaceous genus. He referred the leaves to
Platanophyllum and discussed the phyletic re-
lationships of the species; recently they have been
reassigned to the new genus Macginitiea, named

eo
climatic inferences still remain valid is revealed
in the Chalk Bluffs monograph: “In no case were
ecological considerations given weight in the
choice between two living forms as modern rep-
resentatives of fossil plants... . If identifications
are based on ecology, and then ecology is de-
duced from identified species, the result is a kind
of cyclic reasoning which may lead to consid-
erable error" (MacGinitie, 1941: 96).
While teaching briefly at the University of Col-

NORTHERN ROCKY MOUNTAIN SYMPOSIUM

685

orado, Mac was associated with T. D. A. Cock-
erell, the paleoentomologist who had worked ex-
tensively with insects from the famous lake beds

at Florissant, Colorado. No comprehensive
treatment—systematic or ecologic—existed of the
Florissant flora, and, at the urging of Cockerell
and the vertebrate paleontologist Childs Frick,
Mac began work on the Florissant flora in 1936,
work that culminated in his 1953 monograph of
the flora.

Mac’s research was, however, interrupted by
World War II. Although Mac was 45 when the
United States entered the war, he still decided
to serve. Because of Mac’s expertise in the the-
oretical aspects of climatology and meteorology,
he enrolled as an instructor in the U.S. Army Air

orps. Lt. MacGinitie’s primary responsibility
was teaching crewmen the significance of, and to
recognize, different cloud formations. After the
war, Mac returned to Humboldt State and to his
work on the Florissant.

Above all, the Florissant work involved care-
fully detailed taxonomic work. Mac reduced the
258 species previously attributed to the flora to
less than half that number, but still the Florissant
is the largest flora yet described from the Tertiary
of western North America. One common prac-
tice in Tertiary paleobotany was the erection of
a “fossil” species for fossils that resembled a giv-
en living species. Mac, however, challenged that
practice: “Fossil plants in two different floras may
be likened to the same /iving species without in
any way implying identity ofthe two fossil forms.
Their differences may be great enough in oppo-
site directions to place them in different species

." (MacGinitie, 1953: 79). As Mac noted, the
usual practice led to erroneous concepts of age
and floristic relationships.

In the Florissant monograph, Mac introduced
and elaborated on a number of new ideas and
Observations that have subsequently been well
substantiated. Particularly significant is the idea
that given lineages may persist in a region by
adapting to changing environmental conditions;
he was led to this conclusion because of the shar-
ing of numerous lineages between the older Green
River flora and the Florissant. Mac also clearly
stated (p. 46) that fossil leaf
are biased towards streamside or lakeside vege-
tation and, hence, may not adequately reflect the
regional flora and vegetation; this point has been
increasingly made apparent by many tapho-
nomic studies. In discussing the paleoclimatic
significance of floras then placed in the Oligo-

686

cene, Mac noted: “The point emphasized by this
discussion is that if we accept a Middle Oligocene
age for the Goshen flora, the remainder of that
period must have witnessed an almost complete
revolution in the flora of the region, and more
critical events must be crowded into the Upper
Oligocene than paleontologists have hitherto been
willing to concede” (MacGinitie, 1953: 67); Mac
was the first to recognize the major and rapid
climatic change that is now placed near the end
of the Eocene.

Mac completed the Florissant study in 1951.
Probably because of administrative duties as
Chairman of Natural Sciences at Humboldt State,
Mac did not immediately start on a new project.
In 1954, James Bump sent Mac some leaves from
a new site in Nebraska, and encouragement from
Bump and Harold J. Cook enticed Mac to start
fieldwork on the Kilgore flora, which was com-
pleted in 1958. Following his retirement from
teaching in 1960, Mac finished the systematics
of this assemblage. He had become increasingly
aware of the value of palynology in paleoecolog-
ical interpretations, and, despite no prior paly-
nological experience, he became knowledgeable
of basic pollen morphology with the help of Es-
tella Leopold. The Kilgore paper was the first in
North American Tertiary paleobotany to illus-
trate both megafossils and microfossils and ex-
tract from both types of fossils paleoecological
data. The climatic discussion in the Kilgore pa-
per was based on a wide range of data (paleo-
botanical, sedimentological, vertebrate, and
molluscan) and probably remains the most com-
prehensive discussion of Neogene climates in the
Plains region of North America.

The Kilgore paper also contains an expansion
of the thesis, first expounded in the Florissant
paper, that many plant lineages can persist in a
given region despite significant climatic change.
If, as Mac demonstrated, the bulk of the Kilgore
species were derived from older species in the
. it should be questioned

ry.
dro-Tertiary,’ and the like imply esten els use-
ful concepts if we do not think of these terms as
representing areas or centers from which mass
migrations occurred. They picture to us in a gen-
eral way the vegetation occupying an area, al-
though the particular type of vegetation was
slowly changing" (MacGinitie, 1962: 87). This
statement by Mac represented a clear-cut depar-
ture from the geofloral hypotheses then so widely

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

accepted in North American Tertiary paleobot-
any and historical plant geography. However, as
originally submitted to the Board of Editors of
the University of California Publications in Geo-
logical Sciences, Mac's manuscript was unchar-
acteristically vague about this point. Clyde
Wahrhaftig, one of the editors, asked me what
Mac meant. From numerous discussions with
Mac, I knew what he was attempting to state,
and at Wahrhaftig's request, Mac rewrote his
manuscript to the statement quoted above.

On retiring from Humboldt State, the
MacGinities moved from Arcata to Napa, a
move that was beneficial to Mac and especially
to many students and colleagues. He was made
a Research Associate at the University of Cali-
fornia Museum of Paleontology and had read
access to extensive paleobotanical collections
(many of which he had made) and the University
of California Herbarium and libraries. He was
in an area active in the geological and botanical
sciences, and Mac was often to ve found in deep
discussions gues, to whom
he freely gave of his time. Every thesis in paleo-
botany completed at Berkeley from the 1960s
through the 1980s acknowledges Mac’s assis-

—
£2

nce.
With typical enthusiasm, Mac then began work
on the Eocene floras of the Rocky Mountains to
understand better the floras and vegetation that
preceded the Florissant. The first of these floras,
that of the upper part of the Green River For-
mation in Utah and adjacent Colorado, was, like
the Florissant, a classic flora. The last significant
descriptive work, that by Roland Brown, was
more than 30 years old, and the paleoecological
significance of the Green River flora had never

Eocene localities in Wyoming. In 1968 he com-
pleted work on the Green River flora (Mac-
Ginitie, 1969) and in early 1972 on the Kisinger
Lakes flora (MacGinitie, 1974). He continued
work on other Wyoming Eocene floras, but, un-
fortunately, none were completed for publica-
tion. A major hindrance to completion of these
works was a deterioration of leg joints, which
required surgery and made fieldwork difficult and
painful.

In Mac’s last two works on the Green River
and Kisinger Lakes floras are the same type of
careful and detailed discussions of paleoecology
and paleoclimatology that are present in his ear-
lier publications. Certain parts of these mono-

1987]

graphs should be carefully read by any paleobot-
anist or student in paleobotany, e.g., the chapters
"Certain Aspects of Floristic Evolution" and
"Paleobotanical and Botanical Species"
(MacGinitie, 1969: 68-70, 81-86), and “Distri-
bution of Correlative Living Species” (Mac-
Ginitie, 1974: 29-34). Indeed, Tom Taylor and
Edie Smoot selected the second-listed chapter as
one of eight papers on Tertiary paleobotany in
the “Benchmark Papers in Systematic and Evo-
lutionary Biology Series" (MacGinitie, 1984).
Mac’s discussions of systematic determinations
also had become more sophisticated; he contin-
ually discussed the exact reasons certain generic
determinations were made (not that a fossil tax-
on simply “looked like” an extant taxon) and
illustrated mu ofthe material in detail. Written
with Estella Leopold, who also contributed a
chapter to the Kisinger Lakes paper, the sum-
mary of floristic and vegetational development
in the Tertiary ofthe Rocky Mountains (Leopol
& MacGinitie, 1972) remains a most useful and
concise summary on that topic.

The symposium held in honor of MacGinitie
at the 1983 annual meeting of the American As-
sociation of Stratigraphic Palynologists was a
well-deserved tribute. Palynologists recognized
both the wide scope of Mac's contributions
throughout his career and his bringing together
of palynology and megafossil research in the
gore and later papers. During a tribute listing his
many accomplishments, Mac, sitting in the front
row, nudged Estella Leopold and whispered, “I
never realized that I was that good." Mac had
generally lived in the shadow of R. W. Chaney,
and even when disagreeing with Chaney or other

colleagues, was careful not to cite by name the
originators of hypotheses; instead, Mac would
simply and clearly, through data and logic, refute
the hypotheses. One of the rare instances when
Mac named names was, in fact, at the 1983 AASP
symposium when he stated of a colleague, “He
was a nice man but a terrible paleobotanist,"
much to the amusement of the audience.

One of Mac's major attributes was his ability
to grow intellectually, to accept new concepts and
weld them into his already considerable frame-
work of floristic evolution and vegetational/cli-
matic change in the Tertiary of western North

H
'1

merica. If new and/or refined techniques of

analysis (e.g., palynology and leaf architecture)
appeared, he s among the first to use these
techniques. Mac kept up with advances in pa-
leobotany even after he was no longer an active

WOLFE—NORTHERN ROCKY MOUNTAIN SYMPOSIUM

687

researcher in the mid 1980s. Above all, Mac was
always ready to discuss with colleagues any prob-
lem of mutual interest and to share with them
his vast wealth of knowledge and experience.
Fortunately, part of Mac’s legacy to us—his en-
quiring mind and thoughtful statements—will
endure in his publications. When Mac ceased
being an active researcher, he, in typical generos-
ity, brought down to Berkeley his entire profes-
sional library, leaving for present and future stu-
dents an additional legacy.

Thanks are due to Beatrice Ann Minkler (nee
MacGinitie) for supplying much pertinent bio-
graphical material. Howard E. Schorn supplied
the accompanying photograph and contributed
to some of the content of this memorial. Patrick
F. Fields supplied the list of MacGinitie’s pub-
lications.

BIOGRAPHICAL DATA

Harry Dunlap MacGinitie was born in Lynch, Ne-
braska, on March 29, 1896. After graduating from high
school in Sturgis, South Dakota, he moved to Califor-
nia. He attended and received an A.B. from Fresno
State College in 1926, after which he attended Stanford
University for a year. In 1926-1 1928,

Humboldt State College (now University) in Arcata,
California, where he taught until 1960, except for a
year (1932-1933) when he attended Berkeley full time,
a year (1936-1937) when he taught at the University
of Colorado (Boulder), and two years (1943-1945) when

Paleontology. He was a Fellow of
emy of Sciences and the Geological Society of America
and a member of the Paleontological Society. He died
on January 31, 1987, in Napa, California.

PUBLISHED CONTRIBUTIONS OF
Harry D. MACGINITIE

1933. Redwoods and frost. Science Ser. 2, 78(2018):
90.

The Trout Creek flora of southeastern Oregon.
Publ. Carnegie Inst. Wash. 416: 21-68, 16 pls.
Ecological aspects of the floras of the auriferous
gravels. Geol. Soc. Amer. Proc. for 1933, Ab-
stracts, p. 356. [Abstract.]

Tertic floras of Trinity County, California. Pan-
Amer. Geol., Abstracts, 62: 75-76. [Abstract.]
Tertiary floras of Trinity County, California.

1933.
1934.

1934.

1935.

688

1937.
1937.
1938.
1938.
1941.

1943.
1953.

1958.
i

A
1961.

1962.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Geol. Soc. Amer. Proc. for 1934, Abstracts, p.

390. [Abstract.]

Stratigraphy and flora of the Florissant beds.

Geol. Soc. Amer. Proc. for 1936, Abstracts, pp.

362-363. [Abstract.]

The flora of the Weaverville beds of Trinity

County, California. oo Carnegie Inst. Wash.

465: 83-151,

Stratigraphic — in the Tertiary gravels

of the Sierr vada. Geol. Soc. Amer.

for 1937, aN pp. 246-247. [Abstract.]

Geologic relations along the southwest border
of the Klamath Mountains. Geol. Soc. Amer.

Proc. for 1937, Abstracts, p. 247. [Abstract.]

A middle Eocene flora from the central Sierra

Nevada. Publ. Carnegie Inst. Wash. 534: 1-178,

47 pls.

Central and southern Humboldt County. Calif.

Div. Mines Bull. 118: 633-635.

Fossil plants of the Florissant beds, Colorado.

: 1-198, 75 pls.
Late Cretaceous. Pp. 61-79

bs (editor), pes raphy. Amer.

gocene plants from the upper Ruby River Basin,

southwestern Montana" by H. F. Becker). Ecol-
ogy 42: 853-854.

The Kilgore flora, a late Miocene flora fro

northern Nebraska. Univ. Calif. Publ. Geol. Sci.

35: 67-157

1968.

1969.

1972.

1972.

1974.

1984.

[VoL. 74

Some vegetation types in the Eocene of l. mid-

dle Rocky Mountains. Geol. Soc. Amer. Spec.

Pap. 101: 321, 408-409. [Abstract.]

The Eocene Green River flora of northwestern

Colorado and iesu Utah. Univ. Calif.
Publ. Geol. Sci. 83: 3,

(Wi th E. B. ay Development and affin-

and Paleofloristics of Asia and Eastern Nort
America. Elsevier Publ. Co., Amsterdam
(With R. A. Scott, P. L. Williams, L. C. Cra

age of the palm woods and roots. Amer. J. Bot.
59: 886-896.

An early middle Eocene flora from the Yellow-
stone-Absaroka volcanic pon OV ince, northwest-
ern Wind River AR Wyoming. Univ. Calif.
Publ. Geol. Sci. 108: 210»
The Eocene Green E flora of northwestern
Colorado and northeastern Utah (Systematic
considerations. Paleobotanical and botanical
species). Pp. 262-267 in T. N. Taylor & E. L.
Smoot (editors), Paleobotany, Part II. Triassic
through Pliocene. Van Nostrand Reinhold Co.,
New York

—Jack A. Wolfe, Paleontology and Stratigraphy

Branch, MS-91

9, U.S. Geological Survey, Fed-

eral Center, Denver, Colorado 80225, U.S.A

HERMAN F. BECKER

MEMORIAL TO HERMAN F. BECKER (1907-1985)

Herman F. Becker was born in Duesseldorf,
Germany, on January 10, 1907. After complet-
ing his initial education at the Realgymnasium
in Neunkirchen (1923), he undertook the study
of botany in Dahlem (Berlin) at the Botanical
Gardens with the encouragement of his father
and grandfather. There he completed studies as
a horticulturist. He met his wife, Ruth, in Berlin
and they were married in Frankfurt, Germany
in 1928.

He saw no possibility of continuing his study
of botany in Germany so he decided to go to the
United States and arrived in New York in March
1930. His wife and young son followed about
one year later after he had obtained work as a
Horticultural Assistant (1930-1939) at the
Brooklyn Botanical Garden. Twin sons were born
in New

Beginning in 1939 he taught at Brooklyn Col-
lege as a Lecturer in Geology while he undertook
his B.A. degree, which he completed in 1947. He
continued to teach at Brooklyn College while
earning an M.A. degree from Columbia Univer-
sity in 1952.

ANN. MISSOURI Bor. GARD. 74: 689-691. 1987.

During this time he took part in a geology field
trip to the West with several geology students.
While on this trip he was told, “Becker, you look
for plant fossils," because everyone knew of his
interest in plants. He came back to New York
excited about a shoe box full of fossil plants that
he had collected.

Becker wanted to continue in research but had
little opportunity to do so while teaching a heavy
course load as a Lecturer and later Instructor at
Brooklyn College (1939-1955). However, in the
summer of 1947 he began field studies under the
direction of Professor A. J. Eardley, director of
geologic work at the Rocky Mountain Field Sta-
tion of the University of Michigan. Soon he was
directed to Professor Chester Arnold. He con-
tinued to maintain contact with Arnold and col-
lected fossils in the Ruby River Basin area of
southwestern Montana during the summer of
1949. He then enrolled in a Ph.D. program at
the University of Michigan to work in Paleo-
botany with Professor Arnold. At first he worked
on his degree in the summers. But, after com-
pleting additional fieldwork on the paper shales

690

of the Ruby River Basin in the summer of 1955,
he remained at Michigan, completing his Ph.D.
degree in 1956. He was awarded the Ermine
Cowles Case Award in Geology for his outstand-
ing Ph.D. dissertation.

Herman Becker attempted to teach again at
wasa Cie but the heavy course load gave
him no time for his research on the fossil plants
of the Ruby River Basin to which he was by this
time very committed. Thus, in 1958 he joined
the New York Botanical Garden, first as a Re-
search Associate and later as a Curator of Paleo-
botany. Between 1957 and 1968 he made seven
more major collecting trips to southwestern
Montana, which provided him with the research
material that formed the basis of many of his
publications.

The first two major floristic papers he pub-
lished were on the paper shale beds of the Ruby
River Basin flora (middle Late Oligocene) (196 1a)
and the Mormon Creek flora (upper Middle Oli-
gocene) (1960c). With continued extensive sum-
mer fieldwork in southwestern Montana, he pub-
lished monographic papers on the Beaverhead
Basin (lower Middle Paleocene) (1969c), the
Metzel Ranch flora in the Ruby River Basin
(lower Late Oligocene) (1972b), and the York
Ranch flora of the Ruby River Basin (Upper Oli-
gocene—Lower Miocene) (1973b). In these floris-
tic papers he not only presented the fossil flora
from the specific fossil beds he collected, but he
applied the resulting data to questions of the cor-
relation of their floristic elements to other North
American Tertiary floras, taxonomic age rela-
tionships of the specific floras, and the phyto-
geographical distribution of modern equivalents
for the fossil plants described.

Herman Becker worked at a time when the
questions being asked in the field of paleobotany
were concerned primarily with paleoecological
and phytogeographical issues. However, perhaps
because of his strong botanical background, he
developed a special interest in the relationship
between fossil and extant plants and the evolu-
tion of plants through time. In his paper on the
Beaverhead flora he wrote: “The writer does not
agree to calling Tertiary fossil species ‘identical’
with any extant species, and applying to it the
modern specific epithet, especially where Mio-
cene-Oligocene, or older plants are concerned

. In a strict sense, no individual of a modern
species is morphologically or genetically identi-
cal with another, a fact expressed in major or
minor variability as a phenotype. Genetic, phys-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

iological, and many morphological characters are
usually not available to the paleobotanist to pro-
nounce a specimen identical or NA
equivalent to a modern form" (1969c: 10).

The importance of his research was pa M
by several honors he received after his retire-
ment. In 1977 he was awarded a special Distin-
guished Service Award for his research in paleo-
botany by the Paleobotany Section of the
Botanical Society of America, and in June 1985
a special symposium was held in his honor. This
symposium was sponsored by the Pacific Sec-
tions of the American Association for the Ad-
vancement of Science and the Botanical Society
of America and was titled “Evolution of the
Modern Vegetation of the Northern Rocky
Mountains." Several well-known scientists pre-
sented papers at this symposium and a special
plaque was presented to Dr. Becker. In 1981 he
received a Distinguished Achievement Award
from the Brooklyn College Alumni Association.

Becker was always characterized by an out-
going friendly nature, an interest in people, and
an enthusiasm for plants, living and fossil. His
extensive research papers, which students of pa-
leobotany continue to study and cite, stand as a
record of his persistent hard work in the field and
the laboratory. His research collections provide
an exceptional documentation of the history of
ancient plants that once lived in the area of pres-
ent-day southwestern Montana. To paleobota-
nists of now and of the future, Herman Becker
showed pages of new chapters in plant history
when he dug out and pried open the “books of
paper shale" at the Ruby River Basin. He will
be remembered as a friend, a good colleague, and
a research scholar.

PALEOBOTANICAL PUBLICATIONS OF
HERMAN F. BECKER

Coauthor: W. Donn occurrence and
description of the fossil d noA UE Science
115(2982): 214-215.

1955. A new method of! dli Ë di ving soft
matrix fossils. Asa Gray Bull., N.S. 3: 57-58.
An Oligocene Flora from the Ruby River Basin
in Southwestern Montana. Ph.D. Dissertation.
University of Michigan, Ann Arbor, Michigan.
A new species of Mahonia from the Oligocene
Ruby flora of southwestern Montana. Contr.
1. Paleontol. Univ. "Michigan 15(3): 33-38,

1952.

1956.

1959.

1960a. ES to Eopuntia one Cact. Succ. J.
(Los Angeles) 32: 28-2

The Ruby flora of u SS Montana. Bil-
lings Geological Society, Eleventh Annual Field
Conference Guidebook. September 7-10, 1960.

1960b.

1987]

1960c.

1960d.

196la.

196 1b.
1962a.
1962b.

1962c.

1962d.

1962e.

1962f.

1962g.

1964a.

1964b.

1965a.

1965b.

1966.

DILCHER — NORTHERN ROCKY MOUNTAIN SYMPOSIUM

The Tertiary Mormon Creek flora from the
Upper Ruby River Basin. Palaeontographica,
Abt. B, Paláophytol. 107(406): 83-126, pls. 18-

Additions to the Tertiary Ruby paper shale

flora of southwestern Montana. Bull. Torrey

Bot. Club 87: 386-396.

Oligocene plants from the Upper Ruby River

Basin in southwestern Montana. Mem. Geol.

Soc. Amer. 82: 1-127, 32 pls.

esas ay record of solar change. Ann.
New York Acad. Sci. 95: 684-687.

The resurrection of a landscape. Gard. J. New

York Bot. Gard. 12: 8-1

Roland W. Brown, 1893-1961. Gard. J. New

York Bot. Gard. 12: 71-72.

Two new species of es from the Grant-

Roses of the past. Gard. J. New York Bot.
Gard. 13: 103-105.

Roland W. Brown, i, Bull. Torrey
Bot. Club 89: 260-26

ee 2 P to C Vader det di
erus, comb. . Bull. Torrey Bot. Clu

301-307.
nls a Pp. 150-153 in American
ord Jr.

peepee eg ‘additions to the Oligocene
ora of southwestern Montana. Bull. Torrey
Bot. Club 91: 206-213

Paleobotanical exploits in Colorado and Kan-

sas. Gard. J. New York Bot. Gard. 14: 231-
233.
Stir abi and evolution. Nat. Hist.
74(2): 3

Rare ES "iom Montana (insects). Animals

(England) 8: 9

Additions to 2 revision of the Oligocene
Ruby paper shale flora of southwestern Mon-

tana. Contr. Mus. Paleontol. Univ. Michigan
20(5): 89-119, 6 pls.

1967a.

1967b.

1968.

1969a.
1969b.

1969c.

1971a.
1971b.

1972a.

1972b.

1973a.

1973b.

1976.

691

On western fossils and ss plants. Gard. J.
w York Bot. Gard. 17: 30-33.

Flowers, insects, and irure (1965). R

printed as pp. 392-399 in W. Knobloch (edi-

tor), Readings in Biological Science, 2nd edi-
tion. Appleton, New Yor

A hexasepalous calyx of the fossil Astronium
truncatum (Lesquereux) MacGinitie. Bull.
Torrey Bot. Club 95: 262-263, 1 pl.

ea pis cc forests. Amer. Forests
75(3): 1

Some Mtis Dad fossils. Amer. Forests 75(7):

Fossil plants ofthe Tertiary Beaverhead basins
in southwestern Montana. Palaeontographica,

Abt. B, Palàophytol. 167: 1-144, pls. 1-44

Fossil collecting. Gard. J. New York Bot. Gard.

21: sh

Digging in Montana’s evolutionary past. Amer.

ineft TIO 44-47, 57- Kun

Sanmiguelia, an enigma ounded. Palae-
Tire Abt. B, Paldophytol. 138: 181-
185, p

The md pw flora of the Upper Ruby
River Basin, southwestern Montana. Palae-
ontographica, Abt. B, Paláophytol. 141: 1—61,
pls. 1-

yet Tertiary gramineous fossil. Bull. Torrey
Bot. Club 100: 318-320.

The York Ranch flora of the Upper Ruby Riv-
er Basin, southwestern Montana. Palaeonto-

graphica, Abt. B, Palàophytol. 143: 18-93, pls.
13-40.

Like a phoenix from the ashes, Oregon's long-
vanished forests are revealed in fossils. Gard.

J. New York Bot. Gard. 26: 161-165

— David Dilcher, Department of Biology, Indiana
University, Bloomington, Indiana 47405, U.S.A.

LAND PLANTS OF THE NORTHERN ROCKY MOUNTAINS
BEFORE THE APPEARANCE OF FLOWERING PLANTS

CHARLES N. MILLER, JR.!

ABSTRACT

Bana £1 4 $ £, +h

flowering plants near the end of the Early Cretaceous
is spotty. Seas covered much of the land during the Paleozoic and ea

rly Mesozoic, but certain areas

were exposed, perhaps as islands. Evidence of Devonian vegetation from Beartooth Butte, Wyoming
includes rhyniophytes, iens zosterophylls, and lycopsids. Fossils from several different de-

posits show Late Mississippian to

The floras include lepidodendrids, Eus mites, fer

nnsylvanian e vegetation like that of coal swamps to the east.
, seed fern d PUn dne conifers present at

nder a generally arid but seasonally wet frost-free climate. It was into this environment that the first

u
flowering plants of the region migrate

The object of this paper is to give an overview
of the types of vegetation that occurred in the
northern Rocky Mountain region before the ap-
pearance of flowering plants there. This region
extends along the Rocky Mountains from central
Colorado and Utah north into southern Alberta
and British Columbia. The time span involved
extends from the Early Devonian Siegenian, about
385 million years ago, to the Early Cretaceous
Aptian, about 110 million years ago (Fig. 1).
During the intervening 275 million years both
the vegetation and the land it occupied changed
radically, keeping pace with similar vegetational
and land mass changes elsewhere in the world.

In fact, evidence of vegetation within the pres-
ent northern Rocky Mountain region is spotty
(Fig. 2). The best records of Paleozoic vegetation
come from deposits in Europe and eastern and
midwestern North America. Similarly, early Me-
sozoic, i.e., Triassic and Jurassic, floras are more
abundant and better preserved in Europe and the
southwestern United States than in the northern
Rocky Mountains. It isn't until the Late Jurassic
and Early Cretaceous that records within the re-
gion are as good as those found elsewhere. The
Paleozoic and Mesozoic sites that are known in
the northern Rocky Mountains, however spotty,
show that the types of plants in the region are
similar to those elsewhere and that they probably
were organized in similar communities which
occupied similar habitats. Thus, while the de-

posits within the region fail to provide conclusive
evidence, they nonetheless permit inference based
on our knowledge of nearby vegetation outside
the area.

DEVONIAN

During the Early Devonian, the only part of
the period for which we have evidence, the pres-
ent northern Rocky Mountains was an area of
lowlands and shallow seas along the western
margin of the early land mass Laurussia (Bam-
bach et al., 1980). This continent lay across the
equator, extending from about 10? south latitude
to about 40? north latitude, with the present
northern Rocky Mountains situated at about 12?
north latitude (Ziegler et al., 1981, fig. 7.5). Be-
cause of its position, the area had a tropical cli-

a

rm t
rainfall throughout the year. This, combined with
the location of the northern Rocky Mountains
in the Devonian along the western coast of the
continent where sharp changes in climate were
moderated by ocean currents, resulted in what
must have been ideal growing conditions.

Two deposits of Early Devonian plant fossils
have been recorded in the Beartooth Butte For-
mation in the northern Rocky Mountains. The
classic site at Beartooth Butte, Wyoming (Fig.
1 A) has been known for over 50 years. Five taxa
were described by Dorf (1933, 1934a, 1934b)

! Department of Botany, University of Montana, Missoula, Montana 59812, U.S.A.

ANN. MissouRi Bor. GARD. 74: 692-706. 1987.

1987]

MILLER—LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 693
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FIGURE 1. Index map and P time nia om showing the approximate locations and ages of m D

Early De vonian.— B. Cottonwood Gulch, Ea

anning Canyon
Shale, Early Pennsylvanian. — H r Formation, Early Pennsylvanian.—I Coy Formation, Late Penn-
sylvanian.— J. Spotted Ridge Flora, Pennsylvanian. — K. Cutler Formation, Permian. Wyoming Red Beds,
Triassic. — M. Morrison Flora, Utah, Late Jurassic. — N. Morrison Flora, Wyoming, Late Jurassic. — O. Morrison
T oe Late Jurassic.—P. Kootenay Formation, Late Jurassic.—Q. Douglas County, ei Late
Jurassic. — R. Kootenai Formation, Early Cretaceous. — S. Lower Blairmore, Early Cretaceous. — T. Shasta Flora,
Early yc —U. Burro C

anyon Formation and Cedar Mountain Formation, Early Cretaceous.

based on imprints of plant fragments. More re- are not isolated occurrences and that further

cently, Hueber (1972) and Tanner (1982, 1983)
described additional remains from the locality.

he deposit is regarded as Emsian in age, about
375 million years old.

Tanner (1982, 1983) described fossils from the
new Beartooth Butte Formation locality near
Cottonwood Gulch (Fig. 1 B) in northcentral Wy-
oming, about 50 km east of the original site (Fig.
1A). It is believed to be Siegenian in age, about
385 million years old, and thus somewhat older
than the Beartooth Butte locality. Locating a sec-
ond deposit in the region raises hope that these

search will turn up more deposits.

Five taxa were originally described by Dorf
(1933) from the Beartooth Butte locality. These
include Psilophyton wyomingense Dorf for flat-
tened branch systems bearing numerous spines
(Fig. 2A, B), (?) Psilophyton sp. for an elongated

bearing numerous lateral sporangia (Fig. 2C, D),
Hostimella sp. for flattened branch systems lack-
ing spines, and (?) Broggeria strobiformis for a

cylindrical strobiluslike structure. Hueber (1970)

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FIGURE 2. Examples of plant fossils from the Early Devonian d Butte Formation. A, B. Psilophyton

wyomingense Dorf. — A. Holot
5), x 2.5. —C. Rebuchia ovata (Dorf)
Hueber, USNM 1689
C, D from photographs courtesy of F. M. Hue

changed the name of Bucheria ovata to Rebuchia
ovata and later (1972) described its vegetative
axes linked with the fertile stems (Fig. 2C, D).
These clearly show the affinity of the plant with
the Zosterophyllophytina.

More recently Tanner (1982, 1983) described
additional taxa from both sites of the Beartooth
Butte Formation, bringing the total known to 14.
Six of these are based on vegetative axes with
attached fertile parts, five are known from veg-
etative axes alone, while one taxon is based on
a sporangial pair. Present are rhyniophytes, tri-
merophytes, zosterophylls, and the early lycop-
sids Drepanophycus and Leclercquia (Tanner,
1983). There are also two taxa of undetermined
botanical affinities.

Discussion. Even though the two localities
occur in the same formation, palynological evi-

otype, from Dorf (1933, plate V, fig. 3),

Hueber, USNM 168991B counterpart,

92, x1.3. (Figs. A, B reproduced by permission of the University of Chicago Press; Figs.
er.)

B. from Dorf (1933, plate V, fig.
s 3.—D. Rebuchia ovata (Dorf)

dence indicates that they may be separated by as
much as 20 million years. Such a gap is not ap-
parent in the megafossils or the fish fauna pre-
served at both, although the Cottonwood Creek
assemblage lacks definitive marker species (Tan-
ner, 1983).

n comparison with other Early Devonian as-

posed by Gierlowski (Ziegler et al., 1981) in which
the western North America zone is characterized
by relatively low generic diversity of two to six
taxa per assemblage and a lack or restriction of
zosterophylls. Including the unnamed sporangial

1987]

pair and two morphotypes of sterile axes, the
Wyoming assemblage has ten genera, three of
which belong to the Zosterophyllaceae.

Early Devonian plants share an architectural
plan that with few exceptions consists of rela-
tively naked aerial branches up to a meter tall,
which branch from rhizomes that bear adven-
titious roots or rhizoids (Gensel & Andrews,
1984). This is often interpreted as implying that
these plants grew in marshy habitat. Since many
of the known Early Devonian plant assemblages,
and particularly the Wyoming sites, are from near
equatorial paleolatitudes with equable temper-
atures and moisture, transpiration stress may not
have been enough to require actual growth in
standing water or even saturated soil as would
be found in coastal marshes. Occurrence at more
inland and upland sites may have been possible.

MISSISSIPPIAN

While movement of other continental plates
between the Early Devonian and Mississippian
was significant, the position of Laurussia, and
thus the present northern Rocky Mountains, was
relatively unchanged. There was an apparent in-
crease in mountain building activity west of Lau-
russia and an increase in the amount of shallow
seas and evaporite formation (Bambach et al.,
1980, fig. 11). Much of the northern Rocky
Mountains was an evaporitic basin in which
limestone was deposited. However, some land
was exposed within this zone of shallow seas, for
plant remains are recorded as petrifactions in
northwestern Utah (Fig. 1C; Tidwell & Jennings,
1986), as impressions and as pollen and spores
in northwestern Utah (Fig. 1D, E; Arnold & Sad-
lick, 1962; Schemel, 1950), and as pollen and
spores associated with exceptionally well-pre-
served fish and invertebrates at Bear Gulch (Fig.
1F) near Lewistown, Montana (Cox, 1986).

Deseret Limestone petrifactions. Tidwell &
Jennings (1986) described a new lycopod, Stans-
burya, from the Early Mississippian Deseret
Limestone near Grantsville, Utah, south of Great
Salt Lake. Six stem specimens were included in
a single carbonate concretion. The stems are about
1.4 cm in diameter and are dichotomously
branched. Scalelike leaves are tightly appressed
helically around the stems. None of the axes con-
tain secondary tissues, and this led the authors
to interpret the fossils as representing an her-
baceous plant. Much remains to be learned about
this new genus, but discovery of these fossils in

MILLER—LAND PLANTS OF NORTHERN ROCKY MOUNTAINS

695

the Deseret Li t t further search
may be rewarding.

Daggett County, Utah, Palynoflora. Schemel
(1950) reported several different types of pollen
and spores isolated from a coal sample from Late
Mississippian sediments in northwestern Utah
(Fig. 1E). Over 90% of the palynomorphs belong
to Densosporites. Similar spores have been found
in Pennsylvanian age sediments both dispersed
and in strobili of presumed arborescent lycopods
(Taylor, 1981). Several other types of spores oc-
cur in the Utah coal in small amounts and suggest
that sphenopsids, ferns, pteridosperms, and cor-
daitaleans grew in the vicinity.

Uinta Mountains Flora. The megafossil as-
semblage reported from a locality in the Uinta
Mountains in northeastern Utah (Fig. 1D) is a
small one based on poorly preserved imprints of
vegetative remains (Arnold & Sadlick, 1962). Ar-
chaeocalamites was identified from pith casts.
These show ribs continuous across the nodes
rather than alternate as in Calamites. The oc-
currence of Lepidodendron is patina by im-
prints of characteristic stems, leafy twigs, and
detached leaves. Defoliated axes, probably of seed
fern fronds, identified as Cau/opsis, are present,
as are detached fern or seed fern pinnules of
Fryopsis (formerly Cardiopteris). Lastly, Rhodea,
frond portions of a seed fern characteristic of the
Late Mississippian and Early Pennsylvanian, has
also been identified.

Arnold & Sadlick (1962) interpreted the plant
remains as representing streamside vegetation
with fragments washed into an estuary or bay by
river currents. The plant fragments are small and
are imprints with occasional flecks of carbon.
However, the siltstone matrix contains numer-
ous flecks of black material that presumably rep-
resent macerated plant debris (Arnold & Sadlick,
1962).

The assemblage is regarded as Chesterian in
age based on associated invertebrates (Arnold &
Sadlick, 1962) and is correlated with the Late
Visean or Early Namurian stages in Europe. The
plant assemblage compares well with floras of
similar age in the Appalachian trough.

Bear Gulch palynoflora. Palynomorphs are
also preserved in rocks that have given up ex-
ceptionally well-preserved vertebrate and inver-
tebrate fossils at Bear Gulch, near Lewistown,
Montana (Fig. 1F; Melton, 1971). The palyno-
morphs (Cox, 1986) include: Acanthotriletes,
Anapiculatisporites, Chaetosphaerites, Convolu-
tispora, Cyclogranulatisporites, Denosporites,

696

Endosporites, Lycospora, Procoronospora, Punc-
tatisporites, Raistrickia, Rotaspora, Savitrispo-
rites, Tripartites, and Verrucosisporites. The as-
semblage indicates a Chesterian age for the
deposit and correlates with the Namurian A stage
of Europe.

Furthermore, the palynoflora indicates that a
diverse assemblage of ancient vascular plants in-
habited the land that then surrounded the bay
in which the Bear Gulch sediments were depos-
ited. Although the correlation of the pollen and
spore species found here with the plants that pro-
duced them remains uncertain, other species of
some of these genera have been found in spo-
rangia. By analogy the following megafossil forms
may have been present: lepidodendrids (Denos-
porites and Lycospora), zygopterid ferns (Con-
volutispora), progymnosperms ee
calean fern
(Raistrickia), marattialean ferns ae
rites), and seed ferns (Punctatisporites and Schul-
zospora) (Kosanke, 1950; Millay & Taylor, 1979;
Taylor, 1981)

Discussion. The two palynofloras and the
Uinta Mountain megaflora combine to give solid
evidence that vegetation similar to that occurring
in eastern North America and Europe was pres-
ent in the northern Rocky Mountains during the
Late Mississippian.

rites),

PENNSYLVANIAN

By Pennsylvanian time Laurussia was con-
nected to Gondwana in the process of forming
Pangaea, but the present northern Rocky Moun-
tains remained at about 10° north latitude (Bam-
bach et al., 1980; Ziegler et al., 1981). More of
the present land surface was covered by shallow
seas than earlier, but sufficient land was exposed
to support vegetation characteristic of the period.

Early Pennsylvanian vegetation is well in evi-
dence in the Manning Canyon Shale flora from
northern Utah (Fig. 1G). Similar remains, though
not as extensive, are known from the Weber For-
mation in central Colorado (Fig. 1H). Late Penn-
sylvanian plant remains are known from the
McCoy Formation in central and northern Col-
orado (Fig. 11), and additional fossils have been
reported from a locality in central Oregon (Fig.
1J) of uncertain stratigraphic placement within
the Pennsylvanian.

Manning Canyon Shale. Plant fossils re-
covered from the Manning Canyon Shale in
northern Utah (Fig. 1G) are well-preserved im-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

prints that show considerable variety of plant
form (Tidwell, 1962, 1967; Tidwell et al., 1974).
While the strata were once thought to straddle
the Mississippian-Pennsylvanian boundary, the
flora best correlates with Early Pennsylvanian
Namurian B or possibly late Namurian A stages
of Europe.

The plants are like those of coal swamps to the
east. Arborescent lycopods were numerous (Tid-
well et al., 1974). Lepidodendron (Fig. 3D), Lep-
idophloios, and Si nown to have been
present based on stem imprints. In addition, re-
mains of the presumed reproductive organs of
these plants are represented by Lepidostrobus,
Lepidocarpon, and Sigillariostrobus, with Stig-
maria (Fig. 3C) representing the underground
parts and Lepidophyllum and Cyperites the fo-
lia

age.
Both Archaeocalamites and Calamites (Fig. 3A)
were present, as were three species of calamita-
lean foliage of the Asterophyllites type (Fig. 3B).
A poorly preserved Calamostachys has been
found, but details of its construction ane the type
of spores it produced remain unknow

Ferns and seed ferns are Miei by 11
species of foliage assigned to the genera Adian-
tites, Cornypteris, Crossopteris (Fig. 3F), Diploth-
mema, Mariopteris, Neuropteris, Rhodea,
Sphenopteridium, Sphenopteris (Fig. 3E), and
Zeilleria (Fig. 3G). In addition, the pollen organs
Aulacotheca and Telangium are present, as well
as seeds of Cornucarpus, Holcospermum, La-
genospermum, Rigbycarpus, and Trigonocarpus.

Cordaitales are represented by three species of
foliage, a Cordaianthus (Fig. 3H), and six species
of seeds

Thus, the Manning Canyon Shale flora rep-
resents vegetation similar to that of coal swamps
in the midwestern and eastern United States. The
setting is interpreted as a coastal swamp com-
munity at the margin ofa fresh or brackish water
embayment that underwent several transgres-
sive-regressive cycles (Tidwell, 1967). Some of
the taxa have stratigraphic ranges suggesting a
Mississippian age for the flora, but most indicate
a Pennsylvanian age correlating with the Na-
murian B stage in

Weber Formation, C. blado: Scattered plant

remains are known from the Weber Formation
in Colorado (Fig. 1H), which Read & Mamay
(1964) treated as Early Pennsylvanian Zone 6.
Lepidodendron johnsonii Arnold (1940) repre-
sents the westward extension of typical coal
swamp vegetation into central Colorado. The

Hd
°
ge]

1987] MILLER—LAND PLANTS OF NORTHERN ROCKY MOUNTAINS 697

A!
NOR
"ATTE,
d í

FIGURE3. Example Pennsylvanian plant ye from the northern Rocky Mountains. — A. Calamites, x 0.6.—
B. Asterophylites x1.0.—C. Stigmaria, x0.5.— vio d aire x0.95.—E. Sphenopteris, x0.7.—F. Cros-
sopteris, x 0.6.—G. Zeilleria, x 1.1. —H. Pais naira x0.9.—I. Walchia sp., UMMP 22131, x1.0.—J. Lecrosia

wouldii Arnold, UMMP 21005, x1.0. (Figs. A-H from “wm ss courtesy of W. D. Tidwell.)

698

small amount of coal in the Weber Formation
suggests, however, that the coal swamp was of
limited development and of short duration (Ar-
nold, 1940). Read (1933) reported imprints of
Trichopitys whitei from the Weber Formation
and in 1934 added Lepidostrobus, Stigmaria,
Calamites, Asterophyllites, Adiantites, Dactylo-
phyllum, Neuropteris, Sphenopteris, Diplomema,
Cordaites, and Cordaianthus.

McCoy Formation. Several localities within
the McCoy Formation in Colorado (Fig. 1I) have
yielded imprints of plant remains (Arnold, 1941).
Examples are: Calamites gigas Brongn., Odon-
topteris mccoyensis Arnold, Cordaites anguloso-
striatus Grand'Eury, Samaropsis hesperius Ar-
nold, Lecrosia gouldii Arnold (Fig. 3J), Walchia
stricta Florin, and Walchia sp. (Fig. 3I; Arnold,
1941). Arnold commented on the arid environ-
ment evidenced by the plants and by desiccation
features in the rocks. Thus, the fossils may rep-
resent forms that grew on drier sites than the coal
swamp represented by the plants from the Weber
Formation. The McCoy Formation is regarded
as Late Pennsylvanian Zone 10 by Read & Ma-
may (1964), and the drier setting may correlate
with similar events in coal swamps to the east
(Phillips et al., 1985) or may reflect some early
uplift of the ancestral Rocky Mountains (Pfef-
ferkorn & Gillespie, 1980).

Spotted Ridge flora, Oregon. The position of
this plant megafossil assemblage (Fig. 1J) within
the Pennsylvanian has not been determined; none
ofthe fossils found to date in this flora are reliable
stratigraphic indicators. The flora has the dis-
tinction of being the westernmost deposit of Pa-
leozoic plants in the conterminous United States
and is thought to have grown on an island arc
or microcontinent that was some distance west
ofthe North American craton (Pfefferkorn & Gil-
lespie, 1980). The following taxa have been re-
ported: an unnamed lepidodendrid branchlet,
Mesocalamites hesperius (Arnold) Mamay &

ead, M. crookensis Mamay & Read, Mesocal-
amites sp., Phyllotheca paulinensis Mamay &
Read, cf. Asterophyllites equisetiformis (Schlot-

Schizopteris trichomanoides Goeppert (Arnold,
1953; Mamay & Read, 1956; Read & Merriam,
1940). Of these, Mesocalamites hesperius, Phyl-
lotheca paulinensis, Pecopteris oregonensis, and
Dicranophyllum rigidum are the most common,
with the others represented in the collection by

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

one to a very few specimens (Mamay & Read,
1956). The large size of the collection rules out
sampling bias, and the numbers of specimens are
thought to reflect plant abundances with reason-
able accuracy (Mamay & Read, 1956).

Mamay & Read (1956) commented that the
plant remains occur in sandstone and mudstone
and show little evidence of transport. In fact,
some Calamites stems appear to have been pre-
served in growth position. Most ofthe sediments
are terrestrial in origin or some possibly origi-
nated in a marine or brackish water estuary. Thus,
the vegetation represented does not appear to be
directly comparable with any of the coal swamp
communities of the eastern and midwestern
United States. While the plants probably grew
ina moist habitat, the latter was not a coal swamp.

iscussion. In addition to the fossil assem-
blages described above, fragmentary plant re-
mains are known from subsurface samples in
Montana (Pfefferkorn & Gillespie, 1980), and a
small collection of fossils has been reported from
ee sediments in Wyoming (Sando et
, 1975). These indicate that the type of vege-
n present in eastern AE AS PUE
d land

e Penns yivanl

surfaces in the HUND northern mi Rocky! Mountain
region. Nonetheless, Pfefferkorn & Gillespie
(1980) recognized the vegetation of the region as
belonging to a distinct floral province, the Cor-
dilleran, of which Crossopteris (Fig. 3F) is a
characteristic endemic.

PERMIAN

The Permian was a pivotal period of geologic
time from the standpoint of plant evolution and
from the types of vegetation that occurred. To-
ward the end of the Pennsylvanian, eastern and
midwestern coal ta change in com
munity structure from domination by arbores-
cent lycopods to domination by marattiaceous
tree ferns (Phillips et al., 1985). This perhaps
foretells of changes to come with breakup of Pan-
gaea and the movement of continents poleward
during the early Mesozoic.

Deposits of Permian age plant remains in the
region are few, and none that are known provide
the quantity of well-preserved fossils typical of
earlier periods. One occurrence of Permian age
plant remains has been reported by Mamay &
Breed (1970) from the southern periphery of the
present northern Rocky Mountains. They re-
ported eight specimens from the Cutler Forma-

1987]

tion in southern Utah (Fig. 1K). From these are
identified Taeniopteris sp., Supaia rigida White,
Protoblechnum bradyi Mamay & Breed, and Cal-
lipteris sp. This represents the northernmost oc-
currence of Supaia. Protoblechnum bradyi is most
similar to a species from eastern Asia and sug-
gests the possible affinities of the Utah assem-
blage with the Asian flora (Mamay & Breed,
1970).

Similar remains are found in deposits in Texas,
New Mexico, and Arizona, and we can only spec-
ulate that exposed land surfaces in the present
northern Rocky Mountains supported similar
plants.

TRIASSIC

The only plant fossils reported from the Trias-
sic in our region are an Equisetum, two species
of Pterophyllum, two of Zamites and one Po-
dozamites (Berry, 1924). These come from the
so-called “red beds" in westcentral Wyoming
(Fig. 1L). These fossils are similar to those found
in the Chinle Formation in the Southwest (Ash,
1972; Daugherty, 1941) and suggest that a sim-
ilar vegetation occurred in the northern Rocky
Mountains.

JURASSIC

During the Early and Middle Jurassic, the pres-
ent northern Rocky Mountain region was cov-
ered by an inland sea in which marine limestone
and sandstones were deposited (Imlay, 1984; Sil-
verman & Harris, 1967). Uplift in the Nevadan
orogenic belt south and west of the present north-
ern Rocky Mountains resulted in a broad interior
basin of low relief that was poorly drained by
rivers northward and received sediments from
highlands to the southwest and from the Cana-
dian shield to the east (Brenner, 1983; Walker,
1974). The large size of the interior basin is re-
flected by the widespread deposition of shales
and sands of the Late Jurassic Morrison For-
mation and its equivalents, which are recognized
from southern Utah and Colorado north into
southern Canada (Walker, 1974).

Geologic studies suggest that a somewhat arid
but seasonally wet climate prevailed during the
Late Jurassic (Walker, 1974), supporting a sa-
vannalike vegetation with many shallow lakes
and swamps. There were probably dry periods
during which water tables dropped and exposed
accumulated debris to oxidation and decay. This

MILLER—LAND PLANTS OF NORTHERN ROCKY MOUNTAINS

699

may explain why plant fossils are not more abun-
dant in the region (Walker, 1974).

Plant fossils are known from scattered locali-
ties throughout the region (Delevoryas, 1969;
Tidwell, 1975), but large assemblages of remains
that reflect vegetation are known only from cen-
tral Montana (Fig. 1O) and southern Canada (Fig.
| P). South of the northern Rocky Mountain re-
gion are a number of occurrences of silicified
remains from the Late Jurassic Morrison For-
mation (Fig. 1M). Some noteworthy examples
are: Hermanophyton kirkbyorum Arnold (1962),
a Rhexoxylon-like log, Osmundacaulis wadei
Tidwell & Rushforth (1970), the only species of
petrified osmundaceous rhizome known from the
Jurassic of North America, as well as several
different coniferous woods (Medlyn & Tidwell,
1975, 1979).

Within the region there are reports of foliage
imprints of the two cycadophytes Nilssonia ni-
gricollensis Wieland and Zamites arcticus Goep-
pert (Knowlton, 1916) and of petrified cycadeoid
trunks (Delevoryas, 1960; Ward, 1905), both from
Wyoming (Fig. 1N).

The most extensive Late Jurassic flora (Table
1) in the northern Rocky Mountains is that pre-
served in shales of the Morrison Formation in
central Montana (Silverman & Harris, 1967;
Brown, 1972). Plant remains have been found at
six localities, one near Lewistown and the rest
south of Great Falls near the town of Belt (Fig.
10).

While three of the six localities in central Mon-
tana are close to one another, the most distant
sites are over 170 km apart. Zamites arcticus
(Fig. 4A), Nilssonia cf. compacta (Fig. 4D), Po-
dozamites lanceolatus (Fig. 4G), and Pityophyl-
lum lindstromii (Fig. 4F) occur at all six localities;
and these taxa probably represent the dominants
ofthe regional vegetation. Pagiophyllum sp. (Fig.
4F) and Cladophlebis virginiensis (Fig. 4C) occur
at five of the sites and were thus also widespread.
Sagenopteris elliptica (Fig. 4E), leaflets of a cay-
tonialean seed fern, and Coniopteris hymeno-
phylloides (Fig. 4B), foliage of a dicksoniaceous
fern, are locally abundant at certain localities.
Nowhere are remains of conifers as abundant as
those of cycadophytes. Ginkgophytes (Brown,
1975) are present but rare. Thus, the vegetation
appears to have been a relatively open one with
scattered conifers, more closely spaced cycado-
phytes and tree ferns, occasional ginkgophytes,
and an understory of ferns.

One of the localities clearly represents material

[Vor. 74

ANNALS OF THE MISSOURI BOTANICAL GARDEN

—— LÀ. |
mi P"
fm "er

m. 7

c= n.

k^
* e

FIGURE 4. Typical apt pa from the Late Jurassic Morrison Aeae cer —A. ud eel
—B. C Mp iý Shap e (Brongn. ard, UMPM 1092,
—D. Nilssonia cf. NE (Phill.) seen DUM

G
Cladophlebis virginiensis a. UMPM

1987]

washed in from surrounding areas, and others
provide evidence of an extensive swamp and
swamp margin community stable enough to re-
sult in a thin but mineable seam of low grade
coal (Silverman & Harris, 1967). While coal for-
mation indicates deposition of plant debris in
water having oxygen low enough to retard deg-
radation of remains by aerobic bacteria, some
areas of the swamp were above water level and
supported growth of marchantioid liverworts.

A similar vegetation extended northward into
Canada (Fig. 1P) where megafossils are pre-
served in sediments of the Kootenay Formation
(Bell, 1956), and microfossils are known from
the Upper Vanguard Formation (Pocock, 1962).
However, there is no reason to assume that a
given swamp extended over that distance. Rath-
er, there were probably many swamps that formed
and disappeared over the several million years
represented by the Late Morrison Formation and
the Kootenay and Upper Vanguard formations
in Canada. Thus, it is doubtful that direct cor-
relation of these strata is possible.

Furthermore, the Canadian flora appears more
diverse than that in Montana (Bell, 1956). While
some of this apparent diversity is simply more
species of the same genera and is probably due
to taxonomic splitting, several genera occur in
the Canadian flora that are absent in Montana.
These are the fern K/ukia, the seed fern Czeka-
nowskia, the cycadophytes Ctenis, Pseudoctenis,
and Prilophyllum, and the ginkgophyte Baieria
(Bell, 1956).

There is another Late Jurassic flora close to
but outside the northern Rocky Mountain region
(Fig. 1Q). Over 60 taxa have been reported from
20 localities in Douglas County, Oregon (Ward,
1905), and essentially no research has been done
on this assemblage since it was treated in the
Status of the Mesozoic floras of the United States,
2nd paper over 80 years ago. A modern study of
these fossils has great potential to advance sub-
stantially our knowledge of Jurassic vegetation.

EARLY CRETACEOUS

The broad interior basin in which sediments
of the Late Jurassic Morrison Formation were
deposited persisted into the Early Cretaceous.

MILLER —LAND PLANTS OF NORTHERN ROCKY MOUNTAINS

701

BLE l. Fossil plants from the Morrison Forma-
tion of central Montana

Bryophytes
Marchantiolites sp.
Ferns and fern allies
Equisetum laterale Phillips
Hausmannia fisheri (Knowlton) Oishi & Yamasita
Coniopteris Voies oeil (Brongn.) Seward
Adiantites montanensis aah Brown
Cladophlebis albenia ies Bel
C. heterophylla Fon
C. virginiensis ade waq
Cycadophytes
Nilssonia cf. compacta (Phillips) Bronn.
Zamites arcticus Goeppert
Cycadolepis spp.
Weltrichia sp.
Anomozamites
Sagenopteris dd Fontaine
Ginkgophytes
Ginkgoites cascadensis Brown
G. pluripartita (Schimper) Seward
Conifers
Pagiophyllum sp.
Podozamites lanceolatus (Lindley & n Braun
e lindstromii (Heer) Nathors
Pityocladus
E sp.

Sedimentation was not continuous, however. It
ceased for a time and then resumed. This ces-
sation resulted in a disconformity inpia by a
layer of coarse sandstone which occurs through-
out the region (Walker, 1974). The Teone
is regarded as the boundary between the Jurassic
and the Cretaceous (Brown, 1956).

Sometime after the resumption of deposition,
plant remains were deposited. In central Mon-
tana this was not until what is believed to be the
Aptian stage of the Early Cretaceous (LaPasha &
Miller, 1984) even though invertebrates in the
basal sandstone of the Kootenai Formation in-
dicate a Neocomian age for those strata (Walker,

The plant-bearing layers of the Kootenai For-

—
1093, —E. Sagenopteris elliptica Fontaine, UMPM 1095, x2.7.—F. Pagiophyllum sp. (Pg) and Pityo-
prar de seas evi Nathorst (Pi), UMPM 1096, x1.5.—G. Podozamites lanceolatus (Lindley & Hutton)

Braun, UMPM 1097, x1

702 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

B d

FIGURE 5. Typical plant fossils from the Early apr ep PIER Formation. — A. Klukia canadensis Bell
(K) and Sphenopteris latiloba Fontaine (S), UMP 1.6.—B. Sagenopteris williamsii (Newberry) Bell,
UMPM 1099, x1.3.—C. Cladophlebis nA Fontaine UMPM 1100, x3.5.—D. Sagenopteris mclearnii

1987]

mation represent deposition of sediments and
plant remains in poorly drained swamps that
formed from infilling of lakes. Evidence suggests
that the swamps contained standing water
throughout the year. Some areas were above water
level as indicated by the occurrence of liverworts,
which were probably preserved in place (Brown
& Robison, 1974, 1976; LaPasha & Miller, 1984,
1985). However, the water table at these sites
was high enough to keep the soil saturated.

Certain plants (Table 2) probably grew in the
swamps in which their remains were deposited.
This is indicated by the occurrence of large, lo-
cally abundant remains in low diversity assem-
blages in a wide variety of sediment size classes
(LaPasha & Miller, 1984). Fossils that fit this
category are Marchantiolites, Diettertia, Coniop-
teris simplex (Fig. 5F), Acrostichum, Sagenop-
teris williamsii (Fig. 5B), and Athrotaxites berryi
(Fig. SE). Remains of other plants are generally
found as abundant, small fossils occurring in high
diversity assemblages and in narrow ranges of
sediment size classes, particularly coarse sedi-
ments suggesting slight transport. Examples of
these are Coniopteris, Cladophlebis (Fig. 5C),
Sphenopteris (Fig. 5A), Sagenopteris elliptica, S.
mclearnii (Fig. 5D), Ginkgo (Brown, 1975), Ela-
tides (Fig. SH), and Elatocladus (Fig. 5G). The
same reasoning, combined with the relative scar-
city of fossils, suggests that remains of Equise-
tum, Klukia (Fig. 5A), Hausmannia, Zamites,
and other bennettitaleans were transported some
distance, but exactly how far is unknown

While certain species are common to both the
Morrison flora and the Kootenai flora (Tables 1,
2), the two represent different types of mm
The Kootenai flora appears to have been dom
inated by conifers, especially Athrotaxites, vdih
bennettitaleans relatively rare iler &

84)

certain localities, the stratum of vegetation it oc-
cupied remains unknow

The Kootenai Purus is widespread in
Montana and has equivalents in adjacent states,
but plant fossils are known in abundance only
near Great Falls. Permineralized cycadeoid trunks
have been reported from the Burro Canyon For-

MILLER — LAND PLANTS OF NORTHERN ROCKY MOUNTAINS

703

ABLE 2. Fossil plants from the Kootenai Forma-
tion of central Montana.

Bryophytes

Ar. REX rf. FU

y) Brown & Rob-

À

ison
Megzgeriites montanensis LaPasha & Miller
Diettertia montanensis Brown & Robison

Sphenopsids
Equisetum montanensis LaPasha & Miller
E. cascadensis LaPasha & Miller

Lycopsids
Minerisporites sp. A
Minerisporites sp. B
Pteropsids
Coniopteris keri gr n ) Seward
C. simplex (Lindley & Hut
Klukia canadensis Bell
Hausmannia montanensis LaPasha & Miller
Acrostichum longipennis Fonta
Cladophlebis constricta Aes
C. inclinata Fontaine
C. oblongifolia Fontaine
C. oerstedii (Heer) Seward
C. virginiensis Fontaine
papell dining Bell
S. latiloba F
S. mclearnii m
S. JP e Dunker
Arcellite.
I cascadensis LaPasha & Miller

Cycadophytes
Sagenopteris elliptica Fontaine
i Be

S. williamsii aput] Bell

Zamites arcticus Goepp

Pseudocycas douglasii Linda & Miller
Ginkgophytes

Ginkgo pluripartita (Schimper) Heer
Conifers

Athrotaxites berryi Bell

Elatides curvifolia (Dunker) Nathorst

Elatocladus dunnii Miller & LaPasha

. montanensis Miller & LaPa sis
Masculostrobus montanensis Miller & LaPasha
Conites sp.

—

Berry, UMPM 1101, x2.3.—E. Athrotaxites berryi Bell, UMPM 894a, x1.8.—F. Coniopteris simplex PU

& Hutton) Harris, UMPM 1102, x1.3.—

Elatocladus pres Miller & LaPasha, holotype UMPM 914

x 1.7.—H. Elatides curvifolia (Dunker) Nathorst, UMPM 907,

704

mation (Fig. 1U) in Colorado (Brown, 1950).
Frenelopsis varians and Tempskya are known
from the Cedar Mountain Formation (Fig. 1U)
in Utah (Tidwell et al., 1976), although the upper
parts of both of these formations are regarded
by some as younger than the Kootenai Forma-
tion (Ash & Read, 1976; Tschudy et al., 1984).

An extensive assemblage of plant remains (Fig.
1S) is known from the Lower Blairmore For-
mation of Alberta (Bell, 1956), and the Kootenai
flora compares best with this assemblage. The
Lower Blairmore flora is of special interest be-
cause it contains the earliest convincing flower-
ing plant megafossils in the region, leaf imprints
identified as Sapindopsis angusta (Heer) Seward
& Conway (Bell, 1956). While it is possible that
the sites these leaves come from are in fact some-
what younger than the Kootenai flora, it is also
possible that the leaves indeed represent some
of the earliest flowering plant migrants into the
region.

Another extensive Early Cretaceous assem-
blage is known from outside the northern Rocky
Mountain region, but it is nonetheless of special
interest. Like the Jurassic plants from Douglas
County, Oregon, those from the Shasta Series in
northern California (Fig. 1T) represent a large
and diverse assemblage that has received no fur-
ther work since the flora was described over 80
years ago (Ward, 1905). These fossils should be
the subject of a modern study.

SUMMARY

This report deals with the record of terrestrial
vegetation in the northern Rocky Mountains prior
to the appearance of flowering plants. The span
of time involved covers the Early Devonian 385
million years ago to the Early Cretaceous about

inal and a second collecting site in the Early De-
vonian Beartooth Butte Formation of northern
Wyoming has added considerably to the forms
known. Represented are rhyniophytes, trimero-
phytes, zosterophylls, and early lycopsids. These
compare well with plants of similar age known
from eastern North America and western Eu-
rope, making the Beartooth Butte assemblage one
of the better known Early Devonian floras of the
world.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Several deposits of Late Mississippian and
Early Pennsylvanian fossils show that vegetation
like that of coal swamps of Appalachia and the
North American interior occurred in the north-
ern Rocky Mountain region as well. The latter
vegetation was composed of arborescent lycop-
sids, arborescent sphenopsids, tree ferns, seed
ferns, and cordaites like that of the Interior-Ap-
palachian Province to the east, but the presence
of unique forms is grounds for treating the west-
ern vegetation in the distinct Cordilleran Prov-
ince. Similarly, because of species unique to it,
the Spotted Ridge flora of Oregon is treated as
representing the Oregonian Province.

ur record of vegetation is spotty in sediments

of the Permian, Triassic, and Early and Middle
Jurassic in the northern Rocky Mountains. By
comparison, evidence of Late Jurassic and Early
Cretaceous vegetation is relatively abundant and
informative. Much of the present northern Rocky
Mountains was then a broad interior basin of
low relief that supported an open savanna type
of vegetation interspersed with shallow lakes,
swamps, and rivers. During the Late Jurassic co-
nifers appear to have been widely scattered, cy-
cadophytes and ferns were more closely spaced,
and ginkgophytes and caytonialean seed ferns
were more rare and occasional. This vegetation

were equable, and there is no evidence of freezing
conditions.

While the interior basin persisted into the Ear-
ly Cretaceous and was the site of deposition of
plant fossils, the vegetation represented had an
entirely different aspect. Conifers were abundant
and dominated the vegetation, with cycado-
phytes rare and widely scattered. Ferns formed
the understory and caytonialean seed ferns were
locally abundant. Rainfall and temperatures were
like those of the Late Jurassic.

There is no evidence of flowering plants in the
flora of the Early Cretaceous Kootenai Forma-
tion in Montana, which is regarded as Aptian in
age. However, imprints of flowering plant leaves
are present but rare in sediments of presumably

leaves are
somewhat younger than Aptian, as some believe,
these imprints represent some of the first flow-
ering plants in the northern Rocky Mountain

3
*
=
©
aN
°
B
oO
3
Lond
n
[e]
°
5
e
ie
3
=

ge
[ad
=
[4]

Thus, de their

in the northern Rocky Mountains toward the end

1987]

of the Early Cretaceous, and it is important to
know about the vegetation that occurred on the
land as they migrated into the region. Unlike
problems confronting the first land plants in the
Early Devonian that colonized relatively barren
land, the establishment of flowering plants re-
quired their successful competition with vege-
tation that was already in place. This is indeed
a remarkable feat; however, this survey shows
that replacement of one form of vegetation by
another has been the rule throughout the 275
million years discussed in this paper.

LITERATURE CITED

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Some Paleozoic plants. from central
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o

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[Vor. 74

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Evolution, Volume II. Praeger Publishers, New
York.

ANGIOSPERMS OF THE NORTHERN ROCKY MOUNTAINS:
ALBIAN TO CAMPANIAN (CRETACEOUS)
MEGAFOSSIL FLORAS'!

DAVID R. CRABTREE?

ABSTRACT

Synchronous first occurrences of pollen and leaves indicate that angiosperms entered the Northern
Rocky Mountain (NRM) region during the Middle Albian, approximately 8 Ma later than such shifts
in a from southern Laurasia. The earliest angiosperm pollen and leaf flora Mid s to the
P ac sub-Zone IIB in that it poda a pretricolporate palynoflora and a comparable grade of
e ee based on leaf rank. The Albia un
tanophyll, Protophyll, and Pentalobaphyll mo rphotypes. Sapindophylls are common an
flora contains early North American occurrences of the widespread Upper Cretaceous leaf form genera
Trochodendroides and Cineman Pentalobaphylls (Araliaephylls) appear early and assume a

numerical dominants. Pinnate palms are present by the early Campanian. Higher-level taxa present
in the ji cdi in the Ear ly y Campanian include Magnoliales, Laurales, Chloranthales, Nymphaeales,
Menisperm Platanaceae, H C agales, Rosidae
and sine Dilleniidae.

Mountain (NRM) region based on the megafloral
record. The refinement of descriptive terminol-
ogy for leaf architecture (Hickey, 979;
Dilcher, 1974) and elucidation of the phyloge-
netic significance of leaf morphology (Hickey &
Wolfe, 1975), along with the U.S. National

In 1874 Leo Lesquereux stated:

he plants of the Dakota group, as known mostl

of our time ... and the evident likeness of their : .
facies . . . strikes the paleontologist and may lead Cleared Leaf Collection and the U.S. Geological
him into error... for, really, when we enter into Survey Cleared Leaf Collection, have provided

the framework for the present stu

a more detailed analysis of these Cretaceous leaves, y.
The rich Cretaceous leaf fossil deposits of the

we are by and by forcibly impressed by t

strangeness of the characters ... which seem at
riance With any of those recognized anywhere western interior of North America have long been
in the floras this flora recognized for their importance to angiosper

paleobotany.

of o
does not leave see pew ay tie E to my
ind. (during the late 1800s and early 1900s) mistak-

In this paper I summarize and evaluate the
evidence relevant to the paleoecology and evo-
lution ofearly angiosperms in the Northern Rocky

enly placed many leaf species into Recent genera.
Even investigators who realized their predica-
ment (see Lesquereux quote above) were ham-

!T xc L. J. Hickey, J. U. McClammer, J. A. Wolfe, S. L. Wing, G. R. Upchurch, L. S. Pierce, B. H. Tiffne
and K. R. Johnson for their interest in and Pss of early angiosperms. C. N. Miller provided ie. ic
for the photography and technical detail, as well as much encouragement. The Department of Botany, University
of Montana, funded the preparation of parts of the manuscript. NSF Doctoral Dissertation Improvement Grant
BSR 8400303 funded visits to museums. F. Heuber, S. Wing, and J. Ferrigno made the collections of the U.S.
National Museum available for study. T. Bolton assisted me at the Geological Survey of Canada, Ottawa, and
I. Birker helped at the Redpath Museum, Montreal. C. Blount, Department of Geology, Idaho State University,
loaned me a collection of fossils from the Wayan Formation. C. A. LaPasha made many of the thin sections of
wood from the Vaughn Member. I thank H. Chambers for assistance with Tables 1, 2, and 3 and M. Crabtree
for assistance with some of the figures. Fieldwork i in 1982 was supported in part by a Sigma Xi Grants-in-Aid
of Research to the Author

? Department of Botany, University of Montana, Missoula, Montana 59812, U.S.A.

ANN. Missouni Bor. GARD. 74: 707-747. 1987.

708

pered in their attempts to interpret these leaves
by the absence of appropriate operational para-
digms. Throughout this report, genera of doubt-
ful occurrence in the Cretaceous are marked with
quotation marks. Where possible I have provid-
ed reappraisals of the affinities of these leaves in
my discussions of Cretaceous leaf morphotypes
and individual floras.

I cover the period of about 30 million years
from the apparent advent of angiosperms during
the Middle Albian to the middle of the Cam-
panian. This includes the latter part of stage one
of angiosperm evolution (Vakhrameyev, 1982),
during which the group became established in
restricted areas of a predominantly gymno-
sperm-dominated vegetation; and it includes
most of stage two, in which angiosperms radiated
explosively and displaced gymnosperms from
many parts of the wor

The floras reported here come from British
Columbia, Washington, Alberta, Montana, Wy-
oming, and Idaho (Figs. 1, 2).

Coniferous communities were the dominant
form of vegetation in the world, at least through
the Early Cretaceous (Hughes, 1969, 1976; Kras-
silov, 1981; Miller, 1977; Penny, 1969). Lower
Cretaceous conifers from the NRM region are
regarded as belonging to the Taxodiaceae, Ar-
aucariaceae, Podocarpaceae, Pinaceae, and Chei-
rolepidiaceae (Bell, 1956; Miller & LaPasha,
1984; Singh, 1964, 1971). With the exception of
the Cheirolepidiaceae, which declined during the
Albian, and the addition of the Cupressaceae
which appear to have diversified during the mid-
dle and Late Cretaceous, these groups form the
nucleus for the development in the Late Creta-
ceous of the North Pacific refugium (Vakhra-
meyev, 1982). Compressions and impressions of
foliage and cones of Taxodiaceae, Cupressaceae,
and Araucariaceae occur abundantly in coarser
sediments throughout the region in the Upper
Cretaceous (Berry, 1929a; Knowlton, 1905).

Ginkgophytes decrease in the region during the
Albian, the latest flora in which they form an
important component being the Lower Blair-
more (Dawson, 1886; Bell, 1956), although fo-
liage is reported from the Cenomanian Dunve-
gan Formation in northern Alberta (Bell, 1963),
and Salisburia (cf. Ginkgo) seeds are reported as
late as the Campanian Belly River flora (Dawson,
1886). The Ginkgophyte decline is delayed in the
extreme northern latitudes. Although northslope
Alaskan floras exhibit a notable decline in the
Late Albian (Scott & Smiley, 1979), the group

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

remains important at least into the early Se-
nonian in the Atane beds of Greenland (Seward
& Conway, 1935), and into the Campanian in
the Chignik Formation of central Alaska (Hollick
& Martin, 1930). Ginkgo persists into the Paleo-
gene in the Ft. Union Formation of Montana
and the Willwood Formation of Wyoming (pers.
obs.).

Fern communities of the Recent families Glei-
cheniaceae, Schizaeaceae, Dicksoniaceae, and

mundaceae, and the extinct family Temp-
skyaceae occur abundantly in certain facies, a
characteristic feature of the middle Cretaceous
that Krassilov (1981) interpreted as evidence for
extensive fern marshes. Pteridophytic commu-
nities colonized the widespread upper Albian and
Cenomanian volcanic ash flats, occasionally being
preserved in situ as in the Albino Member of the
Mowry Shale in southwestern Montana (Crab-
tree, 1983; Vuke, 1982). Ferns such as Glei-
chenia, Anemia, Sphenopteris, Cladophlebis,
Tempskya, Coniopteris, and Onychiopsis (but see
Skog, 1985) are found throughout the region in
the Albian and Cenomanian (Andrews, 1948;
Andrews & Kern, 1947; Andrews & Pearsall,

1941: Ash & Read, 1976; Bell, 1956; Knowlton,
1917; Read & Brown, 1937; Seward, 1924). Post-
Cenomanian floras show a decreasing represen-
tation of these genera, concomitant with increas-
ing Polypodiaceae s.l. (pers. obs.).

Cycadophytes such as Nilssonia, Zamites,
Pseudoctenis, Ctenis, and Otozamites decline
rapidly after the Aptian. However, certain species
persist in the Cenomanian Dunvegan flora (Bell,
1963) and the Turonian upper Frontier flora
(Berry, 1929d), and Zamites albertensis Berry is
abundant in the Campanian Allison flora of Al-
berta (Berry, 19292). Cycadophytes also persist
in the Late Cretaceous of Alaska (J. Wolfe, pers.
omm.).

Sagenopteris, foliage of pteridosperms of the
Caytoniales, is abundant during the Aptian and
early Albian in the region (Bell, 1956; LaPasha
& Miller, 1985) but is only rarely reported from
the later Albian or from the Upper Cretaceous
(see Winthrop flora in this report).

The relative abundance of major plant groups,
and the entry of angiosperms into the region dur-
ing the Middle and Upper Albian, can be seen
in the histograms depicting the changing com-
position of the flora during the Middle and Up-
per Albian (Fig. 3). One histogram (Fig. 3A) is
based on published megafossil floras and my own
observations of unpublished floras. The other

e

1987] CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 709
x =
° %
= -
60°N
BRITISH
COLUMBIA ALBERTA
A
, O
D «on
MONTANA
WASHINGTON
459N
250 km
Qn WYOMING

IGURE l. Geographic locations of plant megafossil collections in the northern Rocky Mountain region.
Collection sites are numbered in alphabetical sequence. Names of sites appear in Figure 2. Appendix I contains

additional information on locations.

(Fig. 3B) is based on published palynofloras. Per-
centages shown in the histograms were calculated
by counting species of each major plant group
within individual floras. These raw count totals

were summed and a percentage calculated for
each chronostratigraphic subdivision of the AI-
bian. Megafossil floras show a bias towards pres-
ervation of vegetation adjacent to the deposi-

710 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74
o
NT
oF u PALEOBOTANICAL
ON < STAGE COLLECTION
a
73 =
" 1 ALBINO
c 2 ASPEN
- 3 BADHEART
P (13) 4 BEAVER MINES
ae. 5 BLACK EAGLE
I 6 COMMOTIO
_ A 6a LOWER PART
©) ae MEMBER
B (21) 6b UPPER PART
wp (15) GATES MEMBER
GS) 6c BOULDER CREEK
— SANTONIAN MEMBER
(3) 7 CROWSNEST
875 — 8 DUNVEGAN
885 CQNIACIAN 9 AGLE
= © 10 FALL RIVER
— TURONIAN 11 FRONTIER
— lla LOWER PART
= UR FRONTIER
_ M FORMATION
mo — M 11b UPPER PART
4 FRONTIER
- — i ORMAT I
on (23) (2) 12 JACKASS MOUNTAIN
= 13 JUDITH RIVER -
"c — C1) (22)? LOMA
= UPPER (10) 14 KINGSVALE
-—— 15 MILK RIVER
E 16 MILL CREEK
_ A DEK Q9) 17 PASAYTEN
"B s. 18 — PRE-MUDDY
WE. - D (16) 19 SUMMIT
ín (c) 20 SUN RIVER
E AOOO) 21 TWO MEDICINE
_ " (Ba) Q 22 + WAYAN
7 23 WINTHROP
LOWER
13 —
FiGuRE 2. Chronostratigraphic positions of plant Wicow collections in the sels Rocky Mountain

region. Stage boundary dates according to Harland et al. (1982).
Hickey & Doyle (1977). chy aan Voie of aida is bow on my in
weste h American chronostra

Substage boundaries are base nterpretation of

ye otomac Zones from
terpretation of the literature.
tigraphy. Collections

Ages are in million

of uncertain age are indicated n denis marks (see text ud rises I for discussion).

tionalenvironment; thus Figure 3A approximates
the components of the lowland vegetation. Pal-
ynofloras used in Figure 3B were recovered most-
ly from marine and marginal marine environ-
ments, and thus more closely reflect the regional
flora. From Figure 3A it is apparent that dra-
matic changes occurred in the lowland flora dur-
ing the Middle and Upper Albian. In these en-
vironments angiosperms increased at the expense
of ferns. Because this trend is not evident in Fig-
ure 3B, it is apparent that this early angiosperm
radiation had its greatest impact on lowland
vegetation.

PALEOGEOGRAPHIC AND GEOLOGIC SETTING

The northern Rocky Mountain region during
most of the Cretaceous occupied a strip of land
some several hundred kilometers wide, situated
between the Pacific geosyncline/island arc to the
west and the mid-continent epeiric sea to the east
(McGookey et al., 1972; Williams & Stelck, 1975).
Paleolatitudes may have varied up to ten degrees
from the present range of 41?N in southern Wy-
oming to 60°N in northern Alberta and British
Colombia. The paleocontinent positions of Smith
et al. (1981) indicate that northern Alberta has

1987]

rotated 8—10° to the south since the Cretaceous,
and southern parts of the region are presently
approximately 2—5? farther south.

The first major expansion of the Cretaceous
epeiric sea began in the Late Aptian and contin-
ued into the Albian, attaining its maximum ex-
tent in the late Middle Albian when it reached
unbroken from the Arctic Ocean to the Gulf of
Mexico (McGookey et al., 1972; Vuke, 1984;
Williams & Stelck, 1975). The fluctuations of the
sea initiated a famous series of transgressive—
regressive clastic cycles of deposition that con-
tinued until the end of the Cretaceous (Waage,
1975). Geological formations in the region tend
to parallel the north-south trending western
shoreline of the seaway. Cretaceous sediments
accumulated in terrestrial and marine environ-
ments to great thicknesses: up to 3,500 m in
southwest Wyoming (Rubey et al., 1975) and
2,134 m in northwest Montana (Rice & Cobban,
1977). The source areas in present British Co-
lumbia, western Alberta, Idaho, Washington,
Oregon, western Montana, and western Wyo-
ming are inferred to have had considerable relief
n order to account for the great thickness of
accumulated sediments. Orogenic activity in the
region was of long duration beginning in the Late
Jurassic and Early Cretaceous with the uplift of
the Nevadan Orogeny. The main thrusting of the
Sevier Orogeny occurred throughout the Creta-
ceous, and the uplift of the Laramide Orogeny
took place during and subsequent to the Cam-
panian (Gilluly, 1963; Nichols et al., 1985).

PALEOCLIMATE

Generalized interpretations of paleoclimate,
which include the North American Cretaceous,
have been published by Frakes (1979), Habicht
(1979), Lamb (1977), Schwarzbach (1974), and
Vakhrameyev (1978). They recognized a general
humidification in Laurasian climate from the Ju-
rassic into the Cretaceous. This can be correlated
with the widespread inundation of continental
crust, a worldwide phenomenon in the Creta-
ceous, and the opening of the Atlantic Ocean.

Assessment of precipitation patterns and
amounts during the Cretaceous in the NRM re-
gion is difficult with our present knowledge. Par-
rish et al. (1982) mapped the Cretaceous precip-
itation in the region as low to moderately low
largely on account of the presumed orographic
effect of the Rocky Mountains. However, they
noted that the presence of important high lati-
tude coals in the region, especially in the Late

CRABTREE— NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

711

Cretaceous, may argue for heavy seasonal pre-
cipitation, perhaps a monsoonal effect

Growth rings in fossil wood can be used to
assess environmental influences on plant de-
velopment (Creber, 1977; Creber & Chaloner,
1984, 1985; Fritts, 1976). Presence and degree
of development of latewood and growth rings is
generally correlated with available moisture.
However, there are many examples of growth
rings in trees living in regions of high rainfall.
For instance, Agathis from Fiji, where annual
rainfall is 2,000-6,000 mm, have well-developed
growth rings (Ash, 1985). Nonetheless, I know
of no exception to the — of growth rings
in the wood of trees gro climates,
and conversely, the absence of growth rings al-
pt per-
haps in certain warm-climate eun commu-
nities).

I examined 27 specimens of six species of co-
niferous woods from the Late Albian Vaughn
Member of the Blackleaf Formation in northwest
Montana in thin section for the presence and
configuration of growth rings. Transverse sec-
tions could be grouped into three broad cate-
gories:

Category A. Woodsexhibiting no growth rings.

Category B. Woods exhibiting broad (3-5 mm)

and consistent growth rings with

nsiderable latewood.

Woods exhibiting rings of variable

width, including broad rings (up to
m), pseudorings, and consid-

erable latewood.

seacnnyal

Category C.

The Vaughn Member has been interpreted as

a deltaic swamp deposit (Cannon, 1966) and was
situated at about 55°N paleolatitude (Couillard
& Irving, 1975). Cold-seasonality did not exist
during the Albian at this paleolatitude. Season-
ality as the result of low light during winter
months may be a factor at 50°N, although I know
of no literature on the subject. Broad-leaved ev-
ergreen forests are known to have extended to
65°N during the Cretaceous (Wolfe & Upchurch,
1986). Incomplete seasonal leaf-drop, formation
of latewood, and cambial quiescence might well
e ] with the northern extensions of this
forest. Among several possible explanations for
the mixed assemblage of woods, the following is
suggested as the most plausible: the trees were
growing in a warm, seasonally dry climate, with
some species (Category A) living in areas of year-
round groundwater. The pronounced develop-

712 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

$
4 «e ev? «e
` P o0 Y "d "Y
e
«e e

r of
e N
eo ue ot Nt ANTHOPHYTES ce «e "d CONIFEROPHYTES

lower

pp
Albian

m

°

©
=
a
&
LJ

E Lun

Lower 0 10 20 30
WA Albian % Species

e Cd é
s Fà # z?
° o o
ge ` Pteridophytes ¥ ? "à Coniferales
upper CM
Upper
Albian
lower
Upper
Albian 104
upper
Middle
Albian
106|
middle
Middle
Albian
108
lower Middle
and
Lower Albian NO DES
N as N `
* non-angiospermous 9 10 20 30
% species
FIGURE 3. Relative abundance of major plant groups during the Albian of pe d Rocky Mountain
region. A in million years before present rcentage occurrence of major plant groups based on
megafossils. Occurrences of species for each plant group were tabulated for individual ne These counts were
then sum a percentage calculated for each time interval as list elow. Total species counted for each
interval are 1 e parentheses. Aptian and Lower Albian (119): Kootenai (Miller & LaPasha, 1984

indicated in ntheses. : :
LaPasha & Miller, 1985); Bullhead, Gething, and Luscar (Bell, 1956). Lower Middle Albian (150): Beaver Mines
(Berry, 1929b; Bell, 1956; Mellon, 1967); Lower Gates Member (Stott, 1963). Middle Middle Albian (130):
Upper Gates Member (Mellon et al., 1963); Pasayten (Penhallow, 1907; Bell, 1956); Kingsvale and Jackass
Mountain (Bell, 1956); Fall River and i bw aep Obs.); rd Creek (Bell, 1956; Stott, 1963). Upper
Middle Albian (150): Mill Creek (Dawson, 1886; B , 1929c; Bell, 1956; Mellon, 1967). Lower Upper Albian
(89): Crowsnest (Bell, 1956; Mellon, 1967) ur "Su n River, and Black Eagle (pers. obs.). B. Percentage oc-
currence of major plant groups based on microfossils. Percentages calculated as above. Assignment of spores

1987]

ment of latewood in Categories B and C suggests
seasonality in precipitation. An alternative ex-
planation is that low light availability in winter
induced latewood formation in some species but
not others.

Late Albian or Cenomanian woods of Cu-
pressinoxylon sp. (Andrews & Kern, 1947) and
Paraphyllanthroxylon idahoense Spackman
(1948) from the Wayan Formation in south-
eastern Idaho have no prominent growth rings
and can be seen as evidence for equable year-
round temperatures. These woods suggest either
the absence of seasonality in precipitation or that
the trees grew in swampy or other well-watered
habitats.

Later in the Cretaceous there is further evi-
dence for seasonal or low precipitation in the
region, possibly related to the mountain building
of the Laramide Orogeny. The Campanian Two
Medicine Formation in northwestern Montana
includes caliches, desiccated carbonate nodules,
and sandstone bodies of episodic rivers that in-
dicate a seasonal wet-dry climate, with the dry
season longer (Lorenz, 1981). The abundance of
large (mesophyll and megaphyll size), apparently
deciduous leaves in the Two Medicine flora sup-
ports such a climatic seasonality, although the
leaves can be interpreted alternatively as indic-
ative of successional communities. Low or sea-
sonal precipitation is further indicated by a sub-
stantial notophyllous evergreen component of
coriaceous leaves typically without drip tips. Co-
niferous woods from the lower Two Medicine
exhibit pronounced growth rings of highly vari-
able thickness (Crabtree, pers. obs).

Oxygen isotope ratios are used most accurately
to estimate maximum paleotemperature and to
establish temperature trends of ocean waters
(Frakes, 1979). Oceanic temperatures can be used
to estimate temperatures on nearby land masses.
Since the NRM region during much of the Cre-
taceous was a relatively narrow land mass po-
sitioned between the Pacific Ocean on the west
and the epeiric seaway on the east, it is likely
that oceanic temperatures are significant for ap-
proximation of the land temperatures.

Isotopic ratios from the continental platform

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

713

off the Soviet Union (Douglas & Savin, 1975)
indicate a Cretaceous temperature maximum in
the Albian, followed by cooling in the Late AI-
bian and Cenomanian. Subsequently, a rewarm-
ing occurred in the early Senonian before a Maas-
trichtian cooling (but see also Boersma, 1984).
The Albian maximum is probably the warmest
period that the world has experienced since Pa-
leozoic time (Frakes, 1979). Latitudinal gradi-
ents in surface water temperatures were less steep
during the Cretaceous, and the average temper-
atures of deep water appear to have been signif-
icantly higher than at present (Schopf, 1980). Mid
and Late Jurassic oxygen isotope ratios from
Montana and Wyoming indicate es the surface
water had a maximum of 20°C ( n & Shaw,
1977). Polar ice was unknown das the Cre-
taceous, and Frakes (1979) hypothesized that
north polar surface water was no cooler than 7—
19°C and may have been at the high end of this
range based on isotopic paleotemperatures of
about 15?C for putative North Pacific Deep Water
from the equatorial Pacific (Stevens, 1971). A
computerized model of surface water tempera-
tures for an ice-free Arctic predicts a temperature
increase of 7-10?C over the present 0—-5°C tem-
perature range (Sellers, 1969)

Hermatypic coral reefs occur in tropical and
subtropical oceans. Because modern reefs form
only when minimum water temperatures exceed
18°C, their fossil distribution can be used to es-

Mesozoic distributions of reefs (Newell,
Beauvais, 1973) indicate that oceanic currents in
temperate latitudes were significantly warmer
than at present. A reef from the Jurassic of Wy-
oming (Beauvais, 1973) indicates that they
formed as far north as 50?N paleolatitude, but
there is doubt that the Wyoming reef is herma-

Jurassic and Cretaceous at latitudes 10-20? north
of their present distribution.

Several papers discuss paleofloristic climatic
zonation during the Cretaceous based on the
composition of eastern Asian megafloras (Kras-
silov, 1973a, 1975, 1978) and microfloras

—
and pollen to major plant groups follows the Wisi pans ag E of Singh (1971) and Norris (1967). Numbers in
mber of s

parentheses after age below indicate total num

species counted for that interval. Lower Albian and lower

Middle Albian (179): MeMurray-Clearwater(Vagvolgy & Hills 1969); Loon River (Singh, 1971). Middle Middle

Albian (206): Harmon (Singh, 1971); per Grand Rap
(Singh, 1971); Joli Fou (Norris, 1967). L

s (Norris, 1967). Upper Middle Albian (200): Cadotte

ower Upper Albian (184): Paddy rer 1971); Viking (Norris, 1967).
).

Upper Upper Albian (213): Shaftsbury (Singh, 1971); Upper shale (Norris, 1

714

"c innamomo:
phyll

DD kJ

x Thy
Aud

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Cornophyll A Rhamnophyll | B

FIGURE 4. Cretaceous dicotyledonous leaf morphotypes. Sources of previously published drawings are in-

dicated. SAPINDOPHYLL

M erui obtusilobum Mrs eite Format
on) Bell.—

12, figs. 6, 7 in Lesquereux, 1892).—
Heer, Raritan Formation, Marylan
medium Newberry (pl. 29, fig. 7 in Hollick, 1906).

(Chlonova, 1980). Krassilov's papers propose
four intuitively based, latitudinal climatic zones:
boreal, temperate, warm-temperate, and sub-
tropical. Relative to these zones, the NRM re-
gion is closest to the warm- temperate zone based

the presence of palms and large-leaved deciduous
dicots. A position in the warm-temperate zone
is consistent with Campanian floras in Sakhalin,
Amur, Altai, western Greenland, and Vancouver
Island (Krassilov, 1981). Vakhrameyev (1978)
placed the NRM region straddling the boundary
between hist te-warm | hu-
mid zones for the Early Cretaceous. This place-

“Ficus” beckwithii Lesquereux (Crabtree), Dakota Formation, Golden,
(Peabody Museum Poleibolanite! Collection no. oe —B
Virginia (fig. 17 in Doyle & Hickey, us —

Sapindopsis magnifolia Fontaine, Patapsco For-
y "o t : ]

B. Generalized leaf. CORNOPHYLL: “C ornus" forchhammeri
nd (pl. 82, fig. 1 in Berry, 1916). RHAMNOPHYLL:
— B. Generalized leaf.

— A. “Cinnamomum” inter-

ment is based on the presence of Tempskya and
Cycadeoidea in Montana and South Dakota. The

sarge shifts approximately 100 km to the
ort his Late Cretaceous reconstruction
ae it just north of the present 49th parallel
(about 55°N paleolatitude). The Late Cretaceous
temperate-warm humid zone is based on the oc-
currence of Ni/ssonia and Pseudoprotophyllum,
and the subtropical humid zone is based on palms
and Dewalquea.

Smiley (1967), basing his arguments primarily
on North American high-latitude floras, indi-
cated that the climate gradually warmed from
the Jurassic into the Albian, after which a general
cooling occurred through the Late Cretaceous (at
the same time as Alaska was rotating to the south).

BE.

1987]

Scott & Smiley (1979) reported on the micro-
and megafossil flora from the north slope of Alas-
ka. Their treatment indicates that northern floras

emain predominantly gymnospermous-pteri-
dophytic until the end of the Late Albian, that
they do not exhibit Classopollis pollen in any
significant quantity (but see Herngreen & Chlo-
nova, 1981), and that they are allied to more
southerly Albian floras by abundant spores of
Gleicheniaceae and Schizaeaceae. Vakhrameyev
(1982) recognized the Early Cretaceous as a time
of increasing aridity followed by “humidification

sensitive ferns throughout the Lower Cretaceous
at high latitudes indicates, and did not continue,
since palms in the Senonian indicate a generally
equable year-round temperature. Krassilov
(1973a, 1975, 1981) stated that Late Cretaceous
climates in Asia warm into the Campanian and
subsequently cool through the end of the Cre-
taceous.

THE EARLY ANGIOSPERM FLORA

Hickey & Wolfe’s (1975) synthesis of the sys-
tematic implications of leaf morphology among
extant dicots enables paleobotanists to assess the
relationship of fossil taxa by extrapolation. Kras-
silov (1977) and Hickey (1984) used a series of
informal descriptive names for early angiosperm
leaf fossils that attempt to unify fossil species
with similar morphology. Such leaf morphotypes
are especially appropriate for Early Cretaceous
dicot leaves that have been placed mistakenly in
extant genera. Morphotypes are best viewed as
serving as a descriptive terminology in lieu of
revision.

Most important morphotypes from western
North American mid-Cretaceous floras are il-
lustrated in Figure 4 (see also Hickey, 1984). The
drawings are based in some cases on individual
taxa and in others on an idealized composite of
several different taxa (see figure legend). In all
cases the drawings represent the general form or
range of forms present in the morphotype. Table
1 lists the morphotypes along with the genera
that have been used in the past for species be-
longing to each. Extant genera, such as Populus,
which has been mistakenly regarded as embrac-
ing a large number of disparate fossil taxa, may
be listed under more than one morphotype. Ex-
tant genera cannot be demonstrated from the
mid Cretaceous, with rare exceptions (Doyle,

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

715

1969; Walker & Walker, 1984). Extinct genera
for which the type species is of Cretaceous age
are included under the appropriate morphotype
according to the type description. Fossil genera
for which the type is younger than Late Creta-
ceous have not been considered for inclusion,
seem the inten focus on early angio-
perm hus, Laurophyllum Saporta is named
from ae material; hence, I consider it in-
appropriate as a generic placement for Early Cre-
taceous angiosperms. Morphotypes, since they
are not formal taxonomic classifications, have
no types.

I have chosen to discuss the early angiosperm
flora of the NRM region using the leaf morpho-
types as a descriptive base. For each morphotype
I provide a brief description using the leaf ar-
chitectural terminology of Hickey (1979). In ad-
dition, I provide particulars relevant to the dis-
tribution in time and space for each morphotype
along with observations on relative abundance
and diversity. Possible relationships for each
morphotype are discussed individually and sum-
marized in Table

The occurrence and distribution of early an-
giosperms in the NRM region is presented in
Table 2, which encompasses the first five million
years of angiosperm history in the region. Where
possible, species are grouped according to mor-
photype. All Albian and major floras from the
region are included, in addition to several pre-
viously unreported collections. Names of taxa
appear as originally published except for floras
examined by myself, for which I have provided
identifications. Ages and locations of floras ap-
pear in Figures | and 2 and in Appendix I.

o

Pentalobaphyll (Araliaephyll). Leaves sim-
ple, orbicular, 3-5-lobed. Margin entire. Base +
cuneate. Primary venation palinactinodromous,
with 3 primary veins diverging from above top
of petiole, and 2 subprimary veins branching from
lateral primaries just above base. Secondary ve-
nation eucamptodromous, rather weakly devel-
oped. Tertiary venation reticulate to transverse,
AR to AO. Higher order venation and cuticle
not observed.

This very distinctive group is first recognized
in the region during the Middle Albian. Penta-
lobaphylls are lobate leaves with five principal

veins (Fig. 4). Araliaephyllum obtusilobum Fon-
taine is included in this group on the basis of the

716 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74
TABLE 1. Botanical affinity and important genera for Cretaceous leaf morphotypes.
Morphotypes Important genera! Botanical affinity
Sapindophyll Rosidae

Pentalobaphyll

Fontainea Sade! CM die pure “Androm-
eda," “Diosp ” “Ficus,” “Rhus
‘Salix,” 5 it a. a”

Araliaephyllum Fontaine, Sterculites Dawson, ‘‘Ara-
lia,’ "H edera,” “Liquidambar,” “Sassafras,” "Ster-

Unknown; possibly Rosidae
or Magnoliidae (Laurales)

culia’

Platanophyll * Araliopsis," Araliopsoides Berry, Aspidophyllum Les- | Hamamelididae-Platanales
quereux, nari phyllum ae ioe spido-
phyllum Hollick, “Aralia,” “Plat " “Sassafras”

Protophyll Cissites Debey, Credneri Ae ee Polyphyletic within Hamamel-
Richter, Protophyllum Lesquereux, Pseudoproto- ididae — probably Platanales
phyllum Hollick, Viburnites Lesquereux, “Alnus,” and Hamamelidales
* Betula," “Cissus,” *Parrotia," “Platanus,” “ Popu-
lus,” “

Trochodendrophyll “Cercidiphyllum,” ** Coccu-

dicus sa Berry,
lus

" “Paliurus,” “Populus,” “Smilax,”

o

Menispermaphyll
is," “Cocculus,” “Heder

Nymphaeaphyll

geai tes Lesquereux, oe.
pelop

IE Hollick, Hederaephyllum Fontaine, Nelum-

Trochodendrales and Crecidi-
phyllales

99 66

Cissam- Unknown; possibly Ranuncu-
ii

i
Magnoliid-Nymphaeales

bites Berry, Paleonuphar Hollick, Populophyllum
Fontaine, , Proteaephyllum Fontaine, “Castalia

Hedera
Magnoliaephyll Liriodendropsis Newberry, Liriophyllum Li je acy i some Magnoli-
gnolia Plein (Krasser) Seward, “Bauhin idae, p y Laurales,
*Ficus," "Laurus," “Liriodendron,” u Magnolia," TE
* Persea," mesas S ra
Cinnamomophyll Cinnamomoides Seward, “*Benzoin,” Cinnamo Polyphyletic; some probably
mum," “Cocculus,” “Litsea,” *Oreodaphne," “Pal- Laurales
iurus,” Ene ud
Cornophyll C Eis ipie ewberry, ° <. ” “Cornus,” Polyphyletic; some possibly
“Dio. i pos cus" “Rham Rosidae
Rhamnophyll Polyphyletic; some possibly

>“ Rhamnus,”

us"

Macclintockia Heer, PM " “Ficus,” “Pal-
rus," “Piper, “Smilax,” “Zizy-

palmate Dilleniidae

' See text for criteria used for inclusion of gener.

? Quotation marks indicate extant genera Loma identified in the Cretaceous flora.

characteristic venation, although this and several
other species may have three-lobed leaves. Doyle
& Hickey (1976) include this species in their Plat-
anoid line, based primarily on the palinactino-
dromous primary venation. The palmate lobing
and palinactinodromous venation of the Penta-
lobaphylls is here considered to be insufficient
evidence to establish relationship with the Plat-
anoid line when viewed along with the balance
of leaf-morphological characters. Pentaloba-
phylls show leaf-morphological characters, in-
cluding entire margins, eucamptodromous sec-
ondary venation, and weak tertiary venation,
which serve to distinguish the group from Plat-
anophylls. The tendency towards orthogonal

branching of tertiary and quaternary veins so
characteristic of Platanophylls is absent in Pen-
talobaphylls.

Certain palmate leaves from the Albian flora
cannot be accommodated under either the Pen-
talobaphyll or Platanophyll morphotype. These
include “Liquidambar” fontanella Brown, with
glandular-toothed margins, and “Sassafras”
bradleyi Brown, which has smooth margins but
lacks the suprabasal lateral branches character-
istic of Pentalobaphylls.

Fritel (1914) recognized seven species of Pen-

Cenomanian. He characterized the group based

1987]

primarily on the three basal primary veins, the
lateral two of which give off prominent lateral
veins just above the point of radius, and on the
camptodromous secondary venation. However,
" included dnas E serrate margins,
highe
order venation iens as piae saportanea Les-
quereux, which I assign to the Platanophylls.
Pentalobaphylls are perhaps the most abun-
dant dicots from Albian floras of the NRM re-
gion. Albian Pentalobaphylls from Wyoming (Fig.
5), Montana (Fig. 6), and Alberta (Figs. 7-9) are
representative of the group. Fritel (1914) placed
the Alberta species Araliaephyllum westonii
(Dawson) Bell and “Aralia” rotundata Dawson
into synonymy with Araliaephyllum kowalews-
kiana (Saporta & Marion) Fritel from Europe. A
more thorough treatment of the group is needed
before such conclusions are substantiated. Cu-
ticular detail for the group is unknown with the
exception of the Senonian Araliaephyllum po-
levoi (Krystofovich) Krassilov (1973b) from the
eastern Soviet Union, for which a similarity to
Sassafras and Lindera is suggested. G. Upchurch
(pers. comm.) has suggested a lauralean affinity
for leaves of the A. polevoi group based on cu-
ticular detail, sinus bracing, mesophyll secretory
glands, and possible basilaminar secondary veins.
The secondary and tertiary venation (Figs. 4, 8)
in this group suggests rosid affinity.

"t

Platanophyll. Leaves simple. Margins entire
or serrate, lobed or unlobed. Primary veins usu-
ally 3, palinactinodromous with several pectinal
secondaries on laterals. Secondary veins straight,
forking or exmedially branched, craspedodro-
mous to teeth, brochidodromous if margin en-
tire. Tertiaries and quaternaries forming an or-
thogonal network. Teeth platanoid, glandular
processes often nipple-shaped. Cuticular struc-
ture in Upchurch (1984b), Krassilov (1973b),
Kvacek (1983), Bůžek et al. (1967), Némejc &
Kvacek (1975).

Members of this widespread Laurasian mor-
photype date from the Middle Albian in North
America. The earliest occurrence from the NRM
region is “Platanus” sp. (Fig. 10) from the upper
Middle Albian of Alberta (Bell, 1956). Plata-
nophylls of Middle and Late Albian age occur in
several floras from the NRM region (Table 2). A
particularly rich assemblage of Platanophylls oc-
curs in the Late Albian Blackleaf Formation in
Montana (Figs. 1 1-14). Several poorly preserved
specimens representing the group are reported

CRABTREE— NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

717

from the Middle and Upper Albian of Alberta
(Bell, 1956). As in the Albian flora of Alaska
(Hollick & Martin, 1930) and eastern North
America (Doyle & Hickey, 1976), the group is
most Hio quad encountered in coarse fluvial
sedim

pem cretacea (Lesquereux) Berry is
an early member of this complex and is reported
from the Late Albian in the Patapsco Formation
in eastern North America (Hickey Doyle,

77). Arali d f is al ted from

No

the Cenomanian of Texas (Berry, 1922b), Kansas
(Lesquereux, 1874), and eastern North America
(Berry, 1916; Hickey & Doyle, 1977). In the NRM
region A. cf. cretacea is known from the Late
Albian Summit locality in the Blackleaf For-
mation (Table 2, Fig. 14).

Platanophylls are typically more abundant in
higher latitude floras (above approximately pa-
leolatitude 45°N), and the group may appear in
the NRM region before eastern North America.
Little debate exists over the platanoid affinities
of Platanophylls, but the precise timing of the
origin of the Recent order and family remains
unresolved. Recent Platanus is reported as being
of Senonian age on the basis of pollen mor-
phology (Pactlova, 1978), with pollen of platan-
aceous character recorded from the Albian (Pact-
lova, 1982). Némejc & Kvacek (1975) proposed
on the basis of cuticular analysis that Cenoma-
nian Credneria spp. (here considered Proto-
phylls) from Bohemia are related to the platan-
aceous line. Early Platanophylls are related to,
and convergent with, other groups of the emerg-
ing Hamamelididae, especially the Protophylls.

Fructifications regarded as of platanoid affinity
are widespread in the Late Albian and Ceno-
manian of North America. Notable among these
are records of “Sparganium” from Washington
(Fig. 35) and Wyoming (Fig. 41; Brown, 1933a),
and “Platanus” from Greenland (Seward & Con-
way, 1935) and Kansas (Lesquereux, 1892). For
further discussion of Cretaceous fructifications
of platanoid affinity see Dilcher (1979) and Crane
et al. (1986).

Platanophylls are allied to Protophylls on the
basis of well-defined, orthogonal-reticulate, ter-
tiary and quaternary venation. Palinactinodro-
mous venation and a greater tendency towards
lobation distinguishes the Platanophylls from the
Protophylls, which have pinnate organization
(Fig. 4). I have chosen to segregate these leaves
in the hope that the Cenomanian Cissites-Bet-
ulites-Alnites complex (Protophylls) might be

718 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

TABLE 2. Early angiosperm flora from the northern Rocky Mountain region. 2?

Boul-
der Pasay-
Cree ten
(6c) (17)

Beaver
Mines
(4)

A BC

Mill Creek
(16)

Lower Gates (6a)
Upper Gates (6b)

—
©
—
~
—
o
>
x
=
=

Kingsvale (14)

Pre-Muddy (18)
Jackass (12)

Summit (19)
Black Eagle (5)
3
g

A B C D

>
=
>
=

Sapindophyll
Celastrophyllum acuti-
i X X X X
Celastrophyllum sp. x
cf. Celastrophyllum sp. x
Daphnophyllum dako-

te
Fontainea grandifolia
Newberry x X X X
rues E F. grandi-
folia berry X
. ed sp. X
“Laurus” crassinervis
D

awson
“Myrica” serrata Pen-
hallow

Proteoides daphnoge-
noides Heer X
“Quercus” flexuosa?

Newberry
“Salix” inaequalix?
Newberry
cf. “Rhus”? powelliana
Lesquereux X
Sapindopsis angusta
(Heer) Seward &
Conway
Sapindopsis cf. angusta X
Sapindopsis sp. aff. an-

gusta
yia belvideren-
erry
Sapindopsi brevifolia?
Fon

ontaine X
Sapindopsis cf. brevifo-
li

ia

Sapindopsis magnifolia

Sapindopsis cf. magni-
folia

Sapindopsis sp. X

Sapindopsis sp. X

cf. Sapindopsis sp. X

cf. Sapindopsis sp. X
Pentalobaphyll

“Aralia” rotundata

awson
“Aralia” westonii Daw-

1987] CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

TABLE 2. Continued.

Beaver
Mines

(4)

ABC

Lower Gates (6a)

Upper Gates (6b)

Pre-Muddy (18)

(6c) (17)
AB A B

Kingsvale (14)
Jackass (12)

ABC

Mill Creek
(16)

Summit (19)

g

Sun River (20)
Black Eagle (5)

>
w

Araliaephyllum westonii
(Dawson) Bell

Araliaephyllum? sp.
raliaephyllum sp.
i ar" inte
grifolium Lesquereux
“Sassafras” mudgii?
Lesquereux
cf. “Sterculia? mucro-
nata Lesquereux
“Sterculia” vetustula
Dawson
“Sterculia” sp.
Protophyll
Alnites insignis? Daw-
Cissites? sp.

cf. Cissites sp.
“Platanus” affinis Les-

quereux
“Platanus” affinis var.
ampla Dawson
Populites dawsonii Bell
Populites cf. dawsonii
Populites sp.
cf. Protophyllum sp.
Platanophyll
“Aralia” wellingtoniana
Lesquereux
cf. “Aralia” wellington-
Araliopsoides cf. creta-
cea (Newberry) Berry

cf. Aspidophyllum trilo-
atum Lesquereux

Platanophyllum sp.
“Platanus” heeri Les-

“Sassafras” cretaceum
Newberry
Trochodendrophyll
“Cercidiphyllum” sp.

X X

X

XX

XX

X
x<

720 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

TABLE 2. Continued.

Boul-
der Pasay-
reek te
(6c) (17)

A B A B

Beaver
i Mill Creek
(16)

Lower Gates (6a)
Pre-Muddy (18)

4
A B C ABCD

Kingsvale (14)
Sun River (20)
Black Eagle (5)

Jackass (12)
Summit (19)

jpe potoma-
x
Tr serena am
(“Cercidiphyllum”)
potomacensis (Ward)

Nymphaeaphyll
“Hedera” ovalis? Les-
quereux
Menispermites renifor-
nis Dawson
Menispermites sp. X
Nelumbites sp. X
Nymphaeites sp. X
Magnoliaephyll
“Magnolia” alternans

eer
"Magnolia" magnifica
Dawson

Cornophyll
“Ficus” fontainii?
Berry
Laurophyllum debile
Dawson
Rhamnites sp. X
“Rhamnus” sp. x

Cinnamomophyll

Cinnamomoides ovalis

(Dawson) Bell x x x X x
Cinnamomoides cf.

ovalis (Dawson) Bell x
Cinnamomoides sp. x
Cinnamomoides sp. x
Cinnamomoides sp. x
Macclintockia cretacea

Heer
“Paliurus” ovalis Daw-
son X X
Uncertain
Capparites? sp. X

cf. “Ficus” ovatifolia

1987]

TABLE 2. Continued.

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

Beaver
Mines
(4)

A B C

Lower Gates (6a)
Upper Gates (6b)
Fall River (10)
(18)

Boul-
der
reek
(60 - “ay

AB AB

ae)
zg
°
"<

Mill Creek
(16)

(14)
Jackass (12)
Summit (19)
Black Eagle (5)

A B C D

Juglandites cretacea
Dawson

* Magnolia"? sp.

Menispermites? sp.

Myrtophyllum Pu
Seward & Conwa

* Populus" cf. berggrenii

eer
cod cyclophylla

Quercophyllun? sp.
"Quercus" coriacea

Ne
“Salix”? sp.
"Salix" s
“Sassafras” parvifolium
Fontain x

X

mK mK

! Numbers in parentheses refer to geographic and stratigraphic position of floras as indicated in Figures |

a
2 All species are included as originally cited in the references given below. I have identified the species from

the Summit, Sun
? Vertical rows indicate

Penhallow, 19
1929c. (16)C— aon 1956. (16)D— Mellon, 1967. (7)A—
and (5)— Cra

icis Canada, provided tl

07. (17)B — Bell, 1956. (14) — Bell, 1956. (12)— Bell, 1956. (16)A Denion: 1886. (16

ee, this report. It should be noted that W. A. Bell, in his capac ty | as baleobotan

River, Black Eagle, Fall River, and Pre-M uddy collections.

oe Berry, 1929b. o> Bell, 1956. (4)C—

Bell, 1956. (7)B— Mellon, 1967. (10), (18), (19), (20),
nist with the
t (1963), Mellon (1967),

Geological ^m
and Mellon et al. (1963).

allasa, p

better understood. I suggest that Protophylls are
more similar to mainline Hamamelidales on the
basis of leaf architecture than are Platanophylls,
which are closer to the platanoid line of Ham-
amelids.

Protophyll. Leaves simple. Margin entire to
entate, occasionally lobed. Venation pinnate,
the basal secondaries sometimes strengthened

phylls. Cuticular details in Némejc & Kvacek

(1975) and Krassilov (1973b)
Richter (1905) monographed Credneria Zen-
ker and Paracredneria Richter.

Protophylls can usually be distinguished from
Platanophylls by the basically pinnate venation
and by the more consistent craspedodromy of
the secondary veins (Fig. 4). However, in many
cases the two morphotypes intergrade to the ex-
tent that recognition is difficult. Protophylls are
common in the NRM region but do not exhibit
the diversity or dominance found in the coarse
facies of the Cenomanian Dakota flora of Ne-
braska and Kansas (Lesquereux, 1892). side
species from the Albian floras of Alberta are
placed in this morphotype (Table 2). Seit pen
are present in the Blackleaf flora (Fig. 15). Basal
venation in Platanophylls and Protophylls is
compared in Figures 16 and 17

x

3 ng

EET
= w

Z.
WW
a
a
<
c
<
=
Z.
<
=
°
x
2
e
N
na
=
Ln
x
=
u.
°
N
—
<
Z
Z,
<

1987]

Sapindophyll. Leaves even to odd pinnatifid
or pinnately compound. Leaflets opposite or al-
ternate, sometimes decurrent on rachis. Margins
entire to serrate. Teeth apparently cunonioid.
Petiole stout. Petiolule stout. Primary vein stout.
Secondary veins eucamptodromous or
semicraspedodromous, arched-ascending, often
numerous, diverging at + uniform angle of 35-
50*. Intersecondary veins typically present. Ter-
tiaries weakly percurrent, transverse, AR but
highly variable. Quaternaries irregularly reticu-
late. Cuticle and tooth morphology in Upchurch
(1984b).

This unique Lower and mid-Cretaceous mor-
photype encompasses fossils with pinnatifid or
pinnately compound leaves (Fig. 4). Rachises with
attached leaflets have been reported as Sapin-
dopsis Fontaine and Fontainea Newberry. The
widespread Laurasian species “Andromeda”
a Heer, “Ficus” daphnogenoides (Heer)

a F> Rus Lesquereux, ' *Sapindus
morrisonii Lesquereux, and “Rhus” powelliana
Lesquereux represent i pene leaflets of Sap-
indophylls. Dissected leaves with palmate or-
ganization, cf. Halyserites Sternberg and Cus-
soniophyllum Velenovsky, are excluded from the
Sapindophylls "i have not been observed in
the NRM regio

Arahe are an important component of
many North American Albian and Cenomanian
floras (Berry, 1922a, 1922b; Fontaine, 1889;
Hickey & Doyle, 1977; Lesquereux, 1874, 1883,
1892; MacNeal, 1958; Upchurch, 1984b). The
morphotype is generally considered to be more
abundant in floras from regions to the south of
the NRM region and is sometimes regarded as
representing thermophilous plants. Nonetheless,

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

T23

the morphotype is common in mid-latitude flo-
ras from the NRM region and Greenland (Sew-
ard & Conway, 1935), and occurs in southern
Alaska (*Ficus" daphnogenoides in Hollick &
Martin, 1930) and the north slope (pers. obs.).

This morphotype is represented by several

species in the NRM region (Table 2). A probable
new species of Sapindopsis of Middle Albian age
iS ostate from wicks equivalent to the Ther-
Montana (Fig. 18).
It aoan to be related to the Cenomanian species
“Sapindus” morrisonii and “Ficus” beckwithii.
A Sapindophyll similar to Sapindopsis variabilis
Fontaine is present in the Blackleaf Formation
of Montana (Fig. 19). Sapindophylls are present
in the Albian of Alberta (Table 2, Fig. 20), British
Columbia (Fig. 27), and Washington (Figs. 28-
32). A putative Sapindophyll (cf. Sapindopsis sp.)
from the Upper Albian of Idaho is tentatively
assigned to this morphotype (Figs. 21-23). This
species is allied to the morphotype by admedial
orientation of the tertiary veins (Fig. 21) and
dissected leaves with decurrent leaflets (Figs. 22,
23), but shows the irregular laminar furcations
of Halyserites.

Sapindophylls may be most closely allied to
Recent basal Saxifragales and to other rosid
groups such as the Sapindales and Rutales. This
relationship is based on the pinnately compound
leaves, eucamptodromous venation, admedia
orientation ofthe transverse tertiary veins (Hick-
ey & Doyle, 1977), cunonioid teeth in some
species (Upchurch, 1984b), and cuticular fea-
tures (Upchurch, 1984b). Crane et al. (1986) dis-
cussed possible relationships between Sapindo-

duced tricolpate pollen (viz. Hamamelididae and

—

FiGures 5-58.

Representative dicotyledonous leaves from the Albian through middle Campanian of the
NRM region. All figures with centimeter scales. Specimens from t

he U.S. Nation m are provided with

al
haad field locality numbers, and those from the Geological Survey of Canada are provided with GSC locality
mbers. Types and other cataloged specimens are indicated where appropriate, along wit National
akua (USNM) and Geological Survey of Canada d accession numbe ames applied to illustrated
specimens are as originally published or as identified in paleobotanical collections at the US and GSC. I
have provided names edd for species from the Fall River, Blackleaf, Pre-Muddy, Pasayten, MES Winthrop,
and Two Medicine floras

FIGURES 5-12. Albian age Senet Nps leaves of the Pentalobaphyll and Platanophyll morphotypes. 5-
9. Pentalobaphylls.— 5. cf. “Sterculia” mucronata Lesquereux, Fall River Formation, USGS locality 7404. 6-
8. Araliaephyllum westonii (Dawson) Bell.—6. Blackleaf Formation, USGS locality 9439. 7, 8. Mill Creek
Formation. — locality 1815, GSC 5117, holotype.—8. GSC locality 3066, GSC 5874. —9. “Aralia”
rotundata Dawson, Mill Creek Formation, GSC locality 1815, GSC 5116, holotype. 10-12. Platanophylls.— 10.
“Platanus” sp., Mill Creek Formation, GSC locality 1815, GSC 5106.—11. Platanophyllum sp., Blackleaf
Formation, USGS locality 9437.—12. cf. Aspidophyllum trilobatum Lesquereux, Blackleaf Formation, USGS
locality 9437.

724

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FIG an

Beg age e dicotyledonous leaves ih Platanophyll and Protophyll morphotypes. 13, 14.

Waaa — 13. cf. “Aralia” wellingtoniana Lesquereux, Blackleaf Form U locality l
leas Na cf. mae a (Newberry a erry, Blackle cea 15. Protophyll, cf. Protophyllum sp., Blackleaf
JSGS locality 5986. . Comparison of basal venation of PI land

l
Sub Platanophyllum sp., dubi cue USGS locality 5939. — 17. Protophyll, Protophyllum sp.,

Blackleaf Formation, USGS locality 5986.

Rosidae). Friis & Skarby (1982) and Basinger &
Dilcher (1984) discussed Cretaceous flowers and
pollen of early Rosidae.

Sapindophylls are restricted to Albian and
Cenomanian floras in the NRM region. I am not
familiar with their Late Cretaceous range else-
where. The rosid affinities of Sapindophylls need
to be confirmed by further study, as does the
relationship between the group and the putative

Upper Cretaceous and Paleocene derivative Av-
erhoites.

Trochodendrophyll. Leaves simple, ovate to
reniform. Margin crenate. Teeth convex-convex
(A-1), chloranthoid. Venation actinodromous.
Primary veins usually 3 or 5, sometimes more,
arched apically. Secondary venation semicras-
pedodromous with + prominent brochidodro-

1987]

mousarches present apically. Tertiary and higher
order venation not well known, appearing retic-
ulate in some with tendency towards orthogonal
branching

This oi is characterized by actinodromous
rimary venation and crenate margins bearing
d teeth (Fig. 4). The latter character
distinguishes the group from those Menisper-
maphylls with actinodromous venation. Tro-
chodendrophylls are reported in low frequencies
during the Albian, and are represented in the
NRM region primarily from Alberta and British
Columbia (Table 2). Trochodendroides ('* Cerci-
diphyllum”) potomacensis (Ward) Bell conforms
most closely to the concept of the morphotype
with five primary veins, brochidodromous sec-
ondary veins, and crenate margin (Figs. 24, 25).
Trochodendrophylls have been assigned to
rather unlikely genera (Table 1). Cuticular stud-
d. of Menispermites potomacensis Berry from
Lower Cretaceous Potomac Group (Up-
Y adio l ; pers. comm.) indicate that the
stomatal apace of this species does not con-
form to that of extant Cercidiphyllales and
Trochodendrales. Probable members of the lin-
eage are reported in high frequencies from Seno-
nian and latest Cretaceous deposits, but these
are more advanced in morphology and in some
cases can be appropriately assigned to the Recent
Trochodendraceae and rcidiphyllaceae (see
Krassilov, 1973b). Trochodendrophylls, along
with a series of pinnately veined Lower Creta-
ceous leaves with chloranthoid teeth and open
craspedodromous secondaries (see Upchurch,
1984b; Fig. 36, this report), may represent an
ancestral plexus from which Recent Trochoden-
drales and Cercidiphyllales evolved during the
Cretaceous (but see Wolfe, 1973). The Campani-
n Two Medicine flora (see below) includes four
leaf taxa which have puni reds teeth, but which
and higher order
suggest mosa d elopment ofleaf
characters in Cercidiphyllales, Trochodendrales,
and Chloranthales.

Menispermaphyll. Leaves apparently sim-
ple, broadly deltoid and + trilobed to circular or
ovate-elliptic. Base cordate to obtuse. Margins
entire or lobate forms occasionally with teeth.
Venation acrodromous or actinodromous with
3-9 primary veins. Primary veins craspedo-
drome or becoming camptodrome just before
margin. Secondary veins irregularly spaced,
camptodrome in many but craspedodrome to the

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

725

teeth in nonentire forms. Tertiary veins ran-
dom-reticulate to transverse. Higher order ve-
nation not observed. Cuticle unknown.

This group (Fig. 4) superficially resembles
Nymphaeaphylls but is never peltate and never
has an expanded, multistranded petiole. Cor-
date leaves with major veins unbranched to the
margin are best placed in Menispermaphylls,
whereas such leaves with highly branched pri-

mary and secondary veins are more appropri-
ately placed in Nymphaeaphylls. Trochodendro-
phylls may be distinguished by their crenate
margin.

Menispermaphylls are morphologically con-
vergent with extant Menispermaceae (Ranun-
culiidae). The modern family includes both sim-
ple-leaved and ternately compound members.
The leaves and leaflets are deltoid to ovate with
entire or sparingly toothed margins. Tooth mor-
phology is a modification of the chloranthoid
type (Hickey & Wolfe, 1975). Primary venation
is actinodromous or acrodromous, and major
veins terminate in a fimbrial vein, thus appearing
craspedodromous. Many Recent species exhibit
a mucronate leaf apex; simple, unbranched tri-

homes; and well-developed, orthogonal-retic-
ulate, higher order venation.

Menispermaphylls resemble Recent Meni-
spermaceae in shape, margin, and venation.
However, before relationship to the family can
be established it is necessary to demonstrate the
presence in the fossils of such characters as fim-
brial veins and simple, unbranched trichomes.
Nevertheless, the unusual combination of entire
margin and craspedodrome primary veins is an
important point of agreement between the fossils
and Recent Menispermaceae.

Menispermaphylls show their greatest diver-
sity in the Dakota flora of Kansas and Nebraska
(Lesquereux, 1892). However, here, as in the
NRM region (Table 2), they are a relatively mi-
nor component ofthe flora. The Campanian Two
Medicine flora (see below) includes a species that
is entirely consistent with the leaf morphology
of extant Menispermaceae.

Magnoliaephyll. Leaves simple, ovate to el-
liptical. Margin entire or 2- or 3-lobed. Primary
venation pinnate in entire forms, with 3 primary
veins from base in lobed forms. Secondary ve-
nation brochidodromous or festooned brochi-

odromous. Higher order venation variable.

This morphotype (Table 1) comprises a large
number of problematic taxa that form an im-

ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

1987]

portant part of all but extreme northern Laura-
sian floras. The group is erected primarily for
Te due to the enigmatic positions of
most of its members. Some Potomac Grou
a for which cuticular evidence is
available (Upchurch, 1984a, 1984b) appear to
be related to Laurales and Illiciales. “Magnolia”
magnifica Dawson is illustrated as an example
of the morphotype (Fig. 26). Early members of
the morphotype from the NRM region are listed
in Table

Putative magnoliid infructescences are known
from the mid Cretaceous of central North Amer-
ica (Dilcher, 1979), Japan (Nishida, 1985), and
Greenland (Seward & Conway, 1935). Walker et
al. (1983) reported pollen of Winteraceae from
the Lower Cretaceous of Israel.

Cinnamomophyll. Leaves simple. Margin
entire. Venation pinnate. Prominent suprabasal
secondary veins arching apically. Secondary ve-
nation brochidodromous. Basilaminar second-
ary veins sometimes present. Tertiary veins
transverse, regular or somewhat irregular in
spacing and course. Quaternary veins weak, ran-
dom-reticulate. Cuticle unknown.

This morphotype (Fig. 4) is widely reported
from NRM region floras and is an important
component of the angiosperm vegetation during
the Albian (Table 2). Although not present in all
Cinnamomophylls, basilaminar secondary veins
indicate that some may be related to extant Lau-
rales. Cinnamomophylls are most important in
floras from middle latitudes in Laurasia where
they are consistently represented in the Albian
floras

Nymphaeaphyll. Leaves simple, orbicular to
if Margin entire. Petiole often multi-

venation forming weak brochidodromous arch-

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

TET

es, craspedodromous in “Castalia” and Casta-
liites. Cuticle unknown.

his morphotype converges with Trocho-
E M from which it is distinguished by
its entire margin, and with Menispermaphylls
(see discussion of Menispermaphylls and Fig. 4).

Several species of dique ee exhibit leaf
architecture consistent with those of extant
Nymphaeales and "URN iae ius from
cuticles and fruits is needed to confirm the re-
lationship.

Cordate and peltate leaves with actinodro-
mous venation and entire margins are abundant
and diverse in the North American mid Creta-
ceous, but they are not well represented in the
NRM flora (Table 2). Nymphaeaphylls may rep-
resent an adaptational syndrome for aquatic hab-
itats as suggested by Samylina (1968), Axelrod
(1970), and Krassilov (1977). Hughes (1976) sug-
gested that this leaf morphology may represent
simply an early “experimental” shape that is not
correlated with aquatic habitats.

Rhamnophyll. Leaves simple. Margin entire
or crenate-dentate. Base acute to truncate. Ve-
nation acrodromous. Primary veins 3-5, arising
from within the petiole and arching apically. Sec-
ondary venation brochidodromous or semicras-
pedodromous to teeth, often with several M ipn
and prominent secondary veins originating on
outermost primary veins and arching apically
along margins. Tertiary veins transverse, regu-
larly spaced and usually numerous. Higher order
venation not observed. Cuticle unknown.

Rhamnophylls are rare from floras of Albian
age in the NRM region (Table 2). The group
occurs with greater frequency in Late Cretaceous

tanical affinities of the former remain unknown.

—
FIGURES 18-26.

phyll morphotypes. 18-23. Sapindophylls.

Albian age dicotyledonous leaves 2 ps CC aesan Hoe cp de and Magnoliae-
3. —18. cf. * wellia

a Lesqu , Pre-Muddy collection. —

19. cf. Sapindopsis Sp., Blackleaf Formation, USGS Mat = 6007. ri “Ko nines "erandifolia Newberry, Com-

(Ward) Bell, Kingsvale Form
26. Magnoliaephylls. cress S=.

C collec GSC 6639. 21-23. cf. Sap-

ns),
5. Trochodendrophylls, S ucdendroidis CC ercidiphyllum’’) potomacensis
4. ocality 3449, GSC 6654.—
mas Dawson, GSC locality 290, GSC 5133, holoty

. GSC locality lt GSC 5907.—

728

Cornophyll. Leaves simple. Margin entire.
Primary venation pinnate. Secondary venation
camptodromous, arching apically. Tertiary veins
transverse, regularly spaced, more or less nu-
merous. Higher order venation and cuticle un-
known

Cornophylls occur in limited diversity and
abundance in the Albian floras of the region (Ta-
ble 2). The group assumes a more important po-
sition in later Cretaceous floras. The botanical
affinity of the group is unknown, but some mem-
bers appear to exhibit morphological similarities
to Rosidae.

Dryophyll. Leaves palmately compound.
Leaflets ovate to lanceolate. Margin serrate. Pri-
mary venation pinnate. Secondary venation
craspedodromous. Higher order venation not
observed. Cuticle in Krassilov (1973b) and Né-
mejc & Kvaéek (1975).

This group of palmately compound leaves with
serrate margins is not present in the early angio-
sperm floras. They appear first in the uppermost
Albian and Cenomanian. Serrate forms with
craspedodromous venation and lanceolate leaf-
lets (cf. Dryophyllum Debey) can be shown to

with Fagales, al-
though this Is a a tenuous association for most
Cretaceous forms (Wolfe, 1973).

FLORISTIC PROVINCIALITY OF THE
NORTHERN ROCKY MOUNTAIN REGION
DURING THE ALBIAN

Discussion of floristic provinciality during the
Cretaceous ultimately depends on accurate cor-
relation of stage and substage boundaries be-
tween the proposed provinces. The Upper-Low-
er Cretaceous boundary and the position of the
Albian substages in the NRM region are placed
based on ammonites in the United States (Cob-
ban & Reeside, 1951, 1952a) and on a varied
molluscan fauna in Canada (Jeletsky, 1968;
Kauffman, 1975). Geologists have generally
placed the boundary between the Upper and
Lower Cretaceous in the NRM region at the top
of the Mowry Shale. This boundary coincides
with the uppermost widespread occurrence of
Neogastroplites (Cobban & Reeside, 1951). It also
coincides with the first occurrence of tricolporate
pollen in the region (Nichols et al., 1983; Norris
et al., 1975). Nonetheless, uncertainty exists over
the correlation of the regional boundary to the
European standard. The boundary in the western

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

interior is radiometrically dated as 94 Ma
(Obradovich & Cobban, 1975; but also see Fouch
et al., 1983), whereas in Europe it is dated as
97.5 Ma (Harland et al., 1982)

Brenner (1976) recognized a Northern Lau-
rasian Province and a Southern Laurasian Prov-
ince based on the palynology of Lower Creta-
ceous deposits. The northern province is
recognized by a coniferous palynoflora less di-
verse and more heavily bisaccate than the south-
ern province. The NRM region lies on the
boundary between provinces. Such provinciality
is supported by Srivastava (1981) and by Hickey
& Doyle (1977) and rejected by Herngreen &
Chlonova (1981). Angiosperm pollen has been
used for the recognition of Late Cretaceous pro-
vinciality (Srivastava, 1981). The extension of
angiosperm palynofloral provinces into the Early
Cretaceous seems reasonable in light of the elab-
oration of the timing of the migration of angio-
sperms provided by Brenner (1976) and by Hick-
ey & Doyle (1977).

The poleward migration of early angiosperms
was first recognized from megafossils (Axelrod,
1959) and subsequently refined through paly-
nological studies (Brenner, 1976; Hickey & Doyle,
1977). Early angiosperms appear to have arisen
in northern Gondwana and to have crossed the
Tethys into Laurasia in the Late Barremian and
Aptian (Hickey & Doyle, 1977). aby of mono-
sulcate-prod rthern lat-
itudes of Laurasia was is delayed by several million
years. This may be attributed to climatic or other
environmental barriers, inadequate dispersal
mechanisms, or merely to stalwart pre-existing
vegetation. Tricolpate taxa exhibit a far shorter
lag period. The first major radiation of the tri-
colpate group occurs on a worldwide scale during
the late Middle Albian (excepting at the very
highest latitudes).

Considerations of floral provinciality of early
o common prob-

Sw
hudy & Tschudy, 1986), es-
pecially in the Paleozoic and Mesozoic, and 2)
spurious conspecificity, 1.e., the tendency to as-
sign pollen grains to existing taxa. Both problems
obscure provincialism at lower taxonomic levels.
Penny (1969) commented on the impoverished
appearance of Cenomanian pollen floras relative
to leaf floras of that age. A related problem, which
is more or less confined to the early angiosperm

1987]

pollen floras, is the general absence of differen-
tiation of grains with respect to ornamentation,
shape, and gross morphology. In addition, Me-
sozoic palynologists do not normally elaborate
species frequency as is common for Holocene
workers, a situation that makes it difficult to re-
view existing literature for data which could be
used to determine provinciality based on dom-
inance and importance of species. On the other
hand, a common problem with megafloral tax-
onomy has been overspeciation due to failure to
recognize physiognomic variation in leaf mor-
phology within natural species. Both megafloral
and microfloral treatments are restricted by the
taphonomic biases implicit in any depositional
environment.

The first angiosperms known from the lower
Middle Albian (Potomac subzone IIB) of the
NRM region are two grains of Tricolpites micro-
munus (Groot & Penny) Singh from the upper
Loon River Formation in northwest Alberta
(Singh, 1971) and monosulcate grains from the
Cloverly Formation in Wyoming (Davis, 1963).
Pollen sz megafossils are known throughout the
NRM region with certainty from the middle
Middle hae (Bell, 1956; Davis, 1963; Nichols

& Jacobson, 1982; Norris, 1967; Singh, 1975;
Roberts, 1972). Singh (1971) reported nine species
of middle Middle Albian angiosperm pollen from
northwest Alberta. By upper Middle Albian (Po-
tomac subzone IIB), angiosperm pollen becomes
a persistent element of moderate frequency in
the microflora. This upper Middle Albian surge

IIB). Localities in British Columbia of equal age
yield Sapindopsis and a limited diversity of other
dicot leaves (Table 2). The first diverse assem-
blages of megafossils are of middle Middle and
upper Middle Albian age and include Sapindo-
phylls, Pentalobaphylls, Platanophylls, Cinna-
momoides, Trochodendroides, and Menisper-
mites in order of frequency in collections. The
components of this megaflora continue into the
lower Upper Albian with little apparent change.
The early angiosperm flora of the region is dis-
tinct from more southerly floras in its high rep-
resentation of Pentalobaphylls, Cinnamomo-
phylls, and to a lesser degree Led
and several endemic species are prese

I compared the Lower Cretaceous Ed codon

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

729

of the Patapsco Formation of Maryland (Bren-
ner, 1963) with that of the Loon River and Peace
River formations of northwest Alberta (Singh,
1971) to contrast composition, abundance, and
diversity of Middle and Late Albian floras of
eastern North America with those of the NRM
region. More recent treatments of the Patapsco
palynoflora exist (Doyle, 1969; Doyle & Hickey,
1976; Hickey & Doyle, 1977), but Brenner's work
remains the standard for taxonomic comparison

is likely that some of his taxa represent species
complexes or genera.

Monosulcates first occur in the latest Barre-
mian or early Aptian in eastern floras, consid-
erably antedating their lower Middle Albian oc-
currence in the NRM region. Seven ofthe twelve
Patapsco species of angiosperm pollen are tri-
colpate. The tricolpates appear during the Lower
and Middle Albian and continue upsection to
the top of the formation (middle Upper Albian)
with no addition of new species (Brenner, 1963;
but also see Doyle & Hickey, 1976). Of these
seven species, Tricolpites micromunus occurs in
the lower Middle Albian Loon River Formation,
and six of the seven tricolpates occur in the mid-
dle Middle Albian Harmon Member ofthe Peace
River Formation. Such palynological data indi-
cate close similarity between lower and middle
Middle Albian dicots in the eastern and western
floras of North America, at least at the generic
level.

An endemic flora in the region is indicated by
angiosperm pollen recovered from the Peace
River Group (Singh, 1971), which was deposited

tricolpates, are reported fro
a high degree ofsimilarity, es strikingly less than
that of the earlier flora. Furthermore, five Ca-
dotte taxa have not been reported from Creta-
ceous palynofloras outside the NRM region.
Eastern groups apparently moved north and east
into the NRM region during the Middle Albian
and persisted in the western flora; however, by
the upper Middle Albian, tricolpate and mono-

Liliacidites trichotomosulcatus Singh and the tri-
colpates Retitricolpites maximus Singh, Tricol-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

1987]

pites sagax Norris, T. parvus Stanley, and Frax-
inaipollenites venustus Singh

ndemic character of the flora appears to
result largely from separation of the region from
eastern North America by the Skull Creek Sea-
way for approximately two million years during
the upper Middle and lower Upper Albian. Thus,
shortly after angiosperms entered the region, they
were cut off from their presumed ancestral stock
to the south and east in an area some 20? further
north and potentially subject to substantially dif-
ferent selective pressures. However, it should be
noted that available palynofloras make it difficult
to resolve an east-to-west adaptive radiation
within groups producing small, reticulate, tri-
colpate pollen, since the equivalence of named
species to biological species is debatable. In con-
trast to the simple tricolpate pollen, leaf archi-
tectures and cuticles of Lower Cretaceous angio-
sperms provide an expanded suite of characters
from which biological species can be circum-
scribed. A detailed comparison of North Amer-
ican floras of Albian age is beyond the scope of
this paper.

WINTHROP FLORA

A large collection of fossil foliage from the type
locality of the Late Albian or Early Cenomanian
(Barksdale, 1975) Winthrop Formation in cen-
tral Washington (Figs. 1, 2) appears to be the
latest occurring **archaic" flora in the region. The
flora contains abundant Sapindophylls and Pen-
talobaphylls that are closely related to but not
conspecific i those ofearlier Albian floras from

the NRM regio

Several small p of plant fossils from
the Winthrop have appeared as floral lists in geo-
logical papers (see Barksdale, 1975). Apparently
the collections were sent to the U.S. National
Museum and examined by F. Knowlton and R.
Brown. Reevaluation of these collections in 1986
by S. Wing and J. Wolfe (pers. comm.) indicated
a Paleocene age for the flora, and hence their
origin in the Winthrop Formation must be ques-
tioned. The present report includes only those

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

731

specimens collected from the type locality (Ap-
pendix I) by R. Rau, J. Robison, and myself.
The Winthrop flora is a moderately diverse
assemblage consisting of about 20 species of ferns,
conifers, dicotyledons, and Sagenopteris (Cay-
toniales). Dicotyledonous species include Sap-
indopsis sp. (Figs. 28-32), Magnoliaephyllum sp.
(Fig. 33), Araliaephyllum sp. (Fig. 34), Araliae-
phyllum westonii, “Sparganium” sp. (Fig. 35),
an unidentified dicot with OE r sec-
ondary venation and chloranthoid teeth (Fig. 36),
Nelumbites sp. (Fig. 37), “Ficus” ovalifolia, Men-
ispermites sp., Eucalyptophyllum sp., and several
poorly preserved species

ee

UPPERMOST ALBIAN, CENOMANIAN, AND
TURONIAN FLORAS

Three well-developed floras, the Aspen, Fron-
tier, and Dunvegan (Table 3), provide much of
the megafossil evidence for the angiosperm flora
of the NRM region during the uppermost Albian
and Cenomanian (Fig. 2). The Dunvegan flora
is near the northern limit of the region; the Aspen
and Frontier are at the southern limit (Fig. 1).
Collections from the Albino Member of the
Mowry Formation and the Wayan Formation
(Figs. 1, 2) provide a few additional species.

The Albian angiosperm flora of the region
shows an underlying unity due to the occurrence
in most assemblages of Sapindophylls, Penta-
lobaphylls, Platanophylls, Protophylls, and Cin
namomophylls, which give the flora an “archaic”
appearance when contrasted to the flora of the
post-Albian. The post-Albian flora shows the de-
velopment of north-south provinciality in the
region. However, because of the small number
of floras known from this period (Fig. 2), and the
restricted volcaniclastic facies of the Aspen and
Frontier floras, such provinciality is at this time

al

deposits in the northern half of the region, there
is uncertainty over how widespread and uniform
the early flora was

The development of intraregional provincial-

—
FIGURES 27-37.
Sapindopsis sp. 28-37. Winthrop Formation. 28-32.
Enlarged leaflet of p te in Figure 28.—
joined upper three leaflets. — 33. Magnoliaephyllum sp.—

Dicotyledonous leaves from the Pasayten and Winthrop formations. 27. Pasayten Formation,

Sapindopsis sp.— 28. 9.

. Upper part of compound leaf. — 31. Base of leaf. — 32. Base of
36.

Upper part of compound leaf.—

—34. Araliaephyllum sp.—35. “Sparganium

Unidentified leaf with iuum secondaries and chloranthoid teeth. —37. Nelumbites sp.

732 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

TABLE 3. eo flora from the uppermost Albian, Cenomanian and Turonian of the northern Rocky
Mom region.!

Dunvegan (8)

Lower Upper
Aspen Frontier Frontier
(2) (lla) A B (11b)

Sapindophyll
“Andromeda”? spatula Bel X
“Ficus” eiii irai (Heen Berry X
“Myrica” nervosa K X
Sapindopsis huts it X

Rosidlike Leaves
“Diospyros” nitida Dawson X

“Salix” cumberlandensis Knowlton
“Staphylea”? fremontii Knowlton x

XX

Platanophyll
Ampelophyllites attenuatus (Lesquereux) Knowlton
“Aralia” cf. parvidens Hollick
“Aralia” veatchii Knowlton X
Aspidophyllum dentatum? Lesquereux
"H ereu

Xx

B
Q
[e]
=
S
S
3
&
3
ë
r*
6
ld
z Š
Ë
>
x X X x

stantonii x
omm aspensis dig X

sins a Mp ulum Hollick
"Que

Protophyll
Credneria macrophylla Heer
Credneria truncatodenticulata Bell
“Platanus” affinis Lesquereux
“Platanus” williamsii Bell
Populites cyclophylla Dawson
Protophyllum boreale Dawson x
Protophyllum leconteanum? Dawson
vr multinerve? peni X
tophyllum rugosum Lesquereux X
ED DM bipes ciem Hollick x

XX x x<

Pentalobaphyll
Araliaephyllum rotundiloba (Newberry) Fritel
Araliaephyllum groenlandica? Heer
“Sterculia” towneri (Lesquereux) Berry X

Xx

Dryophyll
Dewalquea pulchella Knowlton

ey
Dryophyllum fesses (Knowlton) Berry x x
Trochodendrophyll

Castaliites cf. cordatus Hollick (in part) x
Trochodendroides (“Cercidiphyllum”) potomacensis
(Ward) Berry

1987] CRABTREE— NORTHERN ROCKY MOUNTAIN ANGIOSPERMS 733

TABLE 3. Continued.

Lowe
Aspen Frontier Dunvegan (5)
(2) (11a) A B

Upper
Frontier
(11b)

Cinnamomophyll
“Cinnamomum” heeri Lesquereux
S r. ` hesperium Knowlton
“Cinnamom sp
Phyllites sp.
Nymphaeaphyll
Castaliites cf. cordatus (in part)
Menispermites reniformis Dawson

n
Nymphaeites exemplaris Hollick
Paleonuphar nordenskioldii (Heer) Bell
Magnoliaephyll
“Ficus” maxima Dawson
“Liriodendron” giganteum Lesquereux

* Magnolia" cf. adu Newberry
* Magnolia" tenuifolia Dawson
“Sassafras” bradleyi Brown
Cornophyll
sni Aba lesquereuxii Knowlton & Cockerell

Knowlton
Laurophyllum debile Dawson
* Magnolia" rhamnoides Bell
Monocots

Sabalites sp.
Uncertain

* Bauhinia”? cretacea? Newberry

“Fa 3" Daane Dawson

PE ied glascoeana Lesquereux

“Ficus” inaequalis Lesquereux
"Ficus"? s -

“Ficus” s

E ca flexuosum oe Bell
Leguminosites spatulatus B

X

»X X

Xx

XX X KK <

x KKK

X

! Numbers in parentheses refer to geographic and stratigraphic position of floras as indicated in Figures 1

an
š ` All species are included as originally cited in the references given belo

vn, 1933a. (11a) —Knowlton, 1917. (8)A—

Diwson, 1883. (8)B—Bell, 1963. (11b)—Berry, 1929d.

ity in the uppermost Albian and Cenomanian is
correlated with the ebb of the Albian epeiric sea-
way. As previously discussed, the seaway may
have influenced the development of endemism

in the flora by restricting movement of eastern

species into the Site rae much of the Upper
bian. / ithdr

at the end of the Albian, venden as from east-

734 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

1987]

ern and southern regions may have altered the
floristics of the southern part of the NRM region.
Although this scenario is hypothetical, the up-
permost Albian and Cenomanian floras of Wy-
oming contain Dewalquea, a genus characteristic
of floras to the south and east of the region.

A large collection of fossil leaves from the
Wayan Formation in southeast Idaho (Figs. 1,
2) is dominated by pteridophytes (Crabtree, 1983)
that are undescribed but appear to be similar to
those from the Frontier Formation in southwest
Wong (Knowlton, a The Wayan con-

p. (Figs.
21-23), which appears to iro a new species. An-
drews (1948) and Andrews & Kern (1947) de-
scribed a large collection of pteridophytic petri-
factions (Tempskya) from the Wayan (Appendix

Another undescribed megaflora is known from
several localities in the Albino Member of the
M ormation in southern Montana (Figs.
1, 2, Appendix I). An extensive fern community
is preserved in the deltaic deposits of the Albino
(Vuke, 1982; Crabtree, 1983). Dicot leaf mats
have been excavated at one locality in the Albino
(Appendix I). Unfortunately, work on these fos-
sils is incomplete and the floristics of this site
remain poorly known. Figure 38 shows a spec-
imen from the Albino that appears to be a pin-
nately compound leaf, possibly of rosid affinity,
but the venation is too poorly preserved to con-
firm such a determination.

The uppermost Albian Aspen.flora (Brown,
1933a, 1933b) from southwest Wyoming (Figs.
l, 2, Table 3) comprises nine species of angio-
sperms. No species from the Aspen are known
from the older floras ofthe region, although some
are recorded from the nearby and slightly youn-
ger Frontier Formation (Table 3). Notable in the
flora is “Liquidambar” fontanella Brown (Fig.
39), which possesses glandular teeth similar to
those of Recent Liquidambar and Altingia. ''Sas-
safras" bradleyi Brown (Fig. 40) is preserved in
a semicoalified state, which makes higher order

°
<

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

735

venation difficult to distinguish. This species
shows smooth-margined, palinactinodromous
leaves with three lobes such as in extant Lau-
raceae and certain palmate Dilleniid families.
“Sparganium” aspensis Brown (Fig. 41) is be-
lieved to be a Hamamelid fructification. Such
fossils are most commonly attributed to Plata-
noids; however, associated leaves of this form
are not present in the Aspen. It is possible that
the fructification was produced by Liquidambar
fontanella. An advanced Sapindophyll, Sapin-
dopsis schultzii Brown, is present in the Aspen,
as well as several other species of putative rosid
affinity (Table 3).

The Cenomanian flora from the Frontier For-
mation of southwest Wyoming (Hall, 1845;
Knowlton, 1917; Andrews & Pearsall, 1941) is
a mixed pteridophyte-angiosperm assemblage
preserved in volcanic sediments similar to those
which yield the Aspen and Albino floras. The
flora is from the lowermost part of the Frontier
Formation and is referred to as the Lower Fron-
tier flora in Figures 1 and 2 and in Table 3.

“Cinnamomum” hesperium Knowlton (Fig.
42) is very similar to Cinnamomophylls from
the Middle and Late Albian, and as such is the
only archaic element in this flora. Platanophylls
are represented by “Aralia” veatchii Knowlton
(Fig. 43), which is advanced over earlier mem-
bers of the morphotype by its deeply divided
lamina. Dewalquea pulchella Knowlton (Fig. 44)
is a toothed, palmately compound leaf of uncer-
tain affinity. The entire-margined, palmately
compound leaf of “Staphylea”? fremontii
Knowlton (Fig. 45), which also occurs in the As-
pen flora (Table 3), shows well-developed, eu-
camptodromous secondary venation such as
characterized ma
pound leaves are restricted to extant Ranuncu-
liidae, Rosidae, and palmate Dilleniidae. Species
with and without entire margins can be found in
each of these groups. More detailed analysis of
the venation and tooth morphology of the pal-
mately compound leaves from the mid Creta-

—

FIGURES 38-47.
Pinnately compound leaf, Mowry Form

ambar” fontanella Brown, USNM 39146, holotype. — 40.
“Sparganium” aspensis Brown, USNM 39140A, holotype. 42-47. F

type.—41.

Dicotyledonous leaves from the Mowry, Aspen, dd and Dunvegan formations. 38.
ation. 39-41. Aspen Formati

, USGS locality 8169.—39. “ Liquid-
“Sassafras” ‘bradle yi Bue USNM 39144, holo-
ontier Formation. —42. “Cin

namomum” hesperium Knowlton, USGS locality 6527, USNM re e 43-45. USGS locality 5049.—

43. “Aralia” veatchii Knowlton, portion Seir ie eaf, USN
t

pulchella Knowlton, m 35255, holot
46, 47. Dunvegan Form
boreale Dawson, GSC i y 4205, GSC 5398, holoty

d holotype. —44. Dewalquea
wlton, USNM 35230, holotype.

UN
46. Pseudoaspidonhllum. tilia Hollick, GSC locality 4197. —47. Protophyllum

736

ceous of the region ii ida pulchella, ** Dios-
= *Hymenaea" agde
Berry, Fauni ie gracile Debey, D. lanceo-
latum (Knowlton) Berry, and m fre-
montii) is necessary before further conclusions
are possible.

Far to the north of the floras just mentioned,
the Dunvegan flora (Figs. 1, 2; Dawson, 1883;
Bell, 1963, 1965) was recovered from coarse flu-
vial deposits in marked contrast to the volcan-
iclastic southern deposits. The Dunvegan flora,
unlike the more southerly floras, contains several
leaf genera that indicate considerable affinity with
early angiosperm floras (Araliaephyllum, Tro-
chodendroides, and “Cinnamomum” ), as well as
new forms. Platanophylls (Fig. 46) and Proto-
phylls (Fig. 47) are numerically dominant in the
Dunvegan assemblage. Entire-margined, pal-
mately compound leaves and Dryophylls relate
this flora to the Frontier and Aspen floras. The
Dunvegan flora contains a suite of large, entire-
margined leaves of the Magnoliaephyll morpho-
type (Table 3).

The Turonian flora (Berry, 1929d) from the
upper part of the Frontier Formation (Table 3,
Appendix I) comprises cycadophytes, taxodia-
ceous conifers, Cornophylls, Pentalobaphylls, and
fragments of the laminae of a putative palm to
which Berry (1929d) applied the name Sabalites,
although the specimens lack evidence of costa-
palmate morphology. In addition, several ro-
sidlike leaves are reported from the upper Fron-
tier (Table 3).

SANTONIAN FLORAS

The lower Santonian Badheart flora from
northeast British Columbia is notable for an
abundance of 0 2 a.
droides arctica (Heer) B yphus"
mcgregorii (Bell)). These occur in elis with
the common Upper Cretaceous araucarian co-
nifer Geinitzia formosa Heer (Bell, 1963).

LOWER CAMPANIAN FLORAS

The floristic record from this time rivals that
for the Albian. Near equivalent floras are known
from the Milk River, Eagle, and Two Medicine
formations in Alberta and Montana (Figs. 1, 2).
Floristics of the individual floras vary within a
context of dominant Hamamelididae. Also im-
portant are Cornophylls and Rhamnophylls.
Some collections contain an abundance of ar-
aucarian, taxodiaceous, and cupressaceous fo-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

liage. The small-leaved aquatic dicot Quereuxia
(cf. "Trapa"? microphylla Lesquereux) is found
in the Two Medicine. The earliest pinnate palm
from the region is present in the Two Medicine.
The woody dicotyledonous flora (Bell, 1963,
1965) of the Milk River Formation in southern
Alberta (Figs. 1, 2) is preserved in coarse fluvial
sandstones and associated sediments. Unusual is
the complete absence of Platanophylls from the
flora. Other than Trochodendrophylls, there is
an overall dearth of Hamamelidids. Trocho-
dendrophylls (Trochodendroides arctica and T.
dorfii Bell), Cornophylls (** Magnolia"? coalvil-
ensis Knowlton and Celastrinites sp.), and
Rhamnophylls (** Ficus" trinervis Knowlton) are
the most abundant dicotyledonous leaf fossils.
The Eagle Formation extends through most of
central Montana and northern Wyoming. A
number of fossil localities are known, and some
collecting has been done (see Appendix I), but
our knowledge of the Eagle flora is based at pres-
ent on the work of Knowlton (1900) in central
Montana, and Bell (1963) in northcentral Mon-
tana. I was not able to locate the original collec-
tions for either ofthese treatments (see Appendix
I), so my discussion is based on the published

nts.

Bell (1963) recognized five dicotyledons from
the Eagle. The composition of the flora is very
similar to that of the Milk River with Trocho-
dendroides dorfii, Cornophylls, and Rhamno-
phylls. In addition, the flora contains a magno-
liid-grade leaf “Ficus?” missouriensis Knowlton,
which was first described from the Eagle in cen-
tral Montana (Knowlton, 1900).

Knowlton (1900) described a small, poorly
preserved collection from the banks of the Mis-
souri River (Appendix I). The flora contains a
Platanophyll (*P/latanus"? wardii Knowlton),
which seems to be reliable to the family level. A
Rhamnophyll (** Ficus” trinervis) and a rosid leaf

veins that terminate near the sinuses ofthe teeth,
margins with rosid teeth and rosidlike sinus brac-
ing, and percurrent, transverse tertiary venation.
“Liriodendron” alatum Newberry, a large leaf of
probable magnolialean affinity, was also reported
by Knowlton (1900). Knowlton's specimen shows
little more than secondary venation, and identity
with the type from the Amboy Clays (Newberry,
1895) must be considered tentative.

A more extensive, well-preserved assemblage

1987]

from the lower Campanian Two Medicine For-
mation in northern Montana (Figs. 1, 2) is cur-
rently under investigation as part of my doctoral
program. The flora consists of about 30 species
of angiosperms, two conifers, and six ferns, and
was recovered in Oilfield Coulee (Appendix I)
from a single fluvial system extending in outcrop
for two kilometers. It records the vegetation along
a coastal plain river.

Pinnate palm fronds (Fig. 37) dominate the
channel margin facies throughout the outcrop.
The fronds are up to two meters long and are
incompletely divided into induplicate segments.
The fossil palm could be assigned to the form
genus Phoenicites (sensu Read & Hickey, 1972)
were it not for the absence of spines at the base
of the frond. Occasional interbedded paludal fa-
cies contain “Trapa” (Quereuxia). Taxodiaceous
and cupressaceous conifers are found in the basal
lag deposits of paludal facies. Nymphaeites sp.
foliage occurs in restricted parts of the outcrop.

Levee and other near-channel facies contain a
diverse dicotyledonous flora that is dominated
by leaves of Hamamelididae. Nonhamamelidid
dicot leaves typically comprise about 10% of the
nego tonsils: Dicot 43 (Fig. 49) is characterized

and weakly
percurrent tertiary venation irregularly spaced,
eucamptodromous secondary veins that origi-
nate almost perpendicular to the midvein and
turn upward rather abruptly near the margin.
The leaf morphology indicates closest relation-
ship to extant Rutales but should be considered
tentative since Rutales have pinnately com-

flora have affinities to Dilleniidae on the basis
of tooth morphology. Two species (Dicots 57 and
19, unfigured) of entire-margined, imperfectly
actinodromous leaves with thin, percurrent, ter-
tiary veins arranged in concentric series relative
to the base of the leaf indicate palmate dilleniids.

A complex of four species (three of which are
illustrated in Figs. 51-53) are related by their
c oranthoid teeth and form an important and

verse-percurrent. The leaf shows closest affinity
with Trochodendrales but cannot be accom-

CRABTREE— NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

737

modated in either of the extant genera Trocho-
dendron or Tetracentron. Another leaf in this
complex (Dicot 12, not figured) more closely re-
sembles extant Trochodendron, especially in de-
tails of tertiary and quaternary venation. Dicot
53 (Fig. 52) is a small, broadly elliptic leaf with
pinnate venation and with secondary veins clus-
tered at the base. Dicot 53 shares with extant
Cercidiphyllum such features as shape, orienta-
tion and behavior of tertiary and higher order
venation, and chloranthoid teeth with a tripod-
like arrangement of accessory veins. Cercidi-
phyllum differs from the fossil in its well-devel-
oped actinodromous venation and greater
tendency for tertiary veins to bifurcate. Dicot 53
is sufficiently like extant Cercidiphyllum that it
can be placed into Cercidiphyllales. Dicot 50 (Fig.
53) has pinnate venation, craspedodromous sec-
ondary veins with the basal pair strengthened,
and acute chloranthoid teeth. Tooth morphology
most closely approaches extant Atherosperma-
taceae; however, the craspedodromous second-
aries and strong basal apd veins of eke
50 make it a problematic species intermediat
in morphology between ae cnn
and Trochodendrales

Dicot 25 (Fig. 54) has a morphology close to
some extant Hamamelidaceae, notably Recent
Fortunearia. The fossil has strong craspedo-
dromous secondary veins that end in scalloped
teeth with spinose tips; the tertiary and quater-
nary venation is strong and branched orthogo-
nally. Dicot 25 is the most abundant and wide-
nis of the To Memomie dicot taxa and

ob abl 1 4 : 41

Ec leaves. Another hamamelid (Dicot
36, not figured) shows leaf morphological fea-
tures convergent with extant Hamamelis and
Fothergilla. Second in overall abundance at Oil-
field Coulee are two species of Platanaceae (Figs.
56, 57). Both are large, rhombic leaves with pal-
inactinodromous venation, pectinal veins, nip-
ple-shaped teeth, and orthogonal tertiary and
quaternary venation

Much of the diverse Oilfield Coulee flora can-
not be treated fully here. One leaf species (Dicot
17, not figured) is entirely consistent with the
leaf morphology of extant Menispermaceae. Sev-
eral taxa of magnoliid-grade leaf morphology are
present (e.g., Fig. 55).

Other poorly known collections from the Two
Medicine exist (Appendix I). Because the Two
Medicine Formation extends upsection to the
top of the Campanian, and I have not revisited

[VoL. 74

ANNALS OF THE MISSOURI BOTANICAL GARDEN

1987]

the collecting localities, it is difficult to assess the
age of these collections. One specimen is illus-
trated (Fig. 47) from these localities. The leaf fits
the Cinnamomophyll morphotype and may rep-
resent Laurales based on the presence of basi-
laminar secondary veins, basal secondaries at a
lower angle than superadjacent secondaries, ad-
medial orientation of tertiary venation, subpar-
allel and irregularly branched quaternary vena-
tion, and possible mesophyll secretory bodies.

MIDDLE CAMPANIAN FLORAS

Floras are known from the Judith River For-
mation in central Montana (Knowlton, 1905) and
the Oldman Formation in southern Alberta
(Penhallow, 1908; Bell, 1963, 1965). The two
floras were recovered from fluvial sandstones and
exhibit a similar composition. Simple leaves with
errate margins and pinnate, craspedodromous
venation allied to Hamamelidaceae predomi-
nate. There is some evidence for the presence of
Trochodendrophylls (Hickey, 1984). Diverse
Platanophylls, including “Platanus” affinis Les-
quereux, are found in the Oldman flora. “Arto-
carpus" sp. (Bell, 1965) from the Oldman is a
pinnately compound leaf, the leaflets of which
are pinnately veined with eucamptodromous
secondary veins and many intersecondaries. This
morphology relates it to rosid groups, including
Sapindales, Rutales, and Rosales. Dawson (1886,
1887) described plants from the Belly River se-
ries, which is equivalent to the Judith River-
Oldman floras. His species appear unusually ad-
vanced for the Campanian, an attribute Knowl-
ton (1900) ascribed to stratigraphic mixing of the
collections. Ramanujan (1972) reported on per-

1977), and the Livingston flora (Knowlton, 1893)

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

739

from southcentral Montana, the Adaville flora
from southwest Wyoming (Dorf, 1955), and the
Point of Rocks flora from southcentral Wyoming
(Brown, 1956; Dorf, 1955; Knowlton, 1900; Les-
quereux, 1878; Newberry, 1898; Ward, 1885,
1887)

SUMMARY

During an approximate three million year pe-
riod of the Middle and Late Albian, angiosperms
displaced gymnosperms as the dominant low-
land vegetation of the NRM region. This oc-
curred shortly after the entry of angiosperms into
the region. Upland vegetation retained a pre-
dominantly gymnospermous composition. Pte-
ridophytes remained important through this
period, although composition changed. Gink-
gophytes, nm and Pteridosperms de-
creased dramatically.

Although "m pec Um of angiosperms can be
seen as the essential agent of change, several en-
vironmental factors correlate with the floristic
changes in the Albian. The Cretaceous epeiric
seaway expanded across the mid-continent for
the first time during the Albian. Climatic sea-
sonality may have increased due to increased
orogenic ee in the Cordillera and increased
temperatur

arly angiosperms with low rank leaves such
as Rogersia and Ficophyllum are not recognized
in the NRM region. The Middle Albian flora is
closely allied to time equivalent floras at lower
latitudes in North America. The similarity is most
pronounced at the level of genus, although a few
species appear to have ranged across North
America during this time. As in the Potomac
flora, Sapindophylls and Platanophylls form a
conspicuous element. Other groups, such as the
Cinnamomophylls and Pentalobaphylls, appear
to have achieved prominence earlier in the NR
region than elsewhere in North America and im-
part a uniqueness to the regional flora.

Leaf morphologies indicate that early mem-

,

FIGURES 48-58. Angiosperm megafossils from the Two Medicine Formation. Specimen numbers correspond
to the research collection housed at the Department of Botany, University of Montana. 48-57. “Oilfield Coulee”

locality.— 48.

Pinnate palm, 23A/M1/003.— 49. Rutales?, Dicot 43, 564/43/001.— 50. Dilleniidae, Dicot 1,

40A/01/001.—51. Trochodendrales, Dicot 11, 50A/11/001.— 52. Cercidiphyllales, Dicot 53, 57A/53/001.— 53.
Indeterminate leaf with craspedodrome secondaries, strong basal secondary veins, and chloranthoid teeth, Dicot
50, 55A/50/001.— 54. Hamamelidaceae, Dicot 25, S174A/25/010.— 55. Magnoliid-grade leaf of uncertain af-
finity, Dicot 27, 39B/27/002. 56, 57. Platanaceae. — 56. Dicot 32, HPC/32/022.— 57. Dicot 6, 24A/06/021.—

58. Laurales, USGS locality 6015.

740

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

HAMAMELIDIDAE

PLATANALES

CONIACIAN

(PLATANALES)
COMPLEX
">>>
Pentolobophylls Gianna)
LATE MIDDLE ALBIAN UPPERMOST ALBIAN
m a
(HAMAMELIOALES 7 GAPINDALES)
K
GRADE
COMPLEX

EUPHORBIALE
PALMATE DILLENIIDAE

@|

T

CAMPANIAN

FIGURES 59-62. Changing diversity and floristic affinity of the pre-M

a:

e
northern Rocky Mountain region. Occurrences of taxa are based on interpretation of leaf biduo ex iod in
the following cases: Chloranthales (Clavatipollenites), Hippuridales (Retitricolpites microreticulatus Brenner),

Fagales (Tricolporopollenites sp.), and Euphorbiales (Erdtmanipollis). Muller (1981) accepted these occurrences
).

as indicative of the presence of the taxa (see also Norris et al.,

bers of several Recent higher taxonomic cate-
gories of angiosperms existed in the Albian flora
(Figs. 59, 60). Sapindophylls appear to be allied
to primitive Rosidae. Hamamelididae is well
represented by Platanophylls, Protophylls, and
Trochodendrophylls. Magnoliidae allied to
Laurales, Magnoliales, Chloranthales, and Nym-
phaeales are probable. Monocotyledons are not
apparent in the megaflora of the NRM region
until the Late Cretaceous.

The early angiosperm flora includes several
prominent groups with no clear relationship to
Recent taxa (Table 1). Pentalobaphylls exhibit

Cornophyll, and Magnoliaephyll are informal
categories based on leaf morphology. The names
do not imply relationship to any modern group,
merely convergence in leaf morphology with

1975; Rouse et al.,

modern forms. Much of the diversity of ined
dicotyledons is manifest in these morphotype
New higher level taxa will probably ve s
to allow for their adequate classificat

Cenomanian floras in the region retain some
older elements (Platanophylls, Protophylls, Cin-
namomophylls, and Trochodendrophylls are
worthy of mention), but on the whole are more
modern in aspect, contain higher rank leaves,
and are more diverse than Albian floras. Latest
Albian and Cenomanian floras from lower lati-
tudes within the region contain Dryophylls and
Dewalquea.

Post-Cenomanian floras continue to diversify,
attaining levels of advancement consistent with

d
gales), palmate Dilleniidae, and Rosidae are a

1987]

persistent part of the flora. Magnoliidae (Lau-
rales, Chloranthales, Nymphaeales, and Mag-
noliales) appear as members of individual floras.
Ranunculiidae (Menispermaceae) is recognized
from the lower Campanian in Montana. Pinnate
palms date from the Lower Campanian. The
group is historically associated with stream mar-
gin environments

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ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

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CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

745

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APPENDIX I
LOCALITY INFORMATION, CHRONOSTRATIGRAPHIC
REFERENCES, AND LOCATIONS OF COLLECTIONS FOR
NORTHERN ROCKY MOUNTAIN REGION FLORAS
Megafossil collections referred to in this report are
listed below in alphabetical order. Numbers i in paren-

used in Figures 2 and 3 and in Tables 2 and 3. Chrono-
stratigraphic references are placed in parent theses i im-

cality information absent or sketchy. All others are
referenced to the publications that provide locality
specifications. Institutions where collections are housed
are indicated.

ALBINO (1). Upper Upper Albian (Roberts, 1972). I
made a small collection from leaf mats in the Albino
Member of the Mowry Formation along Beaver Creek,
Gallatin County, Montana. This locality is in pink and
white volcaniclastic siltstones of the Albino Member
of the — = rmation in the NW bs od 1⁄4, sec. 18,
T.7S,R.4 Department
of Botany, G of Montana. jio extensive fern

ora is also known from the Albino Member (Crabtree,
1983)

N (2). Upper Upper Albian (Cobban & Reeside,
19522; Nichols & Jacobson, 1982). This e is
from the predominantly marine Aspen gon tion at
USGS locality 8168 located in the NW 4, sec. 6, T
24 N, R. 115 b n
County, Wyom
here, and the SUE occur 125 feet from the top (Brown,
ines Collections are housed at the U.S. National

um.

Lower Santonian Pa 1961, 1963;
Bell, 1963). Collections are from the Badheart For-
mation in the vicinity of Belcourt m eastcentral
British Columbia. Collections are housed at the Geo
logical Survey of Canada

BEAVER MINES (4). Lower Middle Albian (Bell, 1956;

Mellon, 1967; Mellon & Wall, 1963; Stott, 1968; Singh,

1971). pix are from the Beaver Mines For-

mation in the vicinity of Crowsnest Pass, southwest

Alberta. Collections are housed at the Geological Sur-
vey of Canada

BADHEART (3).

BLACKLEAF (5, 19, 20). Lower Upper Albian (Cannon,
1966; Rice & Cobban, 1977). Collections come from
several USGS localities in the Vaughn Member of the
Blackleaf Formation in northwestern Montana. Col-

746

lections are described under sh headings which cor-

The single locality i is in the NW
Ys, SE 4, NE 4, sec. 13, T about 5 km
north of Great Falls, TM road Montana. Col-
lections are “itt at the Department of Botany, Uni-
versity of Montan
Summit (19). Collecting Ege are scattered for
ab s Pass alon

ties 5984, 5985, 5986, 6007, and 9439 in Glacier Coun-
ty, Montana. A separate area exists on the west side of
the pass in the S '2, sec. 1, T. 29 N, R. 14 W Flathead
County, Montana. Collections are housed at the U.S.
National Museum with the exception of a small col-
lection from '2 mile west of Marias . which is at
n rsity of Montana.

USGS locality 9437 from the SW
1⁄4, SE “4, NE 4, sec. 36, T. 22 N, R. 13 W on the north
side of the Sun River, Teton County, Montana. Col-
lections are housed at the U.S. National Museum

COMMOTION (6a, 6b, 6c). Collections are from three
members of the Commotion Formation and are housed
at the Redpath Museum and the Geological Survey of
Canada. Each member is treated separately below.

Lower Gates (6a). Lower Middle Albian (Bell, 1956;
Mellon et al., 1963; Stott, 1960, 1968; Singh, 1971).
Collections are from the lower Gates Member in the
vicinity of Pine River, northeast British Columbia.

Upper Gates (6b). Middle Middle Albian es
1956; Mellon et al., 1963; Stott, 1960, 1968; Singh,
1971). Collections are from the upper part of the Gates
Member along Belcourt Ridge, eastcentral British Co-
lumbia.

Boulder Creek (6c). Middle Middle Albian (Bell,
1956; Stott, 1968; Singh, 1971). Collections are from
the Boulder Creek Member in the vicinity of Pine Riv-
er, northeast British Columbia

veteri (7. Lower Upper Albian (Bell, 1956;
Mellon & Wall, 1963; "Mellon: 1967). C ollections are
from the Mill Creek Formation in the vi

ange in southwest Alberta. ee are housed at
the Geological Survey of Cana

DUNVEGAN Cenomanian (Bell, 1963; Stott, 1960,

housed at the Redpath Museum and the Geological
Survey of Canada

EAGLE (9). Lower Campanian (Cobban & Reeside,
1952a; Russell, 1970). The Eagle flora reported by
Knowlton (1900) was collected by Lester Ward in 1883.

Knowlton’s ipe du several of them types, are miss-
ing from the U.S. National Museum

I attem wee to relocate Ward's site and found a small
assemblage from along the Missouri River below Rat-
tlesnake Coulee about 12 km south of nie Banks,
Chouteau County, Montana, in section 24, T. 26 N,
R. 12 E. This paca is at the Department A en
University of M

Bell (1963) E a flora from Buckley and Red

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Coulees, Toole County, Montana, in the vicinity of the
International Boundary south of the town of Milk Riv-
er, Alberta (see Evans, 1930). I was unable to locate
this —— at the Geological Survey of Canada.

In recent years, collecting has been carried out at
USGS “locality 7633 from the Eagle Formation by L.

W. His eve are housed at the U. S. National
Museu

FALL RIVER (10). Upper Albian Reds pers. obs.).
Whileat the U.S. National Museum I examined several
specimens from USGS locality 7312 i in the general
collection. The ledger of field collection sites indicates
that these plants were found in the Fall River Sand-
stone near Pine Creek, Wyoming. This is probably the
Pine Creek mentioned by Ward (1899). As best I can
Judae from Ward’s descriptions, the fossils are from

R. 62 W, T. 55 N in Crook County, Wyoming, from
a level 60 feet below the Dakota ia This locality
could be as old as Middle Albia

FRONTIER (lla, 11b). Two main gaa levels
in the Frontier Formation have yielded plant mega-
fossils. These are treated separately below.

Lower Frontier(1la) Lower omanian (C fus
& Reeside, 1952b; Nichols & Jacobson, 1982). M
— were made from nis lower Frontier For.
matio eldwork Hall (1845)
seco on the earliest collections from the Fr

Char emo

County, Wyoming, in the S '4, sec. 29, T. 19 N, R. 116
.S. Ge ological Survey UM from this area
and the adjacent sec 116 W include

USGS field localities 8676, n i and 5049.
Knowlton (1917) reported on collections primarily from
locality 5049. Subsequent collections from this and
nearby localities were reported by Andrews & Pearsall
(1941). All of these localities are from the same strati-
graphic level about 1,200 feet below the Oyster Ridge
Sandstone and are considered to be of lower Ceno-
manian age. Hall's original collection no longer exists.
Knowlton's published collections are housed at the U.S.
National Museum

versity of Connecticut. Scania collections are at the
U.S. National Museu
Upper Frontier (1 Ib). Turonian (Nichols & Jacob-
son, 1982). Collections from the Frontier near Lander,
Wyoming (Berry, 1929d) are from well above the Oys-
ter Ridge Sandstone. These collections are housed at
the U.S. National Museum.

JACKASS MOUNTAIN (12). Middle Ped eg ee
1956). Collections are from localities th
Mountain Formation in southcentral British ciens
and are housed at the Geological Survey of Canada

(=
£e

JUDITH RIVER-OLDMAN (13). Middle Campanian
(Cobban & Reeside, 1952a; Russell, 1970). The col-
lections from the Judith River Formation described by
Knowlton (1905) were collected at USGS locality 3272
alon 4 km east of old Fort

a
locality information for the Oldman Formation i
southern Alberta, but see Ramanujan (1972) and Bell

1987]

(1965). Judith iia collections are housed at the U.S.
National Muse

KINGSVALE (14). Middle pipe debe (Bell, 1956).
From localities in the Kin mation in south-
central British Columbia. C ollections are housed at the
Geological A of Cana

ir IVER (15). pedido vene (Bell, 1963; Rus-
l, 1970). Localities = in the iver Formation
ie Milk River, s Alberta (Bell 1963). Collec-
Gols are housed at e Geological Survey of Canada.

MILL CREEK (16). Lower Upper Albian (Mellon & Wall,
1963; Mellon, e Collections are from the Mill
Creek Forma ear Crowsnest Pass and north along

tion
the foothills in Glad Alberta. They are housed at
the Geological Survey of Canada
Middle Upper Albian (Barksdale,
the Pasayten Fo tion in

PASAYTEN (17

at the Geological Survey of Canada. specim
d. A - is at the Department of Botany, University
of Montana

PRE-MUDDY (18). Middle Middle Albian (Roberts,
1972). S. Vuke made this collection from beds below
the Muddy Sandstone V Buck Creek, Gallatin
County, Montana, in T. 8 S, R. 3 E. It is housed at the
Department of Botany, University of Montana.

TWO MEDICINE (21). Lower Campanian (Russell, 1970;
Lorenz, 1981). Collections are from the lower
the Two Medicine Formation in “Oilfield Coulee” about
20 m above the contact with the Virgelle Sandstone in

CRABTREE—NORTHERN ROCKY MOUNTAIN ANGIOSPERMS

art of

747

the SW '4, sec. 18, T. 32 N, R. 5 W about 6 km south
of Cut Bank, Glacier County, Montana. These are
housed at the Department of Botany, University of
Montana. Unstudied collections from USGS localities
6009, 6010, 6011, 6013, 6015, 6129, 6388, 6390, 8574,
and 9449 in Glacier County, Montana, are housed at
the U.S. National Museum.
WAYAN (22). Upper Upper Albian or Lower Ceno-
manian (Cobban & Reeside, 1952a; Moritz, 1953).
During the course of many years, an amateur collector
named Thomas Henry made a very large collection of

in the vicinity of Wayan, Caribou County, Idaho. The
exact locality (-ies) is not known, but appears to have

n in sec. 24, T. 5 S, R. 43 E and secs. 19 and 20,

58, » x E. I could not . the sites during
my own visits to the area in 1983. Most of the Wayan
in AURA 1S ae at the Department "e Geology, Ida-
ho State University. The specimens figured in this re-
port (Figs. 21—23) are Dr in the Department of
Botany, University of Mont

WINT P (23). Upper Albian or Cenomanian
(pend e 1975; R. Rau, pers. = Collections
housed at the U.S. National Museum and reputed to
be from the hippies Formation are probably Paleo-
cene in age (J. Wolfe, pers. comm.). Material in T
report was collected by myself, R. Rau, and J. Robis
from the type locality of the Winthrop Formation near
the Boesel Ranch along the Methow River road in the
NW 4, NE 4, SE %4, sec. 14, T. 35 N, R. 20 E, Okanogan
County, Washington. This collection is housed at the
Department of Botany, University of Montana.

EOCENE AND OLIGOCENE FLORAS AND VEGETATION
OF THE ROCKY MOUNTAINS!

Scott L. WING?

ABSTRACT

1 T ^A (1);

Yes of major evolutionary, biogeographic, and vegetational

UE ti
changes in Sepie plants. From a floristic perspective three major trends characterize this period
cky Mo

in the Ro ntain region: 1) development

of more distinct phytogeographic provinces from a

relatively Cana s Paleocene holarctic flora, 2) the early diversification in Eocene upland areas
d

milies, an

) the first appearances of many extant angiosperm
Pale

evergreen forest to high northern latitudes (60°N), possibly ` the greatest geographic coverage suc ch

vegetation everachieved. Thi

more open vegetation of lower stature in many areas. These floristic and vegetational trends ` were of

great conseq

uence for angiosperm evolution and biogeography. They were also significant in creating

the ecological milieu for the evolution of early and mid Tertiary animals.

The Paleogene rocks and fossils of the Rocky
Mountains and adjacent areas provide an excep-
tional window on the flora and vegetation of the
time and place. Eocene and Oligocene rocks cov-
er some 40% of the surface of the state of Wy-
oming (106,000 km?), and their total outcrop
area in the Rocky Mountain region is in the range
of 300,000? km (Fig. 1). Over much of this vast
outcrop, weathering has exposed the strata, mak-
ing them available for paleontological and geo-
logical study. Eocene-Oligocene deposits from a
variety of fluvial, lacustrine, and volcanically in-
fluenced settings have produced tens of thou-
sands of fossils collected at hundreds of localities.
These Paleogene sequences have the potential to
yield one of the most complete and geographi-
cally extensive records of evolutionary and eco-
logical change in a terrestrial biota. A century or
more of paleontological and geological research
has only begun to explore this potential, in part
because of the vast area and great thickness of
the sedimentary sequences.

This paper is intended to serve three goals.
First, to set out the basic data on the location,
stratigraphic positions, and relative ages of the
fossil floras. Second, to summarize published
work on Eocene and Oligocene flora and vege-
tation of the region and discuss previous hy-
potheses in light of current data. Third, to suggest
new directions for future research aimed at a
regional understanding of Paleogene floras that

takes full advantage of the potential of such a
widespread and prolific fossil record.

GEOLOGIC SETTING

The Rocky Mountain region is geologically di-
verse, and its Cenozoic history has been reviewed
in a number of recent volumes (e.g., Robinson,
1972; Curtis, 1975; Flores & ise 1985).
Eocene and Oligocene plant fossils are best known
from the large intermontane basin fill deposits
of the eastern Rocky Mountain area and from
the primarily volcanic deposits of western Mon-
tana, northwestern Wyoming, Idaho, Oregon, and
Washington. The separate depositional histories
of these two areas are important for interpreting
historical change in their floras.

The intermontane basins of the eastern Rocky
Mountain area (Fig. 1) began to develop in the

way. Uplift of local, initially east-west trending,
mountain ranges during the latest Cretaceous-
Paleocene began to divide the foreland basin into
structurally, topographically, and sedimentolog-
ically discrete units. Consequently, each of the
present intermontane basins has a separate Ce-
nozoic history, yet some general trends have af-
fected most or all of the basins.

Fluvial to paludal environments were preva-

l inen to Jack A. Wolfe, Ralph Taggart, and George Rogers for review, and to Mary Parrish for drafting

the figur

: on PEN of Paleobiology, MRC 164, Natural History Building, Smithsonian Institution, Washington,
S.A.

D.C. 20560, U.S

ANN. MISSOURI Bor. GARD. 74: 748-784. 1987.

1987]

120

*
`< x

CARN P

VOLCANIC

FIELD

E Eocene BASIN FILL
OLIGOCENE

ll EOCENE/OLIGOCENE VOLCANICS

FIGURE 1.

lent throughout most of the area in the Paleocene
and were environments of deposition for coals
in the Powder River, Bighorn, Green River, and
other basins. Fresh or brackish water lakes formed
in the Uinta, Wind River, and Bighorn basins
during the latter half of the Paleocene (Johnson,
1985; Keefer, 1965; Yuretich et al., 1984). Be-
ginning in the Clarkforkian? and continuing into

3 Through much of this paper, North American pro-
vincial land m

are known to be door VA i ede ar.
out western North America (Flynn 84) and
because it is easier to Pieces floras S. m mam-
malian sites than with the type age/stages of Europe.
Correlation of NALMA with epochal boundaries and
time are given in Figure 2.

WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

BEARPAW
VOLCANIC*

YELLOWSTONE/ABSAROKA
VOLCANIC FIELD

BIGHORN 234,05.
ASIN -
L POWDER RIVER
BASINI

e
WIND RIVER

GREAT DIVIDE € Il

BASIN Xe

E ANDwASH BASIN u
— C Ma PARK

yea

PICEANCE CREEK
BASIN)

to
t... -NINE MILE
eg VOLCANIC FIELD

SAN JUAN
BASIN

ok]

NM

N

Outcrop area of Eocene and Oligocene rocks in the greater Rocky Mountain area

the Wasatchian, paludal deposition of organic-
rich sediment waned in most of the basins, and
alluvial floodplain sediments began to be mod-
ified by oxidizing pedogenic processes (Bown
1979; Bown & Kraus, 1981). The onset of this
kind of pedogenic modification was probably
controlled by a combination oflocal tectonic and
physiographic factors, as well as by regional cli-
matic change (Wing & Bown, 1985
In the later Wasatchian and Bridgerian, large
lakes again developed in many ofthe basins (Big-
horn, Green River, Uinta-Piceance). In at least
some cases this appears to reflect hydraulic clo-
sure of the basin rather than climatic change
(Johnson, 1985), and the lakes of the Green Riv-
er and Uinta-Piceance Basins are known to have
been saline during much of their later histories.

749

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

RRENEEEESEEZSEXEEREEXEEXGEEEM. Ts
* Ë = O C E N E ‘OLIGOCENE Ë

[ EARLY MID AT EARL | .
^ TRETA | srIoceRIAN | UINTAN | | DUCHESNEAN Ë E ~ ARIKAREEAN |

E 2. Correspondence of North American Land Mammal Ages (NALMA) with epochal and stage
).

FIG
efie pr (after Berggren et al., 198

Much ofthe sediment filling these middle Eocene
lakes and forming coeval fluvial deposits was
derived from newly active volcanic fields in west-
ern Wyoming and Idaho

During the later Bridgerian and Uintan most
of the intermontane basins began to fill with sed-
iment, and during the Duschesnean and Chad-
ronian these sediments began to lap outward onto
the tops of the bordering ranges. During the Oli-
gocene, volcanically derived sediment continued
accumulating at relatively slower rates in the
eastern Rocky Mountains and Great Plains. Oli-
gocene formations over much of the northern
Rocky Mountains and Great Plains are typically
thin, widespread units that were probably de-
posited on a surface of low relief.

The history of the volcanic centers in the west-
ern part of the northern Rocky Mountains is
largely independent of that of the intermontane

the Elkhorn volcanic
field) beginning in the latest Cretaceous and
throughout the Paleocene (Roberts, 1972). How-
ever, the greatest pulse of volcanic activity began
in the late Wasatchian to Bridgerian and contin-
ued through to the end of the Eocene

From about 53 to 38 Ma there was volcanism
across much of western Montana (Bearpaw, Gar-
net Range, Lowland Creek, Gallatin, and Bea-
verhead Canyon fields), northwestern Wyoming
(Yellowstone—Absaroka Field), and Idaho (Chal-
lis Field), and in the Clarno area of Oregon, the
Republic graben of northern Washington, and
southern British Columbia (Chadwick, 1985).
Although the intrusive and sedimentary rocks

m, it is likely that
most of their original extent has been removed
by erosion. Armstrong (1974) suggested that the
Challis volcanic field once covered much of Ida-
ho

The extent and thickness of these broadly co-
eval volcanic deposits in the northern Rocky
Mountain area indicate the region may have been
a rugged, but more or less connected, volcanic
highland during much of the Eocene (Axelrod,

1968; Fritz & Harrison, 1985). The prevalence
of volcanic conglomerates and mudflows, and
analogy with present day volcanic terrains, sug-
gest strong topographic relief, which Fritz & Har-
rison (1985) estimated at 1,000-2,500 m differ-
ence between adjacent valleys and peaks.

Following the end of the Eocene, heavy vol-
canic activity shifted southward to southern Col-
orado, exico, Arizona, and west Texas,
and extensional tectonics began to affect the
northern Rocky Mountains (Chadwick, 1985). A
number of small, fault-bounded basins devel-
oped in western Montana during the Late Eocene
and Oligocene. These basins filled with primarily
fluvial and lacustrine deposits that were derived
from both local sources and volcanics as far away
as the Cascade Range of Oregon (Fields et al.,
1985)

The geological events summarized above are
highly relevant to the floristic and vegetational
history of western North America. The devel-
opment of a volcanic highland region in the
northern Rocky Mountain region during the
Eocene had several major effects on plant life,
including: 1) dividing lowland areas that for-
merly had been continuous from the West Coast
to the eastern Rocky Mountain region, 2) reduc-
ing the flow of Pacific-influenced air into the in-
terior of North America, and 3) changing soil
and groundwater conditions by the influx of air-
borne volcanic ash. Prior to and during the onset
of these effects the low-lying area in the eastern
Rocky Mountains was itself broken into a series
of topographically isolated and somewhat cli-
matically different basins. Thus the overall effect
of geological events during the Eocene and Oli-
gocene was fragmentation of a relatively ho-
g both north-south and

mogeneous ]

Fr

east—west axes.

DISTRIBUTION OF FOSSIL FLORAS

NI Tm a 1 : A
n

of Tertiary plant localities in western North
America. The list of localities given here (Ap-
pendix I) represents an attempt to bring together

1987]

the bulk of Eocene and Oligocene localities of
the greater Rocky Mountain m I estimate
that this list is 60-80% compl

The locality information was m from a
number of published and unpublished sources.
Previous works listing a significant number of
localities include those by Axelrod (1966a, 1966b,
1968), Axelrod & Raven (1985), Brown (1937),
Hickey (1977), Leopold & MacGinitie (1972),
MacGinitie (1941, 1953, 1969, 1974), and Wolfe
(1981, 1985). Unpublished locality information
was derived from collections at the U.S. National
Museum of Natural History (including those of
the USNM, the U.S. Geological Survey, and some
formerly belonging to Princeton University), and
from collections at the Museum of Paleontology
(University of California, Berkeley) and the Yale
Peabody Museum.

Not all of the 240 “localities” compiled here
are strictly equivalent. Some floras commonly
considered as units actually consist ofthe summed
floral lists of a number of separate quarry sites
(e.g., the Ruby paper shale flora of Becker, 1961,
was collected at 1 5 quarries), whereas others were
derived from a single excavation (e.g., Copper
Basin flora of Axelrod, 1966a). Most of the listed
localities are individual quarry sites, but for some
floras listed in the literature it was not possible
to determine how many sites contributed to the
flora.

Criteria used in correlating and determining
the ages of floral localities included mammalian
biostratigraphy, lithostratigraphic equivalence,
magnetic polarity stratigraphy, and radiometric
dates. In order to avoid circularity, floras were
not correlated on the basis of their taxonomic
composition or physiognomic characteristics. The
Tertiary time scale used here is that of Berggren
et al. (1985). Chronological boundaries for North
American Land Mammal Ages are from West et
al. (in press), and Prothero (1985a, 1985b). There
is controversy over the precise age of the Eocene/
Oligocene boundary. The date used here (36.6
Ma) is the most recent opinion, but many authors
(e.g., Wolfe, this volume) use a date of about 34
Ma. Radiometric dates from older publications
have been corrected for new constants using the
tables published by Dalrymple (1979).

Fossil floras of Eocene and Oligocene age have
a highly uneven geographic and stratigraphic dis-
tribution. Stratigraphically, the Wasatchian and
Bridgerian are by far the best-represented time
intervals, with the number of floras diminishing
sharply through the Uintan and Duschesnean,

WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

751

and into the Oligocene (Fig. 3). Geographically,
the northern Rocky Mountain region is far better
sampled than the southern region (Fig. 4). These
inequities in distribution result from at least four
factors. 1) Bias of the author: my work has cen-
tered on the lower Eocene of Wyoming, so I am
particularly aware of these localities. 2) Wa-
satchian, Bridgerian, and early Uintan sections
are thicker and more widely exposed than later
Eocene and Oligocene rocks. 3) Later Eocene and
Oligocene rocks are generally less fossiliferous
than early Eocene rocks, probably reflecting less
favorable conditions for plant preservation as-
sociated with a drying climate in the Western
Interior. 4) Collecting has been more intense in
some areas than in others.

In addition to the uneven distribution of floras
in space and time, there are also changes through
time in environments of deposition. Nearly all
Wasatchian plant assemblages were recovered
from rocks deposited in fluvial or fluvial/paludal
settings. Although some Bridgerian and later flo-
ras are from fluvial deposits, the best known (e.g.,
Green River, Florissant) are from lacustrine sed-
iments. Floral samples drawn from fluvial en-
vironments represent the flora differently than
do those from lacustrine environments.

Fluvial assemblages, particularly those depos-
ited on low-energy flood basins, are largely de-
rived from local vegetation (Scheihing & Pfef-
ferkorn, 1984; Spicer & Greer, 1986; Burnham,
in press). This results in considerable spatial het-
erogeneity in the fossil assemblage that probably
reflects original variation in the local vegetation
(Hickey, 1980; Wing, 1980, 1984)

By contrast, lakes receive input from sur-
rounding lakeshore vegetation and from inflow-
ing rivers (Spicer, 1981). These different ele-
ments may be mixed to various degrees and
scattered across the lake bottom. Although la-
custrine assemblages are overwhelmingly dom-
inated by plant parts derived from nearby vege-
tation (Drake & Burrows, 1980; Spicer & Wolfe,
1987), they tend not to preserve in situ variation
in vegetation. They may, however, preserve more
of the species that were present in the regional
flora. As a result of being derived from a larger
source area and of predepositional irai and
mixing, l
diverse than fluvial assemblages, udi specimens
are more evenly distributed among the taxa pres-
ent.

As a consequence of the inequities in distri-
bution of fossil floras and nonequivalence in their

bly more

ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

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FIGURE 3.

Correlation chart of Eocene and Oligocene floras from the Rocky Mountains and surrounding

areas. Numbers indicate floras listed in Appendix I; letters (A-FF) indicate radiometric dates listed in Table 1.

1987] WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

Green River Washakie Na mam
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FIGURE 3. Continued.

754

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

TABLE |. Radiometric dates of Tertiary floras in Figure 3.
Age Stratigraphic Unit Publication

A, 49.2 + 0.7 Trout Peak Trachyandesite Smedes & Prostka, 1972
B, 50.5 + 1.55 Sepulcher Formation Smedes & Prostka, 1972
C, 43.1 iggins Formation Bown, 1
D, 44.6 Wiggins Formation Bown, 1982
E, 46.7 Wiggins Formation Bown, 1982
F, 47.1 Wiggins Formation Bown, 1982
G, 47.9 + 0.5 Wiggins Formation Bown, 1982
H, 48.5 Wiggins Formation Bown, 1
L 50.4 Aycross Formation MacGinitie, 1974
J, 50.6 Aycross Formation MacGinitie, 1974
K, 42.3 + 1.4 Henry Ranch Member, Wagon Bed Formation Black, 1969
L, 46.2 Wagon Bed Formation, near Badwater, Wyoming Evernden et al., 1964
M, 50.5 Halfway Draw Tuff, Wind River Formation Evernden et al., 1964
N, 50.2 Little Mountain a beu Peak Member,

Green River Form Mauger, 1977
O, 46.7 + 0.9 Washakie Formation, pu 664 of Roehler Mauger, 1977
P, 462 several hundred feet above Mahogany Ledge,

Parachute e Member, Green River Formation MacGinitie, 1974
Q, 26.5 Creede Formation Axelrod, 1987
R, 34 Antero ign Epis & Chapin, 1975
S, 35 Florissant Formation Epis & Chapin, 1975
T, 36-37 Wall Mountain Tuff Epis & Chapin, 1975
U, 36 "Chicken Creek Formation," 5 feet above highest

floral locality Axelrod, 1966b
V, 41+1 Deadhorse Tuff Axelrod, 1966b
W, 43 “Frost Creek m 1,500 feet below lowest

floral loc Axelrod, 1966b
X, 31.1 Williams ced ai Fields et al., 1985
Y, 41.1 + 1.6 Salmon ar Fritz & Harrison, 1985
Z, 44.2 + 1.7 almon area t Fritz & Harrison, 1985
AA, 46.3 + 1.0 rhyolite below “Dewey Beds" Fritz & Harrison, 1985
BB, 47.2 + 1.8 basalt above Germer Tuffaceous member Edelman, 1975
CC, 48.0 + 1.0 Latite-andesite Member, Challis Volcanics Edelman, 1975
DD, 47.0 + 1.8 Klondike Mountain Formation Wolfe & Wehr, 1987
EE, 48.2 + 1.6 Klondike Mountain Formation Wolfe & Wehr, 1987
FF, 50-51 Sanpoil Volcanics Wolfe & Wehr, 1987

depositional environments, there are obvious dif-
ficulties in interpreting patterns of vegetational
and floristic change through time. In spite of these
difficulties, a number of durable generalizations
have emerged concerning the early Tertiary de-
velopment of the Rocky Mountain flora.

FLORISTIC AND VEGETATIONAL HISTORY

Three major floristic trends are indicated by
Eocene-Oligocene fossil assemblages from the
Rocky Mountain area: the modernization of the
angiosperm flora at the generic level, the breakup
of the Paleocene North American province into
distinct phytogeographic subregions, and the
major diversification of many present-day mi-

crothermal lineages (e.g., Rosaceae, Betulaceae,
Aceraceae). There were also three major vege-
tational periods during the Eocene and Oligo-
cene. During the Wasatchian and Bridgerian there
appears to have been a major poleward expan-
sion of subtropical and paratropical forest types
in response to global climatic warming (Wolfe,
1985). Perhaps beginning as early as the Clark-
forkian in some areas, and of increasing impor-
tance during Bridgerian and later time, there is

MacGinitie, 1969). Finally, during the latest

1987]

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WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

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JUNCTION

Map position of Eocene and Oligocene floras mentioned in the text and in Figure 3. Numbers
dix I.

"us ^ n listed in Appen

Eocene and Oligocene, vegetation of the Rocky
Mountain region became dominated by a mix-
ture of conifers and broad-leaved deciduous
forms.

MODERNIZATION

Paleobotanical systematics of the last century
and early in this century was based largely on
superficial characteristics of leaves. This led to
exaggerated estimates of the similiarities be-
tween extinct and modern floras. More recent
systematic work has focused on detailed com-
parisons of leaf venation (e.g., Hickey, 1977) and
on greater use of multiple organs (e.g., Man-
chester, 1986). One result of such work has been

the realization that many Paleocene angiosperms
represent extinct genera or intermediates be-
tween several related living genera.

In contrast, many, if not most, angiosperm
remains younger than Late Eocene can be as-
signed to extant genera with little ambiguity, al-
though sectional or other subgeneric-level affin-
ities may be unclear (see Manchester & Crane,
1983 and Wolfe & Tanai, 1987 for well-docu-
mented exceptions). The temporal pattern of ap-
pearance of extant genera has not been quanti-
fied, nor has it been determined the extent to
which this modernization reflects evolution
within lineages as opposed to extinction of ar-
chaic lines and replacement by modern ones. Al-

756

though this generic modernization of angio-
sperms is a striking systematic pattern, little has
been said about its biological significance. Mod-
ernization at the generic level may reflect a true
radiation of angiosperms, perhaps in response to
changing climatic and topographic conditions in
western North America, or it may be an artifact
of our retrospective view. That is, the appearance
of many modern genera in the Eocene occurred
because angiosperms in a number of independent
lineages accumulated enough recognizable ge-
neric characters to be pigeonholed easily in pres-
ent-day categories. This question could be re-

solved by measuring and comparing rates of

morphological change during the Paleocene and
Eocene.

PROVINCIALITY

The Eocene divergence of Rocky Mountain
and West Coast floras and its relationship to East
Asian-North American disjunct distributions in
a number of living plant groups is probably the
most discussed aspect of the Tertiary paleobo-
tanical record in western North America (e.g.,
MacGinitie, 1941; Leopold & MacGinitie, 1972;
Wolfe, 1972; Hickey, 1977). Consequently, I will
summarize the pattern only briefly.

Paleocene floras from across northern North
America and to some extent of Europe and Si-
beria are relatively homogeneous (Wolfe, 1966).
Leaf assemblages typically are of low diversity
and dominated by a group of taxa including
Ginkgo, Metasequoia, G sb tiu Macgini-
tiea na die s") nobilis, “Carya” antiquorum,

“Ampelopsis” "acea. and ains. of the
Cercidiphyllum complex, among others. Such
floras have been reported from Alaska (Wolfe,
1972), many areas of the northern conterminous
U.S. (Brown, 1962), the Canadian high Arctic
(Hickey et al., 1983), and Greenland (Koch,
1963)

Because some of these Holarctic Paleocene taxa

been described as having “East Asian" affinities
(although they are only East Asian in a modern
context). During the Eocene many of these Pa-
leocene (“East Asian") forms were eliminated
from floras in the eastern Rocky Mountains, pre-
sumably as a result of their inability to withstand
seasonal dryness (Leopold & MacGinitie, 1972).
In contrast, some of the same lineages survived
into the Neogene along the West Coast and in

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

the northern intermountain region of North
America. At the same time that the “East Asian”
forms were being eliminated in the eastern Rocky
Mountains, some of the new genera appearing
there were "Central American"; that is, their
closest living relatives are in the seasonally dry
subtropics of Mexico and Central America
(MacGinitie, 1969; Leopold & MacGinitie, 1972).
This general pattern has been confirmed by ad-
ditional work in the intermontane basins of the
eastern Rocky Mountains, but three modifica-
tions should be noted.

Paleocene megafloral samples may be more
uniform than actual Paleocene floras because the
fossil assemblages have been collected from a
limited and similar array of paleoenvironments
that tend to have low floral diversity. This would
accentuate the impression that a homogeneous
floral province was being broken up in the Eocene.

The regional extinction oftaxa with East Asian
affinities and the appearance oftaxa with Central
American affinities began in the Clarkforkian and
was well under way in some areas by the mid
Wasatchian, considerably earlier than was rec-
ognized by L Id & MacGinitie (1972), who
lacked floras from the Paleocene-Eocene tran-
sition period. Clarkforkian taxa with closest liv-
ing relatives in Central America include Chae-
toptelea microphylla, Woodwardia gravida, and
species of Populus sect. Abaso. By late Graybul-
lian or Lysitean time, some taxa with closest liv-
ing relatives in East Asia, such as Metasequoia
and some members of the Cercidiphyllum com-
plex, were already regionally extinct in basinal
floras of the eastern Rocky Mountains.

The shift from East Asian to Central American
affinities in the flora of the eastern Rocky Moun-
tains was not a uniform process, and some sig-
nificant taxa do not follow the pattern. For in-
stance, the genus P/atycarya, now confined to
East Asia, did not appear in western North
America until the early Wasatchian and achieved
maximum abundance throughout the region in
the latest Wasatchian and early Bridgerian before
going regionally extinct (Leopold & MacGinitie,
1972; Wing & Hickey, 1984). This pattern also
appears to hold true for several undescribed
species in the Icacinaceae, Flacourtiaceae, and
Menispermaceae. Ailanthus, also now endemic
to East Asia, appeared in North America during
the early to mid Eocene (Chalk Bluffs flora, Green
River flora, Rate Homestead flora, MacGinitie,
1941, 1969), and was abundant in Oligocene flo-
ras from southwestern Montana (Becker, 1961,

1987]

1969) before going regionally extinct in the later
Tertiary.

Floristic segregation of northern and southern
areas was probably occurring at the same time
as the better-documented and better-discussed
east-west divergence (Axelrod & Raven, 1985).
Unfortunately, few Wasatchian-Bridgerian flo-
ras have been reported from the southern Rocky
Mountains (Fig. 4). One small Lysitean flora from
the San Juan Basin (Tidwell et al., 1981) shows
that two taxa common in the Gardnerbuttean
and Bridgerian of Wyoming (““Sapindus” den-
tonii and Eugenia americana) appeared two to
three million years earlier in New Mexico. This
suggests the possibility that some taxa were mi-
grating northward during the climatic warming
of the early Eocene. A similar geographic pattern
has been observed in some mammalian species
(Beard, pers. comm., 1986).

DIVERSIFICATION OF MICROTHERMAL LINEAGES

The Wasatchian and Bridgerian floras from the
eastern Rocky Mountain region are a mixture of
taxa now associated with temperate and sub-
tropical to paratropical climates. For instance,
mesothermal to microthermal groups like the
Betulaceae (A/nus, Paleocarpinus), Cercidiphyl-
laceae, and Hamamelidaceae amamelidoi-
deae) are frequently associated with members of
megathermal groups such as the Icacinaceae (Pa-
leophytocrene), Lauraceae (Phoebe), Palmae, and
Cyatheaceae (Cnemidaria). In most of these flo-
ras the ipsae: elements are not diverse,
y be important in terms of

7).

younger age from the volcanic areas farther west
may be strongly dominated in both abundance
and diversity by microthermal taxa. This has
been attributed to the relatively high paleoele-
vation of these floras (Axelrod, 1966b, 1968;
Wolfe & Wehr, 1987). Currently the best known
of these floras is from Republic, Washington (Fig.
4; Wolfe & Wehr, 1987, and unpubl.). The vege-
tation at Republic is inferred to have been a Mixed
Coniferous forest, but the diversity of microther-
al angiosperm groups is striking: Hamamel-
idoideae, 4 species; Fagaceae, 4 species; Betu-
laceae, 5 species; Rosaceae, 19 species; Aceraceae,
7 species (Wolfe & Wehr, 1987; pers. comm.,
1987). Although the Copper Basin flora of north-
ern Nevada (Axelrod, 1966a) is younger and less
diverse, it shows a similar domination by mi-

WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

757

crothermal taxa. Alnus is represented by 2 species,
one of which accounts for 85% of the specimens,
and there are 3 species of Mahonia, 6 of Rosa-
ceae, 3 of Acer, and 3 of Ericaceae.

Although upland floras of Paleocene or greater
age are not known, it seems possible that the first
major diversification of many present-day mi-
crothermal lineages took place during the Eocene
in upland areas like the volcanic highland of the
northwestern United States (Wolfe, 1986, this
volume). Some of these groups were represented
by relatively generalized species in later Paleo-
cene or early Eocene lowland floras. Thus the
adjustment of basically megathermal or meso-
thermal lines to cooler climates initially may have

pers. comm.,
matic cooling, these upland lineages spread and
diversified over much of the Northern Hemi-
sphere, whereas their megathermal or meso-
thermal sister taxa now have relictual southerly
distributions.

One particularly good example of this pattern
is seen in the genus Populus. Species of the prim-
itive section Abaso are common in Clarkforkian,
Wasatchian, and Bridgerian floras over much of
North America (Eckenwalder, 1977; Wing, 1981),
usually in lowland settings, although Populus
adamantea does occur in probable Oligocene up-
land floras described by Becker (1960, 1972,
1973). The extant species of this section, P. mex-
icana, is distributed only in limited parts of
northeastern and northwestern Mexico. The ear-
liest record of the more advanced sections that
account for most of the present-day diversity and
distribution of the genus is in the late Eocene
Bull Run flora (Axelrod, 1966a). A number of
species belonging to the more advanced sections
are common in later Tertiary floras, but there is

no known fossil record of sect. Abaso poplars
ane the early Oligocene.

PATTERNS OF VEGETATIONAL CHANGE

The general pattern of vegetational change
during the Eocene and Oligocene in western North
America has been outlined in a number of pub-
lications (e.g., Axelrod, 1958, 1968; Axelrod &
Raven, 1985; MacGinitie, 1941, 1969; Wolfe,
1971, 1975, 1985). Despite considerable dis-
agreement on details, there is overall agreement
about the large-scale trends. Broad-leaved ev-
ergreen forests were dominant over most of the
area during most of the Eocene, with two main

758

exceptions. Some areas in the eastern Rocky
Mountains may have been dry enough to create
more open, partially deciduous vegetation. Vol-
canic activity in the western Rockies during the
late Wasatchian, Bridgerian, and Uintan gener-
ated uplands where conifers and broad-leaved
deciduous taxa became important components
of the vegetation. Climatic cooling and drying
during the late Eocene and Oligocene brought
about increasing dominance of mixed coniferous
and broad-leaved deciduous forest.

Late Paleocene vegetation in most areas of
western North America was broad-leaved ever-
green forest with an admixture of deciduous ele-
ments (e.g., Golden Valley flora of Hickey, 1977).
During the latest Paleocene and early Eocene
(Wasatchian) world climates were warming (Sa-
vin, 1977; Wolfe, 1971, 1978; Wolfe & Poore,
1982), possibly as a result of elevated levels of
atmospheric CO, and associated effects on cir-
culation patterns (Owen & Rea, 1985; Rea et al.,
1985). At this time evergreen broad-leaved taxa
increased in importance within local floras.

Floras from the Willwood Formation of north-
western Wyoming span most of the Wasatchian
and show an upward increase in the number of
entire-margined (41% to 52%) and thick-tex-
tured leaves, although there are also changes in
sedimentary environment that may be causally
related to this shift (Wing, 1981). Using data on
living vegetation presented by Wolfe (1979), these
leaf-margin percentages correspond with Micro-
phyllous or Notophyllous Broad-leaved Ever-
green Forest. Wolfe (1985) interpreted slightly
older (Clarkforkian) floras from the

of streamside taxa in Bighorn Basin floras from

Fort Union and Willwood formations, or a
relatively higher elevation, has given them a
“cooler” aspect. I think it more likely that the
less entire-margined, presumably more decidu-
ous, floras of the Bighorn Basin were under the
influence of a seasonally dry climate as early as
the middle Clarkforkian. This is also suggested
by the development of oxidized soil horizons in
floodplain sediments of this age in the Bighorn
Basin (Gingerich et al., 1980). A resolution of
this apparent conflict would involve detailed
comparison of depositional settings of the floras
or independent evidence for paleoelevation. Re-
gardless of these somewhat different interpreta-
tons of vegetation in northern Wyoming, it is
apparent that some form of broad-leaved ever-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

green forest extended as far north as 65? during
the Wasatchian (Wolfe, 1985).

A great many Lostcabinian-aged (late Wa-
satchian) fossil localities are known from all over
the state of Wyoming and from the Golden Val-
ley Formation of western North Dakota (e.g.,
numbers 1, 2, 9, 19, 27-30, 32-33, 50-71; Fig.
3, Appendix I). These floras are mostly uniform
in composition and dominated by one of two
species of Platycarya, with important subdom-
inants being Alnus, *Dombeya" novi-mundi,
“Dalbergia,” Zingiberopsis isonervosa, an
species of Icacinaceae, Lauraceae, Magnoliales,
and Palmae. Common pteridophytes are Cne-
midaria magna, Lygodium kaulfussii, Thelyp-
teris weedii, T. iddingsii, and a large-statured
species of Equisetum. The similarity in these flo-
ras may in part reflect similar environments of
deposition occurring ina à number of intermon-
tane basins at app
en, however, that some range of depositional en-
vironments is spanned by these floras, it is likely
that the successional vegetation most likely to be
preserved in fluvial sediments was truly similar
over this large region. This in turn implies the
existence of few sharp climatic differences across
the area, which is in distinct contrast with the
early Bridgerian floras discussed in the next sec-
tion.

the Sa Tit time. Giv-

Vegetation during early Bridgerian time (about
50 Ma). Although the latest Wasatchian and
Bridgerian were the times of maximum poleward
extent of broad-leaved evergreen forests, vege-
tation of the Rocky Mountain region began to
differentiate more strongly during this time in-
terval. This differentiation is illustrated by com-
paring four floras: the Little Mountain flora from
the upper Wilkins Peak Member of the Green
River Formation in southern Wyoming (23 in
Figs. 3 & 4, Appendix I), the Boysen flora from
the upper part of the Wind River Formation in
the northcentral Wind River Basin (36), the Kis-
inger Lakes-Tipperary flora from the Aycross
Formation in the western Wind River Basin (38,
42), and the flora of the lower Sepulcher and
Lamar River formations in Yellowstone Nation-
al Park in northwestern Wyoming (112-118). All
four of these floras are approximately 50-51 Ma
and correlate with Gardnerbuttean (early Bridg-
erian) mammalian faunas

Based on floristic affinities, foliar physiogno-
my, and sedimentological data, MacGinitie
(1969) inferred that the Little Mountain flora was

1987]

derived from open woodland vegetation.
(MacGinitie used the term “savanna woodlan
although he pointed out this might vilsieadinsl
imply that grasses played an important role in
the vegetation.) More recently, Wolfe (1985)
stated that leaf size in the Green River floras is
too large for scrub or savanna vegetation and is
more consistent with semideciduous subtropical
to paratropical forest. The 22 species of dicoty-
ledonous leaves from the Little Mountain flora
are generally small and thick textured, and a
number of the species belong to families or gen-
era typical of seasonally dry subtropical vege-
tation (e.g., Alchornea, Cardiospermum, Populus
sect. Abaso, and a number of microphyllous Le-
guminosae; MacGinitie, 1969: 67-68). Recent
sedimentological work on the Wilkins Peak
Member suggests that deposition took place in
and around the margins of a playa lake that lay
in an orographic desert basin (Smoot, 1983). The
upper part of the Wilkins Peak Member appar-
ently represents the maximum transgression of
the lake and hence the wettest period during the
deposition of this part of the Green River For-
mation (Smoot, 1983).

The Boysen flora is largely undescribed, al-
though it was referred to in a treatment of the
Green River flora (MacGinitie, 1969). This flora
occurs in fluvially deposited, irregularly fissile,
tuffaceous mudstones at the southern edge of the
Owl Creek Mountains. The assemblage has ap-
proximately 15 species and is heavily dominated

y palm leaves, Lygodium kaulfussii, and an en-
tire-margined dicot leaf resembling Sapindus spp.
Other common elements include "Populus"
wyomingiana, Canavalia diuturna, cf. Typha,
Zingiberopsis isonervosa, and Musophyllum
complicatum. The importance of palms, herba-
ceous monocots, and vines suggests low-stature,
perhaps relatively open, floodplain vegetation.
Somewhat similar vegetation may have been re-
sponsible for forming the Lostcabinian-aged
Vermillion pa coal in southern Wyoming
(Nichols, in pre

The Kissinger pm Tipperary flora consists
of 5 ferns, 1 horsetail, 2 conifers (G/yptostrobus
and Chamaecyparis), and 44 angiosperms
(MacGinitie, 1974). Of the 36 well-defined di-
cotyledonous leaf types, 5596 have nonentire
margins, and judging by their closest living rel-
atives, some 60% were deciduous. A number of
the genera in the fossil assemblage are presently
restricted to subtropical or tropical climates (e.g.,
Acrostichum, Apeiba, Canavalia, Dendropanax,

WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

759

ini octal Based on foliar physiognomy and
e distribution of living relatives, MacGinitie

MA inferred that the Kisinger Lakes flora was
derived from a subtropical to tropical, semide-
ciduous forest resembling those native to the
southwest coast of Mexico at elevations of about
1,000 m. These inferences are consistent with
the diverse palynoflora, which in most samples
iS ET by angiosperms and ferns (Leo-
pold, 1

The FU floras of Yellowstone National Park
were first described by Knowlton (1899) and have
not been subjected to a general revision since.
Therefore, most of the published identifications
are probably incorrect. Furthermore, much of
the megafossil material comes from sites that are
imprecisely located and of unknown stratigraph-
ic relationship. It may be that significant tem-
has
remained undetected as a result. In spite of these
problems, most of the Yellowstone assemblages
were derived from vegetation that was quite dif-
ferent from the kind inferred for the assemblages
discussed above

our genera of conifers are known from mega-

fossil remains (Sequoia, Glyptostrobus, Pinus, and
?Podocarpus), and the first three are abundant at
many localities (Dorf, 1960; Aguirre, 1977). As
might be expected, the importance of conifers is
even more strongly indicated by the palynoflora,
where they are diverse (12 genera) and consis-
tently make up about half ofthe noie (Fisk, 1976).
Ferns are also highly diverse and abundant at
many localities in the Sepulcher Formation. Fisk
from the Yellowstone Park palynoflora, and there
are 10-15 named species from the macroflora.
The local abundance of ferns at some localities
(particularly Thelypteris weedii and Allantoidiop-
sis erosa) may be related to the frequency of dis-
turbance by volcanic events. A similar domi-
nance of ferns in the colonizing vegetation has
been noted following eruption and deposition in
the vicinity of EI Chichón volcano in Mexico
(Spicer et al., 5

The foliar physiognomy of the Yellowstone
National Park floras has not been studied in de-
tail, but dicot leaves in the collectionsat the U.S.
National Museum of Natural History are mostly
in the notophyll and mesophyll size categories.
Drip tips are present on a few taxa, and roughly
half of the species have entire-margined leaves.
Although the percentage of entire-margined
species is similar at Yellowstone (approximately

No
oo

760

50%) and Kisinger Lakes (54%), leaf size in the
Kisinger Lakes flora is generally in the micro-
phyll to notophyll range (Wing, unpubl. data;
Wolfe, pers. comm., 1987). These PER aie
aspects of the Yellowstone assemblages, in ad-
dition to the importance of conifers, d that
the vegetation grew under a somewhat cooler and
perhaps less seasonally dry climate. The forest
from which the Yellowstone flora was derived
appears to have been a variety of broad-leaved
evergreen forest that included a substantial ele-
ment of conifers in some local environments.

The question of taphonomic mixing of floral
assemblages is ever present but has been a par-
ticular focus of debate with regard to fossil floras
from Yellowstone National Park (Fisk, 1976;
Fritz & Fisk, 1978; Fritz, 1980a, 1980b, 198 La,
1981b; Retallack, 1981; Yuretich, 1984; Karowe
& Jefferson, 1987). The mixture of "tropical"
and "temperate" elements (e.g., Thuja and pre-
sumed evergreen members of the Lauraceae) in
the Yellowstone floras has been attributed to
transport of plant remains derived from vege-
tation growing at a range ofelevations (Fisk, 1976;
Fritz, 1980a, 1986), and the upright stumps and
autochthonous “fossil forests" described by
Knowlton (1899) and Dorf (1964) have been ex-
plained in part as the consequence of transport
by mudflows associated with volcanic activity
(Coffin, 1976; Fritz, 1980a, 1980b, 1986; Fritz
& Harrison, 1985).

Observations of present-day volcanic systems
have shown that high-energy mudflows can
transport upright stumps from higher to lower
elevations, and that stumps weighted by soil
trapped in their roots may float upright for a time
in lacustrine situations (Fritz, 1980a, 1980b,
1986; Coffin, 1983). Furthermore, sedimento-
logical and stratigraphic studies of the Sepulcher
and Lamar River formations and other Eocene
volcanic units have highlighted the importance
of high-energy deposits that are presumably in-
dicators of steep paleotopography (Fritz, 1980c;
Fritz & Harrison, 1985). In spite of these im-
portant observations, several lines of evidence
suggest that the Yellowstone megafossil assem-
blages are not highly allochthonous

First, although many fossil trees in the Spec-
imen Ridge section may be prone rather than
upright, this is not always the case. The presence
of paleosols around some of the upright stumps
is evidence that these were fossilized in place
rather than transported upright to the site of buri-
al and preservation (Retallack, 1981). Yuretich

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

(1984) and Karowe & Jefferson (1987) have pre-
sented petrographic and sedimentological evi-
dence that the upright stumps are generally in
situ. 1) Tree stumps are rooted in fine-grained
sediments, not in conglomerates. 2) Some con-
glomerates have structures showing that they
flowed around trunks in place. 3) Upper parts of
inclined axes are abraded, but parts contained in
finer-grained rock are not. 4) Petrographic sec-
tions of sediment containing fossil tree roots show
no signs of extensive current bedding but do show
indications of pedogenesis. Many of the leaf
EGER assemblages, which also show mixed

"tempera and "tropical" forms, are derived
from ue fluvial sediments and thus are
unlikely to be highly transported (Wing, unpubl.
data). Some fine- to medium-grained, airfall tuffs
have the potential to entomb plant assemblages
that quite accurately reflect local vegetation
(Burnham & Spicer, 1986).

Second, the presence in the same sedimentary
units of trees showing distinct seasonal growth
with trees lacking distinct growth rings (Wheeler
et al., 1977, 1978) has been cited as evidence
that plants that grew under more than one cli-
matic regime are present (Fritz & Fisk, 1978).
This interpretation is not justified. Tree species
have varying genetic capacities for seasonal
growth, and individual trees are variably influ-
enced by microclimatic and edaphic factors. The
result is that different trees in the same region
may show different patterns of growth (e.g.,
Tomlinson & Craighead, 1972). Fossil log as-
semblages containing specimens with both sea-
sonal and aseasonal growth have been observed
in sedimentary environments that could not have
produced long-distance transport (Bown et al.,
1982).

Third, most of the extreme examples of dis-
sonance in the climatic tolerances of elements in
the flora are generated by comparing the paly-
noflora with the megaflora. Pollen of Abies and
Larix were less than 196 of the assemblages in
which they occurred, which in turn were only a
few of the 20 samples taken by Fisk (1976). Ob-
viously, these could be highly allochthonous pol-
len grains derived from vegetation growing at
higher elevations than the site of deposition. Pic-
ea occurred as 1—3 grains in 12 of the 20 samples,
and reached 296—696 of the flora in three samples
(Fisk, 1976). Cross & Taggart (1982) have noted
that abundance of Picea pollen is generally in
rough proportion to its importance in the source
vegetation, implying that spruce could have been

1987]

a minor part of local vegetation, probably
higher elevations in the watershed containing the
site of deposition.

Fourth, almost all early Eocene floras from
western North America contain some taxa that
presently have mutually exclusive climatic re-
quirements (MacGinitie, 19 74; Wolfe,
1980). In many floras the depositional setting
argues strongly against explaining the presence
of these dissonant elements by transport. For
instance, backswamp compression fossil assem-
blages from the Willwood Formation, perhaps
one million years older than the Yellowstone
flora, have abundant palms, Cnemidaria (cy-
atheaceous tree fern), and A/nus (Wing, 1981).
The Willwood Formation is a mostly fine-grained,
basin-fill formation deposited by low-energy
streams (Bown, 1979); low paleotopography is
further indicated by thin, laterally extensive sheet
sands (Kraus, 1980) and paleosol horizons that
are traceable over many manis n &
Kraus, 1981). Many of the collecting sites for the
Willwood flora were near the center gn e basin,
at least 100 km from the uplifted basin margins.
The assemblages were derived from fine-grained
mudstones (Wing, 1984), and similar assem-
blages are characteristic of this lithology over a
large part of Wyoming (Wing, unpubl. data).
Given this depositional environment, the poten-
tial for long-distance transport of leaves is min-
iscule, and the assemblages must be autochtho-
nous or transported only a short distance.
Therefore there has been change through time in
the climatic requirements of the taxa involved
and/or early Tertiary climates permitted the co-
existence of genera that at present have largely
nonoverlapping ranges

In conclusion, recent stratigraphic and sedi-
mentological studies of volcanic strata in Yel-
lowstone National Park and other Paleogene vol-
canic sequences have demonstrated the
importance of high-energy mudflow deposition
and have made it clear that such deposits are
characterized by a high degree of lateral vari-
ability. However, these studies have failed to
demonstrate that the majority of the megafossil
assemblages from such deposits are allochtho-
nous. Currently, prevailing evidence suggests that
plant megafossils, especially compression assem-
blages of leaves, generally accumulated from lo-
cal sources during quiet periods between violent,
localized, mudflow deposition.

Although the Republic flora (168-170; Figs. 3,
4) is approximately 2 Ma younger than the four

WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

761

floras discussed above, it is similar to the Yel-
lowstone flora in having abundant conifers in
both the megaflora and palynoflora. Wolfe &
Wehr (1987) argued that the Republic paleoveg-
etation was a mixed coniferous forest dominated
by conifers because: 1) Chamaecyparis and Pinus
are abundant at the Graphite Creek locality, 2)
bisaccate pollen dominates the palynoflora, and
3) conifers are quite diverse (including Thuja,
Abies, Picea, Tsuga, and Pseudolarix in the
megaflora). Deciduous angiosperms are the most
abundant and diverse group at the other two lo-
calities, but broad-leaved evergreens are present,
including Phoebe, Photinia, Ternstroemites spp.,
and a new extinct genus. Based on the physiog-
nomy of the assemblage, Wolfe & Wehr (1987)
inferred a mean annual temperature of 12—13°C
and a mean annual range of temperature of 5—
6°C. By comparing these temperature estimates
with those inferred for approximately coeval flo-
ras from the Puget Group, Wolfe & Wehr (1987)
estimated the paleoelevation of the Republic flora
as 725-910 m. The greater importance of coni-
fers and deciduous broad-leaved plants at Re-
public than in the Yellowstone flora probably
does not reflect a higher paleoelevation if esti-
mates for the two areas are even approximately
correct. Therefore, the compositional and phys-
iognomic differences between the two floras
probably result from differing taphonomic pro-
cesses, climatic cooling during Bridgerian time,
or both.

Vegetation during the Chadronian (38-33
Ma). Comparison of the approximately con-
temporaneous Little Mountain, Boysen, Kisin-
ger Lakes, and Yellowstone floras illustrates the
kinds of vegetational and floristic d t
existed over one part of the northern Rockies at
about 50 Ma. The regional importance of conif-
erous and broad-leaved deciduous elements
seems to have increased during the later Eocene
and Oligocene, with dominantly coniferous for-
ests probably becoming established at higher el-
evations. However, later Eocene and Oligocene
floras provide less evidence of the kinds of vege-
tational boundaries implied by the contrasts be-
tween the four floras discussed above. This may
reflect in part a less adequate sample of coeval
floras.

One time interval in the remainder of the
Eocene and Oligocene contains enough fossil flo-
ras to permit an attempt at analyzing vegeta-
tional variability in the Rocky Mountain region.

762

Ten floras in Appendix I are from rocks of prob-
able Chadronian age: Missoula, from westcentral
Montana (245-246; Jennings, 1920); Christen-
sen Ranch, Horse Prairie, and Medicine Lodge
from southwestern Montana (151-152, 153-157,
159-160; the Beaverhead Basins floras of Becker,
1969); uppermost Bull Run from northeastern
Nevada (202; Axelrod, 1966a, 1966b); Florissant
from central Colorado (224—229; MacGinitie,
1953); Red Rock Ranch, Hermosa, and Hills-
boro from the Rio Grande Rift of New Mexico
(232, 233, 234; Axelrod & Bailey, 1976; Meyer,
1986); and a small flora from the White River
Group in the Flagstaff Rim area of central Wy-
oming (2
The Missoula flora was derived from a se-
uence of coal and lacustrine ash beds that Jen-
nings (1920) believed were correlative with the
lower part of the White River Formation in South
Dakota. The flora has not been studied since
1920, and many of Jennings's identifications are
questionable. However the flora appears to be
dominated by Metasequoia, Sequoia, and one or
more species of Betulaceae. Equisetum, ?Thuites,
?Populus sect. Abaso, Acer, and ?Cercidiphyllum
are also present. It is difficult to reach any con-
clusion about vegetation based on such a small
assemblage of relatively ubiquitous taxa.
Becker (1969) considered the Beaverhead Ba-
sins floras to be of latest Oligocene to Miocene
age, but this was based on the thickness of the
local stratigraphic sequence and on floral corre-
lation; the latter was in turn based on an incom-
plete understanding of the taxa involved. More
recent geological and paleontological work sug-
gests that the sections are thinner than originally
thought, and that the “Medicine Lodge Beds,”
from which the floras were collected, may cor-
relate with sediments producing Chadronian
vertebrates (Fields et al., 1985). All three of the
Beaverhead Basins floras represent mixed conif-
erous and deciduous broad-leaved forest, which
Becker (1969) felt reflected a subhumid climate.
The subhumid elements include such taxa as
Mahonia, Juniperus, and various nanophyllous
Leguminosae. Common broad-leaved deciduous
taxa are Fagopsis longifolia, Cercidiphyllum,
Populus, Sassafras, and various Betulaceae and
Ulmoideae. Upsection changes in floral com-
position are relatively modest, although the up-
per two floras have 14 (Medicine Lodge) and 16
(Horse Prairie) species of conifers to the 10 species
found in the lowest flora. There is a pronounced
physiognomic change between the Christensen

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Ranch flora and the upper two in that the per-
centage of species with entire-margined leaves
increases from 18% to 34% and 37%. The sig-
nificance of this shift is cast into some doubt by
concomitant changes in depositional environ-
ment and floral diversity, and by the lower re-
liability of leaf margin data from coniferous
vegetation (Wolfe, 1979), but taken at face value
the increase in percentage of species with entire-
margined leaves would indicate an increase in
mean annual temperature from 7? to 12°C.

Axelrod (1966a, 1966b) stated that the upper
Bull Run floras represented montane forest al-
most totally dominated by conifers, including
Abies, Picea, Pinus, Pseudotsuga, Tsuga, Cham-
aecyparis, and Thuja. The angiosperm compo-
nents are small-leaved and, with the exception
of Mahonia, deciduous genera such as Alnus,
Betula, Ribes, and Zelkova, This low diversity,

igly conifer

markedio “with the Beaverhead Basins floras, and
although the upper Bull Run floras may be slight-
ly older than those of the Beaverhead Basins (the
highest locality is five feet below a tuff dated at
36 Ma; Axelrod, 1966a; Table 1), it seems prob-
able that a difference in paleoelevation is also
involved.

The small flora from the Chadronian of the
Flagstaff Rim area was ted from clastic dike
fillings that also preserve vertebrate skeletons
(Emry, pers. comm., 1986). This peculiar mode
of deposition makes comparisons between this
flora and others questionable, but the low di-
versity (about six forms) and the small leaf size
(microphylls or nanophylls) probably indicate
relatively dry conditions. The identifiable taxa
are Mahonia, ?Ribes, ?UImaceae, Leguminosae,
and an undetermined conifer. This small flora
may represent interfluve vegetation better than
typical collections obtained from rocks deposited
in paludal or lacustrine settings.

The Florissant flora is the most diverse from
the Oligocene of the Rocky Mountain area. Based
on a combination of floristic and physiognomic
criteria, MacGinitie (1953) inferred that the
Florissant fossil d two main
types of vegetation: a mesic, broad-leaved de-
ciduous forest along Ram and lakeshores and
on slopes
and interfluve areas. The ten most common
species in the flora comprise 6096 of the speci-
mens je Fagopsis longifolia, Zelkova dry-

eja, Chamaecyparis, Typha, Populus crassa,
Bhus ae Sequoia affinis, Cercocarpus

a drier Scrub

1987]

myricaefolius, Staphylea acuminata, and Ath-
yana haydenii. The broad-leaved evergreen com-
ponent of the Florissant is not dominant, but the
diversity and abundance of conifers is much less
than in either the Beaverhead Basins or the upper
Bull Run floras.

Chadronian floras from New Mexico (Axelrod
& Bailey, 1976; Meyer, 1986) appear to represent
two different types of vegetation. The Red Rock
Ranch flora is numerically dominated by spec-
imens of Pinus subsection Balfourianae and con-
tains species of Picea, Abies, Zelkova, and pos-
sibly Salix and Rosa (Meyer, 1986). Based on a
list by Farkas (1969), Axelrod & Bailey (1976)
reported several additional taxa including Fa-
gopsis, Halesia, Mahonia, Rhus, and Sapindus,
but, with the exception of Mahonia, these were
not confirmed by Meyer (1986). Although the
Red Rock Ranch flora is small, on the basis of
relatively high conifer diversity and abundance
and low broad-leaf diversity, Meyer (1986) con-
cluded that it most likely represents a mixed co-
niferous forest. This flora was correlated with
Florissant (35 Ma) by Axelrod & Bailey (1976),
but the more recent and direct radiometric date
obtained by Meyer (1986) indicates the flora is
no younger ool > : + 1.1 Ma, or about 2 Ma
older than Flori

The Hermosa nd fiios floras are derived
from sediments associated with the infilling of
the moat of the Emory caldera, and both were
dated at about 32 Ma by Axelrod & Bailey (1976).
New radiometric dates reported by Meyer (1986)
indicate the Hillsboro flora is probably 28.1—30.6
Ma (Whitneyan or early Arikareean) and that the
Hermosa flora is about 33.6 + 1.0 Ma (Chad-
ronian). The floras are similar in composition,
with an overwhelming dominance of specimens
of Pinus subsection Balfourianae, along with a
few small leaves of Mahonia, and possibly Picea
and Crataegus. These floristic and physiognomic
attributes indicate a taiga-type forest growing un-
der a cold temperate climate (Meyer, 1986). Ax-
elrod & Bailey (1976) argued that the difference
between the Red Rock Ranch and Hermosa/
Hillsboro floras was a response to higher pa-
leoelevation of the latter floras; however, Meyer
(1986) pointed out that these floras may bracket
the major decrease in mean annual temperature
and increase in mean annual range of tempera-
ture that occurred at about 33 Ma (Wolfe & Hop-
kins, 1967; Wolfe, 1986).

The pattern of geographic variation in Chad-
ronian vegetation is less obvious than that of the

WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

763

earliest Bridgerian, and this can be attributed to
several causes. First, the Chadronian floras are
less tightly correlated and probably occurred over
a longer interval of time, perhaps 5 Ma. Second,
several of the floras are of low diversity or are
not completely described, with the result that
they are poorer samples of regional vegetation.
Third, there may be a greater altitudinal range
represented by the floras. Fourth, the youngest
floras in the set (Hermosa/Hillsboro) may lie on
the opposite side of a major temperature de-
crease from the other floras. In spite of these
problems, the three floras of presumed inter-
mediate elevation (Beaverhead Basins, Floris-

Rock Ranch) seem to have broadly
similar compositions and to represent mixed co-
niferous and broad-leaved deciduous forest
growing under a seasonally dry climate. The
higher diversity of conifers and mesic taxa in the
Montana floras may indicate higher rainfall and/
or lower rates of evapotranspiration in the north-
ern part of the Rocky Mountains. The subhumid
aspect of all of these floras when compared with
those of similar age from the Pacific Northwest
demonstrates the continuation of the pattern of
regional climatic variation that began during the
early Eocene/latest Paleocene.

The mid Tertiary climatic deteriora-
tion. Much of the evidence for the major de-
crease in mean annual temperature and increase
in mean annual range of temperature that oc-
curred at approximately 33 Ma is derived from
lowland floras from the Pacific Coast of North
America (Wolfe & Hopkins, 1967; Wolfe, 1971,
1986). Isthere unambiguous evidence ofthis ma-
jor climatological change in floras from the Rocky
Mountain re

presented in Figure 3, there are six floras that
closely follow after the 34 Ma date: Mormon
Creek, Metzel Ranch, York Ranch, Ruby paper
shales, Hermosa, and possibly Hillsboro. The
dating of the Montana floras has been uncertain
since their initial descriptions (Becker, 1960,
1961, 1972, 1973), and opinions on the age of
the Mormon Creek flora have embraced some
20 Ma. Recent geological and mammalian bio-
stratigraphic correlations suggest the Mormon
Creek, Metzel Ranch, and York Ranch floras are
of Orellan age (32.2-30.8 Ma) and that the Ruby
paper shale flora is Whitneyan (30.8-29.2 Ma).
These floras were judged by Becker (1960, 1961,
1972, 1973) to represent mixed coniferous and

764

broad-leaved deciduous forest and shrubland
growing under temperate to dry-temperate cli-
mates. Although the Ruby paper shale flora is
inferred to have been somewhat dryer than the
other three, they all bear substantial resemblance
to one another and to the older (Chadronian)
Beaverhead Basins floras. As noted above, the
New Mexican floras are derived from a setting
of some paleotopographic and structural com-
plexity, so that it is difficult to separate potential
effects of elevation and regional climatic change.

Thus the only floras of suitable age do not
provide good evidence of a major temperature
decrease at 33 Ma. This absence of evidence may
result from the confounding effects of changing
elevations and dryer climates in the Rocky
Mountains, or it may simply be a problem of
insufficient data and poor stratigraphic control.

PRESENT METHODS AND FUTURE STUDY

Perhaps the most useful result of summarizing
current knowledge and opinions in a field of study
is that this activity reveals gaps in the data base
and reveals possible directions for future re-
search. At present, research in Tertiary paleo-
botany follows two main themes: the systematic/
evolutionary approach is concerned primarily
with describing new fossil forms and understand-
ing their implications for the evolutionary his-
tory of lineages and relationships among living
groups of plants; the paleoecological/environ-
mental approach focuses largely on understand-
ing habits of extinct species, structure of extinct
vegetation, and patterns of ancient climates. His-
torically these approaches often were combined
in the treatment of a single fossil flora. More
recently, as standards have become higher and
techniques more sophisticated in both ap-
proaches, workers have specialized on one or the
other. This probably reflects a more general sep-
aration of ecology from systematics, but the dis-
junction in viewpoints should not be accepted as

logical data provide the context for understand-
ing the genealogical changes that are inferred from
systematic studies. Furthermore, in paleobotany
both approaches are united at a practical level
by the study of the same sites and specimens.

FIELD DATA

As noted in the section on the data base for
Eocene and Oligocene floras, the stratigraphic

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

and geographic distribution of sites is very un-
even. The total number of sites from which col-
lections have been made is also small, consid-
ering the commonness of plant fossils and the
large outcrop area. The meagerness of the data
set results from there being few paleobotanists,
rom a tendency for the same sites to be collecte
repeatedly, and from relatively small efforts to-
ward finding new sites. The largest “holes” in the
record would be filled by: 1) more late Eocene
and Oligocene floras from the eastern Rocky
Mountain area; 2) more Paleogene floras from
the southern Rocky Mountains; and 3) more flu-
vially deposited late Eocene or Oligocene assem-
blages (or conversely, more Paleocene and early
Eocene lacustrine assemblages).

Field data that would Bien reports of fossil
plants include: number of quarry sites collected;
size of quarry; precise locality information;
stratigraphic unit; detailed lithological descrip-
tion; abundance of fossils and relative abundance
of species; and, where available, biostratigraphic
correlation, radiometric age, and paleomagnetic
correlation. If such data were available, even as
preliminary approximations, the published rec-
ord of Tertiary fossils would be more useful for
interpreting paleovege no paleoclimate, and
possible associations of dispersed organs

SYSTEMATICS

During the last 15 to 20 years several new
methods have brought increased resolution and
rigor to systematic studies of Tertiary angio-
sperms. Comparative studies of the leaf archi-
tecture of living dicotyledons have created a much
firmer basis for interpreting the systematic affin-
ities of fossil leaves (Hickey & Wolfe, 1975). The

structure of fossil pollen (e.g., Crepet et al., 1980),
and from more detailed analyses of fossil flowers
(e.g., Crepet & Daghlian, 1981, 1983). At the
same time, more studies have come to base their
systematic conclusions on multiple fossil organs
belonging to the same species (e.g., Manchester
& Crane, 1983; Wing & Hickey, 1984; Man-
chester, 1986). Finally, in many areas of paleo-
botany, refinements in methods of systematic

the reasoning behind systematic decisions more
explicit (e.g., Stein et al., 4
Although major advances have been made in

1987]

methodology, the vast majority of fossil angio-
sperms from western North America are as yet
very poorly understood. For most times and
places the floras either have not been described
or the only descriptions are those of late-19th
century workers whose goals were more bio-
stratigraphic than systematic. New methods will
have to be applied repeatedly before the botan-
ical relationships of any significant number of
Te estiary fossils will be MASSE ON

the quantification of variability. Comsemtwe y
architecture was an important advance in ana-
lyzing higher-level systematic affinities but has
been much less useful at the level of species. With
few exceptions (e.g., Dolph, 1975; Burnham,
1986a, 1986b; Wing & Eckenwalder, 1987), pa-
leobotanists have been little concerned with
quantifying the variability of their taxa. Yet in-
dividuating taxa in a paleobotanical sample is
the initial step in subsequent systematic, bio-
stratigraphic, and paleoecological syntheses. De-
tection and quantification of low-level morpho-
logical variability is also a key to uncovering
patterns of temporal change in closely spaced
stratigraphic samples.

PALEOECOLOGY

Traditional paleoecological interpretation of
fossil angiosperm floras has been based on flo-
ristic analogy and on leaf physiognomic analysis.
The floristic method assumes that the ecological/
climatic requirements of the fossil taxa were sim-
ilar to those of their closest living relatives. This
kind of direct analogy suffers from several defects
(see Wolfe, 1979). First, it assumes that the bo-
tanical relationships of the fossils have been de-
termined correctly. Second, it assumes that little
evolutionary change has occurred in the climatic
or ecological preferences of the lineages under
study. Justification for both of these assumptions
diminishes as one considers older floras, because
evolution is more likely to have occurred in the
intervening time, making it more difficult to de-
termine close living relatives of older fossil plants.
A third problem with floristic inference is that it
implies that the present-day distribution ofa tax-
on accurately reflects even its present climatic
tolerances. Given the rapid climatic fluctuations
typical of the last two million years, it may be
that the current distributions of many taxa are
strongly influenced by migration rate, plant com-
petitors, or other nonclimatic factors (Davis,
1976

The second commonly used method of pa-

WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

765

leovegetational/paleoclimatic inference is anal-
ysis of leaf physiognomy. Physiognomic analysis
primarily relies on a relationship observed in
living floras; the percent of species in a local flora
that have entire-margined leaves rises with the
mean annual temperature of the site (Bailey &
Sinnott, 1915, 1916). Thus tropical floras have
nearly 10096 species with entire leaves, whereas
temperate floras are dominated by species with
toothed or lobed leaves. This relationship has
been worked out with some precision based on
the humid floras of East Asia (Wolfe, 1979).
Studies based on smaller regions have been used
to question the resolution of leaf-margin infer-
ences (Dolph, 1976, 1978a, 1978b, 1979; Dolph
& Dilcher, 1979), but the basic pattern of climate
change as inferred from Tertiary floras agrees
with data from a variety of other sources (e.g.,
Savin, 1977; Wolfe & Poore, 1982; Hutchison,
1982; Owen & Rea, 1985; Rea et al., 1985). Phys-
iognomic analysis also considers the average size
of leaves in a fossil assemblage, their apparent

number of |
number of species that are probable lianes (those
with cordate-based leaves). Generally these at-
tributes increase with increasing tropicality of
vegetation.

Although physiognomic analysis offers signif-
icant improvement on the floristic method, it has
defects. In addition to their correlation with mean
annual temperature, physiognomic characteris-
tics of leaves are also correlated with water avail-
ability, intensity and angle of incident radiation,
and a variety of other factors. Consequently,
changes in the leaf physiognomy of fossil assem-
blages cannot always be read unambiguously as
changes in mean annual temperature. An in-

daphically Induced water
stress could produce Xn with small, thick,
entire-margined leaves and few lianes. Greater
representation of canopy species in a fossil flora
would produce an assemblage with smaller leaves
(Roth & Dilcher, 1978). This is because canopy
leaves tend to be smaller than interior leaves in
order to radiate heat more efficiently and main-
tain an optimal photosynthetic rate. Perhaps the
most serious factor biasing leaf physiognomic
analysis of fossil assemblages is the probable
overrepresentation of early successional and
stream ges up which grow close to sites of
dep in fluvial and volcanic settings
Maec 1969). Successional and riparian
vegetation in most climatic zones is dominated
by species with lobed, toothed, or compound

766

leaves, probably because these species hold in-
dividual leaves for only a short time, and these
leaf shapes provide a large photosynthetic sur-
face at a small cost of support tissue (Givnish,
1978). Thus a change in the frequency with which
fossil vegetation was disturbed might produce a
change in leaf physiognomy that might be inter-
preted as a change in mean annual temperature.

In spite of their defects, both the floristic and
leaf physiognomic methods produce inferences
about paleovegetation that are generally consis-
tent with paleoclimatic reconstructions based on
other, independent data sets. Furthermore, they
generally agree with one another (e.g., Mac-
Ginitie, 1974; Hickey, 1977). The problem with
these methods is not that they produce grossly
incorrect interpretations of past vegetational
structure, but rather that the inferences lack de-
tail, frustrating the most interesting comparisons
that might be made between extinct and living
forests.

For instance, because of strong seasonality of
light and a low angle of incident radiation, it is
likely that the structure of high latitude, broad-
leaved, evergreen forests in the early Tertiary was
significantly different from that of living broad-
leaved evergreen forests, even though the two
types of vegetation are similar in leaf physiog-
nomy and floristic composition. This hypothesis
can only be examined by finding more ways to
compare fossil and living vegetation. These new
methods of comparison will probably require
collecting data on the distribution of fossils in
the sediment. These distributional data (e.g., al-
pha and beta diversity, relative abundance, spa-
tial heterogeneity) may reflect actual synecolog-
ical characteristics of the vegetation that produced
a fossil assemblage; the difficulty in interpreta-
tion arises from the probability that taphonomic
processes have also influenced the distribution
of fossils. In spite of recent work on the taphon-
omy of fossil plants (Spicer, 1981; Scheihing &
Pfefferkorn, 1984; Spicer & Greer, 1986; Fer-
guson, 1985; Gastaldo, 1986; Burnham & Spicer,
1986; Spicer & Wolfe, 1987), there are as yet no
general recommendations for how leaf assem-
blages can be sampled to reflect best given char-
acteristics of the former vegetation. This kind of
work will have to be done in order to make the
best use of the paleoecological information pre-
served in fossil plant assemblages.

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WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 771

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1987] WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS 777

APPENDIX II.

Additional references on Eocene—Oligocene floras of
h America.

western Nort
Ammons, R. B., S. AM S, G. AMMONS & R. AM
MONS 1980. Cross- arr ni ree of ring signa-
Eocene trees

of Yellowstone, preliminary methodology and evi-
dence. Proc. Montana Acad. Sci. 40: 47.

ANDERSON, R. Y. 1960. Cretaceous-Tertiary paly-
nology, eastern side of the San Juan Basin, New
Mexico. Mem. New Mexico Bur. Mines Min. Re-

ANDREWS, H. N. 1936. A new Sequoioxylon from
Florissant, Colorado. Ann. Missouri Bot. Gard.
23: 439-446.

Notes on the fossil flora of Yellowstone

National Park with particular — to the Gal-
elc latin region. Amer. Midl. Naturalist 21: 454—460.

-E-E-R- E E .LENz. 1946. The Gallatin: nh arent

Hy Hee Hy by 8 (Yellowstone National Park, ng).

99979022 Missouri Bot. Gard. 33: 309-31

PEPPEDS

OUUU

Stratigraphic Unit

ARNOLD, C. A. 1936. Some fossil species of Mahon
from the Tertiary of eastern and southeastern Or-
egon. Contr. Mus. Paleontol. Univ. Michigan 5:
57-66.

White River Fm., below ash D

Huelster Fm.

. 1937. Observations on the fossil sb, e Pali
= > = > 3 s ern and southeastern Oregon, Part I.
Paleontol. Univ. Michigan 5: 79-102.
95]. Fossil capsule valves of Kolreuteria
from the John Day series of Oregon. Palaeobot-
anist 1: 74-78.
Tertiary conifers from the [nicum
Coal Field of British Columbia. Contr. M
leontol. Univ. Michigan 12: 245-258.
& . DAUGHERTY. 1963. The fern genus
Acrostichum i in the Eocene Clarno of Oregon. Contr.
Mus. Paleontol. Univ. Michigan 18: 205-227
& 19 A fossil dennstaetioid fern
from the Eocene Clarno of Oregon. Contr. Mus.
Paleontol. ws Michigan 19: 66-68.
AXELROD, D. I. 75. Tertiary floras from the Rio
Grande Rift. E 85-88 in New Mexico Geol. Soc.
Guidebook, 26th Field Conf., Los Cruces County,
New Mexico.
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293.

NALMA
9

Source

pers. comm
pers. comm.

°

, 1987
1987
1987
1987
1920

od

od,

od,

od,

ings
Jennings, 1920

,
,

ilson

ilson
Wilson, pers. comm.
Leopold & MacGinitie, 1972

Berry, 1919

Wilson, pers. comm.
Wil

w

Axelr

Axelr

Axelr

Axelr

Jenni

Locality

Continued.

BARNETT, J. 1984. Palynology of Tertiary floras of
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C.H 41. The Eocene flora of the Mid-
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[Abstra

ct
BARRINGTON, D. S. 1983. Cibotium oregonense: an

Creede, 5-Mile Bridge; 571A, B, C
Creede, Birdsey Gulch; 574A,
Creede, Dry Gulch; 572A, B,
Creede, Wason Cliffs; 573A, B

Poison Springs palynoflora
Missoula (loc. 165)

Missoula (loc. 196)

LaPorte
Alvord Creek
Cow Creek

APPENDIX I.
unnamed
Bridge Creek

=
>
x=
Z
T
J
"U =
a

Num-
ber
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251

778

APPENDIX II. Continued.

Eocene tree-fern stem and petioles with internal

structure. Amer. J. Bot. 70: 1118-1124.
BASINGER, J. F. 1976. Paleorosa similkameenensis,

gen. et sp. nov, permineralized flowers (Rosaceae)

from the Eocene of British Columbia. Canad. J

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1977. Fossil plants from the Middle 0095
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The Hohe did Rad of d
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G. W. ROTHWELL. 1977. Anatomically pre-
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55: 990.
BEBOUT, Ü Fanon of the Paleocene-Eocene Gold-
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Ph. D. Oats eene une Univer-
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struction for the Golden Valley Formation (Pa-
leocene-Eocene of North Dakota, based o
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Tertiary coniferous woods of west-
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Hx H. F. 1959. A new species of Mahonia from
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33-38.

. 1960. Additions to the Ruby paper shale flora
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87: 386-396.
1962a. Two new species of Mahonia from
the Grant-Horse Prairie Basin in southwestern

l

actes Bull. Torrey Bot. Club 89: 319-330.

. The fossil peepee Hs the genus Rosa.

Bull. ie Bot. Club 90:

96 EE n to the Oli-
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966. Additions to and revision of the Oli-
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1968. A hexasepalous calyx of the fossil As-
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1973. A new Tertiary gramineous fossil. Bull.
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BERRY, E. 924a. An early Eocene florule from
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7-92

1924b. A Sparganium from the middle
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19 A new Salvinia from the Eocene (Wy-
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91-116.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

— ə 1937b.

[VoL. 74

APPENDIX II. Continued.

. 1930. A flora jae aiie River age in the Wind
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— 1932a. poi plants from Wyoming. Amer.
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32b. A etc NN fruit from the lower
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BEYER, A. F. 1954. Petrified wood from Yellowstone
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BowN, T. M. 1982. Geology, paleontology, and cor-
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01-A: 1-75.

. 1962. Chloroplast in Spirogyra from
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967. Two aquatic fungi (Chytridiales) of
Eocene age from the Green River Formation of
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. Eocene algae and plant hairs from the
Green oid Formation of Wyoming. Amer. J
Bot. FUR -78
. 1 a ur tufa from the Eocene Green
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1289-12
BRITTON, E. G. & A. HOLLICK. ids Riiie fossil
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—142.

A new American fossil moss.

Bull. Torrey Ei Club 42: 9-10.

BRowN, J. T. 1975. Equisetum clarnoi, a new species
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BRowN, R. W. 1929, Additions to the flora of the
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934. The recognizable species of the Green
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-7

193 6. The genus G/yptostrobus in North

America. J. Wash. Acad. Sci. 26: 353-357.
1937a. Fossil legumes de e Creek,

Oregon. J. pis Acad. Sci. 27: 418.

rther additions be some Pu floras

of the vestem “United States. J. Wash. Acad. Sci.

27: as "E

—F Fossil leaves, fruits, and seeds of Cer-
siti 4 Paleontol. 13: 485—499,

w species and changes of name in
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30: 344-

—_— rs Temperate species in the Eocene flora
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1946. Alterations in some fossil and living

floras. J. Wash. Acad. Sci. 36: 344-355.

. 1950. An Oligocene Weg cherry from

Oregon. J. Wash. Acad. Sci. 40: 321-324.

1954. Oligocene plants ue correlation. Sci-

ence 119: 350.

1956. New items in Cretaceous and Tertiary

1987]
APPENDIX II. Continued.

floras of the western United States. J. Wash. Acad.
Sci. i ET 4—108.

; . Some paleobotanical problematica. J.
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1959b. A bat and some plants from the upper
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1973. p of O yptostrabus.
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1925b. A record of the presence of Umbel-
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Carnegie Inst. Wash. Contr. Palaeontol. 59: 62.
l . Geology and paleontology of the
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8

1933. Notes on occurrence n nd of fossil
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1936. The succession n and distribution of Ce-

Setchell. Univ. California Press, Berkeley, Cali-

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. 1938. Paleoecological interpretations of Ce-
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Tertiary forests and eiu his-

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1944. A fossil vv dus ur Eocene of

Utah. Amer. J. Bot. 31:

1947. Tertiary centers ese migration routes.

Ecol. Monogr. 17: 139-148.

. Early Tertiary ecotones in western
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1951. A revision of Sequoia and Taxodium

in western North America based on the recent

discovery : Metasequoia. Trans. Amer. Philos.

Soc. 40: 171-263.

. 1952. "aid dominants in the middle Ter-

tiary of the John Day Basin, Oregon. Palaeobot-

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E. I. SANBORN. 1933. The Goshen flora of
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P eieinE ^ D. A. 1906a. The fossil flora and fauna
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. 1906b. rap Lari from the Florissant, Col-

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1906c. Rhus and its allies. Torreya 6: 11-12.
——. 1907a. A Paes described as a moss. Tor-

reya 7: 203-2

1907b. Te genus oe in Colorado.

Univ. ear Stud. 5: 42-43.
enumeration of the localities in
the E basin from which fossils were ob-

WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

APPENDIX II. Continued.

tained in 1906. Bull. Amer. Mus. Nat. Hist. 23:
127-132.

1907d. Description of a new Tertiary fossil
flower from Florissant, Colorado. Torreya 7: 182-
183

1908a. Descriptions of Tertiary plants. I.
Amer. J. Sci. 26: 65-68.
1908b. Descriptions of Tertiary plants. II.
Amer. J. Sci. 26: 537-544.
908c. Some results of the Florissant expe-
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1908d. The fossil flora of Florissant, T
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1909a. Eocene fossils pir Green River, Wy-
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1909b. A new genus of fossil Fagaceae from
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1909c. Fossil Euphorbiaceae, with a note on
Saururaceae. Torreya 9: 117-119.

1909d. Two new fossil plants from Floris-
sant, Colorado. Torreya 9: 184-185.

1910a. Magnolia at Florissant. Torreya 10:

191 Ob. Notes on the genus Sambucus. Tor-
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—— ———. 1910c. A fossil fig. Torreya 10: 222-223.
1910d. pw ing of Tertiary plants. III.
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910e. m Miocene trees of the Rocky
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. 191la. Fossil flowers and fruits. Torreya 11:
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1911b. New names in //ex. Torreya 11: 264.
. 1912. Fossil flowers and fruits. II. Torreya
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. 1913. Fossil flowers and fruits. III. Torreya
13: 75-7

914. Two new plants from the Tertiary rocks
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1915. Equisetum in the Florrisant Miocene.
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922a. A fossil buttercup. Nature 109: 42-

22b. A new genus e fossil Liliaceae. Bull.
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24. A genuine fossil Ophioglossum. Tor-
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. 1925. Plant and insect fossils from the Green
tl. Mus.

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1926a. The supposed fossil Ophioglossum.
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—. 1926b. A Miocene Orontium. Torreya 26:

19 A new oak from the Green River
Eocene. Torreya 27: 94-95.
l A supposed fossil catmint (from Flor-
issant, TEMER Torreya 4.
1935b. A fossil Berberis. Torreya 35: 127.
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CRANE, P. R. & R. A. SrockEv. 1985. An extinct
species of Betula from the Middle Eocene of Brit-
ish Columbia. Amer. J. Bot. 72: 891

780

APPENDIX II. Continued.

CRIDLAND, A. A. & J. R. BUTALA. 1975. Reconsid-
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Cross, A. T. & E. MARTÍNEZ-HERNÁNDEZ. 1974. Pa-
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& 1980. aula ys pollen in early

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a.
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1939. Middle Eocene flora from the vol-
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——. 1974. Early Tertiary fossil forests of Yellow-
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1974b. Palynology of the "Fossil
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that from a modern coast redwood forest. 16th

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

APPENDIX II. Continued.

Annual Meeting Amer. Assoc. SEMIS Pal-
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. AGUIRRE & W. J. Fritz. 1978. Ad-
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,R. A. Cu N, H. P. M & L. H. Fisk.
1984. cries telah and oil dies es in the Fos-
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FREDERIKSEN, N. O. 1981. Comparison of Middle
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983. Late Paleocene and Early Eocene spo-

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Fritz, W. J. 1979. Depositional environment of the
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1980. Areview of Pinus wood from the Eocene

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1982. Geology of the Lamar River Forma-

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984. Yellowstone fossil forests: new evi-

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Fisk. 1979. Paleoecology of petrified

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GAPONOFF, S. L. 1984. Palynology of the quarc
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GASTALDO, R. A., L. C. MATTEN & M. R. LEE. 1977.
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1987]

APPENDIX II. Continued.

GREGORY, I. 1976. An extinct Evodia wood from
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HarL, W. J., E. B. LEoPorp. 1960. Paleocene
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HALL, J. W. & A. M.SwArN. 1971. Pedunculate sida
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A L. Plant fossils in the Clarno

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R. K. PETERSON. 1978. Zingiberopsis,

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52.
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JANSSENS, J.A F. BASINGER.
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1904. A fossil Sequoia from the Sierra

98
geography of the Eocene Chuckanut Formation,

WING—EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

781

APPENDIX II. Continued.

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KIRCHNER, W. C. G.
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KIRN, A. J. & H. B. PARKs. 1936. An Eocene florule
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1898. Contributions to the fossil
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782

APPENDIX II. Continued.

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Love, J. D., . B. LEoPoLD & D. W. Love. 1978.
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ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

APPENDIX II. Continued.

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1987]

APPENDIX II. Continued.

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WING —EOCENE & OLIGOCENE FLORA OF THE ROCKY MOUNTAINS

783

APPENDIX II. Continued.

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784

APPENDIX II. Continued.

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ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

APPENDIX II. Continued.

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AN OVERVIEW OF THE ORIGINS OF THE MODERN
VEGETATION AND FLORA OF THE
NORTHERN ROCKY MOUNTAINS!

JACK A. WOLFE?

ABSTRACT

The present flora of the northern Rocky Mountains has diverse origins. The Late Cretaceous meso-
thermal to megathermal evergreen vegetation of this region had few taxa, even at the familial level,
that live in the northern Ro ok i i
leaved deciduous taxa occupyi
they un nderwent major diversification. Bes early Eocene thermal maximum severely restricted the

areas of l lineages
By the early middle Eocene, volcanic sehen that supported micrathera coniferous forests of
Pinaceae and Cupressaceae had developed in s of the northern Rocky Mountain region. These

ades into newly created T climate. During

Eocene by way of Beringia. Some of the Eocene microthermal lineages s
in the northern Rocky Mountains today, and other extant lineages, although ultimately derived from
taxa in the Eocene uplands, represent morphological types S — ated and diverged in Eurasia,
arriving in North America by migration. Oligocene and Neo
northern Rocky Mountains, although the Columbia Plateaus [s oe t contain m
of these ages. Th l broad-lea . deciduous or coniferous forests,
t lineages that were derived fro
the terminal Eocene temperature deterioration. With the presence of many extant northern Rocky

the mid-Miocen
exchanged between western North sampa, me and Eura x

lineages are known during the Miocene only in areas ne as Alaska and probably represent migrants
into the northern Rocky Mountains during the late Neogene. The present flora of the northern Rocky
Mountains therefore clearly represents a complex overlay of numerous historical biogeographic pat-
terns.

The modern vegetation of the northern Rocky
Mountain region is primarily steppe at low al-
titudes and coniferous forest at higher altitudes
(Habeck, 1987). Temperatures are entirely mi-
crothermal.? The lower and drier part of the co-
niferous forest belongs to the Pinus ponderosa
zone, whereas most of the mesic coniferous forest
at higher altitudes belongs to the Abies grandis
nklin & Dyrness, 1969). The vegeta-

northern Rocky Mountain region are docu-

mented by successive microfossil and megafossil
plant assemblages. In some instances, lack of as-
semblages that represent particular intervals
makes it necessary to infer climate and vegeta-
tion from the fossil record of adjacent regions.

ic) types that have occupied
Mountain region in the past and the history of
the lineages that comprise the modern flora.
Throughout, the focus is primarily on the woody
flora.

! For many helpful discussions, I thank H. E. Schorn, G. R. Upchurch, = and Wesley Wehr. Schorn has

also supplied data from his unpublished studies on the p ú iun

. R. Crane and B. H. Tiffney offered useful comments o

ern American Tertiary conifers.

? Paleontology and Stratigraphy Branch, MS-919, U.S. s EE “sassa Federal Center, Denver, Colorado

80225, U.S.A.

3 Microthermal = mean annual temperature <13°C, mesothermal =
°C

megathermal = mean annual temperature > 2

ANN. MISSOURI Bor. GARD. 74: 785-803. 1987.

mean annual temperature 13-20?C,

786

The extant woody flora is composed of 10 gen-
era and 19 species of conifers and 30 genera and
80 species of woody dicotyledons. The dominant
conifers are Abies grandis, Larix occidentalis, and
other members of Pinaceae, as well as a fe
members of the C upressaceae, The dicotyledons
arc pri Imar ily tor form
streamside communities. Conspicuous dicoty-
ledonous families, as elsewhere in Northern

emisphere vegetation, include Betulaceae, Er-
icaceae, Salicaceae, Rosaceae, and Caprifoli-
aceae, with a few Aceraceae.

The origins of the flora of a region such as the
northern Rocky Mountains are, of course, highly
complex. Each component species, genus, and
Family hasa unique history, REDE some his-

ories pace and time.
indeed ‘the search for patterns to which numer-
ous histories approximately conform is a pri-
mary concern of historical plant geography.
Such patterns might be indicative of similar re-
sponses to specific historical factors, particularly
responses to environmental changes, including
various parameters of climate and topographic
changes resulting from orogenic factors. How-
ever, because two taxa a are now in association or
have similar p timply
similar histories.

The classical methodology for determining the
histories of biogeographic patterns involves re-
construction of phylogenies from phenetic sim-
ilarities of extant and fossil taxa, observing dis-
tribution patterns of extant taxa, and determining
the patterns of distribution of fossils in time and
space. The relative completeness of the fossil rec-
ord is the major problem in using it in biogeo-
graphic analyses. The problem is underscored in
the present case by the almost total absence of
Oligocene and Neogene assemblages of plant
inet in tae northern Rocky Mountains.

ges fill this
gap, the typical taxonomic resolution of palyno-
logical morphology is typically at a generic or
higher level (Muller, 1970) and is of limited as-
sistance in determining histories of lineages on
the scale desirable in biogeographic analyses.
Further, the upland Eocene megafossil assem-
blages from the northern Rocky Mountain region
have been actively collected and studied for only
the last two decades. These assemblages are crit-
ical because they contain the records of diver-
sification of many microthermal groups (see be-
low), and additional collecting and study are
needed. In contrast, the Oligocene and Neogene

E

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

plant-megafossil assemblages from adjoining re-
gions such as the Columbia Plateaus and from
possible source areas such as Beringia are well
known. Estimates based on the geographic and
climatic distribution of fossil and extant taxa in-
dicate that, at least in the Beringian Neogene, the
lineages represented in the fossil record are
thought to be more than 9596 complete for woody
plants.

Inferences also can be made regarding histories
of lineages if a cladistic analysis of a group is
combined with detailed knowledge of history of
the physical environment. This approach, termed

og ick & Nel-
son, 1978) has considerable merit for groups that
have poor or no fossil records. However, it rests
on major assumptions and requirements:

(1) Cladistic analyses inherently rest on the as-
sumption of parsimony in evolution of char-

cters.
(2) Cladistic analyses must be based on valid
determination of polarities of character states.
(3) idi of the physical environments in all
areas m known in detail.
(4) The iid biogeographic analysis also rests on
the assumption of parsimony in dispersals.

In most recent applications, vicariance biogeog-
raphers have either ignored the fossil record or,
at best, have incorporated fossil taxa in cladistic
analyses without reference to stratigraphy, al-
though Grande (1985) emphasized the signifi-
cance of the age relations of fossil taxa, which
provide additional geographic data on distribu-
tions. Another major tendency has been to con-
sider the history of the physical environment
solely in terms of plate tectonics, thus ignoring
particularly climatic changes. Because climates
of a given area generally determine vegetational
types and, in turn, vegetation organizes the en-
vironment into a variety of microenvironments
(Upchurch & Wolfe, 1987) that determine
whether a particular organism can live in a given
area, the history of vegetation assumes para-
mount importance for interpreting the biogeo-
graphic histories of land organisms.

The fossil record can be of major significance
in any analysis in vicariance biogeography, as
Grande (1985) emphasized. Howe b

from the standpoint of icthyology, some contri-
butions of paleontology to vicariance biogeog-
raphy were not stressed or were overlooked:

1987]

(1) The stratigraphic sequence of grades, if cau-
tiously interpreted, can determine polarity
states (Hennig, 1966).
The fossil record can supply evidence of en-
vironments in particular areas at particular
times.
(3) The fossil record can supply evidence of the
existence of dispersal barriers and routes oth-
er than those related to plate tectonics.
The fossil record can supply minimal times
cladogram, and these times
can then be placed in the framework of the
then-existing environments to determine
probabilities of dipersals of the resulting, di-
verging lineages.
The fossil record can independently test the
assumptions of parsimony in both evolution
and dispersal.

~
N
—

~
A
—

h,
wa
—

Unfortunately, almost no cladistic analyses of
groups of concern in the northern Rocky Moun-

tains have been carried out. However, the results `

of a cladistic analysis of Acer (Wolfe & Tanai,
1987) when compared with the known fossil re-
cord of Acer (especially in North America) in-
dicate:

(1) The known relative times of appearances of
sections of Acer generally correspond to those
predicted from the cladogram; those few that
do not correspond probably resulted from
the absence of collections from the appro-
priate environment or areas

Phylogeny reconstructed on a phenetic pa-
leobotanical basis compares well with phy-
logeny reconstructed on a cladistic basis.
Dispersals have not been as parsimonious as
would be predicted from vicariance bio-
geography.

The present distributions of species and sec-
tions of Acer are complex and have resulted
from a variety ofenvironmental, particularly
climatic, factors.

~
N
—

~
Ww
wa

~
+.
—

As more cladistic analyses of extant groups are
completed, they will provide a framework for
evaluating phenetic paleobotanical phylogenies,
but cladistic analyses, because of biogeographic
“noise,” can provide only highly generalized
concepts of histories of biogeographic patterns,
some of which are much more complex than
indicated by cladistics alone and some of which
are invalid (Wolfe, 1981b). In essence, cladistic
analyses test phenetic phylogenies based on the
fossil record, whereas the fossil record tests

WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION

787

models developed from vicariance biogeogra-
h

The divergence that apparently exists between
*vicariance" and “‘dispersalist’”’ schools of bio-
geography has resulted from psychological and
sociological factors. Each school typically has re-
jected the totality of the other’s methods and
results. Both have inherent problems that can be
resolved by reference to the other’s methods and/
or by using the other’s conclusions. Indeed,
Grande’s (1985) discussion of vicariance bio-
geography basically incorporates some dispers-
alist concepts and uses some dispersalist data
that support his conclusions; he argued for sys-
tematic/biogeographic analyses on successive
time planes to attempt to filter out biogeographic
"noise" introduced by subsequent dispersal
events. The approach used in the present report
similarly will involve both cladistic and dispers-
alist concepts and data.

CRETACEOUS

The northern Rocky Mountain region was oc-
cupied by a mesothermal, broad-leaved ever-
green forest during the Late Cretaceous (Dorf,
1942). Some conifers (e.g., Araucariaceae, ev-
ergreen Taxodiaceae) probably were emergents
in this vegetation (Wolfe & Upchurch, 1987b),
and broad-leaved deciduous plants were restrict-
ed largely to successional or disturbed vegeta-
tion, especially along streams. General absence
of drip-tips and a somewhat small leaf size in-
dicate slightly subhumid conditions (Wolfe &
Upchurch, 1986, 1987a). Analyses of woods from
the North American mesothermal region suggest
little seasonality of either temperature or precip-
itation (Wolfe & Upchurch, 1987b). Presence of
extensive marine rocks of Late Cretaceous age
throughout much of the Western Interior indi-
cates generally low altitudes.

Miller (1987) emphasized the great floristic
disparity between known Early Cretaceous flo-
ras, including those of the northern Rocky
Mountain region, and the modern flora of this
region. Even with the rise to dominance of the
angiosperms in the early Late Cretaceous, the

zed ber
dae hese. 1987). Whether any of these ul-
timately gave rise to taxa that now occur in the
northern Rocky Mountains cannot now be de-
termined. By the later part of the Late Creta-
ceous, some families that today have microther-
mal members become recognizable. Aceraceae,

788

for example, occur in the latest Cretaceous of
central Alberta, represented by an extinct genus
interpreted as a sister group to Acer (Wolfe &
Tanai, 1987). Even at high paleolatitudes, the
flora was composed largely of trochodendroids,
hamamelidaleans, and deciduous Taxodiaceae
(Spicer et al., 1987); this high-latitude flora has
some floristic similarities to that of the early Ter-
tiary of the northern Rocky Mountains but no
similarity to the extant flora of this region.

PALEOCENE

In areas such as eastern Montana, the Creta-
ceous-Tertiary boundary is marked by a pre-
sumed fallout layer that has anomalously high
amounts of iridium and shocked minerals (Boh-
or et al., 1984); these are considered to be evi-
dence of an impact by an extraterrestrial body
(Alvarez et al., 1984). This event at the end of
the Cretaceous had a profound effect on the flora
and vegetation of the northern Rocky Mountain
region (Wolfe & Upchurch, 1986). At least 50-
60% of the latest Cretaceous lineages became ex-
tinct (Hickey, 1981). Most extinctions were in
the previously dominant broad-leaved evergreen
element; in addition, araucarians and many oth-
er evergreen conifers became extinct in this re-
gion. Immediately above the presumed fallout
horizon, palynofloras contain almost exclusively
fern spores (Hotton, 1984; Tschudy & Tschudy,
1986). Within a few centimeters above this ho-
rizon, angiosperm pollen occurs with the fern
spores; the associated leaf flora contains both
fern pinnae and a compound-leaved rosid, which
is thin-leaved and probably deciduous. About 10
meters higher in the section, the leaf assemblage
of about 15 species consists almost entirely of
deciduous angiosperms (trochodendroids, ha-
mamelidaleans, Tiliaceae) and deciduous coni-
fers (Taxodiaceae).

The eastern Montana sequence immediately
above the Cretaceous- Tertiary boundary resem-
bles normal secondary succession following a
volcanic eruption (Richards, 1952); in mesother-
mal, broad-leaved evergreen regions, deciduous
plants are typically dominant in secondary
successions (Wang, 1961). However, in the
northern Rocky Mountain region, broad-leaved
deciduous plants dominated north of the Colo-
rado- Wyoming border (Brown, 1962), and the
broad-leaved evergreen element did not return
to dominance until almost the end of the Paleo-
cene, about 10 Ma following the Cretaceous.
Hickey (1981, 1984) attributed the deciduous-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

ness of this Paleocene vegetation to an overall
decline in temperature that persisted through
most of the Paleocene. The occurrence of large
(and presumably ectothermic) reptiles (crocodil-
ians, champsosaurs, large turtles) with this de-
ciduous vegetation, however, indicates the prob-
able absence of low winter temperatures that
would select for deciduousness. That is, the fau-
nal data indicate strongly that the deciduousness
of the vegetation is anomalous; just as signifi-
h

vegetation persisted in the early Paleocene at
about the Colorado- Wyoming border.

Wolfe & Upchurch (1986) suggested instead
that the terminal Cretaceous event resulted in a
brief low-temperature excursion that devastated
the vegetation of both mesothermal and mega-
thermal regions of North America. Some mega-
thermal, broad-leaved evergreen lineages sur-
vived (perhaps in refugia; Tschudy et al., 1984),
and megathermal vegetation continued to be
dominantly evergreen. The megathermal vege-
tation underwent physiognomic change and
gradually increasing diversity during the Paleo-
cene, a phenomenon sess short-term sec-

ondary succession and termed “quasisucces-

Wolfe & Upchurch (1986, 1987a).

broad-leaved evergreen lineages survived, and
replenishment of them would have had to occur

y adaptation of some megathermal lineages to
mesothermal climate on an evolutionary (not
successional) time scale.

The anomalously deciduous character of Pa-
leocene mesothermal vegetation throughout
North America (and in Eurasia) gave deciduous
angiosperms a unique opportunity. These decid-
uous lineages were derived primarily from ele-
ments that were uncommon in Late Cretaceous
mesothermal evergreen vegetation and/or dom-
inant in Late Cretaceous, high-latitude decidu-
ous vegetation (Wolfe & Upchurch, 1986). In
mesothermal regions, dominantly deciduous taxa
(e.g., Hamamelidaceae, Fagaceae, Betulaceae,
Ulmoideae, Juglandaceae) diversified during the
Paleocene (e.g., Nichols & Ott, 1978; Manches-
ter, 1987). By the end of the Paleocene, the
Northern Hemisphere had a considerable diver-
sity of broad-leaved deciduous plants, and many
families of woody angiosperms that now occur
in the Rocky Mountain region were extant.

major increase in precipitation occurred at
the Cretaceous—Tertiary boundary. In the north-
ern Rocky Mountain region this is evidenced by

1987]

a substantial increase in leaf size and by the ini-
tiation of wide-scale peat deposition that typi-
cally marks the early Paleocene in this region
(Wolfe & Upchurch, 1986). Not only would in-
creased precipitation result in an increase in bio-
mass, but water tables also would be raised and
more swamps developed.

Pre-Eocene microthermal vegetation contrasts
markedly with Eocene and later microthermal
vegetation in floristic composition. Pre-Eocene
microthermal vegetation was restricted to high
latitudes and, probably due to low winter light
levels, was almost entirely deciduous (Wolfe,

iS

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ciduous Pinaceae (Pseudolarix), and deciduous
dicotyledons, the great majority of which rep-
resents a few clades of trochodendraleans, plat-
anoids,and other hamamelidaleans (Spicer et al.,
1987; Wolfe, 1987). Although these clades later
contributed some lineages first to the meso-
thermal deciduous vegetation of the Paleocene
and then to the Eocene upland microthermal
vegetation, they comprise a small fraction of the
Eocene and later microthermal flora. The mi-
crothermal vegetation and flora of the Late Cre-
taceous and the Paleocene have few similiarities
to Eocene and later vegetation and flora.

EOCENE

A warming initiated during the latest Paleo-
cene culminated in the early Eocene thermal
maximum (Savin, 1977; Wolfe & Poore, 1982;
Wolfe, 1985). Vegetation in areas such as north-
western Wyoming was dominantly broad-leaved
evergreen and represented warm mesothermal
temperatures (Wing, 1981, 1987). At somewhat
lower altitudes (and particularly in the lowland

acific Northwest), vegetation represented
, 1985).

region was still generally of low altitudes is in-
dicated by the persistence of large lakes from the
early into late Eocene in areas such as south-
western Wyoming and adjacent Utah. The early
Eocene vegetation indicates abundant precipi-
tation, but later Eocene vegetation indicates
increased seasonality of precipitation and de-
velopment of subhumid, seasonal climate
(MacGinitie, 1969; Leopold & MacGinitie, 1972).

e warm mesothermal to megathermal vege-
tation of the early and middle Eocene has few
floristic similarities to the extant flora of the
northern Rocky Mountains. A few extant north-

WOLFE-ORIGINS OF NORTHERN ROCKY MODERN VEGETATION

789

ern Rocky Mountain genera (e.g., A/nus, Populus,
and Acer) are present in this vegetation, but the
species represented are not closely related to ex-
tant species in this regio

Extensive vulcanism and associated tectonism
resulted in a major upland region that extended
from northern Nevada and central Idaho north
into British Columbia during the middle Eocene
(Axelrod, 1966a, 1966b); this upland may have
actually been a series of upland volcanic centers.
Known upland assemblages of early middle
Eocene age (Fig. 1) occur from northeastern
Washington (Republic and associated floras)
northward into central British Columbia (Prince-
ton, Chu Chua Creek, and coeval floras). Exten-
sive vulcanism of early Eocene age became less
intense during the early middle Eocene, and tec-
tonism resulted in a series of grabens, in which
the plant-bearing lacustrine sediments were de-
posited (Pearson & Obradovich, 1977). Altitude
of the Republic basin of deposition is estimated
to have been about 900 m (Wolfe & Wehr, 1987).

Later in the middle Eocene and in the early
part of the late Eocene, major volcanic centers
occurred in central Idaho (Challis Volcanics),
central Oregon (Clarno Formation), and north-
eastern Nevada (rocks containing the Copper Ba-
sin and Bull Run floras of Axelrod, 1966b). Most
Clarno assemblages represent only low altitudes,
but a newly collected assemblage (Sheep Rock
Creek) from the Crooked River Basin is probably
microthermal and indicates the presence of higher
altitudes. By the late Eocene, tectonism in the
central Idaho region resulted in the formation of
grabens and associated lakes on the post-Challis
surface; these lake beds contain floras such as the
Salmon. A similar depositional regime occurred
in southwestern Montana (the numerous floras
described by Becker); to the north in western
Montana are plant-bearing lacustrine beds that
have been collected recently by C. N. Miller and
associates.

The development of this upland region, con-

comitant with a general decline of temperature

following the early Eocene thermal maximum,

tation, such as that at Republic (Wolfe & Wehr,
1987), represents the oldest known coniferous
forest dominated by Pinaceae (Abies, Picea, Pi-
nus, Pseudolarix, and Tsuga) and Cupressaceae
(Chamaecyparis and Thuja). These genera were
of low diversity in the early middle Eocene, but
had increased in diversity by the end of the

790 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

p.
S
7A
S
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$2»
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0 100 200 300 MILES ANA
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l I estern conterminous United

FiGurRE 1. Map showing location of E i

States por adjacent Canada. For each prei o i or group of PEREA sapi ig nae references or
repositories of largely unpublished assemblages are cited. These repositories are: C (Geological Survey of
Canada); OMSI (Oregon Museum of Science and Industry); TMP (Tyrrell M e 5. UAPC
(University of Alberta Paleobotany Collections); UBC (University of British Columbia); UCMP (University of
California Museum of Paleontology); UMPC (University of Montana Paleobotany Collections); USGS

Geological Survey); UWBM (University of Washington Burke Museum). Early middle Eocene: 1. Smithers
(UWBM); 2. Chu Chua Creek (Berry, 1926); 3. Kamloops assemblages (TMP); 4. Princeton assemblages (GSC,
UAPC, UWBM); 5. Republic assemblages (Wolfe & Wehr, 1987; UWBM). Late middle to early late Eocene:
6. Sheep Rock Creek (OMSI); 7. Alvord Creek (Axelrod, 1944b); 8. Elko assemblages (UCMP, USGS); 9. Bull

( ); 16. Beaverhead, Metzel Ranch, Mormon Creek, Ruby, and York Ranch (Becker, 1960,
1961, 1969, 1972, 1973).

1987]

Eocene. As discussed later, major diversification
also occurred in numerous woody dic direi
ous families. This probably resulted from (1) th
areal restriction of microthermal climate Pian
the early Eocene and (2) the opportunistic ex-
pansion of mesothermal lineages into newly cre-
ated microthermal climate in the uplan

reas of microthermal climate were des
highly restricted during the early Eocene thermal
maximum. Indeed, I know of no latest Paleocene
or early Eocene Northern Hemisphere plant as-
semblage that represents microthermal vegeta-
tion; possibly only on mountains in polar lati-
tudes could such vegetation have been present
(Wolfe, 1985, fig. 5). Because of the absence or
extreme restriction of microthermal climates
during the early Eocene, probably few pre-Eocene
microthermal taxa were able to survive this cli-
matic-geographic bottleneck, leaving the newly
expanding microthermal regions available for

l lineages. Further, to-

bokranhie and edaphic diversity, Juxtaposition
of many microclimates, and relative isolation of
one volcanic center from another would all lead
to rapid diversification of lineages in the uplands.
Geographic proximity to lowland mesothermal
vegetation would allow a continuing supply of
new clades.
robable instability of community composi-
tion during the middle Eocene is indicated by
two lines of evidence. First, from one approxi-
mately coeval depositional site to another, species
composition varies markedly; almost all the lo-
calities are in lacustrine shales and presumably
represent similar ecologic settings. Spatial het-
erogeneity more resembled that of mesothermal
or even megathermal vegetation than that of mi-
crothermal vegetation. Second, floristic compar-
isons of early middle Eocene lacustrine assem-
blages with late middle to early late Eocene
assemblages indicate large changes in composi-
tion, some related to evolution of lineages within
microthermal vegetation and some related to ad-
ditions from mesothermal vegetation, as well as
to extinction.

Numerous mesothermal (or mesothermal/
megathermal) evergreen dicotyledonous clades
had adapted to the microthermal climate by the
early middle Eocene, although these typically are

phytocrene (Icacinaceae), and Schoepfia (Olaca-

WOLFE-— ORIGINS OF NORTHERN ROCKY MODERN VEGETATION

791

ceae). None of these genera are known to have
survivedi l vegetation of the late
norm and they can best be considered as un-
S sf although other evergreen
dicotyledons successfully adapted to the mi-
crothermal climate during the Eocene. The pres-
ence of evergreen dicotyledons in microthermal
coniferous forests during the Eocene is consistent
with the low (ca. 5°C) mean annual range of tem-
perature inferred for coeval lowland assemblages
(Wolfe, 1978). Modern microthermal coniferous
vegetation of low-latitude, upland areas (e.g.,
Taiwan, Himalayas) also has many taxa of ev-
ergreen dicotyledons.

Also included in the upland early middle
Eocene vegetation are numerous extant genera
of trees and shrubs that contributed to broad-
leaved deciduous and coniferous forests of the
Neogene and Holocene, although not present in
the northern Rocky Mountain region today. In-
cluded are: Sassafras, Cercidiphyllum, Corylop-
sis, Comptonia, Castanea, Fagus, Tilia, Ulmus,

a, Photinia, Decodon, Rhus, and Aesculus.

Overall, diversity among woody angiosperms
appears to have been higher in the initial phase
(early middle Eocene) of development of the mi-
crothermal coniferous forests than later in the
Eocene. At least 95 genera and 140 species of
woody angiosperms are known in the Republic
and Princeton floras, which are still not thor-
oughly collected. In contrast, about 70 genera
and 110 species of woody angiosperms are known
in the extensively collected and described latest
Eocene coniferous forests from western Mon-
tana (Becker, 1960, 1961, 1969, 1972, 1973).
This general decrease is particularly notable in
presumed large trees, which were perhaps grad-
ually replaced by newly evolved species of co-
nifers.

Very few of the extant northern Rocky Moun-
tain lineages are recognizable by the early middle
Eocene. The oldest known Betula clearly allied
to the B. papyrifera—B. occidentalis complex oc-
curs at Republic, as does an Acer allied to A
negundo. However, the Acer, although possibly
ancestral to A. negundo, is assignable to an extinct
section (Wolfe & Tanai, 1987). Many other ex-
tant native microthermal genera were also par-
ticipants in this early middle Eocene vegetation
(e.g., Alnus, Corylus, Ribes, Rubus, Spiraea, Cra-
taegus, Prunus, and Cornus), but the species can-
not be placed directly in the ancestry of extant
northern Rocky Mountain lineages.

The early middle Eocene microthermal flora

792

of western North America had numerous archaic
elements in terms of Neogene or Holocene mi-
crothermal vegetation. Other than the bond
cessful tl ae
ed, the vegetation had extinct genera of
Trochodendraceae, Cercidiphyllaceae, aie a-
melidaceae, Platanaceae, Fagaceae, Betulaceae,
Rosaceae, and other families; about 40% of the
genera are totally extinct (Wolfe & Wehr, 1987,
unpubl. data

By the late middle to late Eocene, upland mi-
crothermal vegetation had been floristically al-
tered as the result of:

(1) Extinction, e.g., the unsuccessful thermo-
philic experiments.

(2) Gradual evolution in phylads, e.g., in a phy-
lad ultimately leading to Chamaebatiaria
(Rosaceae), the early middle and late Eocene
members represent distinct genera.

(3) Major diversification of early middle Eocene

microthermal clades, e.g., whereas early

middle Eocene Acer comprised 10 species
and 3 extinct sections, late middle to late

Eocene Acer comprised 35 species and 17

sections, 14 of which are extant (including

the first members of sects. Negundo and Ma-
crophylla).

Adaptation of members of previously meso-

thermal clades to microthermal climates, e.g.,

invasion of newly evolved species of Quer-

cus, Mahonia, Salicaceae, and Leguminosae.

—
A
~x

Rosaceae and Aceraceae underwent major di-
versification in Eocene upland microthermal
vegetation. Both are today primarily bee-polli-
nated; entomophily would have been advanta-

angiosperms that are abundantly represented in
the fossil assemblages and that were presumably
of fluviatile habitats: trochodendroids, hama-
melidaleans, Fagopsis, most Betulaceae, Comp-
tonia, and Ulmoideae. However, the fluviatile
habitat and concomitant anemophily/abiotic
dispersal syndrome did not lead to more than
species-level diversification in most of these
groups.

The great majority of extant genera of woody
microthermal angiosperms had evolved by the
end of the Eocene, yet diversification in some
genera and perhaps families had yet to occur:

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

(1) In Salix, all known Eocene species are re-
ferable to subg. Salix.

(2) Ericaceae were of low generic and specific
diversity in the Eocene; thus far only Rho-
dodendron (subg. Rhododendron) has been
found in the upland Eocene vegetation, al-
though several small Eocene leaves that lack
diagnostic characters have been referred to
Vaccinium.

Only a few scattered Eocene palynological
records of Caprifoliaceae s. str. are known,
e.g., Diervilla in the latest Eocene of Alaska
(E. B. Leopold, written comm., 1970). A few
nondiagnostic, microphyllous leaves have
been referred to Symphoricarpos.

~
w
—

OLIGOCENE AND NEOGENE

Wide-scale deposition of lacustrine sediments
apparently ended in the northern Rocky Moun-
tains by the end of the Eocene (ca. 33 Ma)*. This
termination presumably resulted from consid-
erable uplift, leading to downcutting and erosion.
At approximately the same time, a worldwide
major climatic deterioration occurred (Wolfe,
1978), resulting in development of microther-
mal, broad-leaved, deciduous forest at low alti-
tudes of middle latitudes and broad-leaved de-
ciduous and coniferous forests at high latitudes
(Wolfe, 1985). This deterioration involved a de-
cline in mean annual temperature and a major
increase in mean annual range of temperature
(Wolfe, 1978).

Megafloras of Oligocene and Neogene age are
not known from the northern Rocky Mountains.
Wing (1987) and Leopold & Denton (1987) have
therefore largely concentrated on well-known as-
diee from the adjacent Columbia Plateaus

Fig. 2). From the Oligocene into the middle
Miocene (ca. 13-14 Ma), the Columbia Plateaus
were occupied largely by Mixed Mesophytic for-
est; during the middle Miocene, this forest was
replaced by Mixed Coniferous forest (Wolfe,
1981a).

That Miocene basalts of the Columbia River
Group generally lap onto highlands to the east
and north indicates that certainly the region of
the northern Rocky Mountains was higher than

=

4 Many workers place the Eocene-Oligocene bound-
ary at about 37 Ma and would include the latest Eocene
Miss discussed above in the early Oligocene. Which-

ver placeme trelative or actual
ee of the on or paleoclimatic changes.

1987]

the Columbia Plateaus. Based on the tempera-
ture relations of extant vegetational types (Wolfe,
1979), I infer that during the Oligocene through
middle Miocene, the northern Rocky Mountains
had a lower altitudinal belt of Mixed Northern
Hardwood forest (Fig. 3) and an upper belt of

igh Montane Mixed Coniferous forest (the
vegetational type that generally occupies mesic
sites within the present northern Rocky Moun-
tains).

The terminal Eocene temperature deteriora-
tion resulted both in numerous microthermal
lineages migrating downslope from Eocene up-
lands into Oligocene lowlands and in extinction
of many microthermal lineages. For example, the
relationships of most Bridge Creek (early Oli-
gocene) taxa are with Eocene upland taxa
(MacGinitie, 1953) and not with Eocene high-
latitude taxa. Among the Oligocene taxa that il-
lustrate this relationship are: Cercidiphyllum
crenatum, Castanea orientalis, Fagus pacifica,
Quercus clarnensis, Alnus “‘carpinoides,” Betula
aff. papyrifera, “Carpinus grandis," Mahonia
simplex, Plafkeria sp., * Ulmus” brownelli, Ame-
lanchier sp., Crataegus newberryi, Acer (Negun-
do) sp., A. (Macrophylla) osmonti, A. (Lithocar-
pa) sp., Bohlenia sp. (an extinct sapindaceous
genus that = Dipteronia auct.), and * Holmskiol-
dia" speirii. Some lowland Oligocene taxa were
probably derived from mesothermal to mega-
thermal taxa of the Eocene lowlands: Liqui-
dambar, Platanus, Engelhardtia, and Paleophy-
tocrene. Although such taxa are few, both
a and Platanus were vegetationally

mportant during the Neogene. None of the Bridge
os lineages appear to be derived from a north-
ern source.

Although the Oligocene data base is not as

Eocene include Fagopsis, some genera of
ceae, and the sister genus of Acer (" Acer" arcti-
cum group). Some Eocene microthermal lineages
that participated in Oligocene vegetation of the
Columbia Plateaus did not survive into the Mio-
cene (e.g., “Zelkova,” Plafkeria, Bohlenia,
* Holmskioldia"). Clearly, extinction continued
to play a significant role.

The Oligocene vegetation in high-latitude re-
gions such as Alaska was of low diversity (Wolfe,
1972, 1985) and provided a limited reservoir for
mid-latitude lineages. Only near the end of the
Oligocene did taxonomic diversity increase in

WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION

793

high-latitude regions, largely from migration from
lower latitudes and some diversification of im-
migrant clades; the influence at mid latitudes of
the high-latitude flora was generally not felt until
the middle and late Miocene.

By the early to middle Miocene, the flora of
the Columbia Plateaus had increased in diversity
relative to the Oligocene. This diversity increase
resulted from three major factors (Wolfe, 1972,
1977):

(1) Adaptation of mesothermal lineages to mi-
crothermal climates (e.g., Magnolia, some
Lauraceae, and Exbucklandia).

(2) Diversification of Oligocene microthermal
pee (e.g., Fagus, Alnus, Ulmus, Carya, and
Ace

(

o
~x

RE migration of Asian mid-latitude

lineages into Beringia and subsequent south-

ward migration onto the Columbia Plateaus

(e.g., four lineages of Acer belonging to sec-

tions Macrantha, Platanoidea, and Parvi-
ora)

Maximal diversity on the Columbia Plateaus and
adjacent areas was reached during the middle
Miocene (ca. 13-16 Ma)

Notable are the few species shared between
middle Miocene floras on the Columbia Plateaus
and the Kilgore flora of Nebraska (MacGinitie,
1962). Even a putative sh
erodentatum,” in sect. Negundo) is represented
in either region by distinct subspecies. Further,
most Columbia Plateaus species have only a dis-
ant to ofeastern North
America. Divergences between most western and
eastern American lineages had probably oc-
curred during, or at the end of, the Eocene, which
led to a distinctive western American element
(Wolfe & Tanai, 1980: 16-18). By the Miocene
(if not the Oligocene), the Rocky Mountain re-
gion must have formed an effective climatic bar-
rier to migrations of most warm microthermal
pianis (Laopoid & Pentan, R

d species (Acer “
snecies cer =
p ( het

-—

m j

f Acer,
which was apparently able to disperse from east-
ern into western North America. Members of
this series appear suddenly at middle latitudes
of western North America during the early Mio-
cene and have no Beringian record. Cladistic re-
lationships and the fossil record both suggest dis-
persal across the North Atlantic during the
Miocene (Wolfe, 1981b; Wolfe & Tanai, 1987),
and the two lineages of series Saccharodendron
present in the western American Neogene are

ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

& g ve, ç, o h ae AS +< thy, SOS.

* S
ii fi me
N "d
/ \ MONTANA
$
BIN
ay
° 0 100 200 300 400 500 MILES
Se L L L 1 L J
< rey oe
0 100 200 300 400 500 KILOMETERS
Wo

L / yi L l 1
132° 130° 128° 126° 124° 122°

1987] WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 795

Notophyllous broad-leaved evergreen forest

Mixed mesophytic forest

e temperatures
z peli
ateaus

o

Low montane
- mixed coniferous

forest Mixed northern

hardwood forest

Inferred early

in nort
Ocky jos

MEAN ANNUAL TEMPERATURE *C

High montane
_ ene sa
fore

bibi early
ne temperatures

in in alt cs
1

20 25
MEAN ANNUAL RANGE OF TEMPERATURE °C
FIGURE 3. gram showing inferred temperatures for the northern Rocky Mountains during the early
Neogene. C Nora) and temperature relations of forests adapted from Wolfe (1979). Inferred temperatures
bi

for Columbia Plateaus and Beringian assemblages adapted from Wolfe & Tanai (1980) and Wolfe (198 1a).
Arrows indicate direction of Neogene temperature trends.

E 2. Map showing location of some Oligocene and early Neogene (> 13 Ma) plant assemblages in

e

eetnuk River assemblages (USGS); 2. Colorado Creek (USGS); 3. Eagle River (Wolfe et al., 1966); 4. Harriet
Point, Harriet Creek, and Redoubt Point Pe 5. Douglas Mountain (USGS); 6. Sitkinak Island (USGS); 7.
Kukak Bay assemblages (Knowlton, 1904; U nee 8. re i pcd (Wolfe in Lathram et al., 1965);
9. Sooke (LaMotte, 1935; GSC); 10. Gu n untain (UCMP); 11. Cascade Locks (USGS); 12. Sandstone
Creek (USGS); 13. Cascadia (UCMP); 14. end: M 1973); 15. eee (UCMP); 16. Rujada (Lakhanpal,
1958); 17. Yaquina (McClammer, 1978); 18. ap City (USGS); 19. Bridge Creek assemblages (Chaney, 1927;
OMSI, UCMP USGS). Early Neogene: 20. Nenana coal field assemblages (Wahrhaftig et al., 1969); 21. Lower
Cache Creek (Wolfe et al., 1966); 22. Houston (Wolfe et al., 1966); 23. Capps Glacier (Wolfe, 1966; USGS);

i i 25. € :

: ; 33. Clarki
(Smiley et al., 1975); 34. Orofino Creek (Brown, 1940); 35. Whitebird (Berry, 1934); 36. Eagle Creek assemblages
(Chaney, 1920); 37. Collawash and Fish Creek (USGS); 38. Little Butte Creek (USGS); 39. Mascall assemblages
(Chaney & Axelrod, 1959); 40. Baker (USGS); 41. Skull Spring (USGS); 42. Succor Creek assemblages (Graham,
1964, in part; cf. Fields, 1983); 43. '49 Camp (LaMotte, 1936); 44. Rabbit Hole (USGS); 45. Eastgate (Axelrod,
1985); 46. Middlegate (Axelrod, 1985); 47. Goldyke (UCMP); 48. Fingerrock (Wolfe, 1964); 49. San Antonio
(UCMP); 50. Thurston Ranch (UCMP).

796

inferred to have crossed the Rocky Mountains;
consistent with such crossings are the present
distributions of the related 4. brachypterum and
the descendant A. grandidentatum, both of which
live in the Rocky Mountains today. Thus some
probably very limited floristic interchange be-
tween eastern and western North America oc-
curred during the Neogene.

In the absence of actual assemblages, the flora
of the northern Rocky Mountain region during
the post-Eocene can be inferred only from as-
semblages outside the region. Except for floras
to the west and southwest on the Columbia Pla-
teaus, however, floras from adjacent regions are
few. To the north in central British Columbia,
the probable early Miocene assemblage from near
Hanceville on the Chilcotin River appears to rep-
resent a High Montane Mixed Coniferous forest.
In southern Colorado, the late Oligocene Creede
assemblage represents subalpine coniferous
vegetation. In Nebraska, the middle Miocene
Kilgore assemblages largely represent a meso-
thermal gallery forest (MacGinitie, 1962)

Taxa that occurred both in Mixed Northern
Hardwood forest of Alaska and in Mixed Me-
sophytic forest of the Columbia Plateaus repre-
sent taxa that (1) could exist under temperatures
inferred for the northern Rocky Mountain region
and (2) are known to have occurred adjacent to
the northern Rocky Mountains. Such taxa thus
can be reasonably inferred to have been in the
northern Rocky Mountain region. Included in
this category are lineages that are extant in the
northern Rocky Mountains: Alnus aff. incana, A.
aff. viridis, Betula aff. papyrifera, Populus aff.
trichocarpa, Salix aff. lasiandra, Prunus aff. vir-
giniana, Acer aff. negundo, and A. aff. macro-
phyllum. Also included are species of now extinct
lineages: Nordenskioldia ("Cocculus" auricula-

Carya bendirei, Pterocarya nigella, Acer scottiae,
A. septilobatum, A. tigilense, and Nyssa knowl-
tonii. Both lists contain primarily arcto-tertiary
genera, genera that are now disjunct between
eastern Asia and eastern North America or gen-
era that are now characteristic of forests of north
temperate regions (Engler, 1879). These genera
(and many subgeneric groups), however, were
represented in the Eocene upland microthermal
vegetation; their inferred presence in the north-

rn Rocky Mountain region is not necessarily the

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

result of southward migration from high lati-

=
(=

es.

Composition of the northern Rocky Mountain
coniferous element during the Oligocene and
Miocene must also be inferred. In the middle
Miocene of Nevada, the conifers are primarily
those that now inhabit Low Montane Mixed Co-
niferous forest (e.g., Abies aff. concolor, Picea aff.
breweriana, Pinus aff. monticola, P. aff. ponder-
osa, Tsuga aff. heterophylla, and Chamaecyparis
aff. nootkatensis). Most of these conifers were
also dominants of the Columbia Plateaus vege-
tation between 10 and 13 Ma (Fig. 4). In at least
one instance, the fossil species is a sister species
to the lineage that gave rise to the extant species
(Edwards, 1983), suggesting that the fossil may
have been adapted to a different environment
than the extant species.

In the Beringian Neogene, most coniferous lin-
eages were distinct from mid-latitude lineages.
The Abies belong to a group of which the extant
A. sibirica and A. grandis are members. Larix
was a diverse and major element in the Beringian
Neogene; no valid Larix is known during the
Miocene at middle latitudes of western North
America (H. E. Schorn, pers. comm., 1984). The
Beringian Picea either are related to extant Asian
species or represent the P. glauca group (includ-
ing an extinct, large-coned species that survived
into the Wisconsin glacial of southeastern North
America according to Critchfield, 1984). Thus,
the conifers (Abies grandis, Larix, and Picea
glauca) that are now the most distinctive element
of the northern Rocky Mountain region relative
to other regions of the western United States ap-
pear to be derived from a high-latitude source.

Precisely when the Beringian coniferous ele-
ment arrived in the northern Rocky Mountains
is unknown. I suggest, however, that the arrival
was probably post-Miocene. Coniferous lineages
allied to taxa that are now restricted to Low Mon-
tane Mixed Coniferous forest occupied the Co-
lumbia Plateaus and Nevada during the Neogene
and presumably represent ecotypes distinct from
extant relatives (Wolfe, 1964); such extinct eco-

Neogene, many of these northern
tain conifers possibly became extinct as the Be-
ringian lineages migrated south. Perhaps signif-
icant is that the first record of the coastal ecotype
of Abies grandis at middle latitudes is in the Plio-
cene-Pleistocene Sonoma assemblage of Cali-
fornia (Axelrod, 1944).

1987]

Various dicotyledons also may have penetrat-
ed southward from high latitudes during the
Neogene. All North American (including Be-
ringian) Eocene Salix represent subg. Salix. In
Beringia, subg. Vetrix appears in the late Oli-
gocene, approximately coincidental with the first
appearance of other taxa of Asian affinities;
members of sect. Glauceae appear in Beringia by
the early Miocene. Sect. Retusae of subg. Cham-
aetia appear by the end of the Miocene. Skvort-
sov (1968) suggested that subg. Vetrix and subg.
Chamaetia were of Asian origin, based on the
present distribution of species that have inferred
primitive morphologies. Although Asian fossil
data are lacking, certainly the Beringian data are
consistent with Skvortsov's interpretations. Dur-
ing the Neogene, various members of subg. Ve-
trix and sect. Glauceae underwent diversification
at high latitudes. A few lineages penetrated
southward in North America by the late early
Miocene and middle Miocene, but most lineages
of subg. Vetrix and subg. Chamaetia now extant
at middle latitudes of western North America
are probably late Neogene immigrants.

Representatives of Caprifoliaceae such as
Diervilla, Lonicera, and
significant and diverse elements i in the Beringian
Miocene, as were Ericaceae such as Rhododen-
dron (subg. Anthodendron — Azalea) and Vac-
cinium. These are generally absent in Neogene
floras at middle latitudes, although Symphori-
carpos is known in the late middle Miocene of
Nevada (Axelrod, 1956) and the late Miocene of
Idaho (Chaney & Axelrod, 1959). Presumably
most Caprifoliaceae and Ericaceae also represent
late Neogene immigrants into the northern Rocky
Mountain region.

Some extant northern Rocky Mountain Ro-
saceae (e.g., Amelanchier) appear to be derived
from lineages that have lived in or near the north-
ern Rocky Mountain region since the Eocene.
Other Rosaceae, however, probably belong to the
late Neogene Beringian element. Although Ru-
bus was represented in the Eocene upland mi-
crothermal vegetation, the extant R. idaeus has
a probable ancestor in the Beringian Miocene.
Similarly, Prunus subg. Padus has a continuous
record in western North America during the
Eocene and later, but subg. Prunophora first ap-
pears in western North America in Beringia dur-
ing the Miocene, and the extant Pacific North-
west Prunus subcordata appears to represent a
late Neogene immigrant from Beringia.

A pattern somewhat similar to that of Prunus

wUIC also

WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION

797

is indicated for A/nus. Members of subg. A/nus
were present in the northern Rocky Mountain
region as early as late Paleocene, and all the up-
land Eocene A/nus represent this subgenus. The
mid-latitude members of the subgenus differ-
entiated by the Oligocene into a lineage leading
to the extant A. oregona, and the lineage is clearly
recognizable in the early Miocene of Oregon.
High-latitude members of the subgenus are rec-
ognizable as belonging to the A. crispa lineage by
the Oligocene, and this lineage had penetrated
into middle latitudes by the middle Miocene.
Members of subg. A/nobetula are high-latitude
in distribution through the Miocene, and the en-
try of the extant A. viridis into middle latitudes
probably occurred during the late Neogene.

DISCUSSION

The terminal Cretaceous event selected for de-
ciduousness and created a mesothermal region
in the Northern Hemisphere almost devoid of
broad-leaved evergreens, allowing by default oc-
cupation of this region by many deciduous groups.
These groups then underwent major generic-level
diversification. The elimination (or nearly so) of
microthermal climates by the early Eocene ther-
mal maximum and the following creation of mi-
crothermal, mid-latitude uplands of the later
Eocene provided the opportunity for adaptive
radiation of these denuo mesothermal clades
into the new
opportunity for continuing diversification. Fun-
damentally, microthermal ecosystems became
extinct during the early Eocene thermal maxi-
mum and arose again de novo following the ther-
mal maximum. The continuing and major al-
terations in floristic composition during the
Eocene, the apparent high degree of community
instability during at least the early middle Eocene,
and the rapid diversification of groups such as
Rosaceae and Aceraceae during the Eocene can
be viewed as symptomatic of the evolution of
this new ecosystem. Insofar as known, the west-
ern North American volcanic uplands made up
the only major, mid-latitude upland region in
the Northern Hemisphere during the Eocene.
These volcanic uplands thus occupy a central
place in the diversification of many (if not most)
microthermal clades of arcto-tertiary type.

Primarily during the middle and late Eocene,
many microthermal, arcto-tertiary lineages were
able to disperse readily from North America into
Eurasia as adjuncts of a continuous expanse of
coniferous forests (Wolfe, 1985, fig. 10). Partic-

798

ANNALS OF THE MISSOURI BOTANICAL GARDEN

De FS To we

Sy,

Sy District

NORT

of
M
*6kenjj, ~

[Vor. 74

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$ 4 5 KA ud / AN UN 1 Tee
ol / EN el ` P
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rd 1840. 2»?! i d
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$ = MT x
Mrs ri P € 23 EN end
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1987]

ularly with the general cooling that characterized
the later Eocene (Savin, 1977), microthermal cli-
mates were found at increasingly lower latitudes
and produced a greater area available for occu-
pation by microthermal vegetation. As migrating
lineages elaborated over this area, diversification
would continue and some new divergences that
occurred in other regions would probably, in turn,
migrate into North America. Thus, even in the
Eocene, complexities were probably introduced
into historical patterns of a given lineage.

Histories of the component lineages now in
the flora of the northern Rocky Mountains are
varied.

(1) Some lineages can be traced back, more or
less continuously, into microthermal vege-
tation of the Eocene uplands (e.g., Betula pa-
pyrifera complex).

Some lineages have generic representation in
this upland vegetation, but divergences that
led to the extant species probably occurred
in the Oligocene or Neogene at middle lati-
tudes (e.g., A/nus oregona) and some at high
latitudes (e.g., A. viridis).

Some lineages represented in the Eocene up-
lands probably dispersed into Asia, under-
went major divergences into new subgenera,

~
N
—

~
o2
—

en-
tered the northern Rocky Mountain region
in the late Neogene (e.g., species of Salix
subg. Vetrix).

Some lineages represented in the Eocene up-
lands dispersed into Eurasia, underwent di-
versification at the sectional level and re-
turned to North America via long-distance
dispersal from Europe (e.g., Acer series Sac-
charodendron)

~
+
—

WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION

799

(5) Some lineages have long Beringian histories
and entered the northern Rocky Mountain
region during the late Neogene (e.g., Lonic-
era

(6) Another major pattern must be inferred for
many alpine tundra plants; Arctic tundra is
not recognizable until about the Pliocene-
Pleistocene boundary (Wolfe, 1985), and ele-
ments shared between Arctic tundra and al-
pine tundra ofthe northern Rocky Mountain
region may have entered the northern Rocky
Mountains during the Quaternary

(7) Another pattern must be inferred for plants
of subhumid to xeric, southern affinities

(Barnosky, 1984; Leopold & Denton, 1987);

some of these belong to families that did not

originate until about 25 Ma (Muller, 1981).

he Neogene spread of Compositae was rap-

id, probably in relation to the high degree of

dispersibility of their diaspores; however, the

spread appears to be associated also with the
development of dry climates.

Pollen indicates that Sarcobatus (Chenopo-

iaceae) was a member of mesic coniferous
forest during the later Miocene in areas such
as southern Idaho (Wolfe, 1969); adaptation
of Sarcobatus to xeric climate may be a late

Neogene phenomenon.

~
oo
—

No single pattern explains the origin of the
majority of the modern flora of the northern
Rocky Mountain region or even the majority of
the flora of a given vegetational type or zone in
the region. Certainly late Neogene cooling Aprap-
ably along with

mates in the northem Rocky Mountains fy
able to Beringian taxa, but these are intermixed
with taxa whose ancestry goes back to the Eocene
in the northern Rocky Mountains. Further, Be-

—

FIGURE 4.

Map showing location of some late Neogene (<13 Ma) plant assemblages in western North

America. For each assemblage or group of assemblages, either published references or repositories of largely

L. assemblages are cited; see Figure 1 for explanation of reposito

., 1971); 2. McCallum Creek (USGS); 3. Grubstake (Wahrhaftig et al.,

., 1966); 5. Chuitna River (Wolfe, 1966); 6. Tyonek (Wolfe et al., 1966); 7. Assemblages of type and referred
: " W. .

ries. 1. Hoogendoorn

1969); 4.

: gue Bay (USGS);

18. Hidden Lake (USG

nem Tipton id Vinegar Creek (Chaney, 1959; Chaney

cf. Fields, 1983): 27. Thorn Creek (Smith, 1941); 28. Trapper Creek (Axelrod,

1964); 29. Cache Valley ‘ark 1949): 30. Pit River (LaMotte, 1936).

800

ringian taxa entered at different times during the
Neogene and even Quaterna

Despite considerable resemblance at the ge-
neric level to the modern flora of the northern
Rocky Mountains, the microthermal upland
vegetation of the Eocene had many genera that
are now extinct and many genera that are now
exotic to the region. Microthermal vegetation of
western North America experienced major flo-
ristic alteration during the Eocene and later in
response to various extrinsic climatic factors and
intrinsic biotic factors such as competition.

The primary floristic result of the terminal
Eocene temperature deterioration for micro-
thermal vegetation was the elimination of many
dicotyledonous genera, particularly those con-
taining evergreens. Many deciduous dicotyle-
dons, however, also suffered extinction. For ex-
ample, numerous lineages in Acer became extinct,
including all North American members of sec-
tions Acer, Platanoidea, C ampestria, and Mac-
rantha (all of which survived in Eurasia). Pat-
terns of survival among decus taxa differed
markedly between Eurasia and North America,
probably due to intrinsic factors.

The relatively depauperate flora of the early

for radiation and

diversification, with : a general increase in diver-
sity and blurring of the regional Oligocene ge-
neric and subgeneric endemism, especially be-
tween western North America and eastern Asia;
for example, both Platanoidea and Macrantha
reappeared in western North America during the
Neogene. Migrations across Beringia were en-
hanced by the mid-Miocene warm interval, when
broad-leaved deciduous forests were probably
continuous from the Pacific Northwest across
Beringia and into P latitudes of eastern Asia
(Wolfe, 1985, fig. 1

Cooling at about n Ma (late middle Miocene),
concomitant with a general Neogene decline in
summer temperatures and decline in mean an-
nual ranges of temperature (Wolfe, 198 1a), elim-
inated many broad-leaved taxa of trees and shrubs
from the Columbia Plateaus, Beringia, and pre-
sumably the northern Rocky Mountains. Conif-
erous forests probably occupied a continuous belt
from the Columbia Plateaus and the northern
Rocky Mountains north into Beringia. Mixed
Northern Hardwood forest would have been to-
tally eliminated from western North America.
Climates in the northern Rocky Mountain region
during the Neogene may have initially exhibited
the trend toward lower mean annual ranges of

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

temperature. However, by the late Miocene, in-
crease in altitudes of mountains along the Pacific
Coast would have countered this trend, allowing
incursions of Arctic air masses; these masses are
inferred to have influenced eastern North Amer-
ica after 13 Ma (Wolfe, 1985).

During the Neogene a summer-dry precipita-
tion regime gradually encroached from the south
(Wolfe, 1978), and during the late Miocene (ca.
5-11 Ma) increase in altitude of the Cascade
Range created a significant rain shadow east of
the range (Chaney, 1944c; Smiley, 1963). At low-
er altitudes on the Columbia Plateaus, forests
were replaced by steppe vegetation during the
late Miocene (Leopold & Denton, 1987; Wolfe,
1985). An analogous replacement was occurring
in the northern Rocky Mountain region (Bar-
nosky, 1984).

Considering the complexities of climatic
changes and the complexities of orogenic activity
in the northern Rocky Mountain and nearby re-

ions, determination of historical biogeographic
patterns, without recourse to the fossil record of
extant lineages and analyses of successive fossil
biotas, can be highly problematic. As Grande
(1285) emphasized, vicanance methodology as

typically applied
complexities; several patterns can ine artis
on one another in such a region. Numerous pat-
terns have been detected in the modern flora of
the northern Rocky Mountains by recourse to
the fossil record, and different species within the
same genus can represent different patterns. Lin-
eages, some closely related, can be subjected to
the same extrinsic environmental factors and can
respond in different ways, a fact that further in-
creases the general complexity of the biogeo-
graphic tapestry in a region such as the northern
Rocky Mountains.

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Eocene, Wyoming). E. D. Dissertation. Yale Univ.
New Haven, Conne

. 1987 [1988]. Eocene and Oligocene floras and

vegetation of the Rocky Mountains. Ann. Mis-
souri Bot. Gard. 74: 748-784.

Wo tre, J. A. 1964. Miocene floras from Fingerrock

1987] WOLFE—ORIGINS OF NORTHERN ROCKY MODERN VEGETATION 803

Wash, a Nevada. U.S. Geol. Surv. Prof.
Pap. 454-N

1966. PUPA plants from the Cook Inlet
region, Alaska. U.S. Geol. Surv. Prof. Pap. 398-
B: B1-B32.

. 1969. Neogene floristic and vegetational his-
tory of the Pacific Northwest. Madroño 20: 83-
110.

1972. An interpretation of Alaskan Tertiary
floras. Pp. 201-233 in A. Graham (editor), Flo-
ristics and Puleofioristios of Asia and Eastern North
America. Elsevier Publ. Co., Amsterdam

. 1977. Paleogene floras from the Gulf of Alas-

tiary climates in the Northern Hemisphere. Amer
Sci. 66: 694—703.

Temperature parameters of humid to
mesic forests of eastern Asia and relation to forests
of other regions of the Northern Hemisphere and
Australasia. U.S. Geol. Surv. Prof. Pap. 1106: 1-
37

1981a. Paleoclimatic significance of the Oli-

botany, Paleoecology, n. Evolution, Volume
Praeger Publishers, New York.
1981b. Vicariance NEM of the an-

Vicariance oi s; A Critique. Columbia
Univ. Press k.

. 1985. a of major vegetational types
during the Tertiary. Pp. 357-375 in E. T. Sund-
quist & W. S. Broecker (editors), The “Carbon Cycle
and Atmospheric CO,: Natural Variations Ar-

chean to Present. Amer. Geophys. Union Geo-
phys. Monogr. 32, Washington, D.

1987. Late Creta aceous-Cenozoie history « of
deciduousness and the term
Paleobiology 13: 215-226

& R OORE. 1982. Tertiary marine and
nonmarine climatic trends. Pp. 154—158 in W.
Berger & J. C. Crowell (editors), Climate in Earth
History. Natl. Acad. Sci. Studies in Geophysics,
Washington, D.C.

& T. TANAI. 1980. The Miocene Seldovia
Point flora from the Kenai Group, Alaska. U.S.
Geol. Surv. c E 1105.
pores phylogeny, and
distribution A- p (maples) in the Cenozoic of
MM rth America. E Fac. Sci Hokkaido Imp.
., Ser. 4, Geol. 22: A

PCHURCH, ia 1986. Vegetation,
climatic and floral changes at Cretaceous-Ter-
tiary a Nature 324: -152.

& ————. 1987a. Leaf desee across the
C retaceous- Tertiary boundary in the Raton Basin,
New Mexico and Colorado. Proc. U.S. Natl. Acad.
Sci. 84: 5096-5100.
1987b. North American nonmarine

climates and vegetation during the Late Creta-
ceous. Palaeogeogr. Palaeoclimatol. Palaeoecol. 61:
33-78.

EHR. 1987. Middle Eocene dicotyle-
donous plants from Republic, northeastern Wash-
ington. U.S. Geol. Surv. Bull. 1597: 1-25.

D.M PKINS & E. B. LEOPOLD. 1966. Ter-
tiary serenade and paleobotany of the Cook
Inlet region, Alaska. U.S. Geol. Surv. Prof. Pap.
398-A: Al-

PRESENT-DAY VEGETATION IN THE NORTHERN
ROCKY MOUNTAINS!

JAMES R. HABECK?

ABSTRACT

The present-day northern Rocky Mountain vegetation 1s the product of a long history of geologic
ith

and climatic events that have interacted w

General concepts relating to the organization, classification, an

the species populations composing the regional flora.

nd dynamic nature of vegetation are

reviewed. Distributional and structural features of the vegetation cover between the Colorado Rockies
and the Southern Canadian Rockies are discussed. Alpine, upper timberline, subalpine, montane,
lower timberline, and grassland/steppe zones are treated. Climatic, physiographic, edaphic, and geo-

Ulltaill

vegetation. Iti is likely that members of the modern Rocky Mountain flora are not in Fani br Qui with

o
tion. Present

etati t pla re altere
structures and compositions that may represent new ecosystem equilibria acs could be irreversible

under present-day climates.

Vegetation ecology encompasses the descrip-
tion and interpretation of distribution patterns
exhibited by plants and plant communities oc-
cupying a given geographical area (Miles, 1979;
McIntosh, 1985; Walter, 1985). The objective of
this symposium is centered on an interpretation
ofthe evolutionary development ofthe Northern
Rocky Mountain (NRM) flora and vegetation,
including the earliest origins and ages evidenced.
This effort closely complements the reviews of
the origins of the cordilleran flora (Great Basin
and southern Rocky Mountains) provided by
Reveal's (1979) and Axelrod & Raven's (1985)
biogeographic treatments of the western inter-
mountain region, and Peet's (1988) detailed
analysis of structure, distribution, and produc-
tion of forests from throughout the Rocky Moun-
tains. My description of NRM vegetation serves
to orient readers to other symposium contribu-
tions that consider the roles of past events in
shaping this region's flora and vegetation.

THE NATURE OF VEGETATION

Literature on vegetation science reminds us of
the dynamic nature of floras and plant com-
munities (Knapp, 1974; Mueller-Dombois & El-
lensberg, 1974; Miles, 1979; Chabot & Mooney,
1985; McIntosh, 1985). Many botanists engaged

in the study of floras and plant communities agree
that vegetation units associated with any land-
scape arise over time as chance combinations or
assemblages of plant populations, with each
species following more or less independent or
individualistic ecological responses and migra-
tional pathways, as Gleason (1926) hypothe-
sized. Such ideas have been substantiated by
Whittaker (1953), Curtis (1959), McIntosh
(1985), and others over the past half century.

In his recent remarks addressing vegetation
history in the northern Great Basin and Pacific
Northwest, Mehringer (1985: 168) stated, “Plants
responded as vagaries of climate, dispersal po-
tential, competition, selection, soils, topography,
volcanic eruptions, fire, man and chance dictat-
ed. A hazy outline of these responses has been
traced through pictures painted with pollen ex-
tracted from yesterday’s mud. Sketchy as the im-
ages may be, they prove the dynamic nature of
this area’s vegetation.”

Floristic changes over time, whether gradual
or abrupt, underlie the dynamic nature of the
NRM vegetation. Post-Pleistocene pollen se-
quences in this region reveal “unquestionable
fluctuations in vegetation wrought by short, sharp
climatic episodes, by fire and by volcanic erup-
tions” (Mehringer, 1985: 185). Wells (1983),
analyzing wood rat middens throughout western

! Drs. Stephen Arno, Stephen Cooper and Robert Peet provided useful si bad pilus inam technical
vided

review of early stages of the manuscript; valuable suggestions

were also provi by tw onymous reviewers
d.

The contributions of Dr. Charles N. Miller, University of Montana, are also Rios
? Department of Botany, University of Montana, Missoula, Montana 59812, U.S.A.

ANN. MISSOURI Bor. GARD. 74: 804-840. 1987.

1987]

North America, provided descriptions and his-
torical documentations of the ee
ic shifts of western vegetation

Davis (1976, 1981) and mimi ne in their
analyses of forest vegetation stability during the
Quaternary, also addressed postglacial vegeta-
tion dynamics and suggested that eastern North
American plant populations are displaying dif-
ferential rates of range reoccupancy following the
Wisconsin glacial phase. Both Davis and Webb
urged us not to assume that an equilibrium exists
between modern plant distributions and the
ir climate. The NRM assuredly experi-
enced its own Pleistocene stresses and phytogeo-
graphic rearrangements, and it is likely that the
modern flora and vegetation in this area are not
in equilibrium with modern climate.

merous environmental variables have in-
fluenced the NRM flora and vegetation over time.
Lassoie et al. (1985), Smith (1985), and Peet
(1988) provided summary interpretations of the
ecophysiology of Pacific Northwest and Rocky
Mountain conifer forests. Factors such as moun-
tain building, differences in soil chemistry and
physical structure, precipitation and temperature
gradients, seasonal surpluses and water deficits,
evaporation potentials, and wildfire, all inter-
acting with each other and locally conditioned
by physiographic factors, create a multitude of
specific habitats for plant occupancy. Over time,
plant populations sort themselves out differen-
tially among available habitats. The spatial or
geographic pattern (local and regional range lim-
its) of each species is at least partially based on
adaptive features. The physiological studies cited
above emphasize how well the Rocky Mountain
conifer species are adapted and adjusted to their
environment.

Within the context of her eastern North Amer-
ican studies, Davis (1981) supported the view
that species with long histories of co-occurrences
in similar plant communities or habitats have
co-evolved adaptive adjustments to a. in-
terspecific competition. Davis warned, however,
that it is important for ecologists to AA
assess as to whether sufficient time has elapsed
for coevolution to have happened among closely
associated plant species. Often these sorts of as-
sessments represent research that must still be
undertaken on our western vegetation types.

Each plant population is unique in its degree
ofadaptive expression or amplitude of tolerance.
This uniqueness, founded in the genetic consti-
tution ofa species, is reflected in its rates of range

HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

805

extension or retreat, its present ecologic range
limits, and its specific habitat. Differential rates
of migration will affect vegetation distribution
patterns, as well as the presence or absence of a
species in a given vegetation association. The
dynamic nature of plant populations and plant
communities requires that ecologists recognize
that they are dealing with a vegetational phe-
nomenon that exhibits short-term stability but
lacks long-term permanency.

Rocky MOUNTAIN VEGETATION STUDIES

The earliest efforts to provide descriptive sum-
maries of Rocky Mountain vegetation date back
to the turn of the century. C. Hart Merriam (1889),
a prominent figure in the history of North Amer-
ican vegetation ecology, developed a system of
life zones that stressed the importance of tem-
perature in explaining vegetation patterns. He
popularized such life zone terms as Boreal, Tran-
sitional, Canadian, and Hudsonian. ui of
Merriam's scheme is used today. P Rydberg,
an early botanist associated with the New York
Botanical Garden, made major contributions to
our understanding of the phytogeography of the
Roc

Bulletin of the Torre
1913 and 1917). Rydberg's typical reporting for-
mat involved listing the flora of each mountain
zone with its geographic origin and limits.

J. irkwood, pioneer botanist at the Uni-
duit of Montana, published (1918, 1922) some
of the first in-depth ecological interpretations of
northern Rocky Mountain forest compositions
and distributions, applying to this region much
of the basic ecological insight promulgated by
Schimper (1903). Kirkwood provided a detailed
literature review of NRM vegetation studies
available at that time. He clearly emphasized
that vegetation distribution patterns would be
interpreted best when the “‘individualities” of the
“constituent species" were understood, an idea
that Gleason (1926) also advocated. Kirkwood
stated that “the complex interaction of climatic,
edaphic, biotic and other influences that guide
and control the distribution of species is of keen
interest to the student of phytogeography, and
no treatment of the subject is fair that does not
seek to give full weight to the many factors in-
volved and especially to the nature and require-
ments of the individual species" (1922: 15). He
also pointed out that Merriam's life zones often

ANNALS OF THE MISSOURI BOTANICAL GARDEN

110°
A ae A
“ey ake ZA 2 gs
“GE XZ
V
Z A
Z í >
wasife ye # &
ORE Z Z 1! 22 SES r
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1594. MT
445°
IDA
WYO
NEV [UTAH : CO
FIGURE l.

ug
cedar-hemlock-pine reste ae plieaie. Tsuga heterophy a and Pinus monticola).—
il).

[Vor. 74

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Kuchler’s K) n natural vegetation types shee with ionem, ois listed. — A. K-1
western ponderosa yes d astern ponder
ponderosa var. scopul

rum).—

s ponderosa
d ou

Douglas-fir (Abies grandis and Pseudotsuga menziesii

lacked spatial distinction (discontinuity) because
of elevational intermingling of the conifer dom-
inants in the nort

The many contributions made by R. F. Dau-
benmire to our knowledge of northern Rocky
Mountain vegetation must also be recognized. In
a career spanning nearly a half century, Dauben-
mire provided historical interpretations of the
origins of the Rocky Mountain flora, studied the
ecophysiological features of NRM plants, and
was instrumental in developing the habitat type

ern Rockies.

var. ponderos
las-fir forest usine uga menziesii bm pou —C.
D. K-14 = grand fir &

; K-16 =

sa pine forest n"

K-13 =

classification system widely employed in the

Rockies (1943,

1966, 1968, 1969, 1970, 1975,

1978, 1980, 1981). Daubenmire’s classic review

paper, “Veget

ation Zonation in the Rocky

Mountains’* (1943), summarized 175 ecological
studies that bear on the extant vegetation of the
Rocky Mountains. He reviewed floristic origins
and provided interpretations of relationships be-
tween vegetation zonation and mountain cli-

ates.
Following these pioneers, many recent efforts

1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 807

110° 110°
u +
B CANA N
š DA
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DUM. e EN >
WN m ad X
RS (60) S š
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p^ XE PF S s SER
° Aw SS 459. `
45 + as SS RC - MT
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` SR IDA
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WYO
NEV JUTAH A NEV JUTAH , CO
1109 1109
1109
u
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VEZ x &
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x. " E
NSS S
45° SS SN NN M < ia
NAGI Trg
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= S SHE WE SS
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= = WYO
ENSE
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NEV JUTAH n CO

Ficure 2. Kuchler vegetation types.—A. K-15 = western spruce-fir forest (Picea engelmannii and Abies
lasiocarpa). — B. K-50 = fescue-wheatg (Agrop picat d Festuca idahoensis); K-63 = foothills prairie
(Agropyron spicatum, Festuca idahoensis, F. scabrella, and Stipa comata).—C. K-51 = wheatgrass-bluegrass
(Agropyron spicatum, Festuca idahoensis, and Artemisia tridentata), K-55 = sagebrush steppe (Agropyron spi-
catum, Artemisia tridentata). — D. K-56 = wheatgrass-needlegrass-shrub steppe (Agropyron smithii, Artemisia
tridentata, and Stipa comata); K-64 = grama-needlegrass-wheatgrass (Agropyron smithii, Bouteloua gracilis, and

)

Stipa comata).

have been made by resource managers, plant community types most likely to express them-
ecologists, and geograr to describe, interpret, selves in the absence of human influence. Kuch-
map, and classify parts or all of the Rocky Moun- ler realized that the presence or absence of wild-
tain vegetation (Kuchler, 1964; Daubenmire & fire and/or grazing alters the specific nature of
Daubenmire, 1968; Pfisteretal., 1977; Peet, 1981, vegetation cover; this complicated his mapping
1 ; Arno & Hammerly, 1984; Smith, 1985). efforts. Figures 1, 2, and 3 show the general dis-
Kuchler (1964) published a map of what he tributions of Kuchler’s northern Rocky Moun-
termed the “potential natural vegetation" of the tain vegetation types.

coterminous United States. His map portrayed Pfister et al. (1977) and Pfister (1984) provided
the present-day vegetation in terms of those detailed literature reviews of the vegetation clas-

1

[Vor. 74

ANNALS OF THE MISSOURI BOTANICAL GARDEN

777777

NN

N

BZ
Z

110°
u
C CA N
NADA c
u C
WASH
ORE
459J. MT 4t
T2459
IDA 3 8 R3
S b.
=
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š š
ois 3” RE
> = p>
SERES
Il > > a
£ S s P) — STE wyo
RON > = ` a
SENS šs* S SS
.S8 AHg oS E S X
SO MSS = AS SS ju
| s ENS S SA? o
rer & ` IAJN xs
T S NE V UTAHSS a> SVS SS cC
110

Kuchler vegeta

vermiculatus). — B.
Greatbasin sagebrush (Arte
osteosperma, Pinus edulis (e

sification efforts in the R

misia tridentat

tion types.— A
n

ocky Mountains. Clas-

sifications are often individualized by each au-
thor’s philosophy and objectives, and the absence
of universally accepted concepts and principles

tion approach. Many R
tion classifications have

source management, map
storage, and vegetation preservation. These ap-
proaches often assume the existence of repeating
combinations or assemblages of plant popula-

been developed for re-
ping, information

. K-40 = saltbush-greasewood (Atriplex confertifoli
erosa pine-Douglas-fir (Pinus ponderosa and Pseudotsuga menziesii), K-38 =
a).—C. K-23 = juniper-pinyon woodland (Juniperus monosperma,
ast), P. monophylla (west), and Quercus spp.).—D. K-3
oak scrub (Cercocarpus ledifolius, Quercus gambelii, and Acer gran :

a and Sarcobatu

-37 = mountain mahogany-

ss
didentatum)
tions, in association with more or less specific

site qualities.
Pfister (1984) supplied a recent concise sum-

ese hav
categories called “habitat types": physical set-
tings definable in terms of the potential climax
vegetation they are capable of supporting (Dau-

benmire, 1966). Habitat type classifications cov-
er large parts of the Rocky Mountains. However,

1987]

th 1 245 ti affnrtch licited some crit-

ical commentary from other ecologists who have
studied in the same regions (Antos & Habeck,
1981; McCune & Antos, 1981; Baker, 1984;
Cloonan & Habeck, 1985; Crawford & Johnson,
1985; McCune & Allen, 1985a, 1985b).

Baker (1984), working with the Colorado Nat-
ural Heritage Inventory, maintained that vege-
tation classification in the Rocky Mountains is
handicapped by the absence of a standardized
nomenclature. He suggested, however, that no
unique classification may be possible because of
the vegetational variability, as noted above. Ba-
ker provided his own classification of Colorado’s
natural vegetation, a hierarchical listing of plant
community types. Peet (1981) successfully em-
ployed gradient analysis and ordination tech-
niques to the Colorado Front Range vegetation
and derived a workable classification of forests
in that area. An outline of Peet’s results will be
presented later.

My literature evaluation revealed that regional
climatic patterns, physiographic provinces, ele-
vation zonations, soil features, topographic-
moisture gradients, habitat types (potential cli-
max), seasonal moisture regimes, drought stress
categories, plant life forms, community life forms,
ecological interactions between dominants, as
well as human intuition, have all been used to
design classifications of Rocky Mountain vege-
tation. Furthermore, since the natural ecologic
role of wildfire in the Rockies has become better
understood (Habeck & Mutch, 1973; Arno, 1980;
1983; Habeck, 1995), some recent

T disturbed plant community types (Arno et

1., 1985), as well as the multiplicity of seral vege-
ee types created by human activities. Suc-
cessful fire suppression during a century of graz-
ing and crop production has significantly altered
the natural plant cover in the NRM. Fire-de-
pendent vegetation types in the Rockies, espe-
cially, have changed drastically in the past cen-
tury (Gruell, 1983).

Weaver & Dale (1974), Pfister et al. (1977),
Arno (1979), Weaver (1979, 1980), Johnson &
Pfister (1981, 1982), Peet (1981, 1988), Steele et
al. (1981, 1983), and Arno & Hammerly (1984)
contributed recent summaries of the climatic and
physiographic features of the Rocky Mountains
and detailed descriptive interpretations of the
existing vegetation covering this region. These
publications serve as my primary references; they
are based on extensive field studies and in-depth

HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

809

literature reviews covering eastern Washington
and northern Idaho, northeastern Oregon, west-
ern and central Montana, central and eastern
Idaho, Wyoming, the Yellowstone Park area, and
parts of Utah and Colorado.

rno & Hammerly (1984) described and in-
terpreted the lower (warm) timberline and upper

mountainous regions of western No
with supplementary comments on vegetation
types adjacent to or between timberlines. John-
son & Pfister's studies (1981, 1982) centered on
identification and description of naturally oc-
curring geologic and ecologic units in the middle
and northern Rocky Mountains.

Smith (1985) made an effort to provide an
ecophysiological explanation for the distribution
patterns within western montane forests. Var-
ious factors—including moisture, solar radia-
tion, and temperature — exhibiting seasonal vari-
ations within the western plex and
interactive with soil structure and nutrient con-
tent, form the environmental complex that de-
fines plant community distributions and succes-
sional patterns. Smith emphasized the importance
of “biophysical coupling" of site factors, fire, and
various disturbances in dictating successional

mountain

needed before more com
vegetation patterns can be made (Peet, 1988).
Future studies should also emphasize the long-
term importance of human disturbances.

THE ROCKY MOUNTAIN SETTING:
PHYTOGEOGRAPHIC SUBREGIONS

The region typically designated as the northern
Rocky Mountains extends from the Snake River
Plain in southern Idaho to the international bor-
der, joining with the southern Soe ee Rockies
in British Columbia and Alberta. This region
averages 500 km in width, Denice from near
Yellowstone Park (northwestern Wyoming)
westward to Hell’s Canyon on the Ida ho Olan
state lines. Northward, the principal region ex-
tends from central Montana westward to north-
eastern Washington.

Peet's (1988) description and interpretation of
the forest vegetation of the entire Rocky Moun-
tain system recognized the existence of four flo-
ristic regions: far-northern, northern, southern,
and madrean. My review encompasses all of the
northern and part of the southern regions. The
latter closely coincides with Daubenmire’s (1943)

810 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

YP
»\® aa ae C (25) s wo

FIGURE 4. Northern Rocky Mountains with oe P regions (after Arno & Hammerly, 1984;
see text): A—Southern Continental Ranges; B— Northern Continental Ranges; C — Intermountain Ran ges; D—
Inland Maritime Ranges; E—Southern Canadian deris Mountains; F — Middle Rocky Mountains. The Con-
tinental Divide is shown. as a dashed line. States, provinces, Ves national parks, m
features are shown by number and letter symbols keyed as follows: 1 — Canadian “Great Divide" (BC, ALB);
2 — Purcell Mountains (BC, MT); 3—Selkirk Mountains (BC, I MU gie inicium Mountains (BC); 5 — Whitefish

Í A, WYO, ; 8—Swan Range

(MT); 9—Clearwater Mountains (IDA); 10— Bitterroot Mountains (MT, IDA); 11 — Sapphire Mountains (MT);

12— Anaconda-Pintler Range (MT); 13— Little Belt and Big Belt Mountains (MT); 14— a dig Range (MT);

15— Madison Range (MT); 16— Blue Mountains (ORE); 17— Wallowa Mountains (ORE); 18— Idaho Batholith
a :

S ; d
Teton Park (WYO); 27— Wind River Range (WYO); 28— Salt River & Wyoming Ranges ); 29— Wyoming
Basin & Great Divide Basin (WYO); 30— Wasatch Mountains (UTAH); 31 — Uinta Mountains (UTAH); 32—

1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 811

NIVAL ZONE Glaciers and S

owfie
Rock Outcrops & jode m

ALPINE ZONE Alpine Tundra; Meadow ens
(Cold) relifiends, ret bgt ps des ges E
edge Communi s (Treeless) 2700m « UPPER TIMBERLINE
Ipine 1 ;
Whitebark pine - Pinus albicaulis
Mountain hemlock - Tsuga mertenstana - IM b Lodgepole pine (seral)
n d Pinus contorta

SUBALP INE
ZONE

annii
(Cool, Molst) Limber pine - Pinus flexilt 1800-2700m
Bougies tir - Pseudotsuga menziesii Western larch (seral)
ah Lariz oc

and iud - Abies grandis - ctidentalis
MONTANE ZONE ee redcedar - Thuja pl

Mice hemlock - Tsuga heterophylla - IM
Western yew - Taxus brevifolia -

(Temperate-Mesophytic) are apen - Populus tremulo aida vestern seine. = IM
P :
r blrch - Betula papyrifen ($05 BO inus monticola (seral)
Ponderosa pine - Pinus pondero + varieties)
DRY CONIFER FOREST Limber pine - Pinus flexilis oe Hel vpe)
reri pele Juniper - Juniperus
WOODLAND ZONE Bitterbrush - Purshia tridenta ta «

Fidi poids - Cen Ledifoli LOWER TIMBERLINE

(Temperate-Xerophytic) i iia lanl iia 800-1500m

_ Bluebunch wheatgrass - Agropyron spic

SEMI-ARID STEPPE. AND di^ e - Festuca s and ae pene
GRASSLAND ZONE Needlegrass - Sttpa coma

Big sagebrush - Artemisia VT REN (+ varieties)

(Warm, Dry) 1000m
RIPARIAN/FLOODPLAIN Black cottonwood - Populus Dee (* other specles)
QuakIng aspen - Populus tremuloides
Ist) pe ied cae - Betula oaotdentatiai Paper birch - B. papyrifera
(Warm, Molst Alde Willow specie Alnus spp & Salix spp

FiGURE 5. Vegetation zones in the northern Rocky Mountains, typical of Montana, northern Idaho, and
adjacent parts of British Columbia and Alberta. Approximate elevational limits of these zones are given, as are
average positions of the lower and upper timberlines. IM = restricted to Inland Maritime region

"central Rockies," and in my review will be re- as the Middle Rocky Mountains (MRM) is lo-
ferred to as the “middle Rockies,” after Arno & cated south of the Yellowstone River in Montana
Hammerly (1984). Figure 4 shows the geographic and extends southward through Wyoming to parts
units employed in this review of Utah and Colorado; it includes the Uinta and
The Southern Canadian Rocky Mountains Wasatch mountains, Medicine Bow Mountains,
(SCRM), which extend from the international and the Colorado Front Range (Fig. 4). Both the
boundary northward, are closely related physio- SCRM and MRM will be discussed, but empha-
graphically and floristically to the NRM in the sis will be placed on the traditional NRM prov-
United States, as defined above (Rowe, 1959; ince.
Arno & Hammerly, 1984). The region designated The importance of elevation and topographic-

<
Medicine Bow Range (WYO, CO); 33—Park Range (CO); 34—Snake ua Plain (IDA); 35 — Steens Mountains
(ORE); 36— Northeastern Great Basin (NEV); 37 — Great Basin (UTAH); 38— Juniper-Pinyon Woodland (UTAH);
39— Wyoming Basin Sagebrush- oe (WYO); 40— Plains Grasslands on 4] — Palouse Prairie (WASH, IDA);
42—Sagebrush Steppe (WASH). N-BC— Nelson, British Columbia; K-BC— Kimberley, British Columbia; J-ALB—

R-IDA— Riggins, Idaho; S-IDA —Salmon, Idaho; B-IDA — Boise, Idaho; IF-IDA — Idaho Falls, Idaho; P-IDA—
Pocatello, Idaho; TF-IDA — Twin Falls, Idaho; GP-MT —Glacier National Park, Montana; K-MT — Kalispell,
Montana; M-MT-— Missoula, Montana; G-MT—Great Falls, Montana; H-MT—Helena, Montana; B-MT—
(west) — Butte, Montana; B-MT (east)— Billings, Montana; D-MT —Dillon, Montana; YP- WYO- Yellowstone
National Park, Wyoming; C-WYO (north)— Cody, Wyoming; C-WYO (south)— Casper, Wyoming; T-WYO—

Thermopolis, Wyoming; RS-WYO-— Rock E Wyoming; R-WYO— Rawlins, Wyoming; L-WYO-— Lara-
mie, Wyoming; E-NEV —Elko, Nevada; O-UTAH —Ogden, Utah; SLC-UTAH - Salt Lake City, Utah;
V-UTAH — Vernal, Utah; C-CO — Craig, Colorado; en Rocky Mountain National Park, Colorado; B-CO—
Boulder, Colorado.

812

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

NIVAL ZONE

Ice, Snow, Rocks

> 4200m

Sedge/Gra

ALPINE ZONE
(Cold)

lmann spruce -

SUBALP INE

ristlecone pine

(Cool/Mesic)

Wnitebark pine - Pinus albicaulis (N)
- Pinus arista

s Meadows, Fellfields,
De A Tussock, Dl a cla Fields)
edge marshes > 3500m

Q@urrer TIMBERLINE
a engelmannii

bien Lice saya

a (s)
2800m(N)/3500m(S)

rosa pine
MONTANE

FOREST ZONE

(Warm, Mesic)

- Pinus ponderosa

Blue spruce - Picea pungena :

(E E) Open Parklands
w)

t
eei bal
te Moist sites/Canyo
22 pene Sonar

cles oak -

nipers -
FOOTHILLS/STEPPE-
Pinyon

SHRUBLAND ZONE

(Warm, Dry)

“urus gambelii
unipe rus osteosperma,

communis,
pines - Pinus edulis (E);
g

untain Maho ig - age iai us ledt foLiu tus;
ea Greasewood - Sar

«^n

J. monosperma, J. TIMBERLINE

J. horizontalis, J. scopulorum

Pinus monophylla (w)
tum: Lower slopes/Canyons

- Artemista greeny
Shadscale - Atriplex
cobatus vermiculatus

virginiana; Sagebr

1600m-2100m

Grama/Buffalo gra

PLAINS/GRASSLAND ZONE
(Hot, Dry)

grass -
Bluebunch wheatgrass - Agropyron pis uM Bluegrasses - Poa species
«1600

- Bouteloua i dactyloides

Cottonwoods - Popu

Bi rches

RIPARIAN/FLOODPLAIN ZONE
(Warm, Moist)

ulus species; Mill
- Betula species; Alders - Aad i spec

- Saliz species
cles

FiGuRE 6. Vegetation zones typical of the middle Rocky Mountains. Major dominants in each life zone are

iste

moisture gradients in determining Rocky Moun-
tain forest distribution patterns was emphasized
by Peet (1988). He provided a series of “gradient
mosaic diagrams” which portray the latitudinal
shifts in forest composition along the entire length
of the Rocky Mountains. Soil constitutes a third
environmental variable important in interpret-
ing Rocky Mountain vegetation patterns (Weav-
er, 1978; Peet, 1

Figures 5 and 6 show generalized vegetation
zonations in the northern and middle Rockies.
Daubenmire (1980) also documented that NRM
vegetation closely follows environmental gradi-
ents from warm-and-dry to cold-and-wet, re-
gardless of altitude. He stated that elevation above
sea level is of reduced significance in the NRM
because of the interdigitation of vegetation types
within the mountain topography.

Within the NRM, there are significant climatic
gradients which play major roles in modern vege-
tation distribution patterns. In the northwestern
sector of this region (which includ theast
Washington, northern Idaho, and northwestern
Montana to the west slope of the Continental
Divide in Glacier Park), there exists a moist in-

land-maritime zone that features a well-devel-
oped oceanic influence (Kirkwood, 1922; Larsen,
1930; Daubenmire, 1943; McMinn, 1952;
Weaver, 1979, 1980; Lassoie et al., 1985; Peet,
1988). Here the NRM exhibits nearly continuous
forest cover composed of several conifers with
Pacific coastal affinities. These include Thuja pli-
cata, Tsuga heterophylla, T. mertensiana. Taxus
brevifolia, Abies grandis, and Pinus monticola,
as well as coastal shrubs, herbs, and nonvascular
plants (McCune, 1984). The lower slopes and
valleys are also heavily forested, without a well-
defined lower timberline.

As the maritime influence diminishes, much
of the remaining NRM region experiences a sig-
nificantly colder and drier continental climate.
The mountain peaks of this region are also much
higher than those of the inland maritime region,

vegetation
types are caused by the much warmer and drier
continental conditions. Here Pseudotsuga men-
ziesii var. glauca, Pinus ponderosa var. ponder-

1987]

osa, P. ponderosa var. scopulorum, Picea engel-
mannii (and hybrids between P. enge/mannii and
P. glauca), Abies lasiocarpa, Pinus contorta var.
latifolia, P. flexilis, and P. albicaulis are the char-
acteristic tree species.
rno & Hammerly (1984) established several

NRM subregions based on geographic and bio-
climatic criteria. Each of these includes several
mountain ranges (Fig. 4): A. Southern Continen-
tal ranges, B. Northern Continental ranges, C.
Intermountain ranges, and D. Inland Maritime
ranges. The SCRM and MRM ranges were dis-
cussed separately. In their reviews, Johnson &
Pfister (1981, 1982) made use of ‘ ‘physiographic
provinces," as well as forest “climax series" sim-
ilar to those used in the habitat type classifica-
tions cited above

A very comprehensive biophysical land clas-
sification for the southern Canadian Rockies, in-
cluding an in-depth treatment of the be a
is provided by Corns & Achuff (1982). T
study focused on Banff and Jasper National en
but has general usefulness for the SCRM. They
established a series of physiognomic classes: A.
Closed Forest, B. Open Forest, C. Shrub, D. Low
Shrub-Herb, and E. Herb Dwarf-Shrub. Corns
& Achuff used these vegetation types, combined
with climatic and soil factors, to define “‘ecore-
gions” in Banff and Jasper National Parks.

VEGETATION OF THE NORTHERN
Rocky MOUNTAINS

The following description of the present-day
NRM vegetation will make use of the geograph-
ic-biophysical subregions established by Arno &
Hammerly (1984). The information will empha-
size the forested zones as depicted in Figure 5,
with supplementary material on associated non-
forest types.

SOUTHERN CONTINENTAL MOUNTAINS

The southern continental mountains include
those found in eastcentral Idaho and southwest-
ern Montana (Fig. 4, Region A). The Sawtooth
Mountains, Lost River Range, Lemhi Range,
Beaverhead Range, Anaconda-Pintler Range,
Gravelly Range, and Madison Range are includ-
ed. This region is dry and cool; the low moisture
(20-40 cm annually) is due to a rainshadow effect
caused by Oregon's Blue and Wallowa moun-
tains and other Idaho ranges. Steppe and grass-
land types are present on lower slopes and val-

HABECK — NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

813

leys, dominated by several species of Artemisia
and bunchgrasses (mostly Agropyron spicatum
and Festuca idahoensis). The forest cover is con-
fined to a relatively narrow zone between the
lower timberline at 2,000 m and the upper tim-
berline at about 2,700 m

Pinus ponderosa is not well represented in the
lowest forested zones of this part of the NRM
because of the cold climate and short growing
season at lower timberline (Arno & Hammerly,
1984). Instead, Pseudotsuga menziesii is the more
common dominant tree adjacent to the steppe
zone, sometimes mixing with Pinus flexilis at or
near the lower timberline (1,800-2,300 m). In
dry mountain ranges (Lost River Range),
krummholz forms of Pinus flexilis and Picea en-
gelmannii define the upper free line at ^ diis m,
while on r
tooth Range, Beavethead Range, and Madison
Range) Abies lasiocarpa and Pinus albicaulis join
Picea (Arno & Hammerly, 1984). Some southern
continental mountain ranges have such severe
moisture shortages that sagebrush-grassland
vegetation extends up to and through the sub-
alpine zone.

Forest understories in the southern continental
region are low in species richness. Semiarid steppe
and grasslands dominated by Festuca idahoensis,
Agropyron spicatum, Stipa comata, and Arte-
misia tridentata form a mosaic among the Pseu-
dotsuga menziesii forests. At higher elevations,

Vaccinium globulare, Arnica cordifolia, sonia
betulifolia, and Acer glabrum are common
understory associates. Some of these shrubs and
herbs extend into the subalpine zone where they
are joined by Vaccinium scoparium, Linnaea bo-
realis, Smilacina stellata, and Arnica latifolia.

According to Dunwiddie (1977), Young & Ev-
ans (1981), Arno & Gruell (1983), Arno & Ham-
merly (1984), and Butler (1986), fire suppression
since 1900, climatic changes, and livestock graz-
ing have contributed to conifer invasions into
the steppe-grassland zones at lower and middle
elevations throughout the Rocky Mountains. Be-
fore 1900, fires swept through these mountain
ranges at intervals of 20-30 years, killing many
young conifers that were establishing themselves
in grasslands and shrublands. Intensive and ex-
tensive grazing removes or reduces competing
grasses, further facilitating spatial shifts in the
forest-grassland boundaries.

The southern continental region also supports

814

woodlands composed mostly of Juniperus scopu-

orum. This vegetation type is found on dry, rocky
sites, including outcroppings, generally below the
limits of Pseudotsuga menziesii. A common as-
sociate of juniper is Cercocarpus ledifolius. Pres-
ent but much less common is Juniperus osteo-
sperma, which occupies very severe sites and may
be found mixed with Pinus flexilis. Sagebrush-
steppe, dominated by Agropyron spicatum, Ar-
temisia tridentata, Festuca idahoensis, and Pur-
shia tridentata, occurs throughout Region A in
Idaho and Montana (Kuchler, 1964).

NORTHERN CONTINENTAL MOUNTAINS

The northern continental mountains consist of
the central Montana mountain ranges east of the
Continental Divide (including the Lewis Range
and “Rocky Mountain Front”), from the vicinity
of Helena and Harlowton (Big Belt Mountains,
Crazy Mountains, and Little Belt Mountains)
northward to the southern Canadian Rockies in
Alberta (Fig. 4, Region B). The isolated Bear Paw
Mountains and Sweetgrass Hills in northcentral
Montana and the Cypress Hills in southern Al-
berta can be included in this geographic subunit,
although they are located 150 km or more east
of the cordilleran Rocky Mountains.

The valley elevations here are lower than in
the southern continental mountains but the
northern continental ranges experience some-
what less moisture stress (Arno & Hammerly,
1984). This subregion receives 25-30 cm of pre-
cipitation annually, causing the lower timberline
zone to be expressed at 1,200-1,500 m; contin-
uous forest cover occupies a broader elevational
zone. Pinus ponderosa forms the lower timber-
line in parts of central Montana ranges (Arno,
1979) but plays a lesser role north of 47°. Pacific
coastal understory species (see Inland Maritime
Region) are uncommon in central Montana for-
ests but Great Plains species are present, includ-
ing Bouteloua gracilis, B. curtipendula, Andro-
pogon spp., Opuntia fragilis, and Yucca glauca.

Pinus ponderosa is replaced by combinations
of Populus tremuloides, Pseudotsuga menziesii,
Pinus flexilis, and Pinus contorta north of Great
Falls, Montana. Cold winter climates with fluc-
tuating temperatures and 115 kph winds seem
to exceed the tolerance of Pinus ponderosa. Pop-
ulus tremuloides and associated conifers that oc-
cupy this area are often deformed and/or stunt-
ed by the cold winter winds. Good examples of
these effects can be seen on the east side of Gla-
cier National Park. With the reduction of fire,

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Pinus flexilis woodlands have extended into the
prairie communities in this same region.

The montane zone, occupying the middle el-
evations of these central Montana mountains
(1,500-1,800 m), is dominated by Pseudotsuga
menziesii and Pinus contorta in varying amounts.
Picea engelmannii is a dominant in areas tran-
sitional to the subalpine zone, where it is codom-
inant with Abies lasiocarpa and Pinus albicaulis.
The upper timberline (tree limit) occurs between
2,300-2,600 m, with some terrain above 2,400
m supporting alpine tundra.

Associated undergrowth species near the lower
timberline in the northern continental moun-
tains include Agropyron spicatum, Festuca ida-
2 F. oe Bouteloua gracilis, and Sti-

a richardsonii. Symphoricarpos albus, Spiraea
pins pt stellata, Linnaea borealis,
Galium triflorum, Menziesia ferruginea, Vaccin-
ium globulare, and V. scoparium dominate the
understories of higher forest zones

The disjunct Sweetgrass Hills (2,100 m) in
northcentral Montana, although completely sur-
rounded by semiarid Great Plains grasslands and
foothills prairie, support forests composed of
many of the montane and subalpine conifer
species listed above, as well as the hybrid spruce
Picea glauca x engelmannii (Habeck &
Weaver, 1969). According to Thompson & Kuijt
(1976), the Sweetgrass Hills were surrounded but
not covered by the Laurentide glaciers; the hill-
top nunataks may have supported tundra vege-
tation. Later, the Hills were part of a more con-
tinuously forested region in early post-Wisconsin,
but climatic changes isolated the forests from the
Rockies. The post-Hypsithermal shift towards
increased warmth and dryness appears to dis-
favor maintenance of the present-day subalpine
understory species in the Sweetgrass Hills.

INTERMOUNTAIN RANGES

The Intermountain Ranges are found in north-
eastern Oregon, central Idaho, and westcentral
Montana (Fig. 4, Region C). The Blue and Wal-
owa mountains are in Oregon; the Clearwater
Mountains, Salmon River Mountains, and the
western edge of the Bitterroot Mountains are in
Idaho; the Sapphire Range, Anaconda-Pintler
Range (border between Regions A and C), Flint
Creek Range, and the southern parts of the Mis-
sion and Swan ranges are located in westcentral
Montana. The climate is partially influenced by
moist maritime air masses that pass nearby, but
this region receives less total moisture than does

HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

815

FIGURE 7.

the more western Inland Maritime region. On
the other hand, the more severe continental cli-
mate that features temperature extremes, cold
dry winter, and stressful chinook winds is not as
common in this region (Arno & Hammerly,
1984).

The Intermountain Ranges have well-defined
lower and upper timberlines, and the valley bot-
toms (900-1,050 m) and southern aspects sup-
port bunchgrass prairies (Agropyron and Festuca

shrub types were originally interfacing or inter-
spersed with Pinus ponderosa, forming the low-
est forest zone. In presettlement times, the exact
position of the lower timberline was partially
conditioned by wildfires caused by lightning and
aboriginal man (Gruell, 1983).

Frequent low intensity ground fires perpetu-
ated open Pinus ponderosa savannas (Fig. 7), even

Pinus ponderosa forest in Intermountain region, western Montana. Located near Darby, Montana,
at 1,050 m within the Pseudotsuga menziesii zone. This forest type historically was maintained as open savannas
by ground fires. USDA Forest Servic

on mesic sites where Pseudotsuga menziesii and
Abies g Mutch,
1973; Thak. 1976, 1985; Barrett & Arno, 1982;
Freedman & Habeck, 1984). My own analysis of
fire scars on 300- to 350-year-old Pinus ponder-
osa specimens occupying a drainage near Mis-

d 1700s, fro

1750 to 1870 the fire frequency doubled! That
is, intervals between fires were reduced to an
average of five years. Indians practiced routine
burning throughout the Rocky Mountains before
their encounter with Euroamerican influences
(Barrett & Arno, 1982; Gruell, 1985; Arno, 1985;
Lewis, 1985) but may have started more fires
following introduction of horses into western
Montana in the 1730s. Such frequent burning
maintained even moist Pseudotsuga forest sites
as grassland savannas. Remnants of the savanna
bunchgrass, Festuca scabrella, still exist beneath
the Douglas-fir forest canopies that have devel-
oped without fire since 1900.

Above the Pinus ponderosa zone lies the Pseu-

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

< p

ae E- `

Ses
oar Old- ae Larix occidentalis-dominated forest in Inland Maritime region in Flathead National
00 m). Established M a wildfire, the Larix 1s being replaced by the climax species,

For a (1,1
Palas menziesii. USDA Forest Servic

1987]

>
"g^ " gn gef
^ t Yy t w

E9. Subalpine zone exhibiting unev

HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

817

-distributed Abies lasiocarpa. Pinus albicaulis and P. contorta

near ‘Wahoo Pass (2,000 m) in Susa Bitter Wilderness, Montana/Idaho state line. USDA Forest Service.

dotsuga menziesii forest zone, extending from
1,200 to 1,800 m. Within this zone, Larix oc-
cidentalis prevails as a major pioneer species (Fig.
8). This imi dependent species may
achieve diameters of 1—1.5 m (thick basal bark
layers that sauer fire protection contribute to
large girths) and heights over 40 m. Another
common but short-lived associate in the upper
part of the Pseudotsuga menziesii zone is Pinus
contorta; it is also dependent in the NRM on
periodic fires or other site disturbances (Lotan &
Perry, 1983; Muir & Lotan, 198
Common understory species in the Douglas-
fir/western larch forests include Physocarpus
malvaceus, Acer glabrum, Vaccinium globulare,
V. caespitosum, Calamagrostis rubescens, Carex
geyeri, Spiraea betulifolia, Arctostaphylos uva-
rsi, Mahonia repens, Linnaea borealis, and
Symphoricarpos albus.
t from 1,800
m to treeline near 2,700 m (Fig. 9). Pinus con-
torta extends into this zone and mixes with Pinus
albicaulis and Picea engelmannii. Again, fire his-

torically occurred in these higher forest zones
(Fig. 10) but at long intervals, usually more than
100 years and sometimes at 200- to 300-year
intervals (Habeck, 1985)

Dominant understory species associated with
Intermountain subalpine forests include Men-
ziesia ferruginea, Vaccinium globulare, V. sco-
parium, Xerophyllum tenax, Carex geyeri, Ar-
nica cordifolia, A. latifolia, Calamagrostis
rubescens, C. canadensis, Luzula hitchcockii,
Clintonia uniflora, Tiarella trifoliata, Anemone
piperi, and Linnaea borealis.

Insects and other pathogens are known to have
caused periodic ecosystem disruptions in the In-
termountain forests. McCune (1983), however,
has suggested that man’s reduction of wildfires
Montana has predisposed the forests

1982; Anderson, 1985). Their data suggest that

ANNALS OF THE MISSOURI BOTANICAL GARDEN

x
ee
aR

; he

$

|
uy.

Pn ^

A.

^. Wed

“``

suce à UNS p A EE
AR pote Es. Sent
wd. ` -
stel. 2

LE TL

ju det S
—

* % E

NE K (Aes

xt Y os i Z

a af xx à *j
de ee uc M ox, Mors

MEL ge
vere

NN UM
M. à

* n « = ] 4

Melo

FIGUR

fire suppression and logging practices in the Pseu-
dotsuga menziesii forests in western Montana
create conditions that encourage outbreaks of
western spruce budworm (Choristoneura occi-
dentalis) of greater severity and longevity than
those experienced prior to fire control. Without
fire, forest cover has become more structurally

te, "Tr
A "E el y ` 4
- 5 - = « A rdi °
Be n 224. " y " Doi 3

[Vor. 74

on €"

URE 10. Fire-generated, even-aged Pinus contorta in the subalpine zone (2,500 m) in northern Rocky
Mountains region. USDA Forest Service.

uniform, which facilitates insect spread over large
reas.

The upper timberline in the Intermountain
Ranges forms at about 2,700 m and is dominated
by Pinus albicaulis, Abies lasiocarpa, Picea en-
gelmannii and, not infrequently, Larix lyallii.
Each of these exhibits a wind timber develop-

1987]

HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

Ta
Ze.

FIGURE 11.
USDA Forest Service.

ment (dwarfed in size and shape) as each reaches
its individual upper limit of growth. Larix lyallii
is confined to relatively harsh north slopes over
2,100 m and frequently occupies talus slides and
boulder fields. Only occasionally does Larix lyal-
lii grow on level sites with developed soil (Arno
& Habeck, 1972).

Near timberline, Pinus albicaulis, Picea en-
gelmannii, and Larix lyallii form open-grown,
parklike stands. Topography, wind, and snow
deposition paths cause a variety of interesting
spatial arrangements in the timberline zone, such

s "ribbon-forests" and krummholz mats (Bil-
lings, 1969; Fig. 11). Some individual timberline
specimens of Pinus albicaulis, Larix lyallii, and
Picea engelmannii are relatively large (1.0-1.
m in diameter) and old (300 to 500 years). PAyl-
lodoce empetriformis, P. glanduliflora, Cassiope
mertensiana, Vaccinium scoparium, Xerophyl-
lum tenax, and various sedges (Carex spp.) and
rushes (Luzula spp.) are the understory domi-
nants in these timberline forests.

Some bunchgrass communities (Agropyron

Wind-shaped Pinus albicaulis within upper ti

zone (2,500 m) in southcentral Montana.

spicatum, Festuca scabrella, and F. idahoensis),
compositionally similar to lower timberline
grasslands, exist islandlike within the Inter-
mountain subalpine forest zone well above 1,800
m. They are believed to be the results of hyp-
sithermal displacements (Daubenmire, 1968,
1975, 1981; Root & Habeck, 1972) and are pres-
ently maintained by localized xeric conditions
induced by slope, aspect, shallow soils, or un-
weathered parent rock substrates. Although sub-
ject to burning, there is no evidence that fire is
required to perpetuate them. Other grasslands,
or *grass balds," featuring Festuca viridula com-
munities may dominate the subalpine zone in
central Idaho and northeastern Oregon, extend-
ing to over 2,40

Detailed donodan and classifications of the
lower zone Intermountain grasslands (Fig. 12)
have been presented by Mueggler & Stewart
(1980). Some of their 30 grassland and shrubland
habitat types are similar to those documented by
Daubenmire (1970) but others are distinctive for
the east front of the Montana Rockies. Major

820

E 12. Bun

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

PRES A

(Noe
i

nchgrass prairie community on the Deerlodge National Forest (1, 750 m), Montana, Inter-
DA

ou ina EE Pseudotsuga menziesii with groves of Populus tremuloides occupying lower elevations. USD

Forest Servic

Intermountain grassland dominants include
Stipa comata, Agropyron spicatum, Festuca ida-
hoensis, F. scabrella, Deschampsia caespitosum,
Artemisia tridentata, A. arbuscula, Potentilla fru-
ticosa, Purshia tridentata, Cercocarpus ledifolius,
Rhus trilobata, and Sarcobatus vermiculatus.
Other NRM grassland studies have focused on
local expressions of prairies on sites surrounded
by continuous forests. In many instances, dis-
junct grasslands are the result of local mountain
rainshadows, but soil texture (e.g., coarse glacial
till) and past fires have also favored grassland
development (Blinn & Habeck, 1967; Koterba
& Habeck, 1971; Root & Habeck, 1972). Wright
& Wright (1948) have described the grasslands
in southcentral Montana; their five types are list-
ed by community dominants (from most me-
sophytic to more xerophytic): a) Festuca ida-
hoensis, b) Agropyron spicatum, c) Agropyron
spicatum/Carex filifolia/Bouteloua gracilis,
Bouteloua gracilis/Stipa comata/Koeleria cris-
tata, and e) Bouteloua gracilis/Stipa comata.

e.
—

Representative shrubland types in the Inter-
sen region € 13) were described by
Mueggler & Stewart (1980) and by Kuchler
pica "C ercocarpus ledifolius, Purshia triden-
tata, Symphoricarpos oreophilus, and Artemisia
tridentata all form shrub-bunchgrass commu-
nities below lower timberline, occupying dry
rocky sites or river floodplains, although these
shrubland communities may also be found in-
terspersed within the timbered zones. The grass-
es are the same as those that form the prairie
vegetation types in the valleys and lower slopes,
that is, Agropyron spicatum, Festuca idahoensis,
Stipa comata, and Poa sandbergii.

INLAND MARITIME REGION

The Inland Maritime region spans northeast-
n Idaho, and north-
(Fig. 4, Region D). The Selkirk
Mountains of northern Idaho and adjacent parts
of Washington and British Columbia, the north-

1987]

HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

821

FIGURE 13.

ern Bitterroot Range, the Cabinet Mountains, the
Whitefish Range, the northern Swan Range, and
the west slope of the Continental Divide (in Gla-
cier Park) collectively comprise this region.

s its name suggests, this region is relatively

stations in localized rainshadows may record 40
cm or less. Warm, dry weather usually prevails
during July and August. Grassland and Pinus
ponderosa communities, which exhibit drought
tolerance (Minore, 1979; Lassoie et al., 1985),
occur locally in some of the driest rainshadow
areas. The reduced summer moisture sets the
stage for wildfires. Historic fires are believed to
have perpetuated xeric vegetation types in cli-
matically moist parts of this geographic subunit
(Habeck & Mutch, 1973).

Typically, the forest zones in the Inland Mar-
itime region contain combinations of the follow-

Sagebrush shrubland and rangeland vegetation types in upper Big Hole drainage (2,000 m), in
southern continental ranges, Beaverhead National Forest, Montana. Pseudotsuga menziesii forest in distance.
USDA Forest Service.

ing dominants, ranked from highest to lowest in
drought resistance (Minore, 1979): Pinus pon-
derosa, P. contorta, Pseudotsuga menziesii, Picea
engelmannii, Abies grandis, Larix occidentalis,
Abies lasiocarpa, Thuja plicata, Taxus brevifolia,
Pinus monticola, Tsuga heterophylla, and Tsuga
mertensiana (Figs. 14, 15).

Physiological adaptive features of these coastal
conifers were reviewed in detail by Lassoie et al.
(1985). The mesophytic forests dominated by
Thuja and Tsuga resemble in many ways the
forests found on the western slopes of the Coastal
nd Cascade mountain ranges in Washington and
British e E (Daubenmire & Daubenmire,
1968; Habeck, 1978; Williams & Lillybridge,
1983; Cooper et al., 1988)

With increased elevation, even greater amounts
of precipitation occur, often reaching and ex-
ceeding 200 cm annually. A large portion (75-
8596) of this precipitation falls as snow between
September and March. Pseudotsuga menziesii,

£

Abies lasiocarpa, and Picea engelmannii occur

ANNALS OF THE MISSOURI BOTANICAL GARDEN

rae —

T mtm
— JP
` a wem d

LI

$

—

d
1^

d

FIGURE 14. Mature forest in the Inland Maritime region; Deception Creek, Coeur d’Alene National Forest,
Idaho (1,000 m). Tsuga heterophylla, Abies grandis, and Pinus monticola dominate. Note fire-scarred snag on

left. USDA Forest Service.

in abundance on the higher inland Maritime
mountain slopes. Pinus albicaulis is locally abun-
dant on warm aspects that may experience some
July-August moisture shortages.

Between 1,500 and 1,800 m, near the Idaho-
Montana state line, Tsuga mertensiana joins Abies
lasiocarpa in forming the highest closed-canopy
forest type (Habeck, 1967). Tsuga mertensiana
is very intolerant of summer drought and heat
(Minore, 1979) and severe continental winters,
but survives very well in parts of the Inland Mar-
itime Region where suitably mild conditions pre-
vail. Tsuga mertensiana is an abundant timber-
linespeciesin the Coastal and Cascade mountains
of western Oregon, western Washington, British
Columbia, and southeastern; Alaska (Arno &
Hammerly, 1984). The Inland Maritime region's
timberline, formed at 2,000-2,300 m (Fig. 16),
features krummholz of Abies lasiocarpa and Pi-
nus albicaulis; erect or partly wind-shaped Larix
lyallii is also present at times (Habeck, 1969;

Arno & Habeck, 1972). On glacially scoured parts
of the Selkirk Mountains in northern Idaho, Tsu-
ga mertensiana and Larix lyallii are rare or ab-
sent.

Each of the Inland Maritime trees has an in-
dividualistic distribution. They can be ordered
from most restricted to most widely distributed
within this region: A/nus rubra, Tsuga merten-
siana, T. heterophylla, Pinus monticola, Thuja
plicata, Taxus brevifolia, and Abies grandis. It
has been noted that the individuals or popula-
tions occurring at the range limits of these mar-
itime trees, such as Thuja plicata near Missoula,
Montana, are confined to isolated higher (1,400
m) ravine sites (Habeck, 1978; McCune & Allen,
1985b). Summer moisture deficiencies encoun-
tered in westcentral Montana may be counter-
balanced by the cooler temperatures in these
middle elevation ravine sites.

The undergrowth in the mesic and wet-mesic
Inland Maritime region forests is floristically

340636

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HABECK — NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

E
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[^2]
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FIGURE 15.
Montana. USDA Forest Service.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

FIGURE 16.

richer than in any of the other subunits. It in-
cludes Clintonia uniflora, Adenocaulon bicolor,
Disporum hookeri, Tiarella trifoliata, Cornus
canadensis, Aralia nudicaulis, Rubus parviflorus,
Athyrium filix -femina, G iymnocarpium dryopte-

many ed species.

dient exists, west to east, between S
Washington (38 cm/yr., pine-grassland savanna)
and nearby (40 km east) Coeur d'Alene, Idaho
(75 cm/yr., cedar-hemlock-western white pine
forests).

Fires occur at lower frequencies (100-year in-
tervals or longer) in this geographic region (Ha-
beck, 1985). When wildfires do occur, however,
they are often of much higher intensity (“stand

“Snow ghost" forest in the subalpine zone of northern Idaho's Bald Mountain (2,300 m). Snow-
caked trees are Abies lasiocarpa. USDA Forest Service

replacement" fires), due to the greater organic
matter (“fuel”) accumulations in these moist for-
ests. When the right combination of midsummer
"fire weather" occurs—characterized by high
paige gusty afternoon winds, dry ground

potential climax species to make an initial entry
at the same time. Examples of short interval (less
than 10 years) “double” and “triple” burns exist,
and forest recovery on such sites may be retarded
for many decades.

Deep snow accumulations in the Inland Mar-
itime mountains (Fig. 17) cause frequent ava-

often dominated by such woody angiosperms as

1987]

;
`

t
N

_

PL >= 22

HABECK — NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

FiGuRE 17. Snow cornice above Picea engelmannii and Abies lasiocarpa forest at 2,100 m in Panhandle

National Forest, coal Idaho. USDA Forest Service.

]

Alnus sinuata alnifolia, Sorbus spp.,
Acer glabrum, Populus trichocarpa, P. tremu-
loides, Salix spp., and a rich assortment of pe-
rennial herbs that seem well adapted to frequent
avalanches.

VEGETATION OF THE SOUTHERN
CANADIAN ROCKIES

The southern Canadian Rockies are composed
of high, rugged terrain with a long history of
glaciation. The mountain ranges often reach or
exceed 2,400 m, with peaks over 3,000 m. The
southern Canadian Rockies embrace several ma-

jor mountain ranges near the United States/Can-

g

Frigid polar air masses characterize the con-
tinental climate east of the Great Divide in Al-
berta. The dessicating winter winds are similar
to those described for the east front of the Mon-
tana Rockies; these winds are responsible for “red
belt” mortality among the conifer communities
located here. Just west of the Great Divide, the
climate is more like that of the Inland Maritime

826

region in northern Idaho and northwestern Mon-
tana; mild and wet weather prevails as Pacific
coastal storm tracks penetrate eastward (Arno &
Hammerly, 1984).

Southeastern British Columbia, parts of which
lie in the Cascade Mountains rainshadow, has a
warm and dry lower timberline in valley bottoms
where only 25-40 cm of precipitation occurs an-
nually. Open Pinus ponderosa woodlands,
bunchgrass prairies (Agropyron spicatum and
Festuca idahoensis), and sagebrush steppe (4r-
temisia tridentata) form the major plant com-
munities in the semiarid valleys (Tisdale &
McLean, 1957; McLean, 1970; Tisdale, 1974).
Some of the upper slopes of nearby mountains
also have low annual precipitation, and at mid-
elevation slopes a forest zone dominated by
Pseudotsuga menziesii occurs. Above this is the
subalpine zone dominated by Abies lasiocarpa,
Picea engelmannii, and Pinus albicaulis, which
ultimately form a krummholz on upper slopes
above 2,100 m. Past fires in southeastern British
Columbia perpetuated extensive stands of Pinus
contorta at middle and upper elevations (LaRoi
& Hnatiuk, 1980; Arno & Hammerly, 1984).

Those parts of southeastern British Columbia
that receive greater coastal moisture display for-
est vegetation similar to that of the Inland Mar-
itime region in the United States. Warm, dry,
lower timberlines occur only rarely in this part
of the Canadian Rockies, where dominants in-
clude Thuja plicata, Tsuga heterophylla, Abies
grandis, Pseudotsuga menziesii, Picea engelman-
nii, and P. engelmannii x P. glauca hybrids.
Pinus contorta, Abies lasiocarpa, Tsuga merten-
siana, Pinus albicaulis, and Larix lyallii are the
major timberline dominants in the Selkirk and
Purcell mountains. Timberline in these ranges
typically occurs at elevations below 2,100 m;
however, it may occur below 1,500 m in areas
of deep snow accumulations. The highest moun-
tain peaks (over 3,000 m) support permanent
snowfields and glaciers (Shaw, 1916)

The Alberta Rockies consist of the mountains
forming the Great Divide. From Waterton Park
northward to Banff and Jasper National Parks,
the Divide peaks reach from 2,800 m to nearly
3,650 m in height. The continental climate in
Alberta disfavors the occurrence of tree species
adapted to milder maritime conditions. Annual
precipitation is near 150 cm at the higher ele-
vations but less than 50 cm on the east front at
Calgary. The subalpine and montane forest zones
are dominated by the same species listed above

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

for British Columbia, except that maritime trees
like Thuja and Tsuga are missing. At timberline,
Abies lasiocarpa, Picea engelmannii, Larix lyal-
lii, and Pinus albicaulis form the upper treeline
communities, including the krummholz zone
(Moss, 1955; Ogilvie, 1963, 1976; Day, 1972;
Corns & Achuff, 1982).

The lower forest
are mixtures of Pseudotsuga menziesii, Picea
glauca (and hybrids P. glauca x P. engelmannii),
and Populus tremuloides. Local moisture-stressed
sites support Pinus flexilis (Moss, 1944; Rowe,
1959; Stringer & LaRoi, 1970). Pinus ponderosa
is absent, according to Arno & Hammerly (1984),
seemingly unable to tolerate winter desiccation
and rapid temperature changes. As was described
for other areas of the NRM, frequent high in-
tensity wildfires helped develop and perpetuate
extensive stands of Pinus contorta throughout
much of the Alberta Rockies, from the lower to
the upper timberline (Tande, 1979).

The natural grasslands in southern Alberta east
of the Great Divide occur where precipitation
becomes lower, temperatures are warmer, Chi-
nook winds prevail, and evaporation rates are
high (Moss, 1944; Moss & Campbell, 1947). Moss
(1944) described three prairie community types:
Bouteloua gracilis/Stipa comata, Agropyron/Sti-
pa/Carex, and Festuca scabrella/F. idahoensis/
Danthonia intermedia/D. parryi. Much of the
native prairie no longer exists because of modern
levels of grazing and mowing.

+h 1 A

VEGETATION OF THE MIDDLE
Rocky MOUNTAINS

The last part of the Rocky Mountains to be
treated here lies north of Provo and Salt Lake
City, Utah, in the vicinity of 40°N, and includes
the adjacent parts of northwestern Colorado, the
Medicine Bow Mountains in southeastern Wy-
oming, the remainder of Wyoming west and north
of the Wyoming Basin desert, north to south-
central Montana (Beartooth Plateau), and west
to the Snake River Plain in southern Idaho (Fig.
4, Region F). This region coincides with the
"Middle Rockies" as discussed by Oosting &
Reed (1952), Johnson & Pfister (1982), Arno &
Hammerly (1984), and Mutel & Emerick (1984),
and includes the northeastern part of the Rockies
that Reveal (1979) designated “the Intermoun-
tain Region.” Figure 6 shows the typical vege-
tation zones in this part of the Rocky Mountains.

The Utah portions of the middle Rocky Moun-

1987]

tains support a flora that contains many elements
that are geographically centered further to the
south and southwest, characteristic of the pin-
yon-juniper, chaparral, and Great Basin desert
vegetation types. In contrast, western Wyoming
ranges are without these floristic inclusions and
display more elements common to the East Front
of the Rockies in central Montana (Arno & Ham-
merly, 1984). The forest vegetation of this region
is usually dependent on melting of deep winter
snow packs (from Pacific storms) to supply need-
ed summer moisture. The inland continental cli-
mate lacks a maritime component of the sort
described earlier (Baker, 1944).

Arno & Hammerly (1984) described the mid-
dle Rockies as an area of well-separated ranges
diverse in geologic structure and rock types.
Johnson & Pfister (1982) provided detailed in-
terpretive summaries of the geology of this re-
gion. Three geologic mountain types exist (see
Fig. 4 for locations): 1) anticlinal mountains
(Beartooth Plateau, Big Horn—Pryor Mountains,

Uinta Mountains, and Wind River Range); 2)
mountains of the overthrust belt (Tetons, Wy-
oming, and Wasatch ranges); and 3) volcanic
mountains (Yellowstone Plateau and Absaroka
Range). The highest points in the middle Rocky
Mountains reach up to approximately 4,200 m
(Gannett Peak and Grand Teton), although more
typical heights range between 3,000 and 3,600 m.

The Wasatch and Uinta ranges have broken
patches and stringers of timber on their upper
mountain slopes, a result of frequent avalanches
clearing wide tracks through the timber. Several
tree species achieve dominance here, including
Pseudotsuga menziesii, Pinus flexilis, Picea pun-
gens, Abies concolor, and Populus tremuloides.
Pinus contorta is present in the northern part of
the Wasatch Range but Pinus ponderosa is scarce.
The lower Wasatch slopes are covered with
Quercus gambelii, Cercocarpus ledifolius, and
Acer grandidentatum. This mountain shrub or
chaparral type replaces the pinyon-juniper vege-
tation zone found further westward in Nevada
(Hayward, 1945, 1948; Arno & Hammerly,
1984).

The Uinta Mountains, which have an east-

per subalpine zones are stands of Picea engel-
mannii, Abies lasiocarpa, and Pinus flexilis; this
region is beyond the distribution of Pinus albi-
caulis. Extending down to 2,700 m below these

HABECK — NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

827

zones are Pinus contorta forests. A wide band of
Populus tremuloides, serving as a major seral
species in the middle Rockies, occurs below the
Pinus contorta zone. Pinus ponderosa is scattered
on south slopes and found more extensively on
north ones at the lower Uinta timberline down
to 2,100 m. Mid elevations support extensive
stands of Pseudotsuga menziesii; moist drainages
(riparian sites) contain Picea pungens commu-
nities

Arno & Hammerly (1984) and Ellison (1954)
provided additional descriptions of the contrast-
ing climatic and vegetational features between
the Wasatch and nearby Uinta ranges. In contrast
to the Wasatch Range, the Uintas lack the oak-
maple chaparral but support a pinyon-juniper
type. In both areas, the lowest and driest shrub
zones are dominated by a dozen different species
and varieties of sagebrush (Artemisia spp.) and
Atriplex canescens, A. confertifolia, and Cerco-
carpus ledifolius. Variations in summer rainfall,
geologic and topographic features, and contract-
ing patterns of past plant migrations have all
contributed to these differences.

The modern forest vegetation occurring in
Gordo! s Front Range (Fig. 4, vicinity of Rocky
Mountain National Park) has been the subject
of many studies dating back 80 or more years.
The most recent and comprehensive among these
is the detailed treatment provided by Peet (1981),
who reviewed muc the previous vegetation
literature for this ad of the middle Rocky
Mountains. Employing modified gradient anal-
ysis techniques, he derived a classification of the
Front Range vegetation that accommodated both
developmental (successional) and mature (late
seral/climax) community types, relating them to
a moisture-elevational complex. Peet classified
the Front Range vegetation (1981, his fig. 5) into
community series as follows: A. Pinus ponderosa
woodland series; B. Pinus ponderosa/Pseudotsu-
ga menziesii forest series; C. Mesic montane for-
est series (a heterogeneous group that includes
Picea, Abies, Pseudotsuga, Populus, Betula, and
alnus); D. Pinus contorta forest series; E. Picea/
Abies forest series; F. Pinus flexilis forest series;

Alpine transition (krummholz) series. His
classification also delimits community type sub-
units within these series. Subsequently, Green-
land et al. (1985) described a methodology for
defining bioclimatic zones in the Colorado Front
Range utilizing a one-dimensional version of
Peet's vegetation ordination model

The importance of 19th century anthropogenic

828

disturbances within the montane forests of the
Front Range have been investigated by Veblen
& Lorenz (1986). Patterns of forest recovery fol-
lowing severe logging and burning disturbances
inflicted during the Colorado g booms were
studied. Their findings clarify the successional
pathways that forest recovery has followed dur-
ing the last century in this part of the Rocky
Mountains.

The upper-elevation forests in a representative
watershed within the Medicine Bow Mountains
in central Wyoming were discussed in detail by
Romme & Knight (1981), who investigated and
modeled the interactions of fire frequency and
topographic position on vegetation dynamics in
this part of the middle Rockies. Although much
of the high country in the Medicine Bows has a
potential for supporting climax forests dominat-
ed by Picea engelmannii and Abies lasiocarpa,
such expressions of spruce-fir forests are pri-
marily confined at this time to moist sites such
as ravines and valley bottoms. Past occurrences
of short-interval (less than 100 years) wildfires
on open slopes and ridgetops favored develop-
ment and maintenance of Pinus contorta forests
(Lotan & Perry, 1983). Even though fire suppres-
sion has been in operation for more than a cen-
tury, the slow rates of successional processes in
these mountains have tended to impede the re-
turn of spruce-fir forests.

Romme & Knight (1981) included within their
watershed study area a two-dimensional ordi-
nation of all major plant community types oc-
curring between 2,250 and 3,000 m on sites rang-
ing from moist valley bottoms to south slope
ridgetops. Salix, Alnus, and Populus mixtures
occur in the lower, moist habitats; Pseudotsuga
menziesii forests occur on the mesic, lower north
slopes; Artemisia communities occur on most
xeric foothill elevations. At intermediate eleva-
tions of 2,500 to 2,700 m, combinations of Pseu-
dotsuga menziesii, Pinus contorta, and Populus
tremuloides prevail. Only moist sites above 2,700
m currently support mature spruce-fir forests.

The mountains in western Wyoming (Wind
River, Salt River, and Wyoming ranges, as shown

nii and Abies lasiocarpa (Loope,
1976; Steele et al., 1983; Arno & Hammerly,
1984). A krummholz zone composed of these
conifers is found at elevations near 3,050 m
throughout these mountains. Much of the sub-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

alpine forest zone, however, is dominated by ex-
pansive stands of Pinus contorta except where
limestone substrate prevails. This pine, together
with Pseudotsuga menziesii, often forms the up-
per timberline forests at elevations above 2,700
m and then extends downward to the lower zones
at or below 2,100 m (Moir, 1969; Arno & Ham-
merly, 1984). Loope & Gruell (1973) have de-
scribed the natural role of fire in these north-
western Wyoming forests.

Near the lower timberline, Populus tremu-
loides groves comprise a broken belt separating
Pinus contorta forests from the sagebrush com-
munities in the lower valleys (Reed, 1971;
Youngblood & Mueggler, 1981; Arno & Ham-
merly, 1984; Fig. 18). Although Pseudotsuga
menziesii can be found in the western Wyoming
mountain ranges, it does not form a distinctive
forest zone as it does elsewhere in the northern
Rockies, such as in western Montana (Arno,
1979). This is due to the importance of Pinus
contorta as a seral species in zones where Pseu-
dotsuga menziesii is the potential climax domi-
nant. Many ofthe subalpine conifers can be found
at lower timberline, often along mountain stream
courses (Arno & Hammerly, 1984). The valleys
in the western Wyoming mountain ranges are
much cooler than comparable sites elsewhere in
the northern Rockies; for this reason, Populus
tremuloides and mesophytic conifers occupy the
lower timberline zone (Steele et al., 1983). Pin-
yon-juniper and mountain chaparral are missing
from the cold western Wyoming mountains (Arno
& Hammerly, 1984).

The Absaroka Range and adjacent Beartooth
Plateau (Montana-Wyoming border; Fig. 4) dis-
play an extensive alpine tundra zone with many
ice fields at elevations over 3,000 m. Timberline
parkland (open-canopied groves and scattered
individuals) is also present, dominated by Pinus
albicaulis, Picea engelmannii, and Abies lasio-
carpa, similar to those in the Wind River and
Teton ranges.

The Big Horn Mountains, spatially separated
from the other middle Rocky Mountain units,
are located east of the Big Horn Basin in west-
central Wyoming and extend into adjacent parts

reaching about 60 cm in the subalpine forest zone
at 2,700 m. Soils derived from granite, shale,

1987] HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION 829

FiGuRE 18. Old growth Populus tremuloides forest near Scout Lake (3,100 m), Utah. Note conifer invasion
in understory. USDA Forest Service.

830

limestone, and dolomite rocks play a major role
in plant distribution patterns in many of the mid-
dle and northern Rockies east of the Continental
Divide, including the Big Horns. A series of vege-
tation studies has been completed in the Big
Horns (Hurd, 1961; Despain, 1973; Hoffman &
Alexander, 1977).

Big Horn Mountain Pinus contorta forests oc-
re g the coarse and nutrient-poor
granitic soils. Grasslands and shrublands gen-
erally occupy soils from shale and limestone, ex-
cept where forests are supported in areas where
moisture stress is lessened due to slope aspect
(Arno & Hammerly, 1984). Picea engelmannii
and Abies lasiocarpa occur in the subalpine-tim-
berline zone of the Big Horn Range, but Pinus
albicaulis is absent from this area. Pinus contorta
is dominant on granitic soils at elevations below
the subalpine zone which extends from 2,700 m
to 2,100 m. The lower timberline is formed by
Pseudotsuga menziesii near 1,800 m. Juniper
woodlands (Juniperus scopulorum and J. osteo-
sperma) are closely associated with the Pseudo-
tsuga menziesii zone on the west slopes; Pinus
edulis is replaced by Pinus flexilis in this juniper
community type. Pinus ponderosa var. scopu-
lorum becomes important only in this part of the
middle Rocky Mountains (east face of Wyoming
Big Horns and northward into Montana) but does
not display the large growth forms seen among
the P. ponderosa var. ponderosa occurring fur-
ther west. Geographically, P. ponderosa var.
scopulorum occurs eastward into the Black Hills
of South Dakota, to the exclusion of Rocky
Mountain conifers (Johnson & Pfister, 1982).

The middle Rockies support widespread sage-
brush-grass; Tisdale & Hironaka (1981), Blais-
dell et al. (1982), oe < al, (1983), and
West (1983) provided in-dep t view
of the major community ae in the nonforested
areas of this region. Where the Wyoming Basin
and Great Plains meet, there exist sod-forming
grasslands dominated by Bouteloua gracilis and
Agropyron smithii intermixed with Artemisia tri-
dentata and other sagebrush taxa. These com-
munities are the products of the continental cli-
mate, with growth moisture arriving in early
summer rainfall. Elsewhere in southern and cen-
tral Idaho and southwestern Montana, bunch-
grass prairies are present, dominated by Agro-
pyron spicatum, Festuca idahoensis, Stipa
comata, and Poa sandbergii. In these sagebrush-
bunchgrass communities, moisture comes pri-
marily in winter and spring, with droughty sum-
mers.

citec

[^2]

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

The middle Rockies sagebrush-bunchgrass
community types recognized by Hurd (1961),
Beetle & Lud (1982), Johnson & Pfister
(1982), and Hironaka et al. (1983) include the
following, each named after the major dominant:
a) Festuca idahoensis, b) Agropyron spicatum, c)
Bouteloua gracilis, d) Purshia tridentata, e) Sym-
phoricarpos oreophilus, f) Artemisia tridentata,
g) A. tripartita, h) A. nova, i) A. arbuscula, j) A.
longiloba, k) A. cana, 1) A. rigida, in, Atriplex
canescens/A. confer. AUAA. nutta and n)

arcobatus vermiculatus. Shrubland ane
types in the same region include a) Quercus gam-
belii, b) Acer grandidentatum, and c) Cercocar-
pus ledifolius.

Modern (post-1900) vegetation shifts in mid-
dle Rocky Mountain 55) believed to be
induced by fir
(Sauer, 1950; Humphrey, 1962; Steele et al., : 1981,
1983), also involve extensive invasion by Junip-
erus occidentalis into grassland and sagebrush
communities. Tisdale & Hironaka (1981), point-
ing to the sensitivity of Artemisia to fire, sug-
gested that fires in presettlement times must have
been infrequent (long interval) in some parts of
the sagebrush region, such as drier Artemisia

Barney & Frischnecht (1974) and
Young & Evans (1981). Problems attending these
changes were described in a lengthy juniper man-
agement symposium (Martin et al., 1978).
Through wood rat (Neotoma) midden and lake
sediment analysis, Mehringer & Wigand (1986)
documented that western juniper has had a long
history of rapid geographic shifts in response to
climate changes during the past 4,000 years. They
stated, “. . . the spectacular and persistent ex-
pansion of western juniper over the last hundred
years — despite chaining, bulldozing, cutting, poi-
soning and burning—i is not an unusual event nec-
essarily req g explanations unique to the his-
toric period. In fact, the rate and degree of change
in the comings and goings of western juniper over

ant communities historically used as sheep
and cattle range throughout the middle Rockies
have been severely altered as a result of abusive
levels of grazing. Employing paired-stand anal-
ysis techniques, range ecologists have provided
extensive documentation of compositional
changes attributed to grazing. Native species of

1987]

HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

FiGuRE 19. Abies mes (center) and Pinus flexilis (left & right) on King's Hill (2,200 m) in northern
aa ranges, central Montana; Lewis and Clark National Forest, Montana. USDA Forest Service.

Agropyron, Festuca, Poa, and Stipa are classed
as "decreasers" which are reduced in abundance.
In their place, introduced grasses and forbs in-
vaded and, for all practical purposes, have cre-
ated new vegetational equilibria on the western
rangelands. As an example, cheatgrass (Bromus
tectorum), an introduced annual grass, has prov-
en to be a superior competitor in western bunch-
grass communities (Harris, 1967; Mueggler &
Harris, 1969; Daubenmire, 1970; Mueggler &
Stewart, 1980) and has come to dominate large
areas in the middle and northern Rockies.
Other invaders that have become widespread

maculosa, Cirsium vulgare, Tra on dubius,
Euphorbia esula, ypericum perforatum
any of these species, along wit dbeckia oc-

nifer forests severely disturbed by logging and/
or domestic grazing. Man's present-day penchant
for working land on higher and steeper slopes is
contributing to an acceleration of vegetation
change. Furthermore, conflicts between agricul-
ture and livestock, and winter foraging by big

game, frustrate efforts to reverse the loss ofnative
plant components.

OTHER ROCKY MOUNTAIN VEGETATION TYPES

Other vegetation types occurring within the
northern Rockies are described below. Some of
these are distributed throughout the northern
Rocky Mountains, i.e., timberline/alpine tundra
and wetland/riparian types, while others are con-
fined to single mountain ranges or unique local-
ized habitats.

TIMBERLINE AND ALPINE TUNDRA

Upper treeline reaches 2,275 m in southern
Alberta and nearly

ing summer. Sharp temperature fluctuations also
create severe winter stresses. In contrast, inland
maritime timberline/alpine tundra is more moist

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

FIGURE 20. nes and nival life zones in vicinity
Montana. USDA Forest Service.

than the continental areas and experiences less
extreme winter temperatur

-ommon northern ipis Mauntaln umber-
line tree species have been identified earlier; these
intermix islandlike within the lower alpine tun-
dra zone. Slope aspect, microtopography, pat-
terned ground (resulting from cryopedogenesis),
soil chemistry, soil water potential, and wind ex-
posure combine to form an environmental com-
plex (Fig. 20) that dictates the arrays oflow shrubs,
perennial herbs, and graminoids forming the
northern Rockies timberline/tundra ecosystems
(Griggs, 1938; Choate & Habeck, 1967; Bamberg
& Major, 1968; Habeck, 1969; Billings, 1969,
1978; Smith, 1969; Johnson & Billings, 1962;
Arno & Hammerly, 1984; Bliss, 1985; Spence,
1985; Peet, 1987).

Alpine permafrost occurs in some northern
Rocky Mountain ranges, where mean annual air
temperature is at or below 0°C and may, accord-
ing to Pewe (1983), date back to Wisconsin time
when such frozen soil was widespread 1,000 m

of Beartooth Pass (3,300 m), Custer National Forest,

below present-day elevations. Such permafrost
areas coincide with modern treeless areas where
alpine vegetation may favor maintenance of the
frozen soil conditions. Vegetation dynamics on
sites that have experienced recent ice retreat have
been described in detail by Spence (1985) for
parts of Wyoming's Teton Range. He investi-
gated plant invasions on moraines fronting gla-
ciers. Continual disturbances take place on the
moraines, and compositional differences among
his study areas do not seem om related to a
successional chronosequenc

Bliss (1985) reviewed His clivsiolorical ecol-
ogy of alpine and timberline plant communities
in North America, adding to earlier efforts that
emphasized floristic phytogeography. He stated
that complexities of community structure and
composition in the alpine zone, including spatial
discontinuities between mountain ranges, have
permitted only generalized classification units.
Growth and survival in the alpine are closely
related to growth forms. Bliss (1985) listed and

1987]

discussed the following major growth form cat-
egories: 1) annuals (uncommon), 2) cushion and
rosette-forming species occurring on exposed
ridges, 3) graminoids found in all alpine habitats,
4) forbs, and 5) deciduous and evergreen shrubs,
many of which are low mat-forming types.

Based on an extensive literature review, John-
son & Pfister (1981, 1982) listed the following
alpine tundra community types typical of the
Rocky Mountains. The common dominants are
listed for each type:

1. Shrub Communities: Salix rotundifolia, S. re-
ticulata, S. arctica, Betula glandulosa, Dry
octopetala, Kalmia microphylla, Phy llodoce
empetriformis, P. glanduliflora, and Vaccin-
ium scoparium.

. Meadow and Turf Communities: Geum ros-

sii, Deschampsia caespitosum, Carex tolmiei,

C. nigricans, Juncus s Poa alpina, Phleum

alpinum, Polygonum bistortoides, and Oxyria

digyna.

Bog and Fen Communities: Sphagnum spp.,

Carex aquatilis, C. rostrata, C. simulata, Er-

iophorum spp., Juncus spp., Pedicularis

groenlandica, P. contorta, Kalmia micro-
phylla, Ranunculus spp., Calamagrostis can-
adensis, and Eleocharis spp.

Cushion Plant and Fellfield Communities: Si-

lene acaulis, Dryas octopetala, Trifolium nan-

um, Luzula spicata, and Selaginella densa.

. Boulder Field Communities: Geum rossii,

Mertensia spp., Ribes cereum, Polemonium

spp., Aquilegia spp., Penstemon fruticosus, and

Sibbaldia procumbens.

Snowpack Communities: Carex nigricans,

Juncus drummondii, Erythronium grandiflo-

rum, Valeriana sitchensis, Luzula glabrata,

and Senecio triangularis

N

Ww

A

un

on

The nival zones of the Rockies (Figs. 5, 6, and
20) exhibit snowfields that support cold-adapted
floras composed predominantly of algae, but not
vascular plants, although some fungi species may
also be present (Garric, 1965; Vinyard & Whar-
ton, 1978). Snow algae occur on the surfaces of
glaciers and snowfields and may give the snow
a reddish color (or even yellow, green, blue, black,
or purple), depending on the species involved.
These algae have enzyme systems that catalyze
reactions most efficiently at lower temperatures
(Hoham, 1975). In the Rockies, and elsewhere
in western North America, one of the most com-
mon species is Chlamydomonas nivalis, which
is known to cause the pink-red “watermelon

HABECK —NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

833

snow." Red coloration on the snow surface in
summer is caused by an accessory photosyn-
thetic pigment called astaxanthin, which in-
creases in abundance in intense sunlight. The
cryophilic algae encyst during the four to eight
months of winter and break dormancy in sum-
mer.

Other snow algae listed by Garric (1965) in-
clude Scotiella nivalis, S. cryophila, Chodatella
brevispina, C. granulosa, Raphidonema nivale,
R. tatrae, and Romeria elegans var. nivicola.
Snow fungi that may be saprophytic on snow
algae include Chionaster nivalis and Selenotila
nivalis, while another group of fungi, the Chy-
tridiales, has been reported to be parasitic on snow
algae (Vinyard & Wharton, 1978). The snowfield
ecosystems also support ciliated protozoans, ro-
tifers, springtails, and snow worms.

WETLAND TYPES

Wetland ecosystems in the Rocky Mountains
occupy landscape units such as marshes, swamps,
bogs, fens, and other topographic depressions with
high water tables. Water may cover the sites
ephemerally or intermittently and may be shal-
low or deep (under or over 2 meters). The term
“wetland”? also embraces wet meadows, pot-
holes, sloughs, riparian zones (along stream

ses), and river-overflow areas. Permanently-
filled, shallow lakes and ponds less than 2 meters
in depth, supporting emergent vegetation, are also
included in this term. Wetland site definitions
and classifications have been the subject of thor-
ough attention by Windell et al. (1986). Their
review of Rocky Mountain wetlands represents
a comprehensive treatment of the subject and is
the primary source for my synthesis of Rocky
Mountain vegetation.

Rocky Mountain wetlands do not exist as re-
gional climaxes, as in some high latitude arctic
areas, because of the continental climate which
prevails over much of the region. Snowmelt run-
off is rapid, much of the northern Rockies do not
receive high amounts of precipitation, and evap-
oration rates are high in many areas. Thus, wet-
land communities are formed only in certain to-
pographic settings where an abundance of water
occurs seasonally or permanently.

In their ecology of Rocky Mountain wetlands,
Windell et al. (1986) provide detailed informa-
tion on community structure and a general clas-
sification of these ecosystems. Their literature
review covers the entire Rocky Mountains. An

834

overview of their findings is given below, sup-
plemented with information taken from other

regional studies (Pfister & Batchelor,
Youngb

1984;

ood et al., 1985). An outline of wetland

community types will be presented, together with
a listing of plant dominants for each.

l.

N

3.

Permanent Shallow Standing Water (less than
-4 m):

Floating communities: open water surface

dominated by Lemna spp.

b. Rooted submergent communities: domi-
nated by species of /soetes, Nitella, Pota-
mogeton, Najas, and Myriophyllum.

ooted floating-leaved communities:
dominated by species of Nuphar, Nym-
phaea, Sparganium, and Potamogeton.

. Rooted emergent communities: dominat-
ed by species of Carex, Eleocharis, Juncus,
Glyceria, Phragmites, Sagittaria, Scripus,
Sparangium, Typha, Menyanthes, and
Petasites.

Ë tƏ

°

e.

. Seasonal or Permanent High Water Tables,

Without Permanent Standing Water:

Herbaceous wetlands: floating mats dom-
inated by species of Carex and Sphagnum.
On mineral soils: marshes or wet meadows
dominated by grasses, sedges and rushes,

P

. Fe erbaceous wetlands on
organic soils and dominated by sedges
(Carex spp.) and other graminoids such as
species of Juncus, Eleocharis, Deschamp-
sia, and Calamagrostis.
Bog communities: Northern Hemisphere
bogs are dominated floristically by a mat
of Sphagnum spp. and members of the
in the mat. Rocky
Mountain regional bogs include species of
almia, Drosera, Menyanthes, Eriopho-
rum, Potentilla, and Gaultheria.
. Marsh/wet meadow communities with
fresh water: occur on mineral soils and are
dominated by herbaceous species of the
following genera: Carex, Deschampsia,
Danthonia, Juncus, Penstemon, Erigeron,
and Calamagrostis.
Marsh/wet meadow communities with sa-
line water: alkaline sites with sodium con-
centrations over 15%, supporting onl
salt-tolerant species such as
Distichlis, a Puccinellia, Tr. ie
lochin, and Salicor
Forested Wetlands: Rivers and streams in the

=
5
e
°
B
B
£
5
=
©

e

e.

°

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Rocky Mountains that have well-developed
floodplains supporting broadleaf deciduous
forests and woodlands. The term “riparian
wetlands” refers to plant communities oc-
curring adjacent to running water. Rocky
Mountain riparian forests are dominated by
Populus (P. trichocarpa, P. deltoides, P. an-
gustifolia, P. balsamifera, and P. tremu-
loides), as well as Betula papyrifera, Fraxinus
pennsylvanica, Ulmus americana, and Acer
negundo. Needle-leaved evergreens such as
icea engelmannii, Pinus ponderosa, Pseu-
dotsuga menziesii. Thuja plicata, and Abies
grandis also exhibit local dominance on ri-
isa sites extending through elevational
zones. Mixtures of Populus spp., Juniperus
ny pa and/or Pinus ponderosa are also
ound on floodplains. Such sites are subject
to frequent disruptions by flooding and do-
mestic grazing (and historically by fire), and
communities are often in various stages of
successional recover
Shrub-dominated Wetlands (Carrs): Some
wetlands are dominated by shrub species
rather than by forbs, grasses, or trees. Shrub
species listed by Pfister & Batchelor (1984) as
dominants in the northern Rockies are Salix
bebbiana, Cornus stolonifera, Alnus sinuata,
A. tenuifolia, Betula occidentalis, B. glandu-
losa, Prunus virginiana, Sambucus melano-

spp., Symphoricarpos spp., Rhamnus alni-
folia, and Acer glabrum.

. Herbaceous Wetlands along Streams: Along

fast-moving, steep-gradient streams are ri-
parian communities that are dominated by
mosses and/or a variety of herbaceous vas-

elyma, Fontinalis, Funaria, Hygrophynum,
Philonitis, and Oncophorus. Streamside herb-
dominated communities feature species of
Mertensia, Senecio, Mimulus, Heracleum,
Delphinium, Aconitum, Primula, Saxifraga,
Veratrum, and Athyrium

ASPEN COMMUNITIES

Trembling aspen (Populus tremuloides) forms

clonal stands or continuous forests under various
site conditions in all mountain vegetation zones

1987]

from 900 m to 3,600 m throughout the Rocky
Mountains. It intermixes with semiarid shrub-
ands and with wet spruce-fir forests. It is the
most widely distributed native North American
tree; aspenlike trees appear to have been present
in western North America since middle Miocene
times (Harper et al., 1985). Aspen populations
are variable, but no subspecies are recognized.
The most expansive display of aspen in the Rocky
Mountains occurs in the middle Rockies (Reed,
1971; Smith, 1985; Mueggler & Campbell, 1986),
and it forms scattered grovelands in the northern
Rockies (Lynch, 1955; Steele et al., 1981; Arno
& Hammerly, 1984).

Biological and ecological attributes of aspen in
the western United States have been recorded by
DeByle & Winokur (1985). Aspens reproduce
profusely by root shoots (“suckers”), especially
following fire treatment. Establishment from seed
is rare and may occur only during unusually wet
years. As shown in Figure 18, aspen communities
are often fire-induced pioneer or seral stages that
become invaded and replaced by conifers (Lynch,
1955), while under other conditions— beyond
conifer’s limits—the groves may perpetuate
themselves as local uneven-aged climax stands
(Steele et al., 1983; DeByle & Winokur, 1985).

Aspen communities do not readily burn, but
aspen’s thin bark makes it extremely sensitive to
fire. Although it is considered a fire-dependent
community type, aspen stands have low flamma-
bility characteristics (Mutch, 1970). Modern fire
suppression has led to an increase in mature as-
pen; young stands are not common (DeByle &
Winokur, 198

Populus tremuloides communities have been
subject to heavy use by domestic stock as well
as by elk and moose, leading to replacement of

un
—

arvense, Rudbeckia occidentale, Poa pratensis,
and Helianthella spp. (Steele et al., 1983). Pre-
vious severe browsing of aspen shoots by elk and
moose on the Yellowstone Park winter range,
together with fungi and insect damage, led to the
demise of aspen (Krebill, 1972). Recent browse
pressure reduction has allowed vigorous aspen
regrowth (Krebill, 1985, pers. comm.).

MINOR COMMUNITY TYPES

Pinus flexilis has been mentioned as a member
of both the lower and upper timberlines within

HABECK—NORTHERN ROCKY MOUNTAIN PRESENT-DAY VEGETATION

835

parts of the southern Canadian Rockies, north-
ern Rockies (confined primarily to the East Fronts
in Alberta and Montana), and middle Rockies.
It also deserves mention as a dominant pioneer-
ing tree occupying volcanic deposits (cinder cones
and lava flows) in and near the Craters of the
Moon National Monument (Fig. 4, Area 22) in
a Idaho (Eggler, 1941). Some Pseudo-
tsuga menziesii is also found with Pinus flexilis
on these extremely xeric sites, as are Artemisia
tridentata, Chrysothamnus nauseosus, Purshia
tridentata, and Ribes cereum.

communities occupying localized mi-
crosites, such as talus/scree, ponds, bogs, snow
glades, rock outcrops, rock crevices, vernal pools,
and sand dunes have been described by Chad-
wick & Dalke (1965), Daubenmire (1970, 1975,
1978, 1980), and by others. Throughout the
Rocky Mountains, there are numerous special-
ized habitats that support minor communities
often composed of rather unique combinations
of plants. Some of these species have been iden-
tified by Rocky Mountain botanists as rare and/
or endangered. Furthermore, some parts of the
northern Rocky Mountain region support nu-
merous geographic disjuncts and endemics, such
as coastal Cornus nuttallii and Alnus rubra pop-
ulations; these occur in the intermountain ranges
in northern Idaho (Daubenmire, 1943, 1975;
Johnson, 1968; Johnson & Steele, 1974).

CONCLUDING REMARKS

The Rocky Mountain vegetation exhibits vari-
ations in its local and regional distribution pat-
terns as a result of the diversity associated with
this region’s physical setting and a variety of his-

namic nature of vegetation is the basis of its re-
sponses to spanning mil-
lions of years. Any modern description of the
Rocky Mountain flora and plant communities is
really only capturing a “moment” in an ever-
changing phenomenon. We have every reason to
believe that, within an historical context, the
mountain vegetation zones described in this re-
view have shifted altitudinally and latitudinally

adds to the diversity of the Rocky Mountain flora
but may also contribute to the creation of new
vegetation equilibria.

836

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COMPARATIVE AGE OF GRASSLAND AND STEPPE EAST
AND WEST OF THE NORTHERN ROCKY MOUNTAINS!

E. B. LEOPOLD? AND M. F. DENTON?

ABSTRACT

Given the taxonomic and biogeographic differences between dominant species in grasslands of the
Great Plains and the palouse grasslands and steppe of the
two biomes have had separate origi l
of the Rocky Mountain cordillera and pollen data from the cordillera area suggest that three distinct
floristic provinces were in existence by the beginning of mid-Miocene (Barstovian) time. Montane
conifer forest poor in genera typified the Rocky Mountain a and nearby pesi while mixed
conifer-deciduous hardwood forest and Taxodium swamps rich in woody genera occurred in the
Columbia Basin. On the Great Plains the Kilgore flora (Barstovian of Nebraska) iugo deciduous
open forest and prairie dominated by species of eastern and southern affinities. Younger Neogene
floras in the Pacific Northwest suggest that steppe and local grassland were beginning to be important
in the Pliocene of the Pacific Northwest, while grasslands and open forest became widespread on the
Great Plains somewhat earlier (late Miocene or Clarendonian-Hemphillian time). The persistence of
mixed conifer and broad-leaved forest on the Columbia Plateaus suggests that this area was open to
the west through about 8 Ma. Due to the increasing height of the Cascade Rangea nd/or regional up-

According to the regional pollen record, terrestrial herbs become more diverse in the late Neogene.

Recent fossil evidence has helped substantiate
that the grasslands and steppe west of the
tinental Divide in the Columbia Basin area are
about three million years (Ma) old, about ten
million years younger than similar vegetation
types east of the Divide in the Rocky Mountain
foothills and in the Great Plains. Biogeographic
differences between the two areas are reflected in
species composition and in the dominant habits
of grasses. Some climatic differences between
‘east’ and ‘west’ exist, but they do not seem suf-
ficient to account for the vegetation contrasts.

We propose that an examination of the Late
Cenozoic history (Miocene to present) of the
northern Rocky Mountains may help illuminate
the nature of origins of the grassland and steppe
east and west of the Divide. The region to the

America, while the region to the east, the Rocky
Mountain foothills and the Great Plains, have
Mio-Pliocene deposits for which pollen, leaf, and
seed data are now available. To understand the
history of grassland and steppe development, be-

cause they hinge on the present east/west con-
trasts, the following questions need to be an-
swered: 1) When did grassland and/or steppe first
develop in the northern Rocky Mountains area?
2) What were the regional patterns of climate
during the time the grasslands developed? The

Pliocene) vegetation inferred from pollen or leaf
data. Species identifications from the leaf and
seed floras give an index of floristic patterns that
can be compared with the perceived regimes.

It is our thesis that three distinct vegetation
provinces had developed in the northern Rocky
Mountain region by early mid-Miocene time (ca.
16 Ma): 1) a rich deciduous hardwood and mon-
tane conifer forest west of the Continental Divide

mbi

with steppe elements in the Rocky Mountain
foothills; and 3) a deciduous forest with ancestral
grassland elements to the east and in the Great
Plains.

If vegetation types east and west of the cor-
dillera were basically different during the Mio-

! We thank Daniel Axelrod, H. D. MacGinitie, seal s“: Malde, Jack A. Wolfe, and Robyn Burnham for

their comments, and M. Kay Suiter for typing the ma
2 Department of B

3 “Columbia Plateaus”

ANN. MISSOURI Bor. GARD. 74: 841—867. 1987.

Botany, University of Washington, corns “Washington 98195, U.S.A.
is the official physiographic usage by the U.S. Geological Survey (1981).

842

ANNALS OF THE MISSOURI BOTANICAL GARDEN

FIGURE 1. Present distribution of the chief grass-
and vincesin North America according
to dominant a io om Daubenmire, 1978).—A, B.
Andropo. gon scoparius province. — C. Festuca scabrella
province. — D. Bouteloua gracilis province. — E. Agro-
pyron spicatum province

cene, the importance of this massif as a floristic
barrier can be evaluated. Did the same species
occur east and west of the divide or were they
different? How do these vegetation patterns re-
late to the Neogene history of the region and to
the development of grassland and steppe biomes?

PRESENT VEGETATIONAL PROVINCES

Figure 1 illustrates the outlines of ied
grassland and steppe types in the U. each
side of the Rocky Mountain cordillera mus grass-
lands of different character. (1) At the eastern
margin of the mid-continent grassland, there is
the tall-grass prairie, a sod-forming grass asso-
ciation with rhizomatous root habit. The affin-
ities of the dominant taxa, Andropogon scopar-
ius, Andropogon gerardii (little and big bluestem),
Panicum spp., and Sorghastrum nutans (Indian
grass), lie to the south, in Central America, Mex-
ico, and even South America. (2) The short-grass
prairie immediately east of the cordillera, char-
acterized by the dominance of Bouteloua gracilis
(buffalo grass), Aristida spp., and others, is a
province wit izomatous and ESAE
grasses. Most of the dominant taxa have t
floristic affinities with the intermontane dn
of the Rocky Mountain region. (3) To the west
exists the palouse grassland and steppe of Idaho,

[Vor. 74

X — Ns
AT T Sa.
( mM a c
150 Y X \ — ——— 5
I N J j c

Agropyron spicatum |

FIGURE 2.
catum (after Hultén, 1968; Hitchcock et al.,
1972).

Present distribution of Agropyron spi-
1969; Voss,

eastern Washington, and eastern Oregon (Fig. 1),
characterized by bunch grasses and steppic ele-
ments such as Atriplex, Artemisia, Sarcobatus,
and other diverse desert-scrub genera. The bio-
geographic affinities of taxa in the palouse grass-
land and steppe are with areas to the north; for
example, Agropyron spicatum ranges northward
to Alaska and boreal regions (Fig. 2). Festuca
idahoensis has its nearest relatives, the F. ovina
omplex, in the arctic and steppes of North
America, Asia, and northern Europe (Fig. 3). Ar-
temisia cana (Fig. 4) and other members of the
A. tridentata group are mainly arctic or boreal.
Climates east and west of the Continental Di-
vide differ significantly, chiefly with respect to
the amount of summer precipitation. On the
Great Plains summer rain emanates from trop-
ical air masses moving northward from the Ca-
ribbean across this region during June and early
July (Bryson & Hare, 1974). Along the southern
cordillera summer rainfall occurs, especially dur-
ing warm years, along the north-south path of
the “Arizona Monsoon" (Neilson, 1986; Neilson
& Wullstein, 1983). To the west, the palouse
grassland and steppe region is characterized by
l

e

area is dominated by the
Pacific air in summer and is in the rain shadow

1987]

FIGURE 3.
Hultén, 1968; Hitchcock et al., 1969

of the Cascade Mountains; here, the distribution
of grassland and steppe is limited to areas re-
ceiving less than 15-20 inches (380-500 mm) of
annual precipitation, most of which falls in win-
ter (Fig. 5).
The arand provinces east and west of the
Rockies are now dominated by separate taxa dif-
fering in geographic affinity and in growth habit
(Table 1). These grasslands differ in distribution
of C, (warm growing season) and C, (cool grow-
ing season) grasses. C, grasses are always of low
frequency (less than 1896) west of the Rockies
t may dominate in certain areas east of the
Rockies (Mack & Thompson, 1982; Teeri &
Stowe, 1976). In addition, there are fundamental
differences between these two grassland prov-
inces with respect to their carrying capacities for
large, grazing ungulates; for example, the carry-
ing capacity of the palouse grassland dominated
by bunch grasses sensitive to trampling is much
lower than that of grasslands east ofthe cordillera
(Mack & Thompson, 1982). The biogeographic,
physical, and climatic contrasts of these regions
imply that thei t must have
been very different. This is the subject of the
discussion that follows.

DATA BASE

Because our study concerns events after the
early Miocene, we discuss data from sites youn-
ger than 18 Ma (Hemingfordian and younger
stages). The sites are mapped in Figures 6 and 7

LEOPOLD & DENTON —GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

843

Present distribution of Festuca idahoensis and the closely related species Festuca ovina (after
).

and are arranged according to geologic ages in
Table 2

West of the Rocky Mountain cordillera, the
area of the Columbia Plateaus contains a wealt
of well-documented fossil plant sites. We have
studied data from 15 of these, of which four have
yielded pollen data (Fig. 6). Dating is established
by K/Ar isotope ratios for a few of these and by
land-mammal evidence for others.

150

Artemisia cana

E 4. Present distribution of Artemisia cana
(after Hultén, 1968; Hitchcock et al., 1955

844 ANNALS OF THE MISSOURI BOTANICAL GARDEN

20“ ISOHYET -——-—
15“ ISOHYET ------.

==
80

0
SCALE IN MILES

FOREST OR WOODLAND
AREAS øm

[Vor. 74

Relationship of the forest/steppe border to the 15- and 20-inch (380 and 500 n isohyets of

"en ETE in the Pacific Northwest (after Sherman, 1947, and U.S. Weather Bureau data

In the foothills of the northern and e
Ps Mountains, we have four localities (Fi

11

which the main documentation is His

fossil NS although some megafossil dataexist data.

Ln of taxa is on the generic level only).

y scattered, but their
eoi ages were ascertained by land-mammal

120 110 100
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aen | Y | NORTH DAKOTA )
46 . Ellensburg @ ; ` | N
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Hanford Clarkia 7 MSNIANA ! n —2
Cone ; | sme S
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Deschute Ve i
eschutes g Mascall IDAHO N m 4
a permet eee err MM |
ii UNS | SOUTH DAKOTA
lo SS WYOMING |
Stinking Water Í W Horseshoe Bend e Teewinot i
S [e k ^l GI F | cs cu de
Uccor reer: ; enns Ferry —
42 | | OREGON Trout Creek Moonstone s Split Rock |
ul . - 1 oper LP! \
pesi eas ONO _ i NEBRASKA
i T = Saratoga |
! e i
! I “SZ = E a
40 . ! —— nem
I i | Troublesome v7
| | e
! : I i
NEVADA ! UTAH i i
| i COLORADO \ KANSAS
ES ! : :
38 E i :
E : |

FiGuRE6. Locations of Miocene and Pliocene fossil flora sites in the northern and central Rocky Mountains
and Columbia Plateaus. Bi = le ollen

af localities; @ = po

sites; A indicate leaf and pollen data from the same site.
Coal Mine Gulch (not shown) is 50 miles north of Succor Cree

1987] LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

In the region east of the cordillera, there are
several megafossil localities (Fig. 7) containing
small leaf floras and there are abundant fruit and
seed localities (Elias, 1942); recent work with
pollen analysis and electron microscopy of fruit-
ing parts, grass anthoecia, epidermal patterns,
and phytoliths revealed the composition of many
of these assemblages.

MIOCENE VEGETATION PROVINCES OF THE
NORTHERN RocKY MOUNTAIN REGION

Three vegetation provinces had developed by
the mid Miocene in the northwestern and north-
central mid-continent.

I. WEST OF THE ROCKY MOUNTAINS — THE
COLUMBIA BASIN

A. MIOCENE FLORAS

Fifteen leaf floras of mid and late Miocene age
demonstrate that mesic forest vegetation under
a warm-temperate summer-wet climate existed
in the Columbia Basin by about 18 Ma until
Hemphillian time (about 8-4.5 Ma). Difference
in warm- and cold-season average monthly tem-
perature has been estimated at about 20°C Mae
1978). Even though important changes in cli-
mate and vegetation occurred through the main
sweep of Miocene time, the major elements of
the flora were not eliminated. Some chieffeatures
of the vegetation derived from the leaf flora are:

(1) Dominant vegetation (Trapper Creek? of
Clarendonian age is an example; summary in
Table 3) was deciduous hardwood forest and
mixed montane conifer-deciduous forest, with
some broad-leaved evergreen elements and di-
verse (8-26 genera) woody dicots. Shrubs were
important (up to 3096), while herbaceous groups
were few (only four taxa and these were generally
rare aquatics).

(2) Species showed close relationships with
of summer-wet areas in eastern

about equal percentages. For example, at Trap-
per Creek modern affinities seem split between

* Axelrod (1964) considered the age of the Trapper
Creek flora as 15-16 Ma. More recent evidence from
K/Ar dating (Armstrong et al., 1975; Fields, 1983) sug-
gests an age of 10.5-12 Ma.

Chief characteristics of modern grassland types in the U.S.A.

TABLE l.

EAST

Rockv MOUNTAINS

Tall-grass prairie

Short-grass prairie

Grassy steppe

Agropyron spicatum

P

KEY SPECIES:

Andropogon scoparius

Bouteloua gracilis

a

O
Festuca idahoensis

Panicum virgatum

Sorghastrum nutans

Sporobolus

(southern affinities)

(intermontane basin affinities)

(north temperate & boreal affinities)

DOMINATED

MAINLY BY:

rhizomatous/stoloniferous grasses

caespitose (bunch) grasses

summer moist

summer dry

CLIMATE:

845

846 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74
: nm (e mmm (c) Valley forest, including some of the taxa
DAK 1 . :
ar TIT 3 en s : mentioned above and a wide range of woody
| oshin Drank n deciduous groups such as Alnus, Amelanchier,
» IOWA : :
WYO, | Ash Hollow fm2 9 i runus, Parthenocissus, Cornus, Fraxinus, Ul-
BENE € ) mus, Pterocarya, Carya, and Sophora as well as
— NN. ! y
uu e ------ -| an important component of broad-leaved ever-
eWray j S 404 greens, such as Quercus (cf. Q. chrysolepis), Sas-
E s. safras, Berberis, Ilex, and diverse conifers such
ogan Co. ' š ; ;
COLO, |@Wallaceco@ Ellis Co. T as Keteleeria, Picea (cf. P. breweriana), Pseudo-

tsuga, Tsuga, Abies, and Sequoiadendron.
(d) Mountain-slope forest, including many of
the conifers and hardwoods mentioned in the
ae. 9 Beaver I------1 valley forest, also Garrya (G. cf. fremontii), Rhus,
Ghian x Ungnadia, Abies (A. cf. delavayi or A. recurvata),
° | ! Abies concoloroides, Calocedrus (C. cf. decur-
| i ARK rens), and most importantly, Pinus (P. cf. pon-
| derosa, P. cf. monticola), and Cedrus.
Some of the most frost-sensitive forms (Liq-
ane Vi, | Uidambar, Cedrela) seem to disappear during the
' mid Miocene, especially in regions close to the
Location of Miocene and Pliocene fossil Rocky Mountains. However, the occurrence of
flora sites on the Great Plains (High Plains of Axelrod, warm temperate Taxodium swamp vegetation
1979). persisted in eastern Washington until ca. 8 Ma
and in Idaho until ca. 12 Ma.

LJ
Clarendon

NEW MEX.

M ee s

FIGURE 7.

Although Miocene communities typically in-
these three areas (Axelrod, 1964). There were cluded species of eastern Asian and eastern
only minor temporal changes in the role of these American affinities, western American elements
geographic elements through mid-Miocene time seem to dominate in the montane slope forest
(Table 4). communities.

_ GB) Vegetation was as Chiefly woody. Character-

Chaney (see Chaney & Axelrod, 1959: 53) dis-
forthe mid- cussed the possibility that open savanna or prai-
Miocene floras of southern Idaho included: rie vegetation existed at some mid-Miocene sites

(a) Swamp forest, particularly Taxodium where pollen and leaf data are available, e.g.,
swamp with associated Nyssa, Liquidambar, | Mascall and Stinking Water floras of eastern Or-
Persea, Salix, and Alnus.

egon. Part of his rationale was based on the di-
e-border woodland, with Quercus sim- verse fossil mammals whose modern relatives
ulata and species of Acer, Betula, and Popu- livein savanna habitats today ieee horses,
lus.

rodents, lagomorphs, camels, and most abun-

E 2. Stratigraphic ages of Late Cenozoic floral localities of the Columbia Plateaus, northern Rocky
Mountains, and Great Plains.

EASTERN | EASTERN

SOUTHERN|NORTHERN| WYOMING | NORTHERN
Ma OREGON WASH. IDAHO IDAHO COLORADO| NEBRASKA
uaternary | Bruneau
Pliocene ` |Blancan Glenns Ferry
ain Deschutes
è Hemphillian Chalk Hills
E]
_ Hanford Banbury Basalt ering
" Clarendonian | Ellensburg | rappar Creek am Hollow
e 12 Stinking Water oison Creek
Š Trout Vantage
© 3 Barstovian nad Payette Saratoga Kilgore
s _ Jj _ Clarkia _ _ | Sheep Creek
16
> Hemingfordian ist OAE
Ww 18

1987]

Dominant vegetation types in Late Cenozoic floras of the Columbia P

LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

847

t (from Axelrod, 1964;

TABLE 3.
Chaney & Axelrod, 1959; Graham, 1963; Leopold & Wright, 1985; MacGinitie, 1933; Smiley & Rember, 1985;

Taggart et al., 1982)

Barstovian Clarendonian Hemphillian
LJ
°
v S 5 E
S a D S x " °
x x e OQ 8 a Š O E >
o 9 m m z 905 š g D Ble
5 8 S o & w d uc pt y Jat Els
O ` S @ €t Ss qd Bou 9 £ afla Els
= o ` oO S Š ‘9 o A < © o o|8 gje
= Ó GY É uü ^? > à 3 £ 285 SS
e 9 8 6 8 8 S E S S s os Sie
B à D$ > OE 4d doe OF «BO Om
Mixed deciduous evergreen
hardwood forest X
Deciduous hardwood forest X X X X X
cotone X X X
Montane conifer-deciduous
hardwood forest X X X X X
Ecotone X
Montane conifer forest X
Grassland and conifer forest X
Steppe and conifer forest X X
Steppe X
Taxodium swamp forest X X X x x x

dant, the oreodonts). Direct evidence, however,
is limited. Herbaceous plants are an important
elementin modern savanna. Aside from aquatics
and ferns, herbaceous groups are rare in the Mio-
cene leaf record of the Columbia Plateaus. Even
pollen evidence indicates that herbaceous groups
were limited in diversity and abundance (Ap-
pendix I); in the Mascall only two nonaquatic
herb types were identified by Jane Gray (in Cha-
ney & Axelrod, 1959: 43): “Pollen of Gramineae
h

were found, but these, as in the leaf flora, were
all from woody taxa.

The floristic role of herbs in mid Miocene of
the region is illustrated in the pollen lists from
the Succor Creek Formation (14-12 Ma at the
type section; Fields, 1983) in southeastern Ore-
gon (Taggart & Cross, 1980). Six probably
terrestrial herbaceous groups are recorded:
Pachysandra, Ambrosia, Onagraceae, Amaran-
thaceae, Gramineae, and Umbelliferae (Appen-
dix I). Because the pollen diagrams summarize
their abundance according to broad ecological
groupings, the relative importance of herbs is not
documented. In their discussion Taggart & Cross
made it clear, however, that grass and Compos-
itae pollen are sporadically abundant as part of
a successional cycle (see below).

TABLE 4. Percentages of element representation in mid-Miocene floras and age groups, Columbia Plateaus
region (from Chaney & Axelrod, 1959, table 32; Axelrod, 1964, table 5).

Barstovian Clarendonian
uccor Trout Stinking Trappe Lower
Geographic Elements Mascall Creek Creek Water poe Ellensburg
Eastern American 65 61 50 58 58 70
Eastern Asian 50 50 50 53 69 45
Western American 37 37 56 60 62 58
Total taxa 64 46 46 38 61 32

848

A. SUCCOR CREEK
EASTERN OREGON

% total pollen

measured er m
ection section
===

E3
Locustrine F -—

ANNALS OF THE MISSOURI BOTANICAL GARDEN

14-12 Mo

[VoL. 74

B. SUCCOR CREEK MODEL
PALEOASSOCIATIONS

Generalized pollen diagram indicating vegetation phases in a 200—meter section at the Succor
of

E 8.
C 2 "fora type locality
shown on right (from Cross & Taggart, 1982

(4) Vertical arrangements of plant commu-
nities were suggested by pollen data. Many leaf
floras do not demonstrate clearly that an altitu-
dinal zonation occurred in a the Miocene floras,
yet pollen-st |
indeed. At Succor Creek, Oregon, Taggart & Cross
(1980) and Taggart et al. (1982) showed a re-
peated successional cycle of montane conifer for-
est, bottomland/slope associations, and xeric
shrub with steppic elements followed by Pinus
spp.; the sequence then reverts to swamp, bot-
tomland/slope, and then to montane conifers. A
single cycle in a 200-meter section is shown in
Figure 8. The dramatic oscillations can readily
be interpreted as elements from various altitu-
dinal in an altitudinal

tcertain

succession caused by climatic changes. Taggart
et al. (1982) believed the successional changes
are related to disturbance episodes of volcanism,
but it is also possible that these are forced by
small oscillations of climate, or both.

Ata much younger (ca. 8.5 Ma) site at Hanford
in eastern Washington, floodplain sediments
(lowest Ringold formation) of the ancestral Co-
lumbia River record a somewhat similar succes-
sion (Fig. 9a, b; Leopold & Nickmann, unpubl.).
The sediments overlie the Upper Columbia Riv-
er Basalts (10.5 Ma) and an unconformity dated
at 8.5 Ma (Tallman et al., 1981; DOE, 1986: 3-
40). The lithology indicates that local deposi-
tional environments were changing. In a series

mid Miocene age, eastern Oregon (A); a model of the successional sequence (B) is
).

of cycles, fine-grained swamp deposits grade up-
ward to increasingly coarse riparian sediments.
Two pollen diagrams (core holes DC-3 and DC-
7/8), each beginning in fine-grained sediments,
register a rich Taxodiaceae-type (cf. Taxodium)
swamp association. (We infer this identification
because Taxodium is abundant in the underlying
Ellensburg Formation.) This phase is followed

i int tt l d/sl I 1 d and

1

by p ;
finally Cedrus and Pinus dominate. Herbs and
xeric elements are always present in trace
amounts. Presumably in a floodplain area such
as Hanford, the vegetation changes record either
shifting riparian environments (edaphic factors)
or changes wrought by climate. While the ele-
vational relief of each of these forest types oc-
cupied is not clear at either site, it is possible
that as much as 500-1,000 feet of relief existed
at Succor Creek.

The Cascade Range was probably rising during
Miocene time (McKee, 1972; Smiley, 1963), and
its rain shadow eastward eventually changed the
ici of the vegetation from mesic and sum-

r-wet to xeric and summer-dry. The lower
Ellensburg and the Hanford floras suggest that
eastern Washington was open to the west through
ca. 8 Ma. This meant that the Cascades were not
significant enough to block moisture from the
westerlies until some time after the Clarendon-
ian. In part this helps explain the general simi-

1987]

larity of Pacific Northwest floras during the early
and mid Miocene.

B. IMPLICATIONS FROM PLIO-PLEISTOCENE
FLORAS

Given the data above concerning the charac-
teristics of the Miocene forests of the Columbia
Plateaus, when did forest vegetation diminish,
allowing the development of grassland and
steppe?

West of the Rocky Mountains, pollen data from
Idaho demonstrate the decline and impoverish-
ment of the Miocene forests and the develop-
ment of local grassland and steppe. These changes
that occurred from late Pliocene to Quaternary
were surprisingly late. In lake and stream de-
posits of southwestern Idaho a unique and well-
dated composite pollen sequence embraces parts
of the last 11 million years, after Trapper Creek
time through the early Quaternary (Fig. 10; Leo-
pold & Wright, 1985). Fossil pollen in these de-
posits tends to be scarce, and pollen-bearing beds
are hard to find (90% of our collections were
barren). Some megafossils have been identified
at certain sites.

In the sequence (Idaho Group) the lower sed-
iment units are from the Poison Creek, Jenny
Creek, and Chalk Hills formations and Banbury
Basalt of mid and late Miocene age (Armstrong
et al., 1975; Fields, 1983; Leopold & Wright,
1985; Appendix II). The floras record mixed de-
ciduous and conifer forest with declining hard-
woods; these were mainly U/mus but also in-
cluded Pterocarya, Carya, and Juglans. A
holly-leaved oak (leaf evidence from the Poison
Creek Formation) is reminiscent of that recorded
at Trapper Creek (ca. 11 Ma). Wood from the
Chalk Hills Formation records diverse hard-
woods. Younger sediments of the Glenns Ferry
Formation (ca. 3-2 Ma) containing the Hager-
man lake beds record an impoverished pine and
mixed conifer assemblage with rare pollen of ex-
otic hardwoods. Steppe elements (Sarcobatus and
other Chenopodiaceae, Artemisia and other
Compositae) are consistently present and in-
crease sporadically upward in this section. Peaks
(up to 60%) of grass pollen are associated with

clines. Taxa of terrestrial herbs are more diverse
than in the Miocene (Appendix I).

LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

849

Miscellaneous Pleistocene samples from the
area suggest that Artemisia and Chenopodiaceae
are important, if not the dominant, pollen forms.
Above an unconformity (see top of Fig. 10) Ir-
vingtonian g Equus plus K/Ar

evidence date the Bruneau Formation as middle
Pleistocene; a Bruneau pollen sample (D1694)
shows Artemisia to be 50% of the count, sug-
gesting a true Artemisia steppe had developed.
Two other samples rich in Artemisia pollen
(D1120and D1698; Fig. 10), previously reported
as Bruneau (Leopold & Wright, 1985), are now
classified as Yahoo Clay of late Pleistocene age
(Malde, 1982). The top fossil sample (D1697),
showing 80% Artemisia and other Compositae
pollen, is from a late Pleistocene soil above the
King Hill Basalt.

In northern Oregon the Deschutes flora (ca. 4—
5 Ma) suggests low-diversity riparian vegetation
typical of unforested regions and is consistent
with a decrease in summer precipitation in late
Miocene time (Appendix III; Chaney, 1938).

Taken as a whole, the data indicate that steppe
in the Columbia Basin probably did not develop
as a major vegetation unit until after the Hemp-
hillian (4.5 Ma). The Snake River Plain section
places the change from rich (deciduous and) co-
niferous forests to montane conifer forest be-
tween 10 and 3 Ma. In this region steppe and
palouse grassland probably became widespread
for the first time in the Quaternary.

II. THE ROCKY MOUNTAIN FOOTHILLS

The lack of megafossil evidence for northern
Rocky Mountain Neogene sites is unfortunate
(the well-documented Clarkia flora of middle
Miocene age lies in the Columbia Basin floristic
province). Pollen records from the Rocky Moun-
tains and eastern foothills demonstrate the com-
paratively modern aspect of plant communities
there during the Miocene.

At Jackson Hole, Wyoming, the Teewinot lake
deposits (predating the Teton Range) provide a
long record of montane conifer forest with oc-
casional bursts of lowland steppe and riparian
types (Fig. 11; Barnosky, 1984). Presumed plant
communities include:

(1) Saline basins. High percentages of Sar-
cobatus pollen accompany other Chenopodi-
S

These are char-
acterized by pollen of probable phreatophytes,

ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

850

HANFORD RESERVATION
SHINGTON
ate Miocene
£ $
oF Oe RAE
ES

7 TOC
9 kA

Y sd!
JI

E E | |
iM XE ! i
na i L P
IE s E
ES ud pe
de Jn DE
IT tei cold :.

KASS SPS Da a
E" == Walli SILT
ee SILTY - == ae CLAY
SAND

SPORES-
ra

tlla Lbs
from lowermost Ringold Formation, % total pollen (Leopold & Nickman, unpubl. data). Stratigraphic data are
from Department of Energy (1986: 3-16) drill holes DC-3 and DC-78, which are correlative in age and about
12 miles apart. Depths below surface are given in feet. The sediments are dated at 5-10 Ma (probably 8.5 Ma).

FIGURE 9A. Late Miocene pollen sequence, Hanford, Benton County, Washington. Two pollen diagrams
h

1987]
HANFORD FOREST ASSOCIATIONS

8.5 Ma E. Washington

Taxodium Bottomland /

Swamp lope Cedrus Shrubs

Forest Hardwoods & Herbs
+

4
+
3
+
2
+
+
1

+
+

FIGURE9B. Schematic phases of vegetation shown
by Figure 9A pollen diagrams.

Carya, Ulmus-Zelkova, Pterocarya, Sapinda-
ceae, and Salix. _

(3) Montane slope forest. This assemblage is
dominated by Pinus spp. with important amounts
of Abies, Picea, and Cupressaceae pollen. Tsuga,

LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

851

Quercus, Artemisia, grasses, Onagraceae, and ad-
ditional herbs may have been locally abundant
(Leopold & MacGinitie, 1972: 198)

The proportion of broad-leaved Tertiary relict
genera that are now exotic (eastern Asian and
eastern American genera such as Pterocarya,
Carya, and Ulmus-Zelkova) obviously is low
compared with those in pollen and leaf assem-
blages of the same time period west of the Rock-
ies (e.g., Hanford, Poison Creek, and Trapper
Creek). Except for the “riparian” hardwoods, the
flora has a modern aspect indeed, as it compares
well with modern pollen rain (see top of Fig. 11).

Four other pollen sites from widely different
times in the Miocene are in basins along foothills
and in the eastern Rockies of Colorado and Wy-
oming (Fig. 12; localities in Appendix IV; se-
lected pollen counts in Appendices V, VI; Leo-
pold & MacGinitie, 1972). Early Miocene
(Hemingfordian) sites are the Troublesome For-
mation from Grand County, Colorado (Izett,

MODERN: D mw 8)
SAMPLE NO. CONIFERS TERT PEHE! eels ae SHRUBS
01097 ob ET! +
ui
D1694 r:
D-1695-2 £3 s =
11698 aa Xe
D1120 T es CJ 0 Fog
D1671 [ i
11695-4 32a E 2 + | um rm Lan m — Te
. 1 SAGE & | SALT *
01710 L — | iR Q |t [t | WALNUT, |COMPOS- BUSH
z la +O *0* ^u +x HICKORY, GRASS -
E 2
D1715 AAR > € A = +a Š «c |+Ə | LING NUT, o
D1712 ae x + ha a = | OAK E
1: uj + +
uj ore MAPLE 3
D1711 u. + |+ h + q
D)713-1 2 ' + + Je
z
D1692-5 u + +
Di692-4 e +
D169) r + +
D1690-1 + x od |
INANA q P
nisse > g =) +
2 PINE
£ uj + +
I| +
n1208 FX + © pi
OL———3À *
D1699 OY — ——i
(0) 80% 015 o 30 O 80 (0) TO O IOO %

FIGURE 10. C omposite pollen diagram, late Miocene to late Pleistocene, Hagerman area, southwestern Idaho.
old ght,

n is shown

Note that Pleistocene samples D1120 and D1698 are pu EAE fied as from

Sample D1695-2 is Glenns Ferry Formation (Malde,

at top _ a S

& Wri 1985). Fossil wood indicated along
is Quercus, white oak type (Malde & Powers, 1962).
Yahoo Clay of late Pleistocene age.
982). Sample D1697 is from a Late Pleistocene buried

soil on the King Hill Basalt. Sample D1694 is of the ed Formation of middle Pleistocene age. Celtis fruits

(see symbol) are of Late Pleistocene age.

852 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

Lower slopes Valley Basin floor

Samples =]
1 m

S $
š

° E

8 ©

g Š

š Fle- š

- o

š dE E E

v

= >

- =
m m m

io
Total pollen (99

FIGURE 11. Pollen diagram, showing selected types and inferred habitat, upper Teewinot lake beds, Jackson
Hole, Wyoming (% total pollen). K/Ar age is 9 Ma or late Miocene (from Barnosky, 1984).

1968), and the Split Rock Formation from the est, or whether most of these were restricted to
Sweetwater Basin, Natrona County, Wyoming riparian environments cannot be deduced from
(Love, 1961). A mid-Miocene (Barstovian) site the available information, though the latter is
is from the Saratoga Valley, Carbon County, Wy- more proba

oming. The youngest site, the Moonstone For-
mation in the Sweetwater Basin, Wyoming, may
be of Barstovian or younger age (Robinson, 1971;
Love 1961). All are old lake beds near low rolling
granite or bedrock hills that probably existed in
the Miocene. Pollen is moderately abundant to
rare and demonstrates impoverished floras sim-
ilar to that evident at the Jackson Hole, Wyo-
ming site. Only two types of communities can
be conjectured from the data:

Implications. The consistent presence of xe-
ric and desert-scrub taxa that are sometimes
abundant suggests the appearance of steppic
vegetation with a diverse herb flora during widely
separate times during the Miocene. Grass pollen
grains are usually present but never abundant.
They are probably associated with depauperate
conifer forest or woodland. Several other basin
sites in the central and southern Rockies are con-
sistent with the modern aspects of these Miocene
floras (Meyer, 1986; Leopold & MacGinitie,
1972)

The Rocky Mountain data stand in stark con-
trast to the Miocene basin sequences from Idaho
westward to eastern Washington where pollen
and leaf data alike point to rich forest vegetation
containing abundant and diverse deciduous

ardwoods. Broad-leaved evergreen trees are ap-
parently ME in the Neogene of the Rocky
Mountain

(1) Open basin and lakeside environments in
which Artemisia, Sarcobatus and other Cheno-
podiineae, Ephedra, and Eriogonum suggest
steppic and halophytic environments and in
which Salix, Betulaceae, and aquatics imply lake-
margin or riparian environments. Terrestrial
herbs of the Polemoniaceae, Compositae, Ona-
graceae, Umbelliferae, and Polygonaceae may
have grown in these basin environments (Ap-
pendix VII).

(2) Mountain slopes dominated by Pinus with
lesser amounts of Picea, Abies, and Juniperus.
Whether Juglans, Carya, Quercus, and Ulmus- Megafossil floras from the Great Plains region
Zelkova were associated with woodland or for- (Fig. 7) suggest that prairie elements were present

HI. GREAT PLAINS LOCALITIES

1987]
S
q`
E
z
°
=
$
° wawam m+ | o: + + °
<
^ ww mj bos i
ae
a
<
o
e

E ERIOGONUM
x + e
[5] ARCEUTHOB.
° L
ac
š pe
o T

ERIOGONUM
É
š
i POLEMON
s
° gi ae la a mm FETI r? mM
= O 50 0 30

FIGURE 12.

Rock Formation, Wyoming, lower Mi
Wyoming, of younger Neogene age (Appendix V, VI).

and that grassland developed to various degrees
during the Miocene (Elias, 1942; Chaney & Elias,
1938). An excellent summary by Axelrod (1985)
portrays a sequence indicating a generally de-
creasing Lodge Saga regime from about 16 Ma
onw older sites are deciduous hardwood
forest se prairie elements (Fig. 13). According
to Axelrod, the younger sites suggest woodland
or riparian border hardwoods with more exten-
sive grassland (Fig. 13)

From the Neogene sediments of the upper
Arikaree and Ogalalla groups on the Great Plains
and High Plains, Elias (1932, 1935, 1942) made
systematic fruit/seed and leaf collections at al-
most 100 localities from South Dakota to north-
ern Texas. The widespread sediment layers with
occasional volcanic ashes and vertebrate fossils
provided stratigraphy. Elias undertook to define
a sequence of fossil seed zones. Modern dating
indicates that the main part of this sequence
ranges from Hemingfordian through Hemphil-

LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

853

—9$—€9—
— —e-
—e—e-
E End

=]
I]
Pl of PE Eos
r trr m rr? rn FT T T T |
0 40 o 100

Pollen spectra from four Rocky Mountain sites: Troublesome Formation, Colorado, and Split
iocene; Saratoga Valley, Wyoming, mid Miocene; Moonstone Formation,

3? e
1000 SOS ea
Sheep * NE eC "I
Neb. s> Ee ow

e
S
Kilgore, Neb. iJ al ss”
e Eu

Beaver Com ~~

Q
o

Oki. ` rA V
Clarendon; anning},
Tex." al Tx. UY

N
o

Annual Precipitation

Logan Co., Toda
Kan. day
HIGH PLAINS

10 5

million years B.P.

FIGURE 13. Fossil floras from the Great Plains (High
Plains of Axelrod, 1979) suggest a gradual reduction
in precipitation during the late Tertiary

854

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

TABLE 5. Geographic affinities of the Kilgore flora, Nebraska (15-12 Ma, mid Miocene), showing the rep-
resentation of its species in Miocene floras of the Columbia Plateaus (after MacGinitie, 1962).

Modern Distribution Groups

Number of

Species in
Kilgore Number of These
Flora by Species oe

East of Rocky Mountains only 10 l
Mexico and southern Rocky Mountains 7 l
Rocky Mountains only 3 0
West of Rocky Mountains 2 0
Split affinities between distinctly eastern and distinctly western species 3 1?
Cosmopolitan 1? 1?
Eastern Asia 2 2
Total number of species: 27-28 4

6
or ca. 18-21%

lian (ca. 18-4 Ma). At most of his sites in the
Ogalalla Group (Kilgore and younger age) fruit-
ing parts of grasses and prairie herbs were the
most common and in most cases the only fossils,
although arboreal leaves were present at a few
sites (most ofthe latter were summarized by Cha-
ney & Elias, 1938, and by Axelrod, 1979, 1985).

Elias's identifications, corroborated in large
part by the recent work of Thomasson, demon-
strate that ancestral Stipeae appear in the earliest
Hemingfordian strata or earlier, and that Borag-
inaceae (Biorbia, Krynitzka) and Paniceae ap-

Hollow Formation) sites. Diversity of
prairie taxa increases up section

erhaps the most pang feature is the
widespread nature of Elias’s fossil fruit and seed
zones. In Nebraska the following zones range
from Kilgore age to Ash Hollow and younger
units (Kimball Formation) of the Ogalalla Group
(in stratigraphic order; Elias, 1942):

VI. Kimball Formation (calliche).
. Biorbia fossilia with Krynitzka coroniformis
and Stipidium grande (Ash Hollow For-

<

mation).
Krynitzka coroniformis zone with Stipidium

Tune (Valentine Formation).

Elias found the same dominant species at the
same zones in the Ogalalla at Wray, Yuma Coun-
ty, and at many other points in eastern Colorado.
He found his Krynitzka zone underlying the
Biorbia zone at Wallace County, Kansas (asso-
ciated with fossil rhinoceros) and at other sites
in Kansas. Biorbia subzones (not described here)
were found in Beaver and Ellis counties, Okla-
homa.

The presence of prairie elements during the
Miocene has been documented by Thomasson
using scanning electron microscopy to identify
eaves, anthoecia (lemmas and paleas of grass

LE 6. Kilgore species occurring in Miocene floras of the Columbia Piatssi (from MacGinitie, 1962;

TAB
Edwards, 1983). Q
certain

P , but affinity

Species

Related to

Populus washoensis?
Pterocarya oregoniana
Mahonia MIR
C — train

P. grandidentata (E U.S.A., SE Canada)
P. insignis (E Asia)

M. bealii (China)

C. mexicana (NW Mexico

Ace me A. negundo (W & 7 U.S.A. & S Canada)

eas coulteri?

F. oregona (W U

F. americana (E ^ cent U.S.A., S Canada)
F. pennsylvanica (E & central U.S.A., S Canada)

1987]

spikelets), fruits, and seeds (Thomasson, 1978a,
1978b, 1979, 1980a, 1980b, 1980c, 1983, 1984,
1985; Voorhies & Thomasson, 1979). For as-
signment of fossil grasses, epidermal patterns and
silica bodies (phytoliths) were particularly di-
agnostic. The plant fossils occurred frequently
with vertebrate fossils (rhinoceros, elephantids,
horse, and camel). The Ash Hollow Formation
(Ogallala Group) in Nebraska, Clarendonian and

Hemphillian in age, contains a rich assemblage
of prairie plants and aquatics, including repre-
sentatives of eight plant families (Chara sp.,
Characeae; Equisetum sp., Equisetaceae; Pota-
mogeton sp., Potamogetonaceae; Carex gracei
and two other Carex spp., Cyperocarpus pul-
cherrima, and C. terrestris, Cyperaceae; Archae-
oleersia nebraskensis, 12 species of Berriochloa,
Nassella sp., Oryzopsis sp., Paleoeriocoma hitch-
cockii, and Panicum elegans, Gramineae; Bior-
bia, Cryptantha spp., Prolappula sp., Boragina-
ceae; Celtis willistonii, Ulmaceae; and Polygonum
sp., Polygonaceae). In addition, representatives
of the Gramineae (species of Berriochloa) and
Cyperaceae (Cyperocarpus eliasii) have been re-
ported from the Sheep Creek Formation in Ne-
braska (late Hemingfordian), and new species of
the Gramineae (Berriochloa spp.) and Boragi-
naceae (Biorbia sp., Cryptantha spp., and Elia-
siana sp.) have been described from the Keller
site in Ellis County, Kansas (Hemphillian or
Clarendonian).

Although the affinities of most of the fossil
species are speculative, those of the grasses ap-
pear to be rather straightforward. For example,
the fossil grasses are classified in tribes Oryzeae,
Stipeae, and Paniceae. Archaeoleersia appears to
be the forerunner of Leersia (tribe Oryzeae) and
is most similar to living Leersia ligularis and L.
monandra of North, Central, and South America
and to L. triandra of Africa (Thomassan, 1980b).
Berriochloa, Nassella, Oryzopsis, and Paleoerio-
coma are all classified in Stipeae. Berriochloa
shows features that suggest it is ancestral in th
evolutionary series: Berriochloa-Piptochaetium-
Stipa (sect. Hesperostipa) (Thomasson, 1978a)
Paleoeriocoma, likewise, belongs to an evolu-
tionary series: Nassella—Oryzopsis—Stipa; this ge-
nus is found with Nassella in deposits and ap-
pears to be ancestral to species of Oryzopsis (sects.
Eriocoma and Oryzopsis) and possibly to some
species of Stipa (Thomasson, 1980c). Panicum
elegans is most similar to extant species of Di-
chanthelium but has not been transferred to that
genus due to inadequate sampling of the micro-

oO

LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

855

morphological characters of the many species of
Panicum (Thomasson, 1980b). Tribes Oryzeae
and Paniceae, especially, contain grasses of
southern affinities; however, the relationships of
tribe Stipeae remain uncertain (Barkworth, 1981)
even though many extant species have southern
affinities. The other families represented in the
fossil deposits (Boraginaceae, Cyperaceae, and
Ulmaceae) generally are best represented in sub-
tropical or warm regions and, predictably, would
have southern affinities.

he Kilgore flora of Nebraska is central to our
data base since the locality occurs at about the
same latitude as the Columbia Plateaus floras. It
is of mid-Miocene (Barstovian) age and was well
documented by MacGinitie (1962), who provid-
ed leaf and pollen data. Hence it can be compared
with sites of that age on the Columbia Plateaus.
The 28 species MacGinitie identified show the
strongest affinity (57%) with modern taxa that
grow chiefly east of the Rocky Mountains; sec-
ondary affinity is with southerly taxa that now
occur in Mexico and the southern Rocky Moun-
tains (2096). Other relationships are minor and
include a few species that occur west of the Rocky
Mountains: two in eastern Asia, and one cos-
mopolitan group. Importantly, at least three
species are intermediate between distinctly east-
ern and distinctly western North American taxa:
Fraxinus coulteri, Populus gallowayi, and Celtis
kansana (Table 5)

Only six Kilgore species (21%) occur as fossils
in the Miocene floras of the Columbia Plateaus
(Table 6). These include a mixture of geographic
elements, and small east-west differences may

gundo var. negundo, but the Columbia Plateaus
species Acer negundoides is morphologically close
to A. negundo var. californicum (J. A. Wolfe,
pers. comm., 1986). In summary, the floristic
relationship of the Kilgore with the Columbia
Plateaus floras is slight.

I mplications. The mid- Miocene -a (e.g.,
o those
of the present Great Plains grasslands, many with
southern affinity. Woody elements (e.g., Kilgore
flora) are chiefly related to living taxa of the east-
ern U.S. The combined data suggest that the Great
Plains flora of the mid Miocene was floristically
distinct and had little in common with the Co-
lumbia Plateaus Miocene floristic province. The
Kilgore flora of the Great Plains bears a stronger
relationship to Miocene floras of the eastern sea-

ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

856

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1987]

board, e.g., with Calvert Cliffs, than with floras
of the Columbia Plateaus (J. A. Wolfe, pers.
comm., 1987).

Kilgore vegetation was in Axelrod’s (1985) view
probably “a wooded grassland with semi-open
grassy forests and woodlands on the interfluves
as well as patchy grasslands.” The fauna suggests
a frost-free climate. This is consistent with
MacGinitie’s (1962) analysis.

In younger Miocene floras (Clarendonian and
Hemphillian, e.g., from Ash Hollow, Nebraska
and Beaver County, Oklahoma), seeds of prairie
elements are much more abundant and more
widespread. Leaf floras suggest that woody vege-
tation was confined chiefly to valley-bottom and
riparian-border habitats. Together these data
suggest “either parklands or grasslands” on the
interfluves (Axelrod, 1985; 171). Pliocene floras
indicate extensive grassland on the Great Plains,
although, as Axelrod and MacGinitie pointed out,
the evidence is not adequate to establish that
pure grasslands occurred before the postglacial.

SYNTHESIS— CONTRASTS EAST AND
WEST OF CORDILLERA

From the data presented, dominant vegetation
types east and west of the cordillera can be com-
pared for two times during the Miocene. Table
7 summarizes the general picture for the Barsto-
vian/late Hemingfordian (ca. 17-12 Ma) and the
Clarendonian (ca. 12-8 Ma). During these two
intervals, the Columbia Plateaus west of the
Continental Divide consistently maintained de-
ciduous hardwood and montane conifer forests,
both rich in woody genera, while evidence from
the Great Plains east of the Rockies clearly in-
dicated the existence of deciduous valley forest
with grassland elements. Floristically the eastern
floras bear little relation to those of the western
area. During both of these times the eastern de-
ciduous forest and g ted from
forests of the west (1) by the continued existence
z the Rocky Mountain massif, which may have

ecome more elevated during the Miocene
A 1980), and (2) by the continued pres-
ence of an impoverished montane conifer forest
with steppe elements in the cordilleran area.

Through most of the Miocene there appear to
have been floristic provinces distinct from each
other:

(1) The Great Plains province had its primary
affinity with species of the eastern U.S.A. and

LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

857

was floristically distinct from floras of the Co-
lumbia Plateaus. It contained elements that later
became important forest species of riparian val-
lies (e.g., American elm) as well as extinct genera
ancestral to modern grassland elements (Ber-
riochloa and Archaeoleersia).

(2) The Rocky Mountain province was dom-
inated by depauperate conifer forest or woodland
with steppe. Except for infrequent pollen of Ju-
glandaceae, Ulmaceae, and Tsuga, the spectra
resemble modern pollen rain. The presence of
Tsuga suggests that the “maritime conifer forest”
(of Habeck, 1987) now found near the Canadian
border may then have been in a more southerly
position. We do not know what species may have
been involved, but the impoverished generic list
of Pinaceae alone places it in stark contrast with
the rich conifer assemblages of the Columbia Pla-
teaus region during the Miocene.

(3) The Columbia Plateaus province did not
take on a relatively modern character on a ge-
neric basis until at least late Pliocene (Blancan)
time or possibly even Pleistocene time. Although
grassland and steppic elements (e.g., Artemisia,
Sarcobatus) existed in this bewa d the
Miocene, they were unimpor

Plain data, grasses became sporadically abun-
dant in Pliocene (Blancan) time— fully 10 Ma
after grassland may have developed in the Great
Plains region.

The late appearance of grassland west of the
Rocky Mountains may mean that eastern and
western grassland provinces are of different
origins. Support for this tentative conclusion is
derived from the following statement based on
data presented earlier in the paper:

(1) Great Plains grassland taxa with southern
affinities, now considered ancestral forms, ap-
peared during the Miocene.

(2) The Great Plains floristic province was
separated from the Columbia Plateaus during the
Miocene by the orographic barrier and by the
existence of an impoverished montane conifer
forest with steppe on the Rocky Mountain cor-

illera.

(3) Grasses of northern affinities adapted to
summer-dry conditions may have moved into
the Columbia Plateaus from the north after the
major shift from summer-wet to summer-dry
climate about 6 Ma (Hemphillian)— during the
demise of the rich Columbia Plateaus hardwood-
and certain conifer-forest elements. Floristic evi-

858

dence confirms the strength of these barriers since
Barstovian (mid Miocene) time.

BIOGEOGRAPHIC IMPLICATIONS

The existence of montane conifer forest and
steppe on the Rocky Mountain massif through-
out the Miocene means that biogeographic con-
nections between the deciduous forests of the
Columbia Plateaus and those of eastern U.S.A
were cut off since at least the early Miocene.

Eastern hardwood and swamp elements co-
existed on each side of the divide in the Miocene
until they were eliminated in the west by the
ae deterioration during Pliocene time.

mportant shift to a summer-dry climate
came etd rapidly to the Columbia Pla-
teaus, diminishing a vast reservoir of genetic re-
sources; this included many deciduous and co-
niferous taxa with affinities to modern taxa of
eastern Asia and eastern U.S.A. that show ad-
aptations to a summer-wet climatic regime. This
basic change reduced the diversity of the mon-
tane coniferous forests of the interior in the Pa-
cific Northwest except along the Pacific coast. It
further established conditions in which steppic
elements and grassland of northern distribution
probably became established and prominent on
the Columbia Plateaus.

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860

ern Rocky Mountains: a synthesis. Mountain Ge-

grass anthoecia within Miocene rhinoceros skel-
etons: diet in an extinct species. Science 206: 331-
333.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Voss, E. G. 1972. Michigan Flora, Part 1. Cranbrook
Inst. Sci., Bloomfield Hills, Michigan.

Wo re, J. A. 1969. Neogene floristic and vegetational
history of the Pacific Northwest. Madrono 20: 83-
l

. 1978. A paleobotanical interpretation of Ter-
tiary climates in the northern hemisphere. Amer.
Sci. 66: 694-703.

APPENDIX I. Pollen records showing occurrence of shrubs, terrestrial herbs?, and herbaceous aquatics in Late
Cenozoic samples from Idaho, eastern Oregon, and eastern Washington. Numbers show maximum values in

pollen/spore counts; + indicates presence.

MIOCENE
Middle Late
E
aq
£ ` E
- _ = © o
E x 9 S S = x © 8 E > Š
5 š š G ó Š Y. 3 ER š
= Š Š s » £ š ç 9 Š t c st Ze š
S z 5 E B ° 5 & E C 2 £ ug
] 9» Š š š š P 5 3 F š P Sio
EZ ë 2 O à X &k F & $ m= A Wo Bla 3
SHRUBS
Ephedra + 1
cf. nevadensis T + 2 4 +
cf. torreyana + + +
Inus + + + 5 + + 4 + l l
Chenopodiineae + + 1 +? + 7 20 20 3
Sarcobatus + +? + 2 4 3 1 7 l
Compositae
Artemisia + + + 14 30 52 36
Elaeagnus + +
Shepherdia + + +
saceae + 3
Cercocarpus +
Ilex + + + + 4
Rhus 4
Corylus + l + +
rica + +
Malvaceae + + +
Myrtaceae +
Yucca type +
Ericales +
Vaccinium ? ? + +
Caprifoliaceae +
Viburnum +
Symphoricarpos +
Sapindaceae +
Mahonia +
Cornus +
TERRESTRIAL HERBS?
Compositae undet. + + 9 11 2 49
Liguliflorae + + l
Tubuliflorae 3 6
Ambrosia + + 4 +

1987]

APPENDIX I.

Continued.

LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

861

MIOCENE

Middle

Mascall'

Blue Mountain!

Succor Creek?

Clarkia?

Stinking Water!

Miocene, Oregon?

Pliocene, Oregon?

Trapper Creek?

Poison Creek?

Jenny Creek‘

Hanford‘

Banbury Basalt‘

Glenns Ferry”

PLIOCENE

QUATERNARY
Bruneau‘

Late Pleistocene, Idaho‘

Cruciferae
Acalypha type
Onagraceae
Polemoniaceae
Gilia t
Navarretia
Caryophyllaceae
Stellaria
Gramineae
Polygonum
cf. californicum
Eriogonum
Anemone
Aizoaceae type
Pachysandra
Umbelliferae
Ferns

AQUATIC HERBS

Myriophyllum

+?

+

4

+

+

+?

+ ++ ++

+

++

+

+

+

De ee ee ee

—
—

-

! Jane Gray in Chaney & Axelrod, 1959.

z — &
? Gray, 1985.
* Gray, |

1964.
Š oc jud u MacGinitie, 1972.

* This

T enber " Wright, 1985.

Cross, 1980; Graham, 1963.

862 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

APPENDIX II. Selected pollen counts, Snake River Plain, Idaho (Fig. 10). Localities given in Leopold &
Wright, 1985; the Jenny Creek Formation locality was previously considered as “Salt Lake Formation."

Poison Creek Jenny Creek
Formation* Formation Banbury Basalt
USGS Paleobot. Loc.: D1208-3 D1696
Pinus 84.5 30.2 7.2
Picea 1.0 6.0
Abies cf. grandis 5.4 3.6
A. cf. lasiocarpa 1.0 +
Pseudotsuga 1.2
niperus 1.0 4.2
opulus 1.8
Betula 1.8 0.8
Quercus 4.8
us
Subtotal: 92.9 53.6 8.0
Cedrus-type 0.5
Carya 0.5 1.8
Juglans 1.8 +
Pterocarya 0.6
Ulmus- Zelkova 10.1 0.8
Subtotal Tertiary relicts: 1.0 14.3 0.8
Acer 0.4
Ephedra cf. torreyana 0.6 0.8
Ephedra i R. type) 0.5
Sarcobat 2.0 4.2 0.8
Other Chenopodiincae 6.8
Artem 14.0
Other C aie 9.3
Cercocarpus type 0.6
Elaeagnus +
Subtotal shrubs: 2.5 5.4 32.1
udis 0.5
Gramineae 0.6 1.3
Mosecon 1.0 4.8
Cruciferae 0.4
Dicots 2.0 11.9 0.8
Potamogeton 0.6 T
C3P3 56.4
Lycopodium +
Onagraceae +
cf. Parthenocissus +
Fern spores 1.0 8.9 1.0
Subtotal NAP: 4.5 26.8 59.9
Total percent 100.9 100.1 100.8
Total tally (202) (336) (236)
Cysts 0.5 1.2
Botryococcus 12.0

* Fossil leaves from the Poison Creek Formation identified by J. A. Wolfe (pers. comm., 1987) include Quercus
prelobata Condit. and Q. cf. chrysolepis Liebm

1987] LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 863

APPENDIX III. Revised list of the Deschutes flora,

northern Oregon (after Chaney, 1938, with additions
int. A. Wolfe, pers. comm., 1987; Wolfe includes
certain collections made by R. W. Brown from Cha-
ney’s floral horizon but a few miles away).

Acer negundoides
Quercus prelobata

us washoensis
Populus alexanderi (P. aff. trichocarpa)
Prunus irvingii (P. aff. emarginata)
Salix sp. (S. cf. xta of Chaney)
Salix aff. caudat
Ulmus affinis Tod & Wolfe

APPENDIX IV. Localities of selected pollen samples from Wyoming and Colorado (Figs. 6, 12).

USGS
Paleo-
botan
Formation Locality Site Description
Split Rock D1309 Type section: SE '4 NE !⁄4 Sec. 25, T. 29 N, R. 89 W, Natrona Co., Wyoming.
Limestone at top of pumicite, from unit 21 of J. D. Love (1961). he
from the Split Rock type area are of Hemingfordian age.
Split Rock D3319 Upper part of 2,650 ft. EWL 600 ft. SNL Sec. 36, T. 27 N, R. 85 W,

Carbon Co., Wyoming. J. D. Love measured section L63-11. Sample B & Ca
6 and 12 ft. Aus base of his unit 3. Sample D/E from his unit 4. Samples ikher
in this unit contain abundant reworked Cretaceous pollen
Saratoga unit D1540 os Valley, — oil shale. SW !⁄4 NW '% Sec. 26, T. 18 N, R. 84 W,
. D. Love locality L59-49. Associated with vertebrates
of aiden age ponia to McGrew (1951).

Moonstone D1308 Type section; white tuff and white laminated shale. Sec. 17, T. 30 N, R. 89 W,
Natrona Co., Wyoming. Samples | and 2 are from Love's (1961) unit 16 and 17,
respectively.

Troublesome D3473 Rock Creek facies. NW !⁄4 SW 4 Sec. 32, T. 2 N, R. 79 W, Kremmling quadrangle,
Grand Co., Colorado. Glen Izett loc. G-63-303 and -304. Sample A is 18 ft.
above sample B. Vertebrates from the lower and upper parts of the formation
correlate respectively with the lower and upper Hemingfordian Group of Ne-
braska (Izett, 1968).

Troublesome D1905 SW INE IA Sec. 29, T. 2 N, R. 79 W, Kremmling quadrangle, Grand Co., Colorado.
Carbonaceous bed interbedded with silt lake facies containing mid Miocene ver-
tebrates. Izett coll. ra 0153- A

Troublesome D3493 NE % SE % Sec. 28, T. 3 N, R. 80 W, Gunsight Pass, Twin Peaks, Kremmling
quadrangle, Grand i es Izett loc. G-63-305.

864 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74
APPENDIX V. Pollen counts, Troublesome Formation, Grand County, Colorado (Figs. 6, 12). Percent pollen
and spores. Small counts (shown in parentheses) indicate comparative abundance; + records taxa seen but not
in tally
USGS Paleobotany Locality: D3493 DINA
-b -a
Pinus 18.0 72.0 (105) (62)
Abies sp. 1.3 1.3 (1)
A. cf. lasiocarpa 0.7
A. cf. grandis + (1)
Picea sp. 0.7 1.3 (6) (12)
P. cf. lal (27)
Juniperu. e 32.4 0.7
Ephedra P nevadensis 8.0 2.7 (1)
Ephedra cf. torreyana * 0.7 (1)
Ephedra (short axis type) 4
*Juglans + 0.7
*Pterocarya 1.3
Carya + 0.3
*Ulmus-Zelkova 4-5 pored 0.7 1.1 (1)
6 pored +
Acer +
cf. Populus 4
Betula E _0.7

Subtotal trees: 62.4 82.2
Elaeagnus 0.7
Alnus 1.3 3.2

alix 1.3
Sphaeralcea 3 0.7
Caprifoliaceae
onicera 0.3
Symphoricarpos 0.3 0.7
Chenopodiaceae 4.6 PA, (3) (1)
arcobatus
cf. vermiculatus 0.7 (3)
Compositae + (1)
temisia + 0.7 (3)

Xanthium type (1)
Gramineae 1.3 1.1 (1)
Umbelliferae 7
Polemoniaceae 0.3
Dipsacaceae? +
Labiatae 0.3
Arceuthobium 0.3
Apocynaceae +
Claytonia type + 0.7
Sparganium 1.1

icots 3.4 2.0
Monocots 16.6 (4)

Botrychium type (4)

Riccia ty +

Trilete spores 7.4 2.7 _ (2) —
Total percent 100.9 99.1
Total tally (300) (300) (130) (110)

* Genera now exotic to Colorado.

1987] LEOPOLD & DENTON — GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS 865

APPENDIX VI. Selected pollen counts, Natrona and Carbon counties, Wyoming (Figs. 6, 12). Percent pollen
and spores: + records taxa seen but not in tally.

Stratigraphic Unit: Split Rock Saratoga Moonstone
USGS Paleobotany D1309 D3319 D1540 D1308
ocality:
-B -C -D/E -1 -2
Pinus 8.3 39.5 35.3 51.5 80.0 53.0 72.0
Abies spp. 0.8
cf. concolor 2.6
cf. lasiocarpa 2.0
cf. grandis 0.6 1.3
Picea spp. 0.5 1.2 1.1
cf. engelmannii 3.6 5.2
cf. pungens 7.6 4.6
*Tsuga
cf. mertensiana 0.4
cf. heterophylla 0.4
Pseudotsuga
cf. menziesii 0.8 1.2 1.1 0.4
Juniperus type 0.4 20.5 20.7 7.4 1.8
Quercus 1.2 1.0 0.4 0.4?
Populus + 0.5 0.6
*Carya 1.0 0.4?
*Juglans 0.5 0.4
*cf. Liquidambar 1.2
*Ulmus-Zelkova 0.4 1.0 1.6 0.9 12.5 3.1
Betula 0.4 3.2
Salix 0.5 0.4
*Ostrya-Carpinus 0.5 3.2
Subtotal trees: 11.5 65.0 58.4 63.3 95.6 71.4 95.9
Alnus 0.4 0.4 1.1 0.8
Chenopodiineae 5.0 1.0 9.5 4.7 8.5 1.8
Chenopodium type 0.4
Sar t 2.1 6.0 1.8 0.9 7.4 1.2
Ephedra
cf. torreyana + 1.2 0.8 0.4 0.4
cf. nevadensis 0.4 1.5 4.1
Compositae 7.5 0.5 1.2
Liguliflorae 0.5 0.6
Artemisia 42.4 1.2 1.1
mbrosia 0.8 0.5
Sapindaceae 0.5
Ribes 1.0
cf. Ptelea 0.4
Arceuthobium 0.4
Caprifoliaceae type 0.6
Eriogonum 0.4 0.4
Gramineae 0.4 1.5 1.8 0.4 0.4
Umbelliferae 0.5
Cruciferae 0.4
Sparganium 1.5 0.6 0.8
Cyperaceae + 0.5 1.8 0.8
Scirpus 2.4
GP; 4.5

Ç, 15.4 2.5 5.1
Monocots 9.0 13.0 6.6 1.1

866 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74
APPENDIX VI. Continued.
Stratigraphic Unit: Split Rock Saratoga Moonstone
USGS Paleobotany D1309 D3319 D1540 D1308
ocality:
-B -C -D/E -1 -2
Dicots 3.3 4.0 0.4 5.3
Indeterminate 2.9 4.5 0.6 5.7 3.1 3.2 2.4
Fern spores 1.8 3.0 2.4 7.8 1.1
Total percent 99.6 103.0 98.8 100.4 100.0 100.2 102.9
Total tally (245) (208) (169) (257) (223) (188) (251)
Reworked pollen 21 3 28
Botryococcus ++

* Genera now exotic to Wyoming.

1987] LEOPOLD & DENTON—GRASSLAND AND STEPPE OF THE NORTHERN ROCKY MOUNTAINS

APPENDIX VII. Pollen records showing occurrence
of shrubs, terrestrial herbs?, and herbaceous aquatics
in pollen samples from the Mio-Pliocene of the central
Rocky Mountains. Maximum values, % pollen and
spores. + indicates presence or less than 1%.

O
E 4 š -
O ° ° S i
| mm 8 B 2
2 = 2 v 9
€ a 8 d <Š
E N E N 2
SHRUBS
Ephedra
cf. nevadensis 8 4 3
cf. torreyana 1 1 1 l l
Alnus 3 + 2 l
Sphaeralcea l
henopodiaceae 5 10 3 9
Sarcobatus l 6 70 l 7
Compositae
` Artemisia l 42 5 l
Elaeagnus l +
Caprifoliaceae l
onicera
Symphoricarpos l
ibes l
Ptelea type +
Yucca type 3
TERRESTRIAL HERBS?
Composita
Liguliflorae 1 4
ubuliflorae l 8 15
osia 1 +
Xanthium type l
Polemoniaceae l
ilia ty 11
Cruciferae +
Claytonia type 1
ace +
Polygonaceae "P
iog l +
Umbelliferae l 1
Apocynaceae +
Labiatae +
Dipsacaceae +
ramineae l 2 15 +
7 3 l
HERBACEOUS AQUATICS
Cyperaceae 2 +
Sparganium l 2

Scirpus 2 +

867

FOSSIL POLLEN OF SABICEA (RUBIACEAE) FROM THE
LOWER MIOCENE CULEBRA FORMATION OF PANAMA!

ALAN GRAHAM?

ABSTRACT

Fossil bon of Sabicea (Rubiaceae) has been recovered from the lower Miocene Culebra Formation

of Pana
well represented in E tropical moist
it was part - to mo

ada north u lombia. Its association with other m

enus is presently widely distributed in vg lees northern South A
and premontane wet for
erate-altitude insular vegetation p the landscape between

merica; it is
of Panama. In the lower Miocene

embers of the Culebra assemblage

h ern
act tropical paleoclimates similar to those of the present. The genus has not been reported

previously in the fossil record.

During studies on Tertiary vegetational his-
tory of the Gulf/Caribbean region, pollen and
spores are frequently encountered representing
genera with no previous fossil record, or whose
stratigraphic and/or geographic range is consid-
erably extended by the new records. Examples
include Pelliceria (Theaceae/Pelliceriaceae; Gra-
ham, 1977), Mortoniodendron (Tiliaceae) and

Sphaeropteris/Trichipteris (Cyatheaceae; Gra-
ham, 1979), Micractinium ee Gra-
ham, 1981), and Lisianthiu Gra-
ham, 1984). Fossil pollen of Sis (Rubiaceae;
Figs. 1-4) has recently been recovered from the
lower Miocene Culebra formation of Panama,
representing its first known occurrence in the
geologic record.

' The author gratefully acknowledges comments provided by John Dwyer and pollen material provided by

curators at MO.

The research was supported by NSF grants BSR-8500850 and BSR-8619203.

> Department of Biological Sciences, Kent State University, Kent, Ohio 44242, U.S.A.

* iy
ev P
$ noe

p's
SS) PRA
eh: ay -

FiGuRES 1-6. Fossil and modern pollen of e (Rubiaceae). — 1, 2. Fossil pollen, Pan core 456, slide 1,

ESF os D-17.—3, 4. Fossil pollen, Pan
pollen of S. colombiana. All taken at 400 x

cine : ue 3a, ESF Werl U-31,3.— 5, 6. Modern

given in text. Fossil Vg and modern

ctua ay
reference material are deposited in the eae nd silica. Kent State University, Kent,

ANN. Missouni Bor. GARD. 74: 868-870. 1987.

1987] GRAHAM —FOSSIL POLLEN OF SABICEA 869
TABLE l. Modern Rubiaceae pollen examined.
Herbar-
ium
where
Voucher
Taxon Country Voucher Collection Deposited
Amphidaysa ambigua (Standley) Standley Panama Busey 385 MO
Coccocypselum guianense (Aublet) Schum. Honduras “ew & Romero 4263 MO
C. herbaceum Lam nama D'Arcy & D'Arcy 6731 MO
C. lanceolatum (Ruiz & Pavón) Pers. Panama Antonio 1425 MO
Didymochlamys connellii N. E. Br Guyana Maguire et al. 32362 MO
Gonzalagunia brenesii Standley Costa Rica Croat 2659 MO
G. bunchosioides Standley Ferreyra 1660 US
G. panamensis (Cav.) Schum. Panama Johnston 76 GH
Honduras N. Mex. exch MO
G. rosea Standley Panama White 7 GH
Hippotis mollis Standley Colombia Lawrence 505 MO
H. tubiflora Spr Peru Klug 3084 MO
Isertia deamii Sw Guatemala Deam 6016 MO
I. haenkeana A. D rvard exc GH
I. hypoleuca Benth. Guyana Shell Oil exch
Costa Rica Jiménez 4127 MO
na Stimson 5062 MO
I. pittieri (Standley) Standle Colombia St. George Exped. 337 US
Pentagonia brachyotis Cds Standley Panama Dwyer 1385 MO
P. macrophylla Benth. Panama STRI exch. MO
Panama Croat 4646 MO
P. pubescens Standley Panama Croat 4685 MO
P. wendlandii Hook. Panama von Wedel 2018 GH
Raritebe palicoureoides Wernham subsp. Panama Mori et al. 6617 MO
dwyerianum Kirkb
Sabicea brasiliensis Wernham Brazil Irwin et al. 24943 MO
S. colombiana Wernham Colombia Uribe 3041 US
Colombia Gentry et al. 47975 MO
S. panamensis Wernham Panama Dwyer MO
S. paranensis (Schum.) Wernham eru Schunke V. 10548 MO
S. villosa var. adpressa (Wernham) Standley Harvard exch. GH
S. villosa Rose & Standley var. villosa Panama Luteyn et al. 1798 MO
Panama von Wedel 2889 GH
Panama Tyson 3437A MO
Schradera blumii Dwyer & Hayden Panama Mori et al. 6625 MO
Sommera grandis (Bartlett) Standley Panama Allen 1575 GH
Panama Gentry et al. 13581 MO

THE COLLECTING LOCALITY

In 1958 the Engineering and Construction Bu-
reau of the Panama Canal Commission drilled a
well through the Culebra Formation in front of
Gold Hill on the west side of the Canal at latitude
9°02'N, longitude 79°38'W (Hole No. GH-9).
Fifty-seven samples were taken from along the
124-meter core, and 21 contained well-preserved
pollen and spores. The specimens of Sabicea were
isolated from samples at the 456- and 470.6-foot
depths. Other details on the Culebra Formation

are provided by Graham et al. (1985) and Stew-
art & Stewart (1980). The materials and methods
were as described in Graham (1985).

DESCRIPTION

Pollen oblate, amb oval-triangular to nearly
circular; tricolpate/porate (apertures short, slit-
like, ca. 2: 1 length:width ratio), 4-6 x 2-3 um,
equatorially arranged, meridionally elongated,
equidistant, inner margin faintly dentate (due to
overlying sculpture elements), faint costae colpi;

870

wall 2-3 um thick, tectate-perforate; finely retic-
ulate, muri smooth, width about equal to di-
ameter of lumen (ca. 0.5 um); 32-36 um.

DISCUSSION

Sabicea is a genus of about 125 species of
climbing shrubs and vines widely distributed in
tropical America and in Africa and Madagascar
(Dwyer, 1980a, 1980b). In Panama it is repre-
sented by three species: S. panamensis (Guate-
mala to Colombia); S. vi/losa (with var. villosa
widely distributed in Central and northern South
America and with var. adpressa known from
Panama, Colombia, and Peru); and S. stellaris
(Panama). Sabicea villosa var. adpressa grows on
Barro Colorado Island where it is “occasional in
older clearings, on trails and at the margin of the
forest along the lake; less commonly climbing to
the top of the forest canopy and sometimes root-
ing in water," and it “is known from tropical
moist forest in the Canal Zone, Bocas del Toro,
San Blas, Panamá, and Darién, from premontane
wet forest in the Canal Zone, and from premon-
tane wet forest in Panamá and Darién" (Croat,
1978: 827)

Dwyer (1980a: 7) placed Sabicea in the tribe
Mussaendeae with Pentagonia, Sommera, Hip-
potis, Schradera, Amphidaysa, Gonzalagunia,

these (Table 1), and Sabicea can be distinguished
on the basis of pollen characters. For example,
grains of Raritebe examined were smaller (ca. 25
um), thicker-walled, and tetra- to stephanocol-
porate. The pollen of Coccocypselum was con-
siderably larger (45-50 um), oblate-spheroidal,
and scabrate to finely verrucate.

hree species of Sabicea were examined (Ta-
ble 1), including all species and varieties reported
from Panama except S. stellaris (holotype, MO,
not sampled). The pollen showed only minor dif-
ferences between the species (e.g., minute vari-
ations in wall thickness). The reticulum of the
fossil specimens appears slightly more distinct
than in the modern pollen examined. The mod-
ern forms (Figs. 5, 6) having slightly thicker walls
darken more than those with thinner walls during
acetolysis and more closely resemble the speci-
mens in this respect. These are not inherent mor-
phological differences or consistent characteris-
tics of the pollen, however, and the specimens
cannot be referred to any one modern species.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

The specimens were part of a fossil assemblage
that includes bs unos associates (Graham,
in prep.; pre ry identifications): Lycopo-
dium, HUN person yathea, Pteris, Ly-
godium, cf. Antrophyum, Danaea, Gramineae,
Palmae, //ex, Chenopodiaceae/Amaran tha aceae,

pighiaceae, Hampea/Hibiscus, Eugenia/Myrcia,
Rhizophora, Allophyllus, Cupania, Matayba,
Sideroxylon, and cf. Guazuma. The landscape of
present-day Central America consisted of low-
lying volcanic islands at the time the Culebra
assemblage was being deposited during the lower
Miocene, ca. 25 Ma (e.g., Stehli & Webb, 1985).
The association indicates that Sabicea grew in
this physiographic setting and under paleocli-
mates not greatly different from those ofthe pres-
ent.

LITERATURE CITED

Croat, T. B. 1978. Flora of Barro tbe Island.
i iforn

n Flora of DE

GRAHAM, A. . New records of Pelliceria (Thea-
ceae/Pelliceriaceae) a the Tertiary of the Carib-
bean. Biotropica -52.

orton UU (Tiliaceae) and

Sphaeropteris/Trichipteris (Cyatheaceae) in Ce-

nozoic deposits of the Gulf-Caribbean region. Ann.

———. 1985. Studies in neotropical paleobotany. IV.
The Eocene communities of Panama. Ann. Mis-
72: 504-534

souri Bot. Gard. ;
TEWART.

1985. Stud-

bearing sediments. Ann. Missouri Bot. Gard.
485-503.

SrEHLI, F. G. & S. D. Wess (editors). 1985.
Great American Biotic Interchange. Plenum, New
York.

STEWART, R. H. & J. L. STEWART (with the collabo-
ration of W. P. Woodring). 1980. Geologic Map
of the Panama Canal and Vicinity, Republic of
Panama. Scale: 1 : 100,000. U.S. Geol. Surv. Misc.
Invest. Map I-1232

SYSTEMATICS OF THE AMPHI-ATLANTIC BAMBUSOID
GENUS STREPTOGYNA (POACEAE)!

THOMAS R. SODERSTROM? AND EMMET J. JUDZIEWICZ?

ABSTRACT

Streptogyna i is the only herbaceous bamboo genus with an amphi-Atlantic distribution. Streptogyna
Sri Lanka, and India, and S. americana is found throughout the

suggest segregation at the generic level, they are here retained in a single genus because they are quite
similar in spikelet morphology and leaf anatomy. Multicellular microhairs are present on the lodicules

primitive feature in the family as they are com
ffinities in its ligule and leaf anatomy, spikelet structure, caryop-

demb ut differs from the core group ofthe subfamily
in its seedling sal i S and lack of epidermal so "oq en in the two species suggest

of it
1

that neither could have been derived directly from the oth

The grass genus Streptogyna was first brought
to the attention of Western botanists in the late
18th century, when British and Swedish collec-
tors brought back specimens of “rat-catching
grass" from the forests of West Africa. A gath-
ering from Nigeria by Palisot de Beauvois was
used as the basis for Streptogyna in 1812, which
he based on the only species known to him, S.
crinita. The narrow leaves and many-flowered
spikelets of Streptogyna were long taken as in-

Thus,
Bentham (1883), Hackel (1887), a ar
(1936) all considered that the proper disposition
of this genus from the tropical rainforest lay with
this large, temperate-region grass group. But there
were dissenters, and Nees von Esenbeck (1835)
and Steudel (1855), for example, suspected the
bambusoid affinities of the genus. Streptogyna
was briefly revised by Hubbard (1956), who in-
dicated that the group deserved tribal status, but
it was not until Tateoka (1958a) and Metcalfe
(1960) examined its leaf anatomy that the bam-
busoid affinities of Streptogyna became clear.
Recent workers agree that Streptogyna should be
placed in its own tribe in the Bambusoideae
(Calderón & Soderstrom, 1980; Clayton & Ren-

voize, 1986). In a treatment of the herbaceous
bamboos of Sri Lanka, Soderstrom et al. (1987)
offered a detailed descriptive account of the leaf-
blade anatomy in the two taxa. The present study
provides a revision of the genus and attempts to
€ the relationships of the two species by
examining characters that have not yet been
es in detail, such as morphology of lodi-
cules, starch grains, and embryos.

MATERIALS AND METHODS

Specimens of Streptogyna were examined from
the following herbaria: AAU, B, CAY,
CEPEC, F, G, ISC, K, M, MO, NA, NY, P, PDA,
RB, S, US, W, and WIS. For anatomical studies,
spikelets, leaves, and embryos (Table 1) were
dehydrated in dimethoxypropane, infiltrated with
tertiary butanol, embedded in wax, sectioned us-
ing a rotary microtome, and stained in chlorazol
black E. Lodicules were rehydrated with Aerosol-
OT before examination. Starch grains from cary-
opses were cut on a freezing microtome and
stained with I,KI. Observations of living plants
of Streptogyna were made by Soderstrom in Bra-
zil (March 1972, and May 1976) and by Jud-

! We are grateful to Alice R. Tangerini for the illustrations, to Stanley Yankowski for the floret sections, to
C

the curators of the herbaria that lent us specimens for study, an

and to Gerrit Davidse, Lynn G. Clark, and Ma

Sangrey. Part of the research Ris to this paper was performed while the second author was a predoctoral

fellow at the Smithsonian Institutio

? Department of Botany, nds Institution, Washington, D.C. 20560, U.S.A. Died on September 1,
987.

? Department of Botany, Smithsonian Institution, Washington, D.C. 20560, U.S.A.

ANN. MISSOURI Bor. GARD. 74: 871-888. 1987.

872

TABLE 1.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Specimens of Streptogyna for which D (e), floret bases (using the scanning electron micro-

scope) (f), lodicules (1), and starch grains (s) were examine

S. crinita
CAMEROON: Buesgen 530 (1) (US). GABON: without collector, Limbareni, May 1875 (1) (US). GUINEA-BISSAU:
Espirito Santo Expedition 3735 (1) (US). INDIA: Wight 2362 (1, s) (PDA). Ivory Coast: Bamps 2175 (e, s) (BR).
LiBERIA: Baldwin 6305 (1) (US). SIERRA LEONE: Fairchild s.n., Jan. 1927 (e, 1, s) (US). SRI LANKA: without

collector, 3 Jan. 1881, Henaratgoda (1) (

PDA), Senaratna 2700 (1

) (PDA); Gardner s.n., Dec. 1846 (1) (PDA).

ZAIRE: Gilbert 14213 (s) (BR); Lebrun 605 (e, 1, s) (P); Mullenders 1226 (s) (BR); Vanderyst 993 (1) (P).
S. americana
BRAZIL: Eiten & Eiten 8903 (1) (US); Prance et al. 6454 (l, e) (US); Prance et al. 6527 (1, s) (US); Swallen
5089 (1, s) (US); Soderstrom 2193 (e, 1, s) (US). COLOMBIA: Blydenstein 1687 (1) (US); Idrobo & Schultes 608 (1,

s) (US). FRENCH Guiana: Broadway 77 1 (1) (US). GUATEMALA

: Weatherwax 104 (e, s) (US). SURINAM: Maguire

54093 (1) (US). TRINIDAD: Hitchcock 10122 (1) (US). VENEZUELA: Steyermark 86760 (1) (US).

ziewicz in Panama (March 1983). A Cambridge
35 scanning electron microscope was used to ex-
amine the lemmas of Streptogyna species.

RESULTS

Vegetative morphology. Streptogyna species
are herbs of the rainforest understory and are
less than one meter in height. Streptogyna crinita
has long sympodial rhizomes that are densely
covered with striate scale leaves, and the erect
culms are leafy, bearing lanceolate leaves along
most of their length (see figure in Soderstrom et
al., 1987). Streptogyna americana is densely ces-
pitose with the culms produced from short, non-
scaly, sympodial rhizomes, and the linear leaves
are all borne near the base of the plant (Fig. 6A).
The leaves of S. crinita have glabrous sheaths
while those of S. americana are hispid near the
summit (Fig. 6B). At the summit of the sheath
are borne membranous flanges that have been
called eal appendages (Tran Van Nam, 1972),
as they appear to arise from a meristem inde-
pendent of the sheath. These and the oral setae
are moderately well developed in S. crinita but
inconspicuous or absent in S. americana. Both
species also have an indurate abaxial rim at the
summit of the sheath (external ligule), and the
leaf blades are deciduous aboye this structure. A
entin both
taxa. The narrow leaf blades of S. amemus are
strictly glabrous, whereas the broader blades of
S. crinita may have a scattering of long, delicate

short

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panicle of Sirepiasvna
species Is spikelike: but in both species individual
plants have been seen in which an additional
floriferous branch is borne at the base of the main

axis of the inflorescence. The inflorescence of S.
crinita 1s generally shorter than that of S. amer-
icana and appears to be more densely flowered;
this appearance is due in part to the larger glumes
in S. crinita. Both species have loosely several-
flowered spikelets (Fig. 6D) with the uppermost
florets successively smaller and sterile and with
a peculiar downward prolongation of the base of
each floret, which, however, does not appear to
be an elaisome, as no oil was detected within it.
The glumes of S. crinita are broad, elliptic, and
have many nerves, while those of S. americana
have few nerves and are linear to lanceolate (Fig.
6E, F). The florets of both species are indurate
and fall attached to a pointed, persistent rachilla
internode (Fig. 6G) that presumably aids in ex-
ternal animal dispersal. In S. crinita the base of
the floret is pilose and in the scanning electron
microscope (SEM) the bulk of the epidermis,
which appears granular through the light micro-
scope, is composed of short prickles alternating
with square to rectangular cells in a sharply de-
fined pattern (Fig. la, b). The rectangular cells
were observed to deflate when placed under the
electron beam of the SEM. The lemmas of SS.
americana are glabrous and the epidermis is ap-
parently covered by a thick cuticle that obscures
the rounded p and intercalated rectangular
cells (Fig. 1c, d). The lemmas of both species
have long, antrorsely scabrous awns. The paleas
are strongly bikeeled (Fig. 2a).

Flowers. The flowers of both species have three
relatively large lodicules (Fig. 3). Those of S.
crinita are spatulate, of firm texture throughout,
and are strongly nerved until near the summit
(Fig. 3A). Microscopically the epidermis of the
upper portion of each lodicule consists of a uni-
form network of polygonal cells with moderately

on Sierra Leone, 2
Idrob
Soderstrom et al. 2193 (US). Scale bar in a =

thickened walls (Fig. 3B). The apices of the lod-
icules are fringed with about 5-15 thin-walled
microhairs, and each hair contains 4-8 cylindri-
cal cells. The basal cell is slightly longer than the
apical cells and does not collapse when dried as
do the apical cells. The longer lodicules of S.
americana are narrowly lanceolate or linear and

SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA

I : as 3 (US);
o & Schultes 608 (US); e based on Sierra Leone, Afzelius and wasiwi. s.n. (BM); f based on Brazil,
00 u f.)

m for a, c; 20 um for b, d-

often are widest just below the middle (Fig. 3C).
In their lower portions they are strongly nerved
and of firm texture, but at about the middle of
their length the vascular bundles end and the
lodicule tapers gradually to a very delicate, hya-
line apex. Most specimens have the apices and
uppermost margins of the lodicules fringed with

874 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

*005
e
s A (YYY °) \
" p v
% ote
T M o 5

FiGURE 2. Floret anatomy of Streptogyna americana.—a. Transverse section through lower part showing
lemma, palea, three lodicules (1), two stamens (s), and gynoecium (gy) with three stigmatic vascular bundles

1987]

EA
i iy
"ull
EEF
iiid

HE H

ul

FiGure 3. Lodicules of Streptogyna crinita (A, B) a

apex, showing multicellular microhairs
Lodicules. — D. Detail of apex

short cells interspersed with long cells on surface. (Streptogyna crinita based

SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA

— B. Detail of

— A. Lodicules.
and undifferentiated, thick-walled parenchyma cells on surface. — C.
—E. Detail of upper portion of lodicule, showing prickles (mainly marginal) and

nd S. americana (C—E).

meroon, Buesgen 530 (US);
)

S. americana based on Trinidad, Hitchcock 10122 (US). Scale bar = 1 mm for A, C; 0.2 mm for B, D, E

abundant prickles that occasionally grade into
short cilia. The epidermis of the upper portion
consists of alternating, thin-walled long and short
cells, some of the latter modified into prickles

americana. The two stamens of both species are
lateral anterior in position and are free to their
bases (Figure 2a); the anthers are linear (Fig. 6M).
The gynecium of Streptogyna crinita bears a sin-
gle style branching into two stigmas, and the ovary
is pilose near the summit. The stigmas, which
are supplied by each of two lateral posterior vas-
cular bundles in the ovary, are unusual in that
they continue to grow after anthesis, elongating
and producing stout retrorse barbs on their adax-
ial surfaces. Streptogyna americana has three
stigmas (Fig. 6L) produced from three en
bundles within the glabrous ovary (Fig. 2a). T

i grow after ee
g short, papillate pro-
cesses from a meristematic layer near the adaxial
surface (Figs. 2b, 6N

Fruit. Both species have a cylindrical, linear
caryopsis with a linear hilum that extends the
full length of the fruit (Fig. 60, P). The mature
endosperm starch granules of S. crinita are weak-
ly to moderately coherent into masses of 3-6 that
form individual grains 6-10 (rarely 15) um in
diameter (Fig. 1e). The starch grains of S. amer-
icana are nearly round, 10—30 um in diameter,
and are simple or rarely compounded into small
masses (Fig. 1f). There is a prominent lacuna in
the center of many of the grains. The embryo of
Streptogyna species is small, basal, and when
dissected out of the caryopsis is observed to be
about one-half again as tall as it is wide. In me-
dian sagittal sections the embryos of S. ameri-
cana (Fig. 4b) sampled had a prominent epiblast,
no mesocotyledonary internode, and a small cleft
between the scutellum and coleorhiza. In trans-
verse sections the coleoptile had two lateral nerves
and fused margins, the first embryonic leaf had
five nerves and strongly overlapping margins,
and the scutellum had three vascular traces (Fig.
4d). Streptogyna crinita had a similar embryonic

d. — b. Transverse section through upper part showing three stigmas with papillae arising from meriste-

ws).
matic regions on adaxial surfaces. (Based on Brazil, Soderstrom et al. 2193 (US). Scale bar: a —
25 um.)

100 um; b =

876 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

transverse section through plumule show verlapping margins of first embryonic leaf; plumule is attached to
scutellum on bottom. (a based on Zaire, Lae 605 (US); b, d based on Brazil, i a eae et al. 2193 (US); c

1987]

SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA

TABLE 2. Comparison of Streptogyna crinita with S. americana.

Character S. crinita S. americana
Distribution Tropical Africa, Sri Lanka, India Tropical America
H Rhizomatous; leaves spaced along Cespitose from short, knotty rhi-

Ligular area

Morphology of leaf blade
Upper surface of leaf blade
Inflorescence

Glumes

Lemma

Lodicules

Ovary
Stigmas
Starch grains

Embryo

Sheath summit glabrous; sheath au-
ricles and oral setae conspicuous
en x 29(—40) cm long, 1-

2.5c
Nerves say, raised; glabrous or
aringly
l j cm
lappin
Large; first glume 6-8 mm long, the
second 17-26 mm long, 11-17-

id ne spikelets over-

nerve

Base pilose; texture granular, the
epidermal cells distinct in SE

Oblanceolate, 1.5-3.5 mm long;
multicellular microhairs present;
prickles rare or absent; short cells
absent; texture near apex firm

Pilose near s

2, retrorsely a rbed
ompound, individual granules 6—
va s 5) um in diameter

Cleft between scutellum and coleo-

zomes; leaves clustered at plant
base

Sheath summit hispid; sheath auricles
and oral setae inconspicuous

Linear, 50-78 cm long, 0.8-1.6(-2.4)
cm wi

e
Nerves not raised; glabrous

25—40(—67) cm long, the spikelets not
overlapping

Small; first glume 3-12 mm long, the
seco EE 12-16 mm long, 7-9-

en pom texture smooth, the
rmal cells obscure in SEM
m long; a
valine i absent; pri
dant; short cells sain Ld
near apex hyaline
Glabrous
3, with soft, short hairs
Simple, grains 10—30 um in diameter

Cleft between scutellum and coleo-

rhiza not conspicuous

Seedlin

g Unknown
Chromosome number = 12

rhiza distinct
First leaf linear, erect
Unknown

structure except that the cleft between the scu-
tellum and coleorhiza was less conspicuous, often
present merely as an embayment on the lowe
side of the embryo (Fig. 4a).

DISCUSSION

Despite the statement of Jacques-Félix (1962)
that there is little difference between the species,
the Afro-Asian Streptogyna crinita and Ameri-
can S. americana are quite distinct from each

other in a number of characters (Table 2). The
long, scaly, flagelliform sympodial rhizomes of
S. crinita are rare in herbaceous bamboos, a sim-
ilar type being found only in some olyroids such
as Pariana parvispica. The divergence in lodicule
morphology (Fig. 3) between the species is great
and in most grass taxa would imply separation
at the genus level. Microhairs are not found on

the foliage of either species (Tateoka, 1958a;
Metcalfe, 1960; Jacques-Félix, 1962; Renvoize,
1985; Soderstrom et al., 1987), but this study. has

bau those found on the foliage (Metcalfe,
1960; Jacques-Félix, 1962) and lodicules (pers.
Obs.) of the African herbaceous bamboo genus
Guaduella (Guaduellae). Multicellular micro-
hairs are also found on the foliage (Smithson,
1956) and floral bracts (pers. obs.) of species of
Joinvillea (Joinvilleaceae), a group often consid-
ered to be among the closest relatives ofthe grass-
es (Stebbins, 1982; Dahlgren et al., 1985; Camp-
bell & Kellogg, in press). Multicellular microhairs,
consisting of three or more thin-walled cells, and
with a blunt distal cell, have also been noted in
the olyroid bamboos Maclurolyra tecta (Calde-
rón & Soderstrom, 1973), Diandrolyra tatianae

ed on Ivory Coast, Bamps 2175 (BR). Abbreviations: io coleoptile; cr, coleorhiza; ep, epiblast; If, first
um.)

embryonic leaf; ra, radicle; sc, scutellum. Scale bar: a, b =

um; c, d =

878

(pers. obs.), the woody bamboo Arundinaria va-
gans (Metcalfe, 1960), as well as in several other
woody bamboo genera such as Pleioblastus and
in several species of the arundinoid genus Dan-
thonia (Tateoka & Takagi, 1967). The presence
of microhairs on the lodicules but not on the
foliage of S. crinita provides an exception to the
generalization of these authors that only grasses
with microhairs on their leaves also exhibit them
on their lodicules. Tateoka & Takagi also illus-
trated a typical woody bamboo (Sasa species)
lodicule with cilia, prickles, short and long cells,
and bicellular microhairs that in many ways rep-
resents a composite of the features found in the
two Streptogyna species. The lodicules of S.
americana possess abundant marginal prickle
hairs (which occasionally grade into short cilia)
and have surfaces that are differentiated into long
and short cells in the upper portions. In S. crinita,
prickles are rare and cilia and short cells are ab-
sent on the lodicules, but microhairs are present.

The two species differ in stigma number and
morphology. Streptogyna americana possesses
three merely papillose stigmas, whereas S. crinita
has two stigmas that are coarsely armed with
retrorse barbs. However, in both taxa the stigmas
elongate after anthesis, becoming entangled with
each other and with the lemma awns. Reports
of three stamens in at least some florets of both
Streptogyna species (Doell, 1880; Bentham, 1883;
Jacques-Félix, 1962) could not be confirmed in
this stu

Streptogyna crinita consistently has small,
weakly to moderately compound starch grains
(Fig. le). This agrees with the descriptions of
Jacques-Félix (1962) and also Yakovlev (1950),
neither of whom cite specimens. The starch grains
of S. americana are larger and essentially simple
(Fig. 1f). Starch grain characters are not as reli-
able as anatomical or cytological data, but they
have been considered to be taxonomically useful
at the tribal level (Tateoka, 1962). Most bam-
busoids, as well as Joinvillea (pers. obs.), possess
compound starch grains.

The two species of Streptogyna are similar in
embryo structure (Fig. 4). Both possess typically
bambusoid embryos with a formula of F+ PP
(see Reeder, 1962); that is, no internode is pres-
ent between the divergence of the scutellar and
coleoptilar vascular traces (F), an epiblast is pres-
ent (+), a cleft between the lower part of the
scutellum and the coleoptile is present (P), and
the margins of the first embryonic leaf overlap
(P). Jacques-Félix (1962) illustrated an embryo

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

of S. crinita with no internode between the scu-
tellar traces (F), an epiblast present (+), and lack-
ing a cleft between the scutellum and coleorhiza
(F), but in all individual embryos of the speci-
mens that we examined a small embayment was
present at the base of the embryo that could be
interpreted as a cleft, although it was not as
prominent as in S. americana. In transverse sec-
tions the embryos of both species had a first em-
bryonic leaf with strongly overlapping margins
(P). The embryo of both species is unusual among
the Bambusoideae examined in that it is dis-
tinctly taller than it is wide.

CONCLUSIONS

Many grass genera occur in both the Old and
the New World, but only very few tropical-forest
Poaceae have this type of bihemispheric distri-
bution pattern. Among the woody bamboos,
Arundinaria and Bambusa (both Bambuseae),
taken in their widest sense, have been considered
to range across the Atlantic Ocean. However,
studies of these groups by Soderstrom, Ellis, and
collaborators are revealing them to be hetero-
geneous assemblages, and it appears Bambusa
cannot be considered to exhibit a true amphi-
Atlantic distribution. Among the herbaceous
bamboos, only Olyra and Streptogyna occur both
in South America and in the Old World, but
Olyra latifolia is a widespread, weedy species
that was almost certainly introduced by humans
into Africa from South America quite early.
Therefore the only bamboo genera with amphi-
Atlantic distributions are Streptogyna and pos-
sibly Arundinaria.

Streptogyna differs from the bambusoid core
group (Soderstrom & Ellis, in press) principally
in details of seedling morphology (the first ex-
panded blade of S. americana is narrow and ver-
tical, not broad and horizontal as in all other
bambusoids; Soderstrom, 1981) and lack of epi-
dermal papillae, microhairs, and well-developed
arm cells. Of these divergences, the distinctive
seedling morphology is perhaps the most signif-
icant (see Kuwabara, 1960), but the seedling of
S. crinita has not yet been examined. Strepto-
gyna resembles the woody bamboos in possess-
ing many-flowered spikelets, strongly developed
fusoid cells, an adaxially projecting leaf blade
midrib with complex vasculature, multicellular
microhairs (as noted, present in Guaduella and
several woody bambusoid genera), lateral ap-
pendages (Tran Van Nam, 1972), oral setae, and

1987]

especially an external ligule and deciduous leaf
blades. It may be noted that the multicellular
microhairs of Guaduella species occur on bot
the leaf blades (throughout) and lodicules (on
apices only) and that the basal cell is very long,
about half the total length of the hair. Strepto-
gyna crinita, in contrast, has uniseriate micro-
hairs in which the basal cells are not conspicu-
ously longer than the upper cells. Cytologically,
Streptogyna crinita is similar to the — uq
and has a basic chromosome number of n = 12,
as reported by Veyret (1958), REA (BAS
1965), Kammacher et al. (1973), and Dujardin
(1978), who studied material from Sri Lanka,
Uganda, the Ivory Coast, and Zaire, respectively.
Streptogyna americana has not been studied cy-
tologically.

The generalization that microcharacters in the
grass family are often more useful in generic,
tribal, and subfamilial delimitations than are

spikelet phology be-
cause microcharacters are under less intense se-
lective pressure thus finds an exception in the
case of Streptogyna. In this genus, floret mor-
phology, with its special adaptations to external
animal dispersal (Ridley, 1930; van der Pijl, 1982;
Soderstrom et al., 1987) 1s clearly a conservative
feature. The leaf anatomy of the two species is
quite similar, and one of the only consistent dif-
ferences between the two is the presence of more
prominent adaxial ribs over the veins in S. c

nita (Soderstrom et al., 1987) and the ape
presence of adaxial ciliate macrohairs in the lat-
ter species.

We have shown that in the habit, lemma tex-
ture, starch grain morphology, and especially the
lodicule structure the two Streptogyna species are
quite different from each other, to the extent that
segregation at the generic level might be seriously
considered. Based on a comparison with Join-

are probably derived from three, which S
icana retains. It seems likely that both species
evolved from an extinct ancestor and that their
separation is ancient. However, barring strong
differences in leaf anatomy and cytology, grasses
are traditionally segregated into genera on the
basis of gross spikelet morphology, and the
spikelet and floret structures of the two species
of Streptogyna are quite similar, as is the leaf

SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA

879

anatomy. It will be necessary to study the seed-
lings of S. crinita and the cytology of S. ameri-
cana before a final decision can be made on the
taxonomic level at which these two sibling species
should be recognized.

TAXONOMIC TREATMENT

Streptogyna Palisot de Beauvois, Essai Agrost.
80. 1812. TYPE SPECIES: S. crinita P. Beauv.

Streptia Rich. ex Doell in Mart., Fl. Bras. 2(3): 171.
1880. ;

Nomen nudum

Perennial forest grasses; culms solid, un-
branched above the base; leaf sheaths strongly
ribbed, extending upward along both sides of the
pseudopetiole and contiguous with the inner lig-
ule; outer ligule present as a short, indurate rim;
inner ligule short, membranous; lateral append-
ages and oral setae present at summit of leaf
sheath; leaf blades deciduous, linear to lanceo-
late, narrowed below into a short pseudopetiole,
the nerves parallel or very slightly oblique from
the midvein at its base; midvein and prima
nerves manifest only on the abaxial (lower) sur-
face, the secondary lateral nerves and transverse
veinlets inconspicuous; leaf margins antrorsely
scabrous. Inflorescence pedunculate, a spikelike
panicle, unbranched or occasionally with a spike-
like branch at the base; rachis 3-angled, one side
convex and other sides concave and alter-

ately bearing the spikelet pedicels. T
Bos pedi ET greenish, several-flowered, the
lower florets well developed, Hio wet, laterally
compressed, hermaphrodite, the upper ones pro-
gressively smaller and sterile, disarticulation oc-
curring between the fertile florets, each of these
falling attached to the extended curved rachilla
segment above it; glumes 2, membranous, per-
sistent, many-nerved, the first shorter than the
second and attached to the side of the thickened

2-keeled, sulcate between the keels;
elongate, strongly nerved; stamens 2, lateral an-
terior, the anthers basifixed; ovary with a long
style; stigmas 2 or 3, becoming hardened and
persistent, intertwined at maturity with the stig-
mas of other florets in the same spikelet and

880

inflorescence; fruit a linear caryopsis, the hilum
narrow, extending nearly the entire length of fruit,
the embryo small, basal. Basic chromosome
number, n = 12.

KEY TO THE SPECIES OF STREPTOGYNA

la. Plant with long, scaly rhizomes; stigmas 2,
retrorsely barbed; second glume 17-26 mm
long; base of lemma pilose; leaf blades 10-
25(-40) mm wide; — ERE

Stre ii iin crinita
lb. Plant cespitose from short, Seid dip es;

stigmas 3, subglabrous; second glum 12-16

mm long; base of lemma slabrous leaf blades

8-16(-24) mm wide; UE pical

ue [doppie americana

1. Streptogyna crinita P. Beauv., Essai Agrost.
80 + plate 16. 1812. TYPE: Nigeria: [“ prob-
ably gathered in the forests of Oware or Be-
nin" (Hubbard, 1956)] anno 1786-1788,
Palisot de Beauvois s.n. (holotype, P, not
seen).

Streptia crinita Rich., a herbarium name given as a
nonym of Streptogyna crinita by Doell in Mart.,

Fl. Bras. 2(3): 172. 1880.

Streptia secunda Rich., a herbarium name given as a
synonym of “yasa qapya crinita by Doell in Mart.,
Fl. Bras. 2(3): 172. 1880.

— e es Hook. f. in Trimen, Spars
Fl. n: 301—302. 1900. TYPE: Sri Lanka, w
out ii or collector, C. P. [Ceylon Plants] 922
(holot 1).

Culms 55-100 cm tall, each representing the
aerial extension of an upturned sympodial rhi-
zome, the culm itself producing at its base 1-3
additional rhizomes to 25 cm long, these with
short internodes about 1 cm long covered by the
overlapping sheaths; sheaths bladeless, ovate-
lanceolate, strongly striate, 7-10 mm long. Leaves
evenly distributed on culm, not overlapping;
sheaths finely ciliate on the margins, otherwise
glabrous; outer ligule 0.4—0.9 mm long, tipped
by a fringe of ciliate hairs ca. 1 mm long; inner
ligule 1-2 mm long, the upper margin erose, cil-
iate on the abaxial (outer) surface, glabrous on
the adaxial surface; sheath auricles 1-2 mm long;
lateral appendages 0.3-1.2 mm long; oral setae
sparse, delicate, less than 1 mm long; pseudo-
petioles 5-15(-25) mm long; leaf blades narrowly
to broadly lanceolate, 18-29(-40) cm long, 1-
2.5(-4) cm wide, acute at the tip, c) below
to the pseudopetiole, primary lateral veins 4-8
on each side of the midrib; upper blade surface
pale green, glabrous or occasionally with scat-
tered spinelike hairs; lower surface lighter green

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

than the upper surface, glabrous. Inflorescence
11-29 cm long, borne on a peduncle 2-10(-40)
cm long, erect at first, becoming strongly pendent
with age; spikelets (11—)20—30(-42), the glumes
of adjacent spikelets strongly overlapping; rachis
appressed-pubescent. Spikelets on pedicels 0.8—
1.7 mm long, 4—5(-7)-flowered; first glume 6.5-
10 mm long, linear-lanceolate, glabrous, 1—3-
nerved, transversely veined; second glume
(16-)20-23 mm long, 1.7-2.5 mm wide, ellip-
tic-lanceolate, glabrous, 1 1-17-nerved, the nerves
of various thicknesses, not all extending the length
of the glume, with numerous transverse veinlets,
the terminal awn up to 2 mm long, or occasion-
ally the apex emarginate; lemma of lowest floret
18-25 mm long, lanceolate, indurate except for
the broad, scarious margins, 7(—9)-nerved, the
callus 1.5-2.5 mm long, pilose, the terminal awn
(12-)15-26 mm long; rachilla internode attached
to base of lowest floret 3-4 mm long; palea about
as long as the lemma, not conspicuously pro-
truding from it; lodicules firmly membranous,
narrowly obovate or spatulate, with a fringe of
multicellular microhairs at the apex (rarely gla-
brous), the anterior pair 1.5-3.5 mm long, 1-5-
nerved, the posterior pair often slightly longer,
narrowly oblanceolate, 1—3-nerved; filaments
weak and ribbonlike, the anthers pale yellow,
about 4 mm long; ovary fusiform, long-ciliate on
the upper third and lower part of the style; style
long, flattened, with sparse appressed hairs; stig-
mas 2, strongly retrorsely barbed above, the
barbed portions elongating and coiling with age;
caryopsis 12-16 mm long, 0.8-1.2 mm wide,
slightly tapering to a persistent, ciliate beak at
its summit.

Additional d die examined. ANGOLA. CA-
BINDA (MAIO OMBE o Zau "Gossweiler 6557 (BM, K).
BENIN: Djou ` Chevalier 23891 (P.). CAMEROON:
Without aa ras anno 1908-1909, Buesgen 530 (US);
Mbamkin, Letouzey 2652 (BR); Dimako, rive
ine de la riviére Mbonda, Letouzey 2682 (BR); 38

m ESE Djoum, pres Akoafim, Letouzey 8400 (B, K);
x de Yokadouma, Meijer 15146 (MO); N'Kolbrisson,
ca. 8 km W of Yaoundé, de Wilde 1205 (B, BR, K,
MO); Yaoundé, Zenker 533 (U

, Chevalier 10552 (P); oa
Gounda- St. Floris National Park, 8 km S of Cam
Koumbala at confluence of Mbingou and Koumbala,
8°26'N, 21?15'E, Fay 4087 (K); Mbaiki, le Testu 3446
(BM), Tisserant 3446 (K). CONGo: Brazaville a St. Jo-
seph, Chevalier 27341 (K). EQUATORIAL GUINEA. CAMPO
DISTRICT: Bebady, route from Anio, Tessmann 658 (K).
NDO Po: anno 1859, Mann 108 (K, W). ETHIOPIA.
ILLUBABOR: E of Abobo, 7?48'N, 34°37'E, Chaffey 908
(K). GABON: without locality, Griffon du Bellay s.n. (K);

1987]

Limbareni, May 1875, collector’s name wee. (US);
Tchibanga, /e . TEN M). GHANA: 6 chs. from
Akudum, Ankra 5 (US); Amuni, Chipp 53 (K);
Ashanti, C ummins r 164; Pra Suhien E Reserve,
Deaw 363 (F, MO); Kade, Agricultural Research Sta-
tion, Enti s.n., GC-42033 (MO), Ankrah s.n., Ghana
Herbarium No. 20190 (K); Sekodumasi, Kitson 8 (BM);
E of Anyaboni at S edge of Afram Plains, Morton 6095
(K); Dawo Mato Kola, Thomas D-28 (K); Akatri, Tho-
mas D-90 (K); Atuna, Vigne 3522 (BR). GUINEA: Ba-
fing, Adam 13795 (MO); 50 km de Kindia vers Mansu,
Roberty 10682 (G); Télimélé nord, Roberty 10775 (G).
GUINEA-BISSAU: entre Sedengal e Ingone, Espirito San-
to Explorações Botánicas 3735 (US); Bedanda, Pereira
& Correia Eo K). INDIA: without exact locality, Wight
ag (G DA). KERALA: South Travancore, 2,000
yid P (BM, K); Vetillapara forest, Cochin,
jmd 12 (K); Courtallum, Tinnevelly District,
n 1353 (K, W). Ivory Coast: Um Adam
8 (MO); Duékoué-Buyo, fôret N du o, Bamps
ad (BR); near Issia, Daloa-Abidjan uL poA
13609 (K); Bingerville, = hevalier 16036 (P); Amatioré
Forest Reserve, 10 km W of Tiassalé, Fosberg 40539
(MO, US); Yalé, near M Nimba, Geerlin g & Bokdam
1828 (BR, MO); 5 km N of Sassandra, Pu
2261 (BR, MO); N of Sékré, ca. Í ee š

Oldeman 604 (BR, K, MO); 10 k
Oldeman 701 (BR, MO); N’Zidah, sarin 13683 (G,
MO); Dabou, Roberty 15528 (G); N'Zo, Roberty 16049
(G); Adiopodoumé, de Wilde 861 (BR). LiBER1IA: Die-
bla, Webo District, Baldwin 6305 (K, MO, NY, US);
Gbawia, Baldwin 67 14 (MO, NY, US); Gretown, Tchien
District, Baldwin 6924 (MO); Sanokwele, Baldwin 9550
d ina Kle, Boporo District, Baldwin 10570 (K, MO);
Wohmen, Vonjama District, Baldwin 12027 (MO);
hinterland of Monrovia, Dinklage 3375 (BR); Ganta,
Sanokwele District, Harley 959 (MO); Peahtah, Be-
quaert in Hb. Linder 1012 (K, US). NIGERIA: Itu, Cross
River State, Ariwaodo 83 (MO); Onitsha, Barter 1814
(K, W); Obom Itiat, um to Atam Eki, Calabar Prov-
ince, Jones in F.H.I. 6870 (K); Olokemeji Forest Re-
serve, Abeokula Province, Jones et al. in F.H.I. 14564
y; aga Ndoro Di gato Baissa po

: Fore est Herbarium Ibadan
61439 (K); Ilaro Forest Reserve, 45 Ga SW of pees

S.N.
Oban, Talbot 856 (BM);

valier 2390 (P); Bignona, Roberty 6424 (P). SIERRA
LEONE: without exact locality, ee & Smeathman
s.n., anno 1792-1796 (BM, S), J. E. Smith s.n., anno
1791 [*Guinea"] (BM). Mt. Loma, Adam 22422 (MO);
Njala, Dalziel 8420 (US); Kennema, Deighton 397 (K),
Thomas 7822, 7903 (K); Kambia, Deighton 838 (K);
Heddle’s Farm, Elliot 3939 (BM, K, US); near Kambia,
on Scarcies River, Elliot 4389 (BM); Jola, 20 Jan. 1927,
Fairchild s.n. (US); Zimi DM DE Fisher 1 (K); Ma-
T Glanville 58 (K), K. ai Reserve, Lane-Poole

4 (K); Kuntaia, ons 5 (K); Yakala, Thomas
24 (K); Jigaya, Thomas 2719 (K); Kanya, Thomas
2983 (K). SRI LANKA: without definite locality, C.P.
[^T hwaites"] 922 (BM, BR, G, K, W); Henaratgoda, 3
Jan. 1881, Ferguson s.n. (PDA, W); Matale, Dec. 1846,
Gardner s.n., C. P. 922 (PDA), Mar. 1883, Lawson s.n.
(K); Buttala to Sirigala, 3 Mar. 1907, Ro ck s. n. (PDA);
Dolukanda, Senaratna 2700 (PDA); Dulva Kanda, Ta-

SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA

881

teoka 599 (B). SUDAN. EQUATORIA: Talanga, Imatong
Mountains, 4°01'N, 32*45'W, Friis & Vollesen 484 (BR,
K); Lotti forest, Myers 9655 (BM); Sakure, Zande Land,
Wyld 334 (BM š
sawasawa, Haerdi 624 (BR, G). ToGo: Tomegbé, Bru-
nel & Heitz 5837 (B); Cascade de Tomegbé, S of Badou,
Ern 2096 (B); Plateau de Danyi, zwischen Adéta und
Ndigbe-Apédomé, Ern 2703 (B, K). UGANDA: Damba

Island, Kyagwe County, Dawkins 459 (BM, K), Mait-
land 801 (K); Gulu, Zoka forest, Acholi District,
Thomas 4031 (K). ZAIRE: Litendale, Achten 485-A (BR);
Barumbu, Bequaert 969 (BR); Avakubi, Bequaert 1726
(BR); Mayumbe N’Benga, Bittremieux 106 (BR); Men-
kao, Breyne 918 (BR): Mabana, Maluku, Breyne 3292
MO); Mayombi, Brishe 31 (BR); Luni, Brishe s.n.

Delhaye 24 (BR); Nkolo, SNE Devr 54 (BR);
ukolela, Dewevre 544 (BR); B eut Dewulf 331
(BR); INEAC, Luki, Du Bois 333 (BR, K), 334 (BR)

kamba, Bulungu, Dujardin 491 (BR); Bingila, Dupuis
s.n. (BR); Boyabokuda-Bogula (Badangabo), Evrard 310
(BR); Djoa, Bolombo, Evrard 4952 (BR); Tukpwo,
Gerard 2182, 4298 (BR); Ile Esali, Yangambe, Ger-
main 384 (BR, K), Louis 6948 (B, BR), 7900, 13072
(BR); Panza, Inongo, Gilbert 14213 (BR); Kisantu, anno
1900, Gillet s.n. (BR); Yambata, de Giorgi 1669 (BR);
Bolobo, env. Eala, Goosens 2448 (BR); Karawa, Uban-
gi River, Goosens 4123 (BR); Gatanga, de Graer 294

ville, 15 Jan. 1904, Laurent s.n. (BR); Gimbi, Laurent
600 (BR); Bolombo, Lebrun 605 (BR, US); entre Bu-
singa et Banzyville, Lebrun 2040 (BR, US); Moburasa,
p ps 193 (BR); Tambwe-Mvwenza, Dibaya, Liben
4 (BR); Tuzule, riv. Lubi, Liben 2976 (BR); Mu-
s Luluabourg, Liben 3507 (BM, BR); bord de la
cM 40 km N of Kisangani, Lisowski 16480 (BR);
panga, bord de la Mobi, 34 km SE : Kisangani,
Lisowski * 7219 (BR); Lovanium, Kinshasa, Lisowski
18349 (BR, K); 8 km N of Yakusu, Lisowski 86435
(BR); BR Booke wa Mbole, Yangambi, Louis 10780
(BR, NY); entre Ngazi et Aruwimi, Louis 12181 (BR);

dango 3050 (BR); Loata, Meurillon 23, 224 (BR); Yam-
bata, Montchal 136 (BR); Dundusana, Mortehan 634
(BR); Kaniana-Haut Lomami, Mullenders 472, 1226
maka, Luki, Nsimundele 57 (BR);

Pauwels 2360, 2388 (BR); Campus UNAZA (Lemba),
Kinshasa, Pauwels 6378 (BR); Epulu, Putnam 50 (BR);

s.n. (BR); Boguge, prés Mobw y
Luki, be de la Minkudu, Toussaint 2273 (BM, BR,
M); Sonso, Kwango, Vanderyst B-48 (BR); Kiala, Mar.

1907, T wm s.n. (BR);
et Wemba, = r

Sacre-coeur,
BR); Yindu, Puede 93 (BR); Kimuingu, Vanderyst

882 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

AFRICA
=

C
sn
i]
L
P pz
E
"ry.
c.l

(1000 km

Y f M fr A l | ad
\ VTACN
30
L
Y) Y
" U
I ea UN

800 MILES
° 400 800 1200 KILOMETERS |

SINUSOIDAL PROJECTION

| \ |
|
EL \ a \ l l 1 E l | J

Lo 20 O WEST LONGITUDE FAST NGITUDE ( 20 30 _ 40 50 | 60
GOODE BASE MAP SERIES Prepared by Henry M. Leppard
DEPARTMENT OF GEOGRAPHY © 1964 by The University of Chicago

Of
THE UNIVERSITY OF CHICAGO
HENRY M. LEPPARD, EDITOR

FIGURE 5. Distribution of Streptogyna crinita in Africa; inset, distribution in Sri Lanka and southern India.

1987]

313 (BR); Dima, Vanderyst 862 (BR); Iles du Kasai,
Vanderyst 993 (BR, US); Mokaba, Vanderyst 1693,
3606 (BR); Kikwit, Vanderyst 2783 (BR), 2913 (BR,
MO), 9235 (BR); Mukulu, Mee hae 91 (BR); Chen-
al, dei 4502 (BR); Yanga, Apr. 1915, Vanderyst

.H. mpako, Vanderyst 5429 (BR); Kisantu,
Vanderyst 5024, 20399, 29971, 29979, 32082 (BR);

(BR);

= 25681 (BR); Tsanga, Vanderyst 26986 (BR);
u, Kipako, Vanderyst 30696 (BR); Bokoimkori,

E. 32624 (BR); Lemfu, Apr. 1907, Vantolborg

o
name illegible] 21 6 (BR).

Streptogyna crinita is widely distributed in wet
to seasonally dry forests from sea level to 1,000
meters elevation in tropical Africa (Senegal to
southwestern Ethiopia south to Fernando Po,
northern Angola, and central Tanzania), south-
ern India (Kerala), and Sri Lanka (Fig. 5). Based
on abundant collection data from Africa, S. cri-
nita appears to flower all year, with a maximum
during October through January and slight min-
ima (as denoted by a small decrease in the num-
ber of flowering collections and a considerable
increase in the number of sterile gatherings) dur-
ing August and September and to a lesser extent
February and March, although Hens 156 states
that the species flowers "toute l'année" in Zaire.
Several collectors note that S. crinita may be
locally dominant, covering the forest floor in large
rhizomatous clones. The species is used in west-
ern Africa to catch mice and rats, the animals
becoming entangled in inflorescences that are
placed outside their holes (Hubbard, 1956).

zelius & Smeathman s.n., anno 1792-1796,
gave the herbarium name of “‘Aristidoides mu-
ricida" [*awned mouse-killer"] to the plant.

N

. Streptogyna americana C. E. Hubb., Hook.
Icon. Plant. 36(6): 1—6, tab. 3572. 1956. TYPE.
Suriname: trail to Coppename River, rear
of village of Paka-Paka, Maguire 23975 (ho-
lotype, K, not seen; isotypes, F, MO, NY).
Figure 6

Plant cespitose or rarely from a series of short,
knotty, horizontal sympodial rhizomes up to 2.5
cm long, the erect culms representing aerial ex-
tensions of very short sympodial rhizomes,
sometimes becoming decumbent and rooting at
the lower nodes. Leaves clustered at base of plant,
strongly overlapping, usually concealing all the
nodes, often displayed in a fan-shaped arrange-

SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA

883

ment; leaf sheaths glabrous below, ciliate on the
margins, hispid at the summit; outer ligule 0.6-
1.1 mm long, erose or with a smaller apical fringe
of cilia; inner ligule 1.1-2.7 mm long; sheath
auricles not developed; lateral appendages usu-
ally inconspicuous; oral setae not evident, 1-2
mm long; pseudopetiole not well differentiated
from remainder of blade; leaf blades linear, 50-
78 cm long, 0.8-1.6(-2.4) cm wide, glabrous,
oblique, the midrib noticeably excentric, flat but
often becoming inrolled; primary lateral veins 3—
5 on each side of the midrib; upper (adaxial)
blade surface dark green, the lower surface lighter
green; cross-veins inconspicuous. Inflorescence
25—40(-67) cm long, borne on a peduncle 1-8-
(740) cm long; spikelets 14—25(—49), the glumes
of adjacent spikelets not or only slightly over-
lapping; rachis subglabrous below, appressed-
pubescent above. Spikelets 4—6 flowered, borne
on pedicels 1-3 mm long; first glume 3-12 mm
long, linear to lanceolate, a 1-3(-5)-
nerved; second glume 10- m long, 1.1-1.7
mm wide, ovate-lanceolate, aoe 9-nerved, with
scattered inconspicuous transverse veinlets and
an awn up to 3 mm long; lemma of lowest floret
19-24 mm long, narrowly lanceolate, completely
glabrous, the (5—)7(—9)-nerves green, evident on
the inner surface but not visible on the granular
outer surface except near the summit ofthe body;
calluslike prolongation of lemma 1-2 mm long,
the terminal awn 12-21 mm long, arising from
between 2 inconspicuous teeth at the summit of
the body of the lemma; rachilla internode at-
tached to lowest floret, persistent, 4-6 mm long;
palea slightly longer than the lemma, usually pro-
truding from it by 0.5-3 mm; lodicules narrowly
lanceolate, firmly membranous and strongly
nerved below, often abruptly widening about '4
of the way from the base, then tapering in the
upper 75 to an attentuate, nerveless, hyaline apex,
this usually fringed with prickles or occasionally
with a few cilia, microhairs absent; anterior lod-
icules 3.2-6 mm long, 0.5-1 mm wide, 1—3(-5)-
nerved, the posterior lodicule often slightly
shorter and narrower, | (—3)-nerved; anthers 2.5-
3.5 mm long, narrowly linear; ovary glabrous;
stigmas 3, lacking coarse barbs, glabrous below,
at maturity hispidulous-papillose adaxially near
the summits; caryopsis 14-17 mm long, 1-1.2
mm wide, glabrous.

_ Addit nd specimens examined. BELIZE: Cohune

n road, Gentle 8121 (BM, F, G, NY, S, US);
Cohune riae. ill slope, Hummingbird Highway,
Genile 8682 (BM, F, G, NY, S, US); 40 miles section

884 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

A 4 — =
POP
f

FIGURE 6. Streptogyna americana. — A. Habit of plant, showing deciduous leaf blades.—B. Ligular region,
showing hispid sheath summit, outer ligule (left), and inner ligule and oral setae (right). — C. Section of abaxial
surface of leaf blade, showing midrib (left) and absence of transverse veinlets. — D. Spikelet.—E. First glume. —
F. Second glume. — G. Base of floret, showing basal prolongation of lemma and persistent rachilla internode.—

. Lemma, unrolled, showing inner surface. — I. Palea, ventral surface. — J. Palea, profile. — K. Lodicules, with
posterior member in center. — L. Androecium of two stamens and gynoecium. — M. Stamen. — N. Stigma, showing

1987]

Hummingbird Highway, Gentle 9005 (S, US). BOLIVIA.
BENI: Alto Ivon, 11?45'S, 66°02'W, Boom 4792 (NY),
4827 (MO, NY, US); 18 km E of Riberalta, 11°05’S,
65°50'W, Solomon 6169, 7801 (MO). BRAZIL. ACRE:
125 km from Rio Branco on road to Pórto Velho,
9°45'S, 66°20'W, Calderón & Soderstrom 2301 (US).
AMAPÁ: Rio Jari, near Cachoeiras das Guaribas, 0?24'N,
53°07'W, Egler & Irwin 46417 (NY, US); road to Ama-
pá, vicinity of km 108, Rio Pedreira, Pires & Caval-

, 2.5 km sommet, Sastre 1656 (CAY
nte Pascoal, p S, 39°24’ W,

2410 (B, NY o
101 at point 13 km N of Itama arajú, Soderstrom et al.
2193 (CEPEC, US). E : Reserva Florestal
de Linhares, 19°24’ S. 40°04 W, Martinelli & Soder-
strom 9757 (RB), Soderstrom & Sucre 1883 (CEPEC,
US). MARANHÃO: Fazenda Bacaba, Doctor Haroldo, 5
km S of MA-119 from entrance 3 km NW of Lago do
Junco, 4?26'S, 44°58’ W, Daly et al. 469 (MO, NY); Rio
Pindaré, Monção, Fróes 20312 (US); Caxias to Barra
do Corda, before Curador [Pres. Dutra], Swallen 3583
(US). MATO GRosso: Fazenda Az de Ouro, 14?13'S,
57*02'W, Amaral 9 (RB); 260 km along road NNE of
Xavantina, a few miles W of base camp at 12°51’S,
51?45'W, Eiten & Eiten 8903 (US); 5 km NW of base
camp, 12?49'S, 51?46'W, Harley & Souza 10271 (US);
8 km NE of base camp, Ratter et al. 974 (NY, US); 4
km N of base camp, Ratter et al. 1820 (K, NY, US);
Serra do Itapirapuan, Alfonso, Lindman A-3347; Sar-
aré, 15?05'S, 59°50'W, Pires & Santos 16316 (MO,
NY). PARÁ: Belterra, Black 47-936 (NY); inter Sào
Joao et Santa Anna, Burchell 9201 (BR, US); Rio Part
de Oeste (TIRIOS), Cavalcante 824 (US); Gorotire vil-
lage at Rio Fresco, 7°47’S, 51°07'W, tidie QA ad -
sey 17-22183, 32-24183 (MO); Curuá Al r,
Kuhlmann yp iud S); Maicuru, s Francisco, Pires

Y, US); Serra Buritirama entre B-2 e
Marabá, ese 12320-A (US); range of low hills
ca. 20 km of Rendencao, near Córrego pad João
eresa, 8°03’S, 50°10'W,

Rendencáo on road to Barreiras dos Campos: Fazenda
Inajapora between Rio Inajazinho and Rio Inajá, ca.
8945'S, 50°25’W, Plowman et al. 8883 (F, MO, NY,
US); Belém-Brasilia Highway 17 km S of Ligacao do
Para, near km 1,509, ca. 4°17'S, 47°32'W, Plowman et
al. 9409 (F, MO, NY, US, WIS); 6 km N of Ligação
do Para, near km 1,532, 4°05'S, 47°32'W, Plowman et
al. 9522 (MO); 12 km E of Représa Tucurui (Rio To-
cantins), 3°45’S, 49°40'W, Plowman et al. 9799 (NY);
Jari, estrada do Munguba, km 14, Si/va 2198 (MO,

Y); Lageira, airstrip on Rio Maicurt, 0°55'S, 54°26' W,
Strudwick & Sobel 3109 (F, MO, NY); Sete Varas air-
strip, Rio Curua, 0°59’S, 54?29'W, Strudwick & Sobel
4289, 4313 (F, MO, NY); Santarém, Swallen 3281

SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA

885

(US); Japanese concession 35 km N of Monte Alegre,
Swallen 3411 (US); Obidos, Swallen 5089 (US ).

Cordeiro 924 (MO); 1 km N of Riberáo, road Abuná-
Guajará-Mirim, Prance et al. 6454 (MO, NY, US),
6527 (F, MO, NY, S, US); in sylvis umbrosis ad flumen
Guaporé, Riedel 1248 (G, NY); Mineração Mibrasa,
setor Alto Candeias, km 128, 10?35'S, 63°35'W, Teix-
eira et al. 625 (MO). RORAIMA: Conceição, Rio Blanco,
Luetzelburg 21383 (M). COLOMBIA. CAQUETÁ: Entre

3629 (F). META: E artin,
del Cano Camoa, í. 1687 ); Monte de
Caño oa, Herman S); sabanas de Sa
Juan de Arama, margen (Ee. ] iiejar, ater-

rizaje “Los Micos,” Idrobo & Schultes 608, 1217 (US);

margen izquierda del Rio Sansa, Sierra de la Macarena,
Idrobo 2160 (NY); Cano Ciervo, Sierra de la Macarena,
Philipson et al. 2023 (BM, S, US); margen derecha del
Rio Guayabero, Raudal de la Macarena (Angostura 1),
Pinto & Bischler 334 (US). VICHADA: ca. 35 km from
Las Gaviotas on road to Santa Rita, Davidse & Llanos
5211 (MO); Gualandayas, ca. 100 km E of Gaviotas,
Wood 4220 (K). CosrA RICA. PUNTARENAS: Finca Los
Helechales, between Buenos Aires and Cerro Pittier,
Hatheway 1686 (US); Los Tejares de Buenos Aires,
Pittier 10602 (BR, M, US, W); entre le Rio del Con-
vento et Buenos Aires, Tonduz 3643 (BR, W). FRENCH

Cremers 4546 (CA
pann-Mitaraka (frontiére) P.K. 7.5, Granville 1139,
1417 (CAY, US); versant N des Monts Galbao, 10 km
WSW de Saül, 400 m, Granville 1621 (CAY); 14 km
de Dégrad Claude, Granville 2267 (CAY); Sommet Ta-
bulaire, ca. 50 km SE Saül, Granville 3586 (CAY, MO,
: min des Emérillons, 1 km de Saut Verdun,
Granville B- 5037 (CA Y); Cayenne, Chemin du Moulin
Vidal, 13 July 1955, Hoock 1187 (K, P); Saül, 30 June
1956, Hoock s.n. (NY); Cayenne, anno 1839, Leprieur
s.n. (G). anno 1866, Jelski s.n. (W); Montabo, Herb.
L. C. Richard s.n. (Wy; Karouany, Sagot 1076 (BM,
W). GUATEMALA. ALTA VERAPAZ: ca. 6 km E of Sebol
on San Luis Road to Achiote, Contreras 4486 (US).

MEXICO. CHIAPAS: Javalinero, Palenque, Matuda 3637

, US). VERACRUZ: Sanborn [ca. 17?34'N, 95?07' w],
Orcutt 2933 (K, MO, US). NICARAGUA. ZELAYA: region
of Braggman's Bluff, E nglesing 218 (F); Miguel Bikan,
aspám, P.

Lake, Miller 2045 (US yh y
27480 (MO, US); forest eee telephone cable trail

—

adaxial papillae near tip.—O. Caryopsis, ventral surface showing linear hilum.—P. Caryopsis, dorsal pid

showing small, basal embryo. (Based on uie
for CF, L J, L, O, P; 2 mm for B, G, H;

zil, Soderstrom et al. 2193 xa Scale bar = 24 mm for A; 4
mm for K, M; an N.)

nd 0.5 mm

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

Ratte d
O 100 200 300 400 500 600 mies
Prepared by Hendrik R Rypkema \

30

|

FiGuRE 7. Distribution of Streptogyna americana.

between S-16 and S-49, Rio Indio, Steyermark & Allen
17459 (BR, G, MO, S, US). DARIEN: Cerro Pirre, Gentry
& Clewell 7147 (MO), Mori & Kallunki 5374 (MO,
; 0.5-2.5 km NE of Manené, Hartman 12175 (MO).
PANAMÁ: San José Island, Johnston 433 (BM, US), 728,
1115 (US). PERU. MADRE DE pios: Rio Tambopata,
Lago 3 Chimbadas, ca. 65-70 river km SSW of Puerto
Maldonado, ca. 10-15 km air SW eff. Río La Torre,
12°49 9°17'W, P 5762 2: SURINAM:
without locality, Weigelt s.n. (W); Zuid River, 3 km
above confluence with Lucie River, /rwin et - 55900
(B, MO, NY, US); ab Wia wia-bank e Zwie-
belzwamp, Lanjouw & a 1153 (NY) ad
prope Moengo, Lindeman F); Brownsberg, near
Is fall, Lindeman ATES (K, ih Lucie River, ca.
2 km below affluence of Oost River, Maguire et al.
54093 (NY, US); Avanavero oever, Stahel & Boon-
acker 4579 (US). TRINIDAD: without locality, Botanical
Garden Herbarium No. 3367 (US), Crueger 362 (US
fragment), Finlay 3 (K, W); Caparo forests, Broadway
4932 (US); Tabaquite, edge of High Woods, Hitchcock
10122 (BM, US). VENEZUELA: Alto Orinoco, Rusby &
Squires 349 (K, NY). AMAZONAS: vicinity of Culebra,
Río Cunucunuma, 3?40'N, 65?45'W, Steyermark &

Delascio 129185 (MO); Mavaca, Alto Orinoco, Aris-
teguieta & Lizot 7380 (NY). BOLIVAR: alrededores km
88, carretera El Dorado, Aristeguieta 3713 (MO); 17
km W of Río Caura on road between Caicara and
Ciudad Bolivar, Davidse 4443 (MO, WIS); 10 km SW
of Río Aro on road between Caicara and Ciudad Bo-
livar, Davidse 4476 (MO); km 28, S of El Dorado,
Davidse 4966 (MO); 20-35 km SE of Monteco on road
to San Pedro de las Dos Bocas, 7?10'N, 62*55'W, Lies-
ner & Gonzalez 5851 (MO); along pica 105, 40 km S
of Tumeremo, 29 km N of El Dorado, Steyermark
86572 (NY); Pica La Lira, at km 27 S of El Dorado,
l m E of hi ave ages 86638 (NY); woods
bordering savannah by Rio

S : hee Par ke Steyermark 86760 (U
AMACURO: 3 k Piacoa [p E in the state of
Delta yen ancl Steermark tag :

Rio Cuyubini, Cer a,
(NY). SUCRE: S ies n in Imp
Cedeno and Boca del Tataricual, along Quebrada Im-
posible, Steyermark 62845 (F, US).

a, above raudal Cotua,
S). DELTA

Streptogyna americana is found in shaded,
well-drained sites in moist forests below 500

1987]

(-800) meters from Veracruz, Mexico, and
Trinidad south to northern Bolivia and Espirito
Santo, Brazil (Fig. 7). Most common on the mar-
gins of the Guyana Highlands and in easternmost
and southernmost Amazonia, this species is rare
or absent in the central portion of the Amazon
Basin. Collectors in Panama (Judziewicz 4440),
Venezuela (Davidse 5211), and Surinam (Irwin
55900) noted that the leaf blades become inrolled
during hot dry weather or soon after collection.
Most Central American collections were made
November through April, indicating a dry season
peak of bloom, while Guyanan and eastern Am-
azonian collections have been made principally
June through August; the five Atlantic Brazilian
gatherings were made from March to May. Com-
of S. americana include “barba de
azil) and "barba-
tigre" (Cabrera 3629, CE. suggesting ex-
ternal animal dispersal.

LITERATURE CITED

BENTHAM, G. 1883. Gramineae. In G. Bentham & J.
. Hooker, Genera Plantarum, Volume 3, Part 2.

R. SODERSTROM. 1973. Mor-
phological and carpets considerations of the
grass subfamily Bambusoideae based on the new
genus Maclurolyra. Smithsonian Contr. Bot. 11:
i-i, 1—

1980. The genera of the Bambu-
soideae (Poaceae) of the American continent: keys
nd comments. Smithsonian Contr. Bot. 44: 1

CAMPBELL, C. S. & E. A. KELLOGG. c: press. Sister
group EENE om the Poaceae. /n T. R. So-

l. (ed tors), Una cedet s and
Wash-

Symposium on Grass Systematics and Evolution.
Washington, D.C., 27-31 July 1986.]

CLAYTON, W. D. & S. A. RENVOIZE. 1986. Genera
Graminum. Her Majesty's Stationary Office, Lon-
don

DAHLGREN, R. M. T., H. T. CLIFFORD & P
1985. The Families of the vetula ua
Springer-Verlag, Berlin.

DoELL, J. C. 80. Tribe 11, Bambusaceae. 7n C. F
P. von Martius (editor), Flora Brasiliensis 2(3):
161—220, pls. 44—56. [Fascicle 83.]

DuJARDIN, M. 1978. Chromosome numbers of some
tropical 'African grasses from western Zaire. Ca-
nad. - Bot. 56: patient

ERR 1887. Gra

Peer Die a in. spa] sa ea pex E

9

HOOKER, : 1900. In H. a teri Hand-
book an Flora of Ceylon, V e

HUBBARD, pi E. 1936. Pp. 475- 602 in} Hutchinson

SODERSTROM & JUDZIEWICZ—SYSTEMATICS OF STREPTOGYNA

887

& J. M. Dalziel (editors), Flora of Tropical West
Africa. Crown Agents for the Colonies, London.
1956. Streptogyna crinita Beauv., Gramin-
eae, Tribus Streptogyneae. Jn Hook. Bot. Icon.
36(6): 1-6, tab. 3572.
É 62. Les graminées d'Afrique
cale. I. Généralités, classification, descrip-

Viviéres, Bull. Sci. (Paris) 8: xi + 1-345.
. ANOMA, E. ADJANOHOUN & L. AKE
3. Nombres chromosomiques de Gra-
ées de Cóte d'Ivoire. Candollea 28: 191-217.
SERE Y. 1960. The first FUE, leaf in grass
systematics. J. Jap. Bot. 35: 5.
METCALFE, C. R. 1960. . of the Monocoty-
ledons. I. Gramineae. Clarendon Press, Oxford.
NEES VON ESENBECK, C. G. D 35. Bambuseae Bra-
silienses. Recensuit, et alias i in India orientali pro-
4.

. 1812. Es-
une Nouvelle Agrostographie: ou Nouveaux

Gen nres des Graminées; avec Figures Sali in
i: d de Tous les Genres. Fai

PIL, 1982. Principles of olde in
te "Plane 2nd edition. Springer-Verlag, Ber-
lin.

REEDER, J. C. 1962. The bambusoid embryo: a reap-
praisal. k J. Bot. 49: 639-641.

RENVOIZE, S. A. 85. A survey of leaf-blade anat-
omy in — V. The bamboo allies. Kew Bull.
40: 509- cy

RIDLEY, H. 1930. The Dispersal of Plants
=a i the World. L. Reeve, Ashford, Kent,
England.

nglan

SMITHSON, E. 1956. The comparative anatomy of the
Flagellariaceae. Kew Bull. 3: 491-501.

SODERSTROM, T. R. Some evolutionary trends
in the Bambusoideae (Poaceae). Ann. Missouri Bot.
Gard. 15-47.

R. P. ELLIS. In press. The position of bam-

and Evolution. Washington, D.C., 27-31 July
86.]

E. J. JUDZIEWICZ. ded pes Phar-

eae and Streptogyneae of Sri orpho-

logical-anatomical study. Hu meu Cae Bot.

65: 1-27.

STEBBINS, G. L. 1982. Major trends of evolution in
the Poaceae and their possible significance. Pp. 3-
36 in J. R. Estes et al. (editors), Grasses and Grass-
lands. Univ. Oklahoma Press, Norman, Okla-
homa.

SrEUDEL, E. G. 1853 [1855]. ped: ii T
Glumacearum, Part 1. J. B. Metzler, Stu

n the genus [oed (Po-

: 364-366.

omatic chromosomes of Leptaspis
and Streptogyna (Poaceaz). Nature 182: 1619-

Starch grains of endosperm in grass
systematics. Bot. Mag. (Tokyo) 75(892): 377-383.

888

1965. Chromosome numbers of some East
African grasses. Amer. J. Bot. 52: 864-869.
—— & T. TAKAGI. 1967. Notes on some grasses.
XIX: systematic significance of microhairs of lod-
icule epidermis. Bot. Mag. (Tokyo) 80(952): 394—

4

TRAN VAN Nam, MME. (TRAN THI TUYET Hoa). 1972.
Les "oreillettes" des Graminées. Bull. Soc. Bot.
France 119: 441—462.

VEYRET, Y. 1958. Observations caryologiques chez
quelques Graminées tropicales. J. Agric. Trop. Bot.
Appl. 5: 308-310.

YAKOVLEV, M. S. 1950. Struktura endosperma 1 za-
rodysha zlakov kak sistematicheskii priznak.
Moskva , Lenin ngrad: Akad

seriya orfologiya i Anatomiya Rastenil) 1:
122-218. [Endosperm and embryo structure of
grasses as a taxonomic feature. Moscow, Lenin-

grad: Transactions ofthe V. L. Komarov Botanical
Institute of the Academy of Sciences of the USSR,

series 7 (Morphology and Anatomy of Plants) 1:

121-218. Unpublished English translation by the
Indian National Scientific Documentation Centre,
prepared in 1975.]

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

APPENDIX I.

LIST OF TAXA CITED

Arundinaria vagans Gamble
am h

Joinvilleaceae Tomlinson
Maclurolyra tecta Calderón & Soderstrom
Feds latifolia L
na parvispica R. Pohl
Pleiobla stus Nakai
Sasa Makino & Shibata
Streptia crinita Rich. ex Doell
Streptia secunda Rich. ex Doell
Streptogyna P. Beauv
S. americana C. s Hubb.
S. crinita P. Bea
S. gerontogaea Hook. f. in Trim
Streptogyneae C. E. Hubb. ex ^p E & Calderón

WOOD ANATOMY OF NOTEWORTHY SPECIES OF
LUDWIGIA (ONAGRACEAE) WITH RELATION TO
ECOLOGY AND SYSTEMATICS!

SHERWIN CARLQUIST?

ABSTRACT

Ludwigia anastomosans, a tree to 10 m, is studied because it is unusual in the genus in its arborescent
habit. It proves to vods a = vessels; it also has interxylary phloem, hitherto reported for

only one species of the

they may be more

a are the first observed in Onagrac
hos

. Wood anatomy of Ludwigia peduncularis and L. torulosa shows that
eae uc Bud don so far indicated. Vestigial bars on perforation plates of L.
aceae and are believed to represent an instance of paedomor-

phosis, but also retention of a primitive feature. Other indicators of paedomorphosis in Ludwigia are

abundance
grouping is related to ecology in taxa having fi

of erect ray cells and — long vessel elements. The hypothesis that degree of vessel
ber -tracheids Or libriform fibers i is validated by Ludwigia,

which has the lowest degree of the
y, combined in ‘the Mesomorphy ratio, present a not dissimilar pattern

features reflective of ecolog

1dilill

c. Other eyes e

that can be integrated with hn given by

HF

rates and temperature regimes as well as water availability.

Data on wood anatomy of eight species of Lud-
wigia have been presented earlier (Carlquist,
1975, 1982a). That number seems small unless
one takes into account the fact that Ludwigia is
predominantly herbaceous; the most familiar
species are nonwoody herbs of very wet habitats
such as ponds, ditches, and streams. One of the
species in the present study is a notable excep-
tion: L. anastomosans (DC.) Hara is a tree. The
data on the collection studied here describe it as
a tree 10 m tall with a trunk 15 cm dbh. Wood
anatomy is of special interest because of this hab-
it. In fact, the results obtained below demon-
strate that the wood of L. anastomosans differs
appreciably from that of other Ludwigia species.

Dr. Elsa Zardini, who collected the material

see if wood anatomy demonstrated the degree of

relationship between them. Dr. Zardini has con-

templated the idea that L. peduncularis (Griseb.)

Gómez may be closely related to L. torulosa (Arn.)
ara.

The wood anatomy of Ludwigia is of consid-
erable interest with respect to ecology because
Ludwigia characteristically grows in very wet
places. Ludwigia anastomosans was collected in
bamboo clumps by a blackwater stream in the

Parque Natural de Caraca, Minas Geraes, Brazil.
The material of L. peduncularis came from ditch-
es near Havana, Cuba. The L. torulosa specimen
was collected in a natural pond 17 km south of
Tumeremo, Distrito Roscia, Estado Bolívar,
Venezuela. In taxa with libriform fibers such as
Onagraceae, Carlquist (19842) hypothesized de-
gree of vessel grouping to be in direct proportion
to adaptation to dry conditions. In this case Lud-
wigia species ought to exhibit a low degree of
vessel grouping. Although figures for vessels per
group were developed in the earlier survey o

woods of the family (Carlquist, 1975), no com-
parisons were made between those figures and
ecological regimes occupied by the various
species.

Ludwigia is a group of interest with relation
to paedomorphosis in wood anatomy. This is, in
turn, related to habit and ecology. The species
in the present study were examined in this con-
text to see if woodiness in Ludwigia is primary
or secondary.

Although a study on wood anatomy can be
expected to reveal new records for anatomical
features, two in the present study proved of es-
pecial interest and worthy of discussion: occur-
rence of interxylary phloem and presence of ves-
tigial bars on some perforation plates.

' Grants from the National Science Foundation to Peter H. Raven made possible the collection and selection

of the material for this researc!
? Rancho Santa Ana Botan
Pomona College, Claremont, California 91711,

ANN. Missouni Bor. GARD. 74: 889-896. 1987.

c Garden, Claremont, California 91711, U.S.A., and Department of Biology,
U.S.A.

890

MATERIALS AND METHODS

Voucher specimens are located at the Missouri
Botanical Garden. Appreciation is expressed to
Dr. Elsa Zardini for providing dried wood sam-
ples suitable for study. For L. peduncularis and
L. torulosa, samples represented basal portions,
but diameter was small (2 and 4 mm, respec-
tively. The material of L. anastomosans was
from a branch about 1.5 cm in diameter. Al-
though this is much less than the 15 cm diameter
reported for trunks of this species, the branch
material is considered here to represent an es-
sentially mature pattern.

Woods were boiled in water, stored in 5096
ethyl alcohol, and sectioned on a sliding micro-
tome. Sections prepared in this way were, in part,
satisfactory, but cell collapse on account of thin-
ness of wood cells was excessive in some in-
stances. Therefore an alternative method in-
volving further softening, embedding in paraffin,
and sectioning on a rotary microtome (Carlquist,
1982b) was employed. Sections prepared by both
techniques were stained in a safranin-fast green
combination. Macerations were prepared with
Jeffrey's Fluid and stained with safranin.

Means are based upon 25 measurements (few-
er if feature is scarce) except for vessel wall thick-
ness, libriform fiber diameter, and libriform fiber
wall thickness, in which a few typical cells were
measured. Vessel diameter includes the wall, al-
though lumen diameter may be preferable for
some purposes and may be calculated by sub-
tracting wall thickness from the data presented
here. Mean values for vessel grouping are ob-
tained on the following basis: a solitary vessel —
1.0, a pair of vessels in contact — 2.0, etc. Bark
was not observed specifically in the present study,
although sections of the stem of L. torulosa in-
cluded portions of the spongy stem covering that
proves to be aerenchyma like that figured for
“Jussiaea repens" L. (now a Ludwigia) by Met-
calfe & Chalk (1950), and studied in detail by
Ellmore (1981) for L. peploides (HBK) Raven.

ANATOMICAL DESCRIPTIONS

Ludwigia anastomosans (DC.) Hara, Zardini &
Gentry 2175 (Figs. 1—6).

Growth rings inconspicuous, and probably re-
lated to water level of the riparian habitat (Fig.
1). Mean number of vessels per mm?, 51. Mean
number of vessels per group, 1.24; vessels tend-
ing to be grouped into radial multiples (Figs. 1,

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

3). Mean vessel diam., 109 um. Mean vessel wall
thickness, 2.5 um. Mean vessel element length,
458 um. Perforation plates simple (Figs. 2,

Lateral wall pitting of vessels alternate, pits
crowded and circular to polygonal in outline,
about 12 um diam. on vessel-vessel interfaces
(Figs. 4, 5). Vessel-axial parenchyma and vessel-
ray pitting alternate to scalariform, pit apertures
long (sometimes scalariform), pit apertures wide
(“gaping”). Pitting with relatively conspicuous
vesturing on vessel-vessel pits (Fig. 5, upper
right), vesturing less pronounced on vessel-axial
parenchyma and vessel-ray pits. Occasional ves-

loses abundant (Figs. 1, 3). Imperforate (cheap
elements all libriform fibers because pits appar-
ently simple, although a few exceptional pits with
small borders also observed. Many libriform fi-
bers septate. Mean libriform fiber diam., 28 um.
Mean libriform fiber wall thickness, 2.3 um. Mean
libriform fiber length, 588 um. Numerous libri-
form fibers with gelatinous walls (Fig. 6) and
therefore probably reaction wood. Axial paren-
chyma vasicentric scanty. Bands of phloem-con-
taining axial parenchyma present in marginal po-
sitions (end of growth rings) or scattered without
any relation to growth rings (Figs. 1, 3). Rays
multiseriate and uniseriate, the former slightly
more frequent (indicated by relative heights in
Fig. 2). Ray cells predominantly erect and square
(Fig. 2), a few procumbent cells present in mul-
tiseriate portions of multiseriate rays. Mean mul-
tiseriate ray height, 2,707 um. Mean uniseriate
ray height, 349 um. Mean width multiseriate rays
at widest point, 3.95 cells. Ray cell walls mod-
erately thin, lignified. Wood nonstoried. Raph-
ides present in phloem-containing axial paren-
chyma strands.

Ludwigia peduncularis (Griseb.) Gomez, Ekman
13416 (Figs. 7, 8).

Growth rings absent (portion studied probably
less than one year's accumulation). Mean num-
ber of vessels per mm’, 62. Mean number of
vessels per group, 1.30. Mean vessel diam., 50
um. Mean vessel wall thickness, 1.8 um. Mean
vessel element length, 484 um. Perforation plates
simple. Lateral walls of vessels with crowded al-
ternate elliptical pits about 4 x A. Emo with
pointed ends l-vessel
contacts). ensis pi mE and vessel-
ray pitting similar but with pits longer (appearing
pseudoscalariform) and wider, and with wider

1987] CARLQUIST—WOOD ANATOMY OF LUDWIGIA 891

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FiGURES 1-4. Wood one of Ludwigia anastomosans, Zardini & Gentry 2175.—1. Transection; termi-
nation of growth ring occurs in the middle of the photograph. —2. Tangential section; multiseriate rays are wide,
more numerous than uniseriate rays.— 3. Portion of transection, showing strands of interxylary phloem (arrows)
and vessels in radial multiples.—4. Portion of vessel wall from tangential section; intervascular pitting and
A dopa: plate rim, below; axial parenchyma cells, upper left. Figures 1, 2, i5) r siapa scale above Figure

finest divisions = 10 um); Figure 3, divisions = 10 um; Figure 4, divisions = — JOu

892 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

*
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"Vac

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FIGURES 5-8. Wood sections of Ludwigia. 5, 6. L. anastomosans, Zardini & Gentry 2175. —5. Intervascular
pitting from tangential section, showing transition between circular and polygonal pit shapes; vesturing evident
in pits at upper right. —6. Portion of vessel and associated cells from tangential section; note aberrant pit and
pit aperture configurations and splits in gelatinous fiber walls (right). 7, 8. L. peduncularis, Ekman 13416.—
7. Transection; vessels solitary; dark-colored deposits in ray cells. — 8. Tangential section; all rays are uniseriate.
Figures 5, 6, magnification scale above Figure 4; Figures 7, 8, magnification scale above Figure 3.

1987] CARLQUIST—WOOD ANATOMY OF LUDWIGIA 893

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FIGURES 9-13. Wood sections of Ludwigia torulosa, Holst, Steyermark & Manara 2257.—9. Transection;
libriform fibers are thin-walled.— 10. Tangential secti Itiseriat „l juent tł iseriat
rays.— 11. Portion of radial section showing vestigial bars on a perforation plate.— 12. Portion of a radial section
showing vessel-ray pitting.— 13. Portion of a radial section showing rodlike crystals in two ray cells. Figures 9,
10, magnification scale above Figure 1; Figures 11-13, scale above Figure 4.

894

apertures. Pits of vessels vestured, less so in ves-
sel-axial parenchyma and vessel-ray contacts
than on vessel-vessel pitting. Imperforate tra-
cheary elements all libriform fibers, the pits mi-
nute, apparently simple. Libriform fibers mostly
septate. Mean libriform fiber diam., 18 um. Mean
libriform fiber wall thickness, 1.2 um. Mean li-
briform fiber length, 778 um. About half of the
wood consisting of gelatinous fibers (Fig. 7),
therefore apparently reaction wood. Axial paren-
chyma vasicentric scanty, very few strands ad-
jacent to each vessel (or some vessels with no
adjacent axial parenchyma). Interxylary phloem
absent. Rays almost all uniseriate (Fig. 8). Ray
cells all erect. Mean uniseriate ray height, 1,566
um. Ray cells moderately thin-walled but ligni-
fied (Fig. 8). Wood nonstoried. No crystals ob-
served. Axial parenchyma and ray cells contain-
ing droplets or massive accumulations of a
brownish substance (Figs. 7, 8).

Ludwigia torulosa (Arn.) Hara, Holst, Steyer-
mark & Manara 2257 (Figs. 9-13)

Growth rings absent (portion studied probably
less than a year's accumulation). Mean number
of vessels per mm?, 62. Mean number of vessels
per group, 1.55. Mean vessel diam., 55 um. Mean
vessel wall thickness, 1.6 um. Mean vessel ele-
ment length, 374 um. Perforation plates mostly
simple, but some with modified or vestigial bars
(Fig. 11). Lateral walls of vessels with alternate
polygonal pits about 5 um diam. on vessel-vessel
contacts. Vessel-axial parenchyma and vessel-
ray pits alternate, opposite, or scalariform, larger
and with wider apertures than the vessel-vessel
pits (Fig. 12). Vestures dense within cavities of
vessel-vessel pits, somewhat less abundant on
vessel-axial parenchyma and vessel-ray pits.
Imperforate tracheary elements all libriform fi-
bers, because the pits apparently simple. Libri-
form fibers nonseptate, but starch present in them.
Mean libriform fiber diam., 25 um. Mean libri-
form fiber wall thickness, 1.5 um. Mean libriform
fiber length, 538 um. Libriform fiber walls some-
what gelatinous. Axial parenchyma vasicentric
scanty, often only one strand adjacent to a vessel.
Rays multiseriate and uniseriate, uniseriates more
abundant (Fig. 10). Ray cells mostly erect, a few
square cells present. Mean multiseriate ray height,
1,188 um. Mean uniseriate ray height, 270 um.
Mean multiseriate ray width at widest point, 2.2
cells. Cell walls moderately thin but lignified.
Wood nonstoried. Large rodlike crystals (one tip

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

concave, the other convex, suggesting paired
crystals) present in a few ray cells (Fig. 13).

CONCLUSIONS
HABIT AND ECOLOGY

In a given floristic area, wider vessels char-
acterize trees as compared with shrubs and sub-
shrubs (Carlquist & Hoekman, 1985). In a genus
or family that ranges into diverse habitats, trop-
ical tree species tend to have the widest vessels
(e.g., Vliet, 1979). Ludwigia anastomosans has
notably wide vessels for the family Onagraceae.
Mean vessel diameter in this species equals the
widest mean diameter reported in the earlier sur-
vey (Carlquist, 1975), a fact very likely related
both to its arboreal nature and tropical habitat.
The figure recorded for vessel diameter in L. an-
astomosans may be conservative, because the
EE studied is relatively small compared with

e large diameter of trunks of these trees. In
trees, vessel diameter is greater at the ica
of older stems (e.g., Carlquist, 1984

The abundance oferect ray cells in comparison
with procumbent ray cells in Ludwigia, even in
L. anastomosans, suggests that Ludwigia may
represent some degree of secondary woodiness.

his idea was entertained for Ludwigia earlier
(Carlquist, 1975). The presence of a few vestigial
bars on perforation plates in L. torulosa (Fig. 11)
is pertinent in this regard. Presence of a few such
plates does suggest retention of primitive fea-
tures in the primary xylem (the "refugium" the-
ory of Bailey, 1944), but it also suggests that this
feature is carried forward into the secondary xy-
lem by virtue of paedomorphosis. Paedomor-
phosis is indicated by erect ray cells and the rel-
atively long vessel elements (Carlquist, 1962),
two features well displayed in Ludwigia. Pres-
ence of occasional scalariform perforation plates

of the Asteraceae (Carlquist, 1983a), or Patrinia
of the Valerianaceae (Carlquist, 1983b). The
presence of scalariform perforation plates is,
however, more primitive than presence of simple
plates exclusively.

In an earlier paper (Carlquist, 1984a), vessel
grouping in taxa with libriform fibers (as in On-
agraceae) or fiber-tracheids was held to be pro-
portional to adaptation to dry conditions. This
hypothesis is worth testing in Onagraceae, be-

1987]

cause of the wide range of ecological circum-
stances occupied by its members. Data on vessel
grouping were prepared earlier for the family
(Carlquist, 1975), and indices designed to enable
comparison with ecological conditions were de-
vised (Carlquist, 1977). These data can now be
analyzed in the light of the vessel grouping hy-
pothesis. Vessel grouping figures were not offered
for six Ludwigia species studied earlier (Carl-
quist, 1982a), and so are presented here: L. bul-
lata rers! Hara, 1.08; L. elegans saad
Hara, 1.36; L. peruviana (L.) Hara, 1.10; L. s
icea rae Hara, 1.12; L. sonémtosà s
bess.) Hara, 1.04; L. sp. (aff. L. longifolia), 1.23.
The mean value for vessel grouping in the 12
collections of Ludwigia, sole genus of tribe Jus-
sieeae, is e next lowest value, by tribe,
occurs in Hauyeae (1.67), followed by Fuchsieae
(1.80) and Epilobieae (1.85); data for Lopezieae
and Onagreae were subdivided into habit cate-
gories. If one considers that the aquatic habitats
characteristic of Ludwigia species represent the
most mesic ecological situations of the family,
vessel grouping is a more accurate indicator of
ecology within Onagraceae than the indices

morphy figure for Hauyeae (1,242), which seems
high compared with that for 12 collections of
Ludwigia (415), is related to the wide vessels in
that tribe, which in turn is doubtless character-
istic for a species transpiring large volumes of
water in a warm, moist tropical forest. Both of
these figures are much higher than for groups in
the family occupying drier habitats, such as an-
nuals of tribe Onagreae (161) or caudex peren-
nials of Onagreae (48).

SYSTEMATICS

The data provided are consonant with the idea
that L. peduncularis and L. torulosa may be
closely related. These species have an unusual
feature in common, the presence of rays exclu-
sively or nearly aerial at the outset of sec

L. torulosa is not considered decisive. Such crys-
tals have been reported in L. bullata, L. octo-
valvis, and L. peruviana (Carlquist, 1975, 1982a).
In turn, these crystals resemble the large, sty-
loidlike crystals of Hauya; these latter crystals
occur in axial parenchyma rather than in rays,
as in Ludwigia. Data from wood anatomy should

CARLQUIST—WOOD ANATOMY OF LUDWIGIA

895

not be interpreted in too detailed a way here:
variability within species is not known and not
all species of all sections of Ludwigia have been
studied. Wood anatomy is not often a decisive
indicator of relationships at species and section
(subgenus) levels

e occurrence of interxylary phloem in L.
anastomosans is noteworthy. Interxylary phloem
has been reported previously for the genus only
in L. sericea (Carlquist, 1982a), although it oc-
curs elsewhere in the family (Carlquist, 1975).
The significance of interxylary phloem seems to
lie with seasonality in translocation of photo-
synthates, as noted earlier (Carlquist, 1975). Be-
cause L. anastomosans sa tree, presumably with
an annual flowering season, production of in-
terxylary phloem would be adaptive, whereas in
an annual or short-lived perennial Ludwigia plant,
the phloem from a single year, undiminished b
crushing at the end ofa season as in longer-lived
plants, would probably suffice to channel pho-
tosynthates into flowers and developing fruits.

LITERATURE CITED

BAILEY, I. W. 1944. The development of vessels in
angiosperms and its significance in morphological
research. Amer. J. Bot. 31: 421-428.

CARLQUIST, S. 1962. A theory of paedomorphosis in
dicotyledonous woods. Phytomorphology 12: 30-

1975. Wood anatomy of Onagraceae, with
notes on alternative modes of photosynthate
movement in dicotyledon woods. Ann. Missouri
Bot. Gard. 62: 386-424.

1977. Wood anatomy of Onagraceae: addi-
tional rd and concepts. Ann. Missouri Bot.
Gard. 64: 627-637.

. 1982a. Wood anatomy of Onagraceae: fur-
ther species; root anatomy; significance of ves-
tured pits and allied structures in b E ai
Ann. Missouri Bot. Gard. 69: 755-769.
2b. The use of eibvlenediantinb] in soft-
ening hard plant structures for paraffin sectioning.
Stain Techn. 57: 311-317.

83a. Observations on the vegetative anat-
omy of Crepidiastrum and Dendrocacalia (Aster-
aceae). Aliso 10: 383-395.

1983b. Wood anatomy of Calyceraceae and
Valerianaceae, with comments on aberrant per-

of dicotyledons. Aliso 10: 413-425. j
ssel grouping in dap adus ue wood:
s mperforate tra-

i anatomy of Tiimenitiosds Plant

Syst. Evol. 144:

& bg 1985. Ecological wood

anatomy of the s Aa southern Californian flora.
IAWA Bull. 6: 319-347

896 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

ELLMORE, G. S. 1981. Root dimorphism in Ludwigia "napua sere stem in = family ipi p arcis
eploides Onagraceae): structure and gas content S. str. ation to the evolution of life for
aa roots. Amer. J. Bot. 68: 557-568. Bot. Z dier ics te dean a 65: 627-639.
METCALFE, CR .& L. CHALK. 1950. Anatomy of the [In Peka
Dicotyledons. Clarendon Press, Oxford. VLIET, G. J. C. M. vAN. 1979. Wood anatomy of the
SHULKINA, T. V. & S. E. Zikov. 1980. The anatomical Combretaceae. Blumea 25: 141-223.

THE ORIGAMI OF BOTANY: A GUIDE TO COLLECTING
AND MOUNTING SPECIMENS OF CYCLANTHACEAE!

BARRY E. HAMMEL?

ABSTRACT

Guidelines for preparing and mounting specimens of Cyclanthaceae, as well as a key to the genera,

are presented in order to facilitate collecting of a difficult and

often neglected family. In general,

specimens should be prepared so the depth of the leaf division is apparent and the lower surface of

the basal part of the leaf is visible or accessible.

The family Cyclanthaceae has far fewer species
than the other related families oflarge intractable
monocots for which guides to the preparation of
herbarium specimens have recently been pub-
lished (Croat, 1985; Dransfield, 1986; Stone,
1983). Neverthel in much ofthe wet lowlands
of the Neotropics the cyclanths form a very con-
spicuous element of the understory and epiphyte
flora (Figs. 1, 2). Careful collecting in almost any
area of wet primary forest often yields new species
because the family is poorly represented in her-
baria and many species are narrowly endemic
This paper is a call for more collections as well
as a guide to help insure that they be properly
prepared.

SPECIMEN SELECTION AND PREPARATION

How to find them. Cyclanths are often left
uncollected due to their large size, epiphytic hab-
it, ephemeral and seasonal flowering, and green
fruits. Moreover, the fertile structures are born
low on the plant and are overtopped by obscuring
leaves (Fig. 3). However, even in a population
that is out of season, examination of numerous
plants will often reveal fertile structures. This
effort is important. Both staminate and mature
fruiting materials are essential for describing new
species and often for identifying known species.

Dramatic—though nearly microscopic — dif-
ferences in floral and fruit structure between veg-
etatively similar species are easily overlooked by
the nonspecialist collector who may tend to “see”
(and collect) only one common species of a genus
in an area where three or four occur. Different
species may occupy the same habitat on adjacent
ridges or along different branches of a single

stream. Differences in depth of lamina division,
presence/absence and position of lateral costae,
phyllotaxy, habit, and even subtle differences in
lamina and petiole texture and color help reveal
different taxa.

What to do to them. The artful folding (ori-
gami) of whole plants of large-leaved monocots
may in itself be reward enough for a specialist
in the particular family, but one can hardly ex-
pect specialists in other groups or even generalist
collectors to spend so much time on one gath-
ering. The final specimen is usually better for
data recovery when redundant material has been
removed and the essential properly folded. Thus,
the first need in collecting large-leaved monocots
is to know what parts ofthe plant to collect. Most
of the following comments pertain to vegetative
parts because most of the difficulties stem from

PES
=

em.

Since the leaves of cyclanths are bilaterally
symmetrical, they can be split down the middle
(Fig. 4). In general, the whole petiole and a picce
of stem with attached inflorescence or infructes-
cence should be included. Even when whole or
half leaves are collected, two sources of infor-
mation are often obscured or lost: the depth of
division of the leaf and an abaxial view of the
blade. It is not sufficient simply to press leaves
so that the division is visible, since it often splits
deeper on drying. On large leaves, which must
be split to fit the press, the point of division can
be indicated by cutting a notch at this point and
folding the leaf so that the notch shows (Fig. 5).
For leaves small enough to press entire, the best
way to avoid ambiguity is to cut off one leaf lobe
at the point of division (Fig. 6). The label should
briefly record this notching and lopping of lobes,

' Supported by so BSR-8508463 from the United States National Science Foundation. Many thanks also

to John Myers for
draft of the manu

ne drawings and to G. Wilder and an anonymous reviewer for helpful comments on a

? Missouri l. Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

ANN. Missouni Bor. GARD. 74: 897-902. 1987.

FIGURES |, 2.—

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

| p H / ba
h | "ç I 9
, M j

1. Dense understory of Asplundia uncinata Harl. in the wet lowlands of northeastern Costa

Rica. Plants ca. 1.5 m tall.—2. Evodianthus funifer (Poit.) Lindman, a very widespread root-climbing epiphyte.

Leaves ca. 50 cm long.
e.g., “notch marks point of division,” or “lobe
cut off at point of division.”

The presence/absence and position of lateral
costae is taxonomically meaningful at species and
higher levels. Although these costae are very con-
spicuous on the lower surface, they are not visible
at all from above. Thus, the blade should be
folded to show at least the basal portion of the
abaxial surface. This is true in all genera except
Carludovica, and a few species of Asplundia,

GURE 3. The

F most ice auris fertile mace:
when the staminodia are exserted, is ephemeral and
hidden among the leaves. Leaf blades ca. 65 cm long.

where features on the upper surface become im-
portant.

SPECIFIC (AND GENERIC) CONSIDERATIONS

The following key to the 10 genera of Cyclan-
thaceae is presented to facilitate discussion of
generic characters and the requirements for col-
lecting and mounting each genus. Pseudoludovia,
no longer accepted, is not included in the key,
nor are two new genera (R. Erikson, pers. comm.;
Hammel & Wilder, in prep.), which require no
special techniques.

ere 4. Asplundia aii n ready for press-
g.—A. Section to be discarded. — B. Notch to mar
Fn of division. — C. Lateral costae.

1987]

HAMMEL—COLLECTING AND MOUNTING CYCLANTHACEAE 899

KEY TO THE GENERA OF CYCLANTHACEAE WITH EMPHASIS ON VEGETATIVE CHARACTERS

la. Leaf blades with the 2 major lateral costae running nearly the entire length of the blade, usually deeply

=

bifid nearly to the base; spadix a cylinder of rings or TE or both formed by alternating staminate
and pistillate units; plants terrestrial, usually short-stemm Cyclanthus
Leaf blades with the 2 lateral costae (if present) always w era well below the tip of the blade,
entire to deeply divided; spadix a cylinder or sphere of tightly packed and variously connate to free,
oh more typical staminate and pistillate flowers; plants epiphytic, lithophytic, or terrestrial, often long-

em

. Petioles mostly 1.5-3 m ied - blades palmately divided into 4 segments, the segments deeply
toothed, the lateral costae and far removed from the margin; plants terrestrial; surface of
€ mature spadix Brrr i to reveal the pubes orange seed pulp and rachis; seeds

Carludovica

I Petioles mostly less than | m long; leaf € piem entire or bifid, the segments rarely toothed

but then the lateral costae long and running in t

N
c

argin; plants terrestrial, epiphytic, or litho-
ete.

phytic; surface of mature spadix not splitting irregularly, seeds flat or ter

3a. Leaves with 2

ta (oft Asplundia

rarely in Dicranopygium, always in the monotypic Schultesiophytum).

4a. e thes mostly dispersed along the peduncle; fruits dehiscent by apical caps, fused at the
bas Asplundia
4b. pem ena ne just below the spadix; fruits indehiscent.
bruising and drying black; fruits completely separate U. WADE
5b. eiie remaining green; fruits connate in basal !⁄ or more u... Dicranopygiu

w
q

. Leaves without conspicuous lateral costae.

6a. Pg clustered immediately below the sp
ds

nts lo

6b. Spathes stig rinse along the peduncle.

ichous.
9a.

ng-stemmed and openly eT usually climbing; stems and € leaves
rous; fruits completely separate; seeds fla
7b. Plants zd short-stemmed and
g on rocks along streams; stems and dry leaves smooth; fruits connate in basal
er

Evodianthus
clumped, rarely climbing a short distance, usually

Dicranopygium

Leaf ae crenate at most, neve

r bifid; fruiting spadix nearly smooth, the

pistillate flowers connate throughout; plants lianas or short-stemmed epiphytes

Ludovia

9b. Leaf blades of mature individuals bifid; fruiting spadix not smooth, the pistillate
flowers completely or partly free; plants terrestrial or epiphytic, usually short-

stemme

10a. Seeds mostly rounded at both ends
Seeds bile oe appendages at both ‘ends

8b. ed spirally arra

Sphaeradenia
Stelestylis

la. Petioles urs jie broadly channeled false petioles (sheaths) extending to the

TO, leaves of climbing stems a undivided; mature fruits tan; s
plants lianalike canopy clim

spathes 8-
Tho oracocarpus

11b. petioles present i in most species, the. false petioles narrowly channeled when

pre
debui 2-8; plants low trunk climbers or terrestrial

A J. Ay a Fis

p pyg t for over
half the species in the family, and it is within

the lower leaf surface applies especially to these
genera. For the small-leaved species of Asplundia
it is important to insure that more than one leaf
is preserved and that both surfaces of the basal
part of the leaf are visible. This is necessary in
order to see adaxial scars at the distal end of the
false petioles, characteristic of several species in
subgenus Chaonopsis. In Asplundia and Dicrano-
pygium, whether or not the sheath splits up into

OF pte
And

ly orange

fibers and the color and quality (dull vs. shiny)
of its surface are necessary data points for iden-
tifying or characterizing species. Here, specimens
should preserve a portion of the stem together
with the leaf at its point of attachment.
Although Sphaeradenia consistently has only
the median and no lateral costae, nothing is lost
by preparing specimens of this genus (and of most
others) as for Asplundia and Dicranopygium. In
any case, for Ludovia, Sphaeradenia, and Ste-
lestylis one should preserve part of the stem with
attached sheaths in order to verify the distichous
arrangement of the leaves and to show internode
length (Fig. 6). Asplundia, Dicranopygium, and

900 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

FiGuRE 7. Carludovica specimen ready for press-
e 2 sections to be discarded cut at point of
div

Sphaeradenia account for most of the species in
the family so that the general rules for collecting
these genera apply to most of the species one
encounters.

Carludovica needs special attention in con-
nection with the very large size and unusual shape

FIGURE 5. Asplundia specimen properly folded and of its mature leaves. In order to represent the
mounted with string. essential features (the depth of the teeth and the

FIGURE 6. Sphaeradenia specimen showing blade FIGURE 8. Carludovica specimen properly folded
cut off at point of division (A) and distichous phyllo- and i ga with string, showing hastulae (A) on up-
taxy (B). per surfac

1987]

depth of the divisions) only one of the four leaf
segments need be preserved. The best approach
is to cut off the two lateral segments and one of
the central segments at the point of division and
then fold the blade so that the teeth and the
adaxial surface of the basal portion of the leaf
are visible (Figs. 7, 8), thus showing the presence/
absence of hastulae (Wilder, 1976). Sometimes
even this reduced leaf must be split to fit on one
sheet. Variation in petiole and peduncle length
appears to be of little use for et arn species
of Carludovica, but the length of these structures
should at least be recorded in the field notes.
Finally, for Carludovica, variation between
species in shape of juvenile and immature foliage
makes leaves from young plants worth preserv-
ing.

FIELD NOTES

abit. Approximately 60% of the species of
Cyclanthaceae are either strict epiphytes or root-
climbers, which may eventually lose connection
with the ground. Habit is quite diverse (and
sometimes variable within species) in Asplundia
and Dicranopygium. The longest-stemmed
species are usually found only as climbers, some
with shorter stems climb or grow free-standing,
and the shortest-stemmed species never climb.
Strict epiphytes occur only in Ludovia, Sphaera-
denia, and Stelestylis, and terrestrial species oc-
cur in all of these genera. Carludovica is always
terrestrial; Cyclanthus rarely climbs. Only species
of Dicranopygium grow on rocks along streams.
For these reasons one should always make care-
ful note of the habit: epiphytic, lithophytic, ter-
restrial, or root-climbing. If the collector 1s cer-
tain that a species is both terrestrial and climbing,
it is very useful to write that in the notes, e.g.,
“apparently same species also climber here but
not fertile." If the stem is not preserved, the leaf
arrangement should be noted. The approximate
length of the stem or height above ground for
climbers (e.g., “climbing to 10 m") should also
be recorded.

Other d rini characters. Even though care
has been taken to record on the specimen the
depth of the us du it is also a good idea
to record in the field notes the approximate range
of this cl based on the plant or population
collected, e.g., “leaves divided from !^—/." The
leaf segments of all species of Carludovica and a
few species of Asplundia develop teeth in the
unopened leaf. Unfortunately, with age, drying,

HAMMEL- COLLECTING AND MOUNTING CYCLANTHACEAE

901

or other trauma some large Asplundia leaves can
split into teeth, which on herbarium sheets re-
semble those arising developmentally. Thus, for

Asplundia one should indicate whether such teeth

originate developmentally or by trauma.

As noted above, subtle differences in leaf and
petiole surface textures and colors often separate
species and should be recorded. In a few species
of Asplundia and Dicranopygium the leaf folds
are not adaxially keeled. Because all leaves on
herbarium specimens appear to have variously
keeled folds, it is best to record exceptions in the
field notes.

The cross-sectional petiole shape is less vari-
able in the Cyclanthaceae than in certain genera
of Araceae (Croat, 1985), but again this feature
is difficult to assess using dried herbarium ma-
terial and should be described for unusual cases.
Most petioles are basically terete and barely to
obviously flattened above (D-shaped) and may
have a small median adaxial groove. In Thora-
cocarpus and a few species of Asplundia the sheath
reaches all the way to the base of the blade so
the leaves lack a petiole. The petioles of ScAul-
tesiophytum are asymmetrically D-shaped with
one margin rounded and the other sharply an-
gled. The color of the stem cross section may
also be taxonomically significant; in certain
species of Asplundia and Dicranopygium cut
stems rapidly turn reddish brown om in-
dicating high concentrations of tan

Fertile structures. Rui n of cy-
clanths are monoecious and protogynous. Their
most conspicuous stage— when the staminodia
are exserted—is also the most ephemeral. Vary-
ing in size and color, the vermicellilike stami-
nodia may harbor taxonomic structure but have
not been used because they are so seldom seen
and poorly did on drying. When an inflo-
rescence with dia is collected, field
notes should incline color and approximate
length of staminodia. The number of spathes
serves to distinguish taxa but is often difficult to
count on herbarium material without damaging
it. Field notes should mention the number, color,
and texture of spathes. Most of these time-con-
suming measurements can be delayed (and many
more characters accurately preserved) by pre
serving inflorescences in liquid. Dried pu
of these extremely succulent parts so misrepre-
sents the living condition that, except for well-
known species, collectors are strongly urged to
preserve inflorescences or parts thereof in 70%
Et

902

ae ome | A ak 1d

The color of
be noted. They may be bite: red, yellow, or
green at maturity. In Sphaeradenia the color of
mature seeds is also variable and should be re-
corded.

MOUNTING THE SPECIMENS

All of the foregoing discussion, especially on
artful folding, is to no avail if the dried material
is mounted upside down. Collectors always as-
sume that their material will be mounted the way
they pressed it, with the numbered side of the
newsprint equal to the up side of the specimen.
However, once the connection between news-
paper and plant is broken, as it can easily be with
so many steps between collecting and mounting,
and as it always is in the end, the decision must
be made: which side is up? Responsibility for
this final important step rests upon the mounter.

In the end, a liberal use of string and judicious
use of glue can obviate even this decision. Gen-
erally, one need glue down only the first layer of
leaf fold and then tie the stem and petiole(s) at
appropriate places (see Figs. 5, 8). In this way

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

the specimen can be unfolded partly or com-
pletely, and both sides of the leaf can be easily
examined

Mounters, of course, will continue to receive
material gathered by those who have missed, ig-
nored, or forgotten this guide, or who are not
artists. If parts are missing, or no mark records
the point of division, nothing can be done. But
when the leaves have been folded improperly,
this guide is especially for the mounters, who
with a little care can salvage the specimen and
enjoy the art.

LITERATURE CITED
CROAT, T. B. 198
mens of Araceae.
252-258.
DRANSFIELD, J. 1986

5. Collecting and preparing dien
Ann. Missouri Bot. Gard.

A guide to collecting palms.
: 166-176.

A guide to collecting Pandanaceae
(Pandanus, Freycinetia, and Sararanga). Ann.
Missouri Bot. Gard. 70: 137-1

WILDER, he ir 1976. Structure and development of
leav 7
reference to other Cyclanthaceae and Palmae.
Amer. J. Bot. 63: 1237-1256.

MESOAMERICAN SISYRINCHIUM (IRIDACEAE):
NEW SPECIES AND RECORDS, D
NOTES ON TYPIFICATION'

JAMES E. HENRICH? AND PETER GOLDBLATT?

ABSTRACT

This paper supplements our treatment of Sisyrinchium for Flora Mesoamericana in which 13 species

are recognized. One new species is described, S. su
and Costa Rica. Sisyrinchium trinerve, previously known

from T

alpinum, from Mexico (only Chiapas), Guatemala,
from Andean South America, S. dimorphum

Mexico, S. longispathum from Oaxaca, Mexico, and S. subcernuum from

northern Mexico are reported in Mesoamerica for the first time. The type material for S. convolutum,

S. tenuifolium, S.
designated for each species.

In preparation for a treatment of Iridaceae for
Flora Mesoamericana (Henrich & Goldblatt,
in press), it has become clear that previous re-
gional treatments of Sisyrinchium are to varying
degrees inadequate or erroneous. The same
species are treated under different names in some
floras and few of the species have been matched
with their types. Notes are thus presented here
on the typification of several widely recognized
species to supplement the formal treatment. There
is also apparently one new species in Mesoamer-
ica, S. subalpinum, a yellow-flowered species
of section Echthronema Bentham. This is a dwarf,
high-altitude relative of the common S. tincto-
rium. We also report for the first time the oc-

TABLE 1.

mandonii, S. tinctorium, and S. bogotense has been examined and a lectotype

currence in Mesoamerica of the Andean S. tri-
nerve, the northern Mexican S. subcernuum, the
southern Mexican S. /ongispathum, and the pre-
dominantly Texan S. dimorphum. Finally, the
species previously identified as S. iridifolium
Kunth in Guatemala and Honduras corresponds
closely to the Oaxacan S. exalatum, a tall,
branched species with narrow leaves and thick-
ened roots sometimes swollen terminally.

We recognize 13 species (Table 1) of Sisyrin-
chium in Mesoamerica, which is regarded for the
Flora as including all the territory from the
southern Mexican states of Yucatán, Campeche,
Tabasco, Quintana Roo, and Chiapas south ward
to Panama. The species are numbered as they

The species of Sisyrinchium recorded from Mesoamerica with their general distributions.

S. chiricanum Woodson

S. convolutum Nocca

Mexico Pisas Guatemala, El Salvador, Costa Rica,

Panam
Chiapas, Mexico to Panama, possibly the north coast of

South Ameri

k p a kiya

dimorphum R. Oliver
exalatum Robinson & Greenman

. Johnstonii Standley

longispathum Conzatti
mandonii und

micranthum

subalpinum See & Goldbl.

subcernuum Pon Henrich & Goldbl.
mb. &

tenuifolium H
tinctorium on
trinerve Baker

Bonpl. ex Willd.

U.S.A. a Sy T" Guatemala
southern m (Oaxaca, Chiapas), Guatemala,
Hondur:

Mexico POM Guatemala, Costa Rica
Mexico (Oaxaca, Chiapas)

Guatemala, Costa Rica, Panama, Peru, Bolivia
Mexico to South America, West Indies

Mexico (Chiapas), Guatemala, Costa Rica

Mexico, Belize

Mexico, Guatemala, Panam

southern Mexico to ind and Venezuela

Costa Rica, Colombia, Venezuela, Ecuador to Bolivia

iS

! Support for this research from U.S. National Science Foundation grants DEB 81-19292 and BSR 85-00148

quad acknowledge
ouri Botanical Garden
? B. AK

, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.
Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri
63166, U.S.A.

ANN. Missouni Bor. GARD. 74: 903-910. 1987.

904

appear in the Flora treatment. Species 1—3 belong
to section Sisyrinchium (= section Bermudiana

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[VoL. 74

Bentham) and 4-12 to section Echthronema
(Bentham, 1883).

KEY TO SISYRINCHIUM IN MESOAMERICA

la. Flowering stems unbranched, consisting of 1 long internode without cauline leaves, but sometimes
with a terminal bract subtending the opposed spathes of the 1-3 apical inflorescence units (rhipidia)
Apos rhipidia with a short stalk, more often sessile).

s filiform wide; inflorescence units 1—3
T nie a to lanceolate, at least 1.5 m
tuberous, fleshy (Fig. 1A); flowers blue to w

.. 8. S. longispathum

mm wide; Aue ha units ] or 2.

cR 3. S. johnstonii

3b. Root ots pois rous OT nerd. but not tuberous (Fig. 2B. "DX flowers yellow.
4a. Flowering stems subterete and not winged; ovary included in the upper part of the spathes
7.

S. trinerve

4b. Flowering stems flattened and winged; ovary exse
5a. Spathes (20-)25-50 mm long, the outer euh EE twice) than the inner

11. S. tinctorium

5b. Spathes 11-16 mm long, usually subequal.
6a. Plants 5-12 cm tall; roots slender but somewhat thickened (Fig. 3) ..

12. E subalpinum

6b. Plants 15-25 cm tall; roots fibrous (Fig. 2D)
. Flowering stems EA of more than | internode, branched or at least with | or more cauline
le

c

eav
7a. Roots fibrous or thickened and cylindric, sometimes with persistent hairs (Fig. 1B, 2.
Is 5-15(-25) cm tall; capsules 3 mm long, globose

8a. Annua

S. subcernuum

9: eig dé

8b. Perennials (15-)20-60 cm tall; capsules 4-5 mm long and globose, or 8-11 mm long a
ob

ovoid.

9a. Flowers blue; spathes 14-18 mm long .....

9b. Flowers yellow, spathes 26-32 m

. S. dimorphum

Oa. Roots with persistent hairs throughout or at least near the base (Fig. 1B)
5.

10b. Roots smooth, not bearing persistent hairs 1C)
7b. Some roots tuberous and fleshy, either close to o

lla. Basal leaves 3-8 mm

^u chiricanum
OCURRE: S. exalatum
m the p i (Fig. Y C).

falca

wide, narrowly to broadly eerie to

2a. Plants usually less than 30 cm high and with falcate leaves; ue strongly 3-lobed
6.

sually with swollen fusiform us thickened from the base (Fig. 2A
ore m high and with lanceolate elio capsules hae
d; roots ceri if at "all. distant from the base (Fi js
wide, linear; capsules not strongly 3- ne
De 5-7 mm long ....
13b. Capsules oblong, 12-16 mm long .

lobe
llb. Basal Testa 0.5-2 m
13a. Capsules glo

NOTES ON SELECTED SPECIES

1. Sisyrinchium dimorphum R. Oliver, Ann.
Missouri Bot. Gard. 55: 397. 1969. TYPE:
United States. Texas: Val Verde Co., San
Felipe Springs, Del Rio, Warnock & Cam-
eron 9894 (holotype, SMU).

A species of Sisyrinchium from Chiapas and
Guatemala sometimes identified as S. scabrum

ate size with relatively short leaves 1-2 mm wide,
a branched stem, and persistent fibrous leaf bas-
es, distinctive in Sisyrinchium. It clearly belongs
in section Sisyrinchium and has the blue (to white)
campanulate perianth and connate (or nearly
connate) filaments that characterize the group.
The Mexican S. scabrum, typified by Schiede

S. patinen

exalatum

—-— . 9. S. tenuifolium
10. S. mandonii

1020 (HAL), is similar in general morphology
yet it is smaller in stature, has a smaller flower
with an unusually small, sparsely pubescent ovary
less than 1 mm long, and globose capsules ca. 2
mm in diameter, whereas S. dimorphum has a
glabrous ovary ca. 2 mm long and larger, some-
what obovoid capsules 4-7 mm long.

4. Sisyrinchium exalatum Robinson & Green-
man, Amer. J. Sci. 50: 166. 1895. TYPE:
Mexico. Oaxaca: Cuilapan Mountains, 7,000
ft., Smith 52 (not seen). Figure 1C

When reported from Guatemala and Hondu-
ras (Standley & Steyermark, 1952; Molina, 1975),
Sisyrinchium exalatum was treated as S. iridi-
folium Kunth. We concur in regarding it as dif-
ferent from all other Mesoamerican species but
it does not correspond with S. iridifolium. This

1987]

FIGURE 1.

Root systems n stem bases.
Scale bars = 2 cm; top bar fo B.

is a South American species, the type from Car-
acas, Venezuela, which is similar to S. exalatum
in leaf and stem morphology, but the roots are
fibrous and not thickened as in the latter and the
capsules are relatively small and nearly globose,
rather than obovoid-ellipsoid and somewhat
trigonous.

In Mesoamerica Sisyrinchium exalatum oc-
curs in Mexico (Chiapas), Guatemala, El Sal-
vador, and Honduras. It grows on grassy to light-
ly wooded slopes at elevations of 1,700-3,700
m

Additional specimens examined. XICO. CHIAPA
north end of San Cristóbal Las ae sap wa 6045
(F). EL SALVADOR. MORAZAN: Slopes of La Montañita,

HENRICH & GOLDBLATT—MESOAMERICAN SISYRINCHIUM

905

—A. Sisyrinchium johnstonii. — B. S. chiricanum. —C. S. exalatum.

1,700 m, Williams & Molina R. 10429 (F, MEXU,
ICH, MO). aar EMALA. HUEHUETENANGO: between

Tojqu iá and Caxín bluff, Steyermark 5014 1 (F); Sierra

Cuchumatanes, Sur 1243 (F)

anto Tomás,

COMAYAGUA: abierto de La Piramide, Molina R. 14357

F).

6. Sisyrinchium convolutum Nocca, Ticin, Hort.

l . TYPE: Guiana “Cap. B. Spei,"

cultivated in “Hort. Parolin,” collector un-

known (lectotype, BASSA, here designated).
Figure 2A

Sisyrinchium vg crois (Baker) Standley & Stey-

eld Mus. Nat. Hist., Bot. Ser. 23:

39. 1944: s of Guatemala, Fieldiana, Bot. 24:

906

73. 1952. S. alatum var. guatemalense Baker,

andbk. Irideae 180. 1892. TYPE: none cited nor

locaied but authentic material at K, annotated by
Baker

Sisyrinchium convolutum is one of the more
common Mesoamerican species of Sisyrinchium.
It can generally be recognized by its moderate
height, more or less falcate leaves, winged and
branched stems, and large, inflated, trigonous
capsules 8-10 mm in diameter. The capsules are
glabrous, in contrast to the related and some-
times similar S. tenuifolium, in which they are
sparsely pubescent. The flowers are yellow and
relatively large with tepals to 15 mm long, and
the spathes of the inflorescence are distinctively
broad, giving it a somewhat inflated appearance.
The roots are fleshy and tuberous (Fig. 2A

Typification of Sisyrinchium convolutum has
posed some difficulty. The illustration in the pro-
tologue matches the species to which the name
has been applied here and elsewhere, except for
the swollen, almost bulbous base and the roots
figured as forming a fibrous mass. A specimen
in the herbarium at Bassano del Grappa, Vicen-
za, where Nocca’s types are believed to be housed,
matches closely the illustration but lacks a base
and roots. Given the stylized depiction of these
organs, it seems likely that the artist drew them
from imagination. We have chosen the specimen
rather than the illustration as lectotype.

Sisyrinchium convolutum was confused with
S. alatum, a strictly South American species, by
Baker (1892) amongst others. Baker used the
name S. alatum var. guatemalense for the plant,
as did Standley & Steyermark (1952) in the Flora
of Guatemala and Molina (1975) in an enumer-
ation of the plants of Honduras. Sisyrinchium
alatum is probably not closely related to S. con-
volutum and does not have fleshy tuberous roots
as do the latter and related species.

7. Sisyrinchium trinerve Baker, J. Bot. 14: 267.
1876. Sisyrinchium bakeri Klatt, Abh. Na-
turf. Ges. Halle 15: 378. 1882, nom. superf.
pro S. trinerve. TYPE: Bolivia. La Paz: vicin-
ity of Sorata, Mandon 1218 (lectotype, K,
here designated); syntypes: Mandon 1220
(K); Pearce 87 (K). Figure 2B

Known until recently only from the high An-
des of Bolivia, Peru, Ecuador, Colombia, and
Venezuela, Sisyrinchium trinerve has now been
found in the páramo of Costa Rica above 3,000
m. It is readily recognized by its long, un-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

branched, nearly terete stem; rigid, narrow, pre-
dominantly three-veined leaves; yellow perianth;
and ovary at least partly included in the spathes.
A lectotype has been designated, since the pro-
tologue includes three collections, none desig-
nated as a type (Baker, 1876), and no herbarium
is mentioned, although the Kew Herbarium is
assumed to house the specimens seen by Baker.
The collection Mandon 1218 was chosen, since
it comprises the best-preserved and most com-
plete material.

Additional specimens examined. CosTA RICA.
LIMON: Cordillera de Talamanca, Kamuk massif, 3,000-
3,300 m, a & Herrera Ch. 29375 (MO). CAR-

AGO-SAN JOSE: Cerro de la Muerte, paramo, along
stream, Heithaus 239 (MO).

8. Sisyrinchium longispathum Conzatti, Flora
Taxonomica Mexicana 2, 2: 124. 1947. TYPE:
Mexico. Oaxaca: Conzatti 4203 (not seen).

Known from only a handful of collections, all
from mountainous areas of the southern Mexi-
can state of Oaxaca, Sisyrinchium longispathum
has been recorded from a single site in Chiapas,
Breedlove 27156, 22 km north of Tuxtla Gut-
tierrez. The species appears to be restricted to
limestone outcrops and cliffs. Sisyrinchium lon-
gispathum is distinctive among the Mesoamer-
ican species of Sisyrinchium in having filiform
leaves and 2-3 inflorescence units per flowering
stem, all crowded at the apex and each subtended
by a short leaf. The yellow flowers are typical of
section Echthronema. Sisyrinchium longispa-
thum has no close relatives among the Mesoam-
erican species of Sisyrinchium.

9. Sisyrinchium tenuifolium Humb. & Bonpl. ex
Willd., Enum. Pl. Hort. Berol. 2: 691. 1809,
Hort. Berol. 2: 691. 1809. TYPE: Mexico.
Without precise locality, Humboldt & Bon-
pland s.n. (lectotype here designated, B—
Herb. Willdenow — only microfiche seen).

Sisyrinchium tenuifolium is one of the smaller
species of the genus and can usually be recog-
nized by its branched stems, narrow basal leaves
0.5-2 mm wide, and swollen tuberous roots. The
bright yellow flowers are more or less typical for
section Echthronema. It is particularly variable,
with plants ranging from slender and unbranched
with very narrow leaves to robust with broad
leaves and many-branched stems. The latter
forms resemble S. convolutum fairly closely and

1987]

FiGurE 2. Root systems and pa bases.
D. S. per ee Scale bar = 2c

can be distinguished by their minutely pubescent
ovary and smaller capsule.

The specimen in the Willdenow Herbarium at
Berlin is chosen as the lectotype. Other Hum-
boldt and Bonpland specimens of this species
may be duplicates of the type collection and thus
isolectotyes, as is the illustration in the Hortus
Berolinensis.

10. Sisyrinchium mandonii Baker, J. Bot. 14: 269.
1876. TYPE: Bolivia. La Paz: vicinity of Sora-
ta, Rancha de Cochipata, Mandon 1217 (lec-
totype, K, here designated; isolectotypes, B,
not seen, MO photo); syntypes: New Gran-

HENRICH & GOLDBLATT —MESOAMERICAN S/SYRINCHIUM

—A. Sisyrinchium convolutum. —B. S. trinerve. — C. S. mandonii. —

ada (Brazil), Jurgensen id o New Gran-
ada, Purdie s.n. (K). Fig

Sisyrinchium mandonii has distinctive small,
swollen, tuberous roots (Fig. 2C); branched stems;
relatively narrow leaves 1-2 mm wide; and un-
usual large, ellipsoid capsules. It is predomi-
nantly Andean, represented by a few collections
from Mesoamerica (Steyermark 30513, Guate-
mala; Allen 698, Costa Rica; Woodson & Schery
427, Panama), all from relatively high altitudes.
It was first reported for Mesoamerica by Wood-
son (1945) from Panama.

Of the three collections cited by Baker in the
protologue, Mandon 1217 is selected as the lec-

908

totype as it is in good condition and Baker an-
notated it as the "Type." There is no indication
whether Baker saw any duplicates of this collec-
tion but this 1s certainly possible. The two other
collections cited, both at Kew, appear to be the
same species, although they are from New Gra-
nada in eastern Brazil where Sisyrinchium man-
donii does not grow. Mislabeling seems likely
here.

11. Sisyrinchium tinctorium Kunth, Nov. Gen.
Sp. 1: 324. 1815. TYPE: Venezuela. T.F.
Amazonas: banks of the Orinoco near Es-
meralda and the confluence of the Sodo-
monis, Humboldt & Bonpland s.n. (holo-
type,

su Lag id Kunth, Nov. Gen. Sp. 1: 323.

. TYPE: Colombia. Cundinamarca: Bogotá,

En. Sins between Suba and Suacha, Humboldi

& Bonpland s.n. (holotype, P).

—

Sisyrinchium tinctorium is a common and well-
known species in Mesoamerica. It also occurs in
northern South America but is apparently less
common there. It is distinguished by unbranched
and often broadly winged flowering stems with
terminal spathes, pale yellow flowers, and pen-
dent, more or less pyriform capsules. Specimens
in fruit are collected most often. The leaves have
a characteristic thin, almost membranous tex-
ture when dry, and the roots are fibrous. Me-
soamerican collections match the type of a sec-
ond species, S. bogotense, particularly well. Our
examination of the types of this and the Vene-
zuelan S. tinctorium showed the two to be con-
specific. The type specimens of S. tinctorium have
comparatively narrowly winged stems and cap-
sules smaller than most collections ofthe species.
However, a few similar specimens have been col-
lected in Mesoamerica, and variation is contin-
uous from typical S. tinctorium to typical S. bo-
gotense.

A diminutive species clearly allied to Sisyrin-
chium tinctorium is recognized here as S. sub-
alpinum, which is shorter and has narrower leaves
and smaller capsules than those of S. tinctorium.

12. Sisyrinchium subalpinum Henrich & Gold-

blatt, sp. nov. TYPE: Costa Rica. Limón: Cor-
dillera de Talamanca, Atlantic slope, Ká-

,30
d Herrera Ch. 29294 (holotype, MO). Fig-
e 3.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74
Plantae 5- 12 cm altae, radicibus aliquantum in-

m longis iai pr filamentis 1.5
mm longi infra liberis antheris ca. 2 mm longis, ramis
styli ca. 1.5 mm longis, capsulis nutantibus glabris.

Plants 5-12 cm tall; the roots slender, some-
what thickened but cylindric, not fleshy or tu-
berous. Leaves several, all basal, '^ to as long as
the flowering stems, 1-2 mm wide. Flowering
stems less than | mm wide, unbranched, com-
prising | internode, flattened but barely winged;
spathes 11-16 mm, subequal or the outer to !^
longer than the inner. Flowers small, yellow, stel-
late; tepals 4-5 mm long, subequal; filaments 1.5
mm long, free at least above (not seen below);
anthers ca. 2 mm long. Ovary about 3 mm long;
style branches ca. 1.5 mm long, slender, extend-
Ing between the stamens. Capsules nodding, ma-

t seen, these probably somewhat pyr-
ton and glabrous; seeds unknown.

Sisyrinchium subalpinum has been recorded
at high elevations in Mexico (Chiapas) and in
Guatemala and Costa Rica. It 1s restricted to
paramos and moist, high, pine forests.

Additional spiel examined. MEXICO. CHIAPAS:

. Male near Porvenir, 3,200 m, Matuda 4629 (MO).

GUATEMALA. SAN MARCOS: along quebrada Canjula be-

tween Sibinal and Canjula, Volcán oq 2,200-

2,500, Steyermark 35970 (F). Costa RICA. :

within 200 m of the summit of Cerro Chirriposillo.
3,400 m, Weston 1574 (MO

13. Sisyrinchium M (Bickn.) Henrich
& Goldblatt, comb. nov. Hydastylis sub-
cernuus 2 Bull. Torrey Bot. Club 27:
385. 1900. TYPE: Mexico. Baja California
Sur: Sierra i Laguna, Brandegee s.n. (ho-
lotype, CAS). Figure 2D.

Sisyrinchium subcernuum belongs to what may
be called the S. tinctorium complex, the species
of which share similarly textured leaves, un-
branched and winged flowering stems, and
drooping capsules. The type is from Baja Cali-
fornia Sur, and several more collections are
known from higher elevations in Mexico. A few
ofthe collections from Belize match the northern
Mexico plants closely and must be treated as the

tepals about 6 mm long, and small capsules 6
mm long and 3 mm wide. The outer spathe of

1987]

FIGURE 3.

HENRICH & GOLDBLATT — MESOAMERICAN SISYRINCHIUM

Type specimen of Sisyrinchium subalpinum (scale

909

MISSOURI
BOTANICAL GARDEN
HERBARIUM

N° 3212055

olo! o IS rin ehiu. ubo pin |
— r Sisyrinehui gu end Goldblatt

Ref

Sisyrinchium subalpinum Henrich ¢ Goldblatt
qc ang by jJ. E. Henrich & P. Goldblatt 1986

un Botanical Garden

COSTA RICA
IRIDACEAE
Sisyrinchyum
LIMON: Cordillera de Talamanca, Atlanti
slope nen uk massifs í paramo north
east the main Kamuk peak; elev
3000- $300 m; 9°16'-9°17'Ne
83*00'-03*02'W.
Blechnum-shrub association in Chusquea-
Hyper icum pårano.
Flowers yellow
aoe 19 Ser 1984
G. Davidse & G. Herrera 4
(MO)

MISSOURI BOTANICAL. GARDEN HERBAR TUM

— 5 cm).

910

the inflorescence is longer than the inner, some-
times as much as twice as long.

Additional specimens eS BELIZE. CAYO: near
Millionario, , roadside, Gentry 7694 (MO);
between Millionario and ren Dwyer 10821 (MO);
vicinity of Cuevas, south of Millionario, ca. 1,900 m,
Croat 23602 (MO

LITERATURE CITED

BAKER, J. G. 1876. New Aristeae and Sisyrinchia. J.
B 69.

. Handbook of the Irideae. George Bell
Sons, London.

BENTHAM, G. 1883. Irideae. /n G. red & J. D.
ooker, Genera Plantarum 3: 681-710.
Bei E. P. 1900. Studies in ele hium VIII.
Sisyrinchium californicum and related species of
the neglected genus H ydast ylus. Bull. Torrey Bot.

Club 27: 373-387.

HENRICH, J. E. & P. GOLDBLATT. Iridaceae. Jn G. Da-
lees et al. (editors), Flora Mesoamericana, Vol-

me 1 (in press).
ia F.W. 1882. Ergänzungen und Berichtigungen

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

zu Baker's Systema Iridearum. Abh. Naturf. Ges.
Halle 15: 337-404.

KuNTH, C. S. 1815. Voyage de Humboldt et Bon-
pland. Sixiéme Partie. Botanique. Nova Genera et
Species Plantarum (= Novae Genera et Species
Plantarum quas.
Latine-Allemande, Paris. Gide Fils,

MOLINA, R. A. 75. Enumeración de las sies de
Honduras. Ceiba 19: 1-

NOCCA, 1800. Ticinensis Horti Academici Plantae

T I

. Sisyrinchium dimorphum pis

rege a new species from Texas and Mexi
. Missouri Bot. Gard. 55: 397.

eon B. L. & J. M. GREENMAN. 1985. New and
noteworthy plants chiefly from Oaxaca collected
by Messrs. C. G. Pringle, L. C. Smith and E. W
Nelson. Amer. J. Sci. 50(296): 150-168.

STANDLEY, P. C. & J. A. STEYERMARK. 1944. Studies
of Central png iet plants. M Publ. Field Mus.
Nat. Hist., Bot. Ser. 23: 31-109.

1952. aoe In Flora of Gua-
temala. _Fieldiana, Bot. 24: 159-178.

WOODSON 45. Iridaceae. /n Flora of Panama.
Ann. Ve Bot. Gard. 32: 34-42.

NOTES

THE MESOAMERICAN NEOMARICA (IRIDACEAE),
N. VARIEGATA HENRICH & GOLDBLATT, COMB. NOV.

The genus Neomarica (Sprague, 1928) com-
prises some 15 species of evergreen herbs of trop-
ical forests and woodlands. Ravenna (1976) ap-
parently merged it with Trimezia Herbert, a
treatment we do not accept. The genus is dis-
tinctive in having a broadly winged flowering
stem, the branches ofthe inflorescence often sub-
sessile and crowded at the stem apices, and a
single large cauline leaf subtending and extending
well beyond the inflorescence. Neomarica is cen-
tered in eastern Brazil but occurs widely in South
America, with a single species extending through
Mesoamerica into southern Mexico. In several
regional floristic treatments for countries in Me-
soamerica the native Neomarica has been treated
as N. gracilis (Herbert) Sprague. The type of N.
gracilis, a figure in Curtis' Botanical Magazine
(66: tab. 3713. 1839), is believed to have been
found in Brazil and it matches an eastern Bra-
zilian species. It does not correspond with the
Mesoamerican species in morphology of inflo-
rescence, flower, or seed.

The single Mesoamerican species of Neomar-

ica was validly described in 1843 by Martens & ,

Galeotti as Marica variegata, based on Galeotti
5370, collected in Veracruz, Mexico. The type
at the Herbier du Jardin Botanique National de
Belgique (BR) matches other collections of Neo-
marica from Mesoamerica in their subsessile in-

ANN. MissouRi Bor. GARD. 74: 911. 1987.

florescence spathes and flower morphology. Var-
iegata 1s the only epithet available for the
Mesoamerican Neomarica and the combination
is made below.

Neomarica variegata (Martens & Galeotti) Hen-
rich & Goldblatt, comb. nov. Marica var-
iegata Martens & Galeotti, Bull. Acad. Roy.
Sci. Belgique 10: 112. 1843. TYPE: Mexico.
Veracruz: Zacuapan, Galeotti 5370 (BR, ho-
lotype).

LITERATURE CITED
Martens, M. & H. G. GALEoTTI. 1843. Enumeratio
synoptica plantarum phanerogamicarum ab H
Galeotti in regionibus Mexicanis collectarum. Bull.
cad. Roy. Sci. Belgique 10: 110-134.
RAVENNA, P. 76. Neotropical species threatened
and endangered by human activity in the Irida-

Extinction is Forever. New York Botanical Gar-

, New York.

SPRAGUE, T. A. 1928. Marica and Neomarica. Kew
Bull. 1928: 278-281.

—James E. Henrich and Peter Goldblatt, B. A.
Krukoff Curator of African Botany, both at Mis-
souri Botanical Garden, St. Louis, Missouri
63166, U.S.A

TWO NEW XYRIS (XYRIDACEAE) FROM THE
AMAZON BASIN OF BRAZIL

Among undetermined Xyridaceae sent to the
senior author for definitive treatment during
1980-1982 are two from northern Brazil that we
agree are new species to be added to the flora of

Xyris pectinata Kral, Smith & Wanderley, sp.
nov. TYPE: Brazil. Amazonas: Estrada
Transamazónica, campina aberta, terreno
arenoso, Proj. RADAM, | June 1976, T.
Bahia 35 (holotype, INPA; isotypes, US,
VDB). Figure 1.

erba perennis densicaespitosa, tenella, glabra; ra-
dices graciles. Folia line nearia, 4-6 cm longa, erecta vel
sc lora inae

satis, lateribus valde longitudine ideo n
vel Intense ferrugineae, me ibus in lamai na ne
ada

longam fascientes, infime as dilatatae. ane

retes, filiformes, r ter torti,

i 5 mm crassi, olivacei, distaliter iu

bicostati, costis laevibus. ipie subglobosae is late

obovoideae, 4.5-6 mm longae, e, pluriflorae, d at-
n

tae, la t atae, suborbiculatae, aut reni-
formes ae, convexae et leviter carinatae,
obtusae vel subt fir ad apicem utrinque erosae
et ciliatae, scariosae, minute tuberculato-rugulosae, a
medio ad basim lto crassiores, nitidae, brunneolae,
marginibus effuse et pectinate rigidofimbriatis; arca
dorsalis ovata, ca. 2-2. , glau la la-

teralia ca. ^ connata, ca. 2
lobis acutis scariosis, ala carinali angusta, integra
Laminae petalorum bov , Ca. mm lon-
gae, ad apicem rotundatae, laceratae, luteolae. Stami-

m longae, loculis parallelis dis-
tinctis; filamenta ca. 0. 5 mm longa. Capsula dorsali-
ventraliter compressa, oblongo-cylindrica, tenuissima,
1.2-1.3 mm longa; placenta basalis. Semen solitarium,
lenticulariter oblongo-ellipsoideum, 1-1.2 mm lon-
gum, translucidum, pallide luteo- brunneolum, longi-
tudine subtiliter striatum.

Delicate, smooth, cespitose perennial; roots
slender. Leaves linear, 4-6 cm long, erect or

ANN. Missouni Bor. GARD. 74: 912-916. 1987.

somewhat spreading, longer than the scape
sheaths; blades 3-5 times longer than the sheaths,

plane or slightly twisted, 0.9-1.2 mm wide, lon-
gitudinally distinctly multinerved, strongly flat-
tened, ferrugineous to olive-green; apices con-
tracted, incurved-acute; margins thickened,
minutely ciliate; sheaths carinate, with carinas
minutely red ciliate, incrassate, the sides strongly
longitudinally nerved, pale to deep red-brown,
the margins gradually converging into the blade,
at apex producing an acute ligule 0.5 mm long,
below gradually dilating. Sheaths of scapes lax,
mostly open, twisted, shining toward the base,
carinate at the middle, with blades either similar
to those of principal leaves or shorter. Scapes
subterete, filiform, + spirally twisted, 1.2-2 dm
high, ca. 0.4—0.5 mm thick, olivaceous, distally
with sharp, smooth costa. Spikes subglobose to
broadly obovoid or short-cylindric, 4.5-6 mm
long, several-flowered, short-attenuate; sterile
bracts 2(-4), 2-2.5 mm long, with dorsal areas
linear and as long as bract, the lowest pair oblong;
fertile bracts tightly spirally imbricate, broadly
ovate, obovate, suborbicular or reniform, ca. 3
mm long, convex and slightly carinate, obtuse to
subtruncate at apex on either side, erose, scari-
ous, minutely rugulose-tuberculate, much thick-
er from the middle to the base, shining, brown-
ish, with margins effusely, dini and rigidly
fimbriate; dorsal area ovate, ca. 2-2.5 mm long,
gray-green. Lateral sepals ca. p connate, ca. 2
mm long, inequilateral, the lobes acute, scarious,
the carinal keel narrow, entire. Petal blades nar-
rowly obovate, ca. 1.5 mm long, apically round-
ed, lacerate, yellow. Staminodia somewhat re-
duced, bibrachiate, the branches at apex
short-plumose with moniliform hairs. Anthers
oblong, ca. 0.3-0.4 mm long, the locules parallel,
distinct; filaments ca. 0.5 mm

1 ids: lenticularly pres ellipsoid,
long, filling capsule, translucent, pale yellow-
brown, finely longitudinally striate.

Additional material examined. BRAZIL. AMAZONAS:
Transamazonas às anes y,53km W ee River;
bus na" regio MARE in open campina of white
27 june 1979, Cleofé E. Calderón, O. P.
a & J. Guedes 2696 (INPA, US, VDB); Muni-
cipio de Borba, acima de Terra Preta, campina do rio

1987]

Surubim, afluente do rio Abacaxis, dos S, 58?*33'W.
Campina aberta, areia branca. Erva de 10 cm de altura;
flores SAN 4 July 1983, C. A. d 4026 (INPA,
NY, VDB).

This species is easily distinguished by its fringe
of strong though slenderly tapering rigid bristles
on the margins of the fertile bracts, nearly sep-
arate anther locules, and reduced staminodial
condition. It is unusual in its particularly small
and thin-walled capsule, this tightly filled by a
single large seed.

Xyris calderonii Kral, Smith & Wanderley, sp.
nov. TYPE: Brazil. Amazonas: Transama-
zonas Highway, 9 km W of Rio dos Pombos,
ca. 1.5 km E of Igarapé dos Pombos, and
ca. 64 km E of the Aripuanà. Common in
white sand campina, flowers yellow, 18 June
1979, Cleofé E. Calderón, O. P. Monteiro &
J. Guedes 2549 (holotype, INPA; isotypes,
US, VDB). Figure 2.

Planta humilis, annua, praeter inflorescentiam gla-
bra. Radices filiformes. Folia linearia, solum basal

à
carinatae, porphyreae, pluricostatae, alesis um sca-
osae £ aminas gra-

curtam latam fascientibus, infime pean ee expansae,

apertae,
foliorum aut brevibus. Scapi che ree plus
minusve spiraliter torti, 5-10 cm m cras-
si, distaliter leviter multicostati, ied levibus Spicae
subglobosae vel late ovoideae, 3-5 mm longae, pluri-
florae, obtusae, involucratae. Bracteae steriles 2-4,

m
"0
c

vel acuminatum, profunde

Ideen: areis dorsalibus valde papillosis, vulgo
sine laminis. Bracteae fertiles late ovatae vel subor-
biculatae, 2.5-3 mm longae, valde rotundato-convex-
ae, villosociliati, areis dorsalibus ovatis, valde granu-

NOTES

913

lato-papillosis. Sepala lateralia libera, oblonga vel ovata,
2-2.5 mm longa, valde inaequilatera, ala carinali lata,

a integ d istante ciliata,
picem versus lacerata. Laminae petalor late ob
atae, ca ngae, luteolae, ad api late ro-

MAR don ca. 0.5 mm lon

longa, pallide brunneola, plus minusve
reticulata.

Low annual, smooth except for the inflores-
cence. Roots filiform. Leaves linear, strictly bas-
al, 3-7 cm long, spreading flabellately, com-
monly longer than the scape sheaths. Leaf blades
2-4 times longer than sheaths, 0.5-1 mm wide,
flat, straight, longitudinally few-costate, flattened
from base to apex, brown to yellow-green, the
tips abruptly narrowed, incurved-acute, the mar-
gins entire, not thickened; sheaths carinate,
brown, many-ribbed, scarious except for the ribs,
with the thin edges stramineous, gradually nar-
rowing into the blades or apically producing a
m scarious, broad ligule, gradually dilating

base, the edges entire. Sheaths of scape
lax, Sici open, straight, carinate, with blades
similar to those of foliage leaves or shorter. Scapes
subterete, filiform, + spirally twisted, 5-10 cm
high, 0.4-0.5 mm thick, distally with many low,
smooth costae. Spikes subglobose to broadly
ovoid, 3-5 mm long, several-flowered, obtuse,
involucrate. Sterile bracts 2-4, xubdecussite: vil-
lous-ciliate, the lowermost pair foliaceous, rigid,
2-5 times longer than the spike, lanceolate to
oblong, 2-3 mm long, carinate, the dorsal areas
linear, green, with blades similar to those of fo-
liage leaves but triangulate; inner pair (if present)

mm long, strongly rounded-convex,
villous-ciliate, the dorsal areas ovate, strongly
granular-papillose. Lateral sepals free, oblong to

ovate, 2-2.5 mm long, very inequilateral, with
keel broad, proximally entire, at middle distantly
ciliate, toward apex lacerate. Petal blades broadly

FIGURE 1.
Blade and sheath junction.— e.
blade, stamen, staminodial min j. See

FIGURE 2.
mid bla
bract.—f. Petal blade and stamen.—g. Stylar

Xyris pectinata i R. Bahia 35).—a. Habit sketch.—b. Leaf tip.—c. Sector of mid blade.
eaf base. — f. Spike. — g. Fertile bract.
d.

—d.
— h. Lateral sepals.—i. Stylar apex, petal

Xyris calderonii Kral, Smith & Wanderley (Calderon et al. 2549).—a. Habit sketch. —b. Leaf at
de.— c. Leaf blade-sheath junction, side (left) and ventral (right) views. —

— d. Mature spike.— e. Fertile

r apex, staminode.—h. Lateral sepal, capsule. — i. Seed.

914 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 74

1987]

NOTES

915

916

obovate, ca. 3 mm long, yellowish, broadly
rounded and strongly erose at apex. Staminodia
bibrachiate, the branches sparsely long-penicil-
late. Anthers oblong, adi ca. 0.5 mm long;
filaments ca. 0.8-1 mm long. Mature capsule
broadly obovoid, a, ca. 1 mm long,
the placenta basal. Seed broadly ellipsoid, ca. 0.3
mm long, pale brown, + reticulate.

There is no question that the affinities of this
little plant are with X. u/eana Malme; in the
production of narrow leaf blades it is most sim-
ilar to var. angustifolia Lanj., which also some-
times produces long-tipped basal bracts. How-

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

ever, the lateral sepals are smaller and show a
somewhat different keel configuration; the dorsal
areas are consistently long excurrent to produce
acicular blades several times longer than the sub-
tended spike. The scapes are uniformly terete.

— Robert Kral, Herbarium, Department of Biol-
ogy, Vanderbilt University, Box 1705, Station B,
Nashville, Tennessee 27235, U.S.A.; L. B. Smith,
Department of Botany, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.; and Sra. Dra.
Maria das Graças de Lapa Wanderley, Instituto
Botánica, Caixa Postal 4005, 01000 Sao Paulo,
razil

A NEW SPECIES OF PIPER (PIPERACEAE)
FROM CENTRAL AMERICA

Piper calcariformis M. Tebbs, sp. nov. TYPE:
Costa Rica. Alajuela: Finca Los Ensayos,
NW of Zarcero, 850 m, Croat 43546 (ho-
lotype, MO; isotype, BM). Figure 1.

, Frutex I- 2 m altus. Folia oblongo- ‘ovata apice acuta

Petiolus va-

ginatus spica pendula, bracteis triangularibus, basin
versus longe calcariformibus

Shrub 1-2 m high, the stems shortly pubescent.
Leaves 16-24(-28) cm long, 10-17(-21) cm wide,
ovate-oblong, glabrous or with short sparse hairs
on upper surface, pilose on veins beneath, the
apex acute, the base cordate and sometimes with
one lobe slightly longer than the other; secondary
veins 4—6, arcuate-ascending towards apex; pet-
iole sheathing, pubescent, 4—9 cm long. Inflores-
cence pendulous, 10-18 cm long, 7-10 mm wide
in fruit, the peduncles 2-4 cm long. Floral bracts
2 mm long, triangular, sparsely to densely pu-
bescent with lower part elongated into a long
spurlike process. Anthers 0.8-1 mm long, de-
hiscing laterally; filaments 1-1.2 mm long. Style
3-4 mm long; stigmas 3, recurved. Fruit 1-1.5
mm long, round-oblong

Additional specimens examined. | PANAMA. BOCAS
DEL TORO: McPherson 7362, 8658 (MO). CHIRIQUÍ:

ANN. MissouRi Bor. GARD. 74: 917-918. 1987.

aiias ge Vodicka 5519 (MO); Correa et al. 2113 (MO).
VERAGUAS: Croat 27695, 27726 (MO); Mori & Kallunki
2586, 3891, 3896 (MO).

Distribution. In moist forest from 400 to
1,200 m in the Cordillera Central in Costa Rica
and in the Serranía de Tabasara in Panama. Only
one collection from Costa Rica, the majority of
specimens coming from Panama.

Piper calcariformis is most closely related to
P. sagittifolium C. DC. and shares similarities of
inflorescence, bract and fruit shape. However, it
can be separated easily from the latter by its ovate-
oblong as opposed to sagittate leaves, its much
longer, pendulous inflorescence, and by its styles
with three stigmas rather than two as in P. sagitti-
folium. Both of these species can be separated
from the rest of Piper by their distinctive inflo-
rescences with unusually shaped bracts. They are
related to the large-leaved pipers with long pen-
dulous inflorescences of section Macrostachys
Miquel.

— Margaret Tebbs, Department of Botany, Brit-
ish Museum (Natural History), Cromwell Road,
London, England.

FIGURE 1.

ANNALS OF THE MISSOURI BOTANICAL GARDEN

MT.
Lu

Piper calcariformis (Croat 43546). — A. Habit. — B. Bract. — C. Fruit. — D. Stamen.

[Vor. 74

A NEW SUBSPECIES OF QUARARIBEA FUNEBRIS
(BOMBACACEAE) FROM NICARAGUA

Recent monographic work has clarified the re-
lationships among members of Quararibea Au-
blet and Matisia Humb. & Bonpl. (Bombaca-
ceae) in the northern Neotropics (Alverson, 1986).
Taken together, these two genera consist of at
least 60 species of trees found in wet or moist,
primary forests from central Mexico to Brazil.
Fieldwork in Mexico, Nicaragua, and Costa Rica
has resolved the taxonomy of some of the more
problematic species belonging to Quararibea
sensu stricto (subgenera Archiquararibea and
Lexarza of Vischer, 1920), including Quararibea
funebris (Llave) Vischer. This widespread species,
named from Mexican material (La Llave & Le-
xarza, 1825; Standley, 1923), includes two sub-
species in Nicaragua, subsp. funebris and subsp.
nicaraguensis. The typical subspecies occurs in
moist to wet lowland and mid-altitude forests
from central Mexico to northwestern Costa Rica,
including forests at altitudes of 300-800 m with-
in Nicaragua. The diminutive a first
described here, occurs above 1,200 m in high-
land forests of a restricted area of s ia A
Nicaragua.

Quararibea funebris (Llave) Vischer subsp. ni-
caraguensis Alverson, subsp. nov. TYPE:
Nicaragua. Matagalpa: Pon "gel María
de Ostuma, Cordillera Central de Nicara-
gua, 1,400 m, 18 Jan. 1965 (fr), L. O. Wil-
liams, A. Molina R., T. P. Williams, D. N.
Gibson & C. Laskowski 27978 (holotype,
WIS; isotypes, F, NY, US). Figure 1.
rbor, 5-25 m alta. Folia obovato-elliptica, 7-22

cm longa, 2-10 cm lata, pilorum caespitibus densis

infra in venarum secundarium axillis exceptis glabrata

16-22 mm longa, 24(-30?) thecis in apice fasciculatis.
Stylus 17-26 mm longus. Fructus drupaceus, subglo-
bosus, in sicco 20-25 mm longus et 14-18 mm dia-
metro, calyce persistenti accrescenti in sicco rugoso et
11-13 mm longo per 3—2 longitudinis inclusus. Sub-
species typicae similis sed praeter pedicellos saepius
longiores in omnes partes minor.

Tree, 5—25 m tall, 25-30 cm dbh, monopo-
dial; trunk smooth and slightly fluted; branches
verticillate, diverging horizontally from trunk and

ANN. Missouni Bor. GARD. 74: 919-922. 1987.

drooping along distal half; bark pale and rela-
tively smooth, not conspicuously peeling; stip-
ules decid narrowly triangular, 2-5 mm long,
1 mm wide, densely lepidote with fimbriate-pel-
tate trichomes. Leaves simple (unifoliolate?), al-
ternate, entire; petioles 7-18 mm long, including
the inconspicuous proximal and distal pulvini,
green, densely lepidote; b/ades entire, obovate to
elliptic, the apex acute to acuminate, the base
acute to rounded, 7-22 cm long, 2-10 cm wide,
dark green, lustrous and glabrate above, medium
green, duller, and glabrous to sparsely pubescent
below, firm-chartaceous; veins prominent below;
secondary veins 5—7 per side, pinnate, arched,
loosely brochidodromous, bearing conspicuous
domatia in their axils; tertiary veins reticulate.
Flowers solitary or few together; pedicels 11-15
mm long, densely puberulent with pale yellow-
ish-brown echinate-stellate trichomes, bearing 3
bracteoles, these broadly triangular, ca. 1.5 mm
ong, 1.5 mm broad at base; calyx infundibuli-
form, irregularly lobed at summit, 9-11 mm long,
pale green, densely pubescent with pale stellate-
echinate trichomes without, densely sericeous
with long, pale, ascending trichomes within, the
apical lobes bluntly triangular, to 3 mm long;
petals spathulate, recurved at anthesis, 15-16 mm
ong, 5-6 mm wide, white when fresh, usually
becoming sepia-brown when dried, moderately
to densely pubescent with pale, lax, stellate tri-
chomes on both sides; staminal column cylin-
dric, dilated towards the toothed summit, 16-22
mm long, 1.5 mm diam., densely pubescent with
pale, stellate trichomes for most of its length,
becoming puberulent towards apex, bearing 24(-
30?) thecae, the 5 apical teeth inconspicuous, ca.
0.5 mm long; thecae distinctly paired, ca. 2 mm
long, 1 mm wide; style filiform, 17-26 mm long,
exceeding the staminal column by 1-2 mm, to-
mentose; stigma capitate, ca. 2 mm diam. Fruits
drupaceous, broadly elliptic to slightly obovate,
bluntly mammillate and briefly apiculate with
persistent style base, 20-25 mm long, 14-18 mm

—

brown color when dried; mesocarp fibrous-fleshy,
without conspicuous taste or odor in fresh, un-
ripe fruits; endocarp bony-fibrous; pedicels of

920

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vor. 74

4 a— ee EE ee eS i
Ñ ha i
aj uararibea funebris (Llave) Mos
T m | ssp. nicaraguensis Alver
ç y [ HOLOTYPE! - |
^ | Det.: W.S. ace peach 3 1986 |
L L Her ba rium, Un jf Wisconsin | (WIS)

lw, g. PME; e [fo

N
Chicago Natural History Museum
Esc

uela Agricola Panamericana

Quararibea funebris (Llave) Vischer
det. AmolinaR. 1967

Mente - fores or cloud forest, Finca Sta. Marta de Ostuma, Cor-
HERBARIUM

alt. 1400 m
January 18, 1965.
To
MADISON (WIS) a e. wa "- Melina R., Terus P. Derothy N. Git
— V — and Chester Laskowski,

FIGURE |. Holotype of Quararibea funebris subsp. nicaraguensis.

1987]

fruits slender, 10-19 mm long, bearing 2-3 per-
sistent bracteoles near middle; ca/yx persistent,
accrescent, cupulate, enclosing fruits for ^-^ their
length, longitudinally finely rugose when dry,
wingless, green, with vestiture as in flowering
condition, the summit erose or irregularly
toothed. Seeds 2, or 1 by abortion, ca. 13 mm
long, 8 mm diam., consisting mostly of pale cot-
yledonal tissue; testa thin, dark brown. Seedlings
unknown when young, probably hypogeal and
cryptocotylar, when older bearing leaves smaller
and more than those of mature plants.

Paratypes. NICARAGUA. ESTELÍ: El Zacatón, “El De-
lirio” camino a la laguna de Miraflor, 13?13'N, 86°14' W
1,400 m, 11 Jan. 1984 (fr), Moreno 22676 (MO, WIS).
JINOTEGA: entre Santa Lastenia y entrada a Aranjuez,
13°02'N, 85?55'W, 1,200-1,250 m, 19 Jan. 1984 (fr),

Sandino 4704 (MO); Finca Aventina, in sierra east of

Jinotega, 1,400-1,500 m, 23 June 1947 (st), Standley
10002 (F); region of Las Mercedes, sierra east of Jino-
tega, 1,200-1,500 m, 3 July 1947 (st), Standley 10746
(F); vicinity of Finca San Roque, sierra east of Jinotega,
300-1,500 m, 5 July 1947 (fr), Standley 10856 (F,
US). MADRIz: Cerro Volcán Somoto (Volcán Tepeso-
moto), 13?26'N, 86?35'W, 25 Sep. 1980 (fl), Moreno
2892 (MO). MATAGALPA: vicinity Santa María de Os-
tuma, on flanks of Cerro El Picacho, immediately east
of Hwy. 3, ca. 9 km north (by air) of Matagalpa, 1 3300'N
85°55'W, ca. 1,500 m, : ies 1982 (fl buds, fr). Alver-
son & Moreno 1982 , CR, F, MEXU, MO, NY,
US, WIS); obice 1984 (MO, NY, WIS); (st),
420 m, 5 Jan. 1984 (fr), Gentry, Stevens
O

aL rarer carretera a Jinotega, 8
1,400-1,450 m, 7 Oct. 1980 (fr), MEE 3381 (MO,
WIS); 13°00'N, 85°55'W, 1,400-1,480 m, 1 Dec. 1982
(fr), 18951 (MO) (same locality as . Alverson & Moreno
: 982); Cordillera cards Santa M
m N of Matagalpa, 1, m, I
Ness (MO); 26 Nov. 1977 (fr), 3005 (BM, ed
MO); 1,500 m, 15 Jan. 1 (fr), b nti
A Ub ndn 23952 (F, G, us en Hda.

95°53" m 1,200-1,300 m,
22574 (MO).

B Dec. 1983 (fr), Stevens

Quararibea funebris subsp. nicaraguensis is
known only from the highlands of northwestern
Nicaragua in the departments of Estelí, Jinotega,
Matagalpa, and Madriz. It is common to loca ally

between 1,200 and 1,600 m. Flowering occurs
during August and September, and fruiting takes
place October through January, indicating a phe-
nology similar to that of the typical lowland sub-
species.

Although very similar, this new subspecies is

NOTES

921

BLE l. Comparison of the subspecies of Quara-
ribea funebris in Nicaragua. Material is from Nicaragua
with the exception of flowers of subsp. funebris, which
are from northwestern Costa Rica

Subspecies
nicara-

Character guensis funebris
Leaf length 71-221 145-289
Leaf width 27-104 62-125
Petiole length 7-18 15-27
Floral pedicel length 11-15 13-14
Floral calyx length 9-11 18-20
Petal length 15-16 33-40
Staminal column length 16-22 33-34
Style length 17-26 35-36
Fruiting pedicel length 10-19 9-17
Fruiting edi length 11-13 18-26
Fruit leng 20-25 25-30

Measurements in mm.

morphologically distinct from Quararibea fu-
nebris subsp. funebris, being smaller in nearly all
respects (Table 1). Its fruiting pedicels, however,
tend to be longer than those of subsp. funebris.
The two subspecies can be easily distinguished
by their floral measurements. When in fruiting
condition, the calyces provide the most obvious
key character; they cover half or less of the ma-
ture fruit of subsp. nicaraguensis, as opposed to
more than half in subsp. funebris.

Thanks are due to D. A. Kolterman for help
with the Latin description, to C. Lipke for the
photograph, to A. Pool for providing specimen
data, and to H. H. Iltis, A. H. Gentry, and W
D. Stevens for comments on the manuscript. I
gratefully acknowledge the O. N. and E. K. Allen
Fund : i ,

Wisconsin for financial support of field research
in 1982

LITERATURE CITED

ALVERSON, W. S

A emiri aes Aubl. s.l. (Bom-
bacacea -

America and the

. 12 in Novorum Vegetabilium Descriptiones,
Volume 2. Mex

STANDLEY, P. C. 1923. Pp. 787—788 in Trees and

922 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 74

Shrubs of Mexico, Part 3. Contr. U.S. Natl. Herb. — William S. Alverson, Herbarium, Botany De-
23(3): 517-848. artment, University of Wisconsin, Madison

VISCHER, W. 1920. Surles Quararibea Aubl., un genre a ds 53706 ey ie Discs
de Bombacées à ovaire infére. Bull. Soc. Bot. Ge- COMAN ae
néve 11: 199-210.

ANNALS OF THE
MISSOURI BOTANICAL GARDEN

VOLUME 74
1987

The Annais, published quarterly, contains papers, primarily in systematic
botany, contributed from the Missouri Botanical Garden, St. Louis. Papers origi-
nating outside the Garden will also be accepted. Authors should write the Editor
for information concerning arrangements for publishing in the ANNALS.

EDITORIAL COMMITTEE
GEoncE K. Rocers, Editor

Missouri Botanical Garden

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Missouri Botanical Garden

GERRIT DAVIDSE

Missouri Botanical Garden
Jonn D. DWYER

Missouri Botanical Garden & St. Louis University

PETER GOLDBLATT

Missouri Botanical Garden

HENK VAN DER WERFF
Missouri Botanical Garden

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(O Missouri Botanical Garden 1988

ISSN 0026-6493

VOLUME 74

ALVERSON, WILLIAM S. A New Subspecies of Quararibea funebris (Bom-
bacaceae) from Nicaragua

AUERBACH, MICHAEL & Avi SHMIDA. Patch Formation among Israeli Cru-
cifers: How Do They Get Away with It?
AVERETT, JOHN E., PETER H. RAVEN & ELSA ZARDINI. Flavonoid Systematics
of Seven Sections of Ludwigia (Onagraceae)

BARNETT, Lisa C. An Unusual New Species of Helmiopsis H. Perrier (Ster-
culiaceae) from Madagascar

BENTLEY, BARBARA L. Nitrogen Fixation by Epiphylls in a Tropical Rain-
forest

Benzinc, Davip H. Vascular Epiphytism: Taxonomic Participation and
Adaptive Diversity

BERNHARDT, PETER. A Comparison of the Diversity, Density, and Foraging
Behavior of Bees and Wasps on Australian Acacia

Brown, KEITH S., JR. Chemistry at the Solanaceae/Ithomiinae Interface
Brown, KEITH S., JR. (See Boyce A. Drummond III & Keith S. Brown,
J

r.

BRYSON, CHARLES T. (See Robert Kral, James Manhart & Charles T. Bry-

son)

BURMAN, ALASDAIR G. (See Gerrit Davidse & Alasdair G. Burman) ...............
CARLQUIST, SHERWIN. Wood Anatomy of Noteworthy Species of Ludwigia
(Onagraceae) with Relation to Ecology and Systematics
CARR, GERALD D. (See Elisabeth Rabakonandrianina & Gerald D. Carr)
CaTLING, PauL M. Notes on the Breeding Systems of Sacoila lanceolata
(Aublet) Garay (Orchidaceae)

CLARK, LYNN G. Two New Mesoamerican Species of Chusquea (Poaceae:
Bambusoideae)

CLARK, LYNN G. New Combinations in Chusquea (Poaceae: Babusoideae)

CRABTREE, Davip R. Angiosperms of the Northern Rocky Mountains: Albian
to Campanian (Cretaceous) Megafossil Floras

Croat, THoMAS B. & MICHAEL H. GRAYUM. New Combinations in Central
American Araceae

D'Arcy, WILLIAM C. & JOHNNIE L. GENTRY, JR. Witheringia folliculoides
(Solanaceae): A New Species from Costa Rica

D'Arcy, WiLLIAM G. & Davip N. SMITH. Saracha spinosa—A New Com-
bination in Peruvian Solanaceae

DAVIDSE, GERRIT. Four New Species of Axonopus (Poaceae: Paniceae) from
Tropical America

1987

DAVIDSE, GERRIT. (See Robert Kral & Gerrit Davidse)
DAVIDSE, GERRIT. (See Ahsan A. Vahidy, Gerrit Davidse & Youji Shige-

nobu)

DAVIDSE, Gerrit & ALASDAIR G. BURMAN. A New Species of Thrasya
(Poaceae: Panicoideae) from the Mosquitia of Nicaragua and Hon-
duras

DavipsE, GERRIT & R. P. Erlis. drundoclaytonia, a New Genus of the
teyermarkochloeae (Poaceae: Arundinoideae) from Brazil 0...

DE Nevers, GREGORY C. The Genus Attalea (Palmae) in Panama ...........
Denton, M. F. (See E. B. Leopold & M. F. Denton)

DIETRICH, WERNER & WARREN L. WAGNER. New Taxa of Oenothera L.
Sect. Oenothera (Onagraceae)

DIETRICH, WERNER & WARREN L. WAGNER. A New Combination and New
Subspecies in Oenothera elata Kunth (Onagraceae)

DiLcHER, Davip. Memorial to Herman F. Becker (1907-1985) oo.
DopsoN, C. H. (See Alwyn H. Gentry & C. H. Dodson)

DRUMMOND, Boyce A., II] & KrrrH S. Brown, JR. Ithomiinae (Lepidoptera:
Nymphalidae): Summary of Known Larval Food Plants

Dwyer, JoHN D. Book Review
Erus, R. P. (See Gerrit Davidse & R. P. Ellis)

FADEN, RoBERT B. & D. R. Hunt. Reunion of Phaeosphaerion and Com-
melinopsis with Commelina (Commelinaceae)

GENTRY, ALWYN. Book Review

GENTRY, ALWYN H. & C. H. Dopson. Diversity and Biogeography of Neo-
tropical Vascular Epiphytes

GENTRY, A. H. & J. STEYERMARK. A Revision of Dilodendron (Sapindaceae)
GENTRY, JOHNNIE L., JR. (See William G. D'Arcy & Johnnie L. Gentry)

GOLDBLATT, PETER. Chromosome Cytology of Oldenburgia (Asteraceae—
Mutisieae)

GOLDBLATT, PETER. Systematics of the Southern African Genus Hexaglottis
(Iridaceae—Iridoideae

GOLDBLATT, PETER. Notes on the Variation and Taxonomy of Watsonia
borbonica (W. pyramidata, W. ardernei) (Iridaceae) in the South-
western Cape, South Africa

GOLDBLATT, PETER. (See James E. Henrich & Peter Goldblatt) u.
GOLDBLATT, PETER. (See James F. Henrich & Peter Goldblatt) .....
GOLDBLATT, PETER. (See James E. Henrich & Peter Goldblatt) 20000000...
GOLDBLATT, PETER. (See Porter P. Lowry II, Peter Goldblatt & Hiroshi

Tobe)

GOLDBLATT, PETER & JAMES E. HENRICH. Notes on Cipura (Iridaceae) in
South and Central America, and a New Species from Venezuela .............

121

GoLDBLATT, PETER & Jon WILLIAMS. Notes on Chromosome Cytology of
Rutaceae—Diosmeae

GRAHAM, ALAN. Fossil Pollen of Sabicea (Rubiaceae) from the Lower Mio-
cene Culebra Formation of Panama

GRAYUM, MicHAEL H. (See Thomas B. Croat & Michael H. Grayum) .......
HaBECK, JAMES R. Present-day Vegetation in the Northern Rocky Moun-

HAMMEL, Barry E. The Origami of Botany: A Guide to Collecting and
Mounting Specimens of Cyclanthaceae
HENRICH, JAMES E. (See Peter Goldblatt & James E. Henrich) U...
Henrico, JAMES E. & PETER GorpBLATT. A Review of the New World
Species of Orthrosanthus Sweet (Iridaceae)

HENRICH, JAMES E. & PETER GorpBLATT. Mesoamerican Sisyrinchium (Iri-
daceae) New Species and Records, and Notes on Typification .............
HENRICH, JAMES E. & PETER GorpBLATT. The Mesoamerican Neomarica
(Iridaceae), N. variegata Henrich & Goldblatt, Comb. Nov. .. u...
HERRERA, JAVIER. Flower and Fruit Biology in Southern Spanish Mediter-
ranean Shrublands

Hoim-NIELSEN, L. B. & J. E. LawEssoN. New Species of Passiflora Sub-
genus Passiflora from Ecuador

Hunt, D. R. (See Robert B. Faden & D. R. Hunt)
Jupziewicz, EMMET J. (See Thomas R. Soderstrom & Emmet J. Judziewicz)

KRAL, ROBERT. A new *Viorna" Clematis from Northern Alabama ..........

KRAL, Robert & Gerrit Davipse. A New Species of Bulbostylis (Cyper-
aceae) from Tropical America ..

KRAL, ROBERT & ARMANDO URQUIOLA. Two Cuban Novelties in Xyris ......
KRAL, ROBERT, JAMES MANHART & CHARLES T. Bryson. A New Carex Sect.
Oligocarpae (Cyperaceae) from Western Arkansas and Eastern Okla-
homa ......
KRAL, ROBERT, L. B. SMITH & MARIA DAS GRAÇAS DE Lapa WANDERLEY. Two
New Xyris (Xyridaceae) from the Amazon Basin of Brazil ecco

Kuyt, Joa. Novelties in Mesoamerican Mistletoes (Loranthaceae and Vis-
caceae) .
Lawesson, J. E. (See L. B. Holm-Nielsen & J. E. Lawesson) .....
Leopo.p, E. B. & M. F. Denton. Comparative Age of Grassland and Steppe
East and West of the Northern Rocky Mountains

LIESNER, RONALD. (See Julian A. Steyermark & Ronald Liesner) U.
LITTLETON, JAN. (See C. J. Webb & Jan Littleton)
LonENCE, Davip H. The Fruits of Decarydendron (Monimiaceae) ..............

Lowry, Porter P., II, PETER GorpBLATT & HiRosui Tose. Notes on the
Floral Biology, Cytology, and Embryology of Campynemanthe (Liliales:
Campynemataceae) "

MAcLEISH, NANDA F. F. Revision of Eremanthus (Compositae: Vernonieae)
MANHART, JAMES. (See Robert Kral, James Manhart & Charles T. Bryson)
MEEROW, ALAN W. Biosystematics of Tetraploid Eucharis (Amaryllidaceae)

MiLLER, CHARLES N., JR. Contributions to a Symposium on the Evolution
of the Modern Flora of the Northern Rocky Mountains: Introductory
Remarks

MiLLER, CHARLES N., JR. Land Plants of the Northern Rocky Mountains
before the Appearance of Flowering Plants

MILLER, JAMES S. Two New Species of Cordia (Boraginaceae) from Central
merica

Mort, Scorr A. Eschweilera costaricensis (Lecythidaceae): A New Species
for the Floras of Costa Rica and Nicaragua

Mort, Scorr A. & GHILLEAN T. PRANCE. A Guide to Collecting Lecythidaceae

NEILL, Davip A. Trapliners in the Trees: Hummingbird Pollination of Fr-
ythrina Sect. Erythrina (Leguminosae: Papilionoideae)

ORNDUFF, RoBERT. Reproductive Systems and Chromosome Races of
Oxalis pes-caprae L. and Their Bearing on the Genesis of a Noxious

Weed

OYEWOLE, S. O. Cytotaxonomic Studies in the Genus Urginea Stein in West
Africa. II. Karyotype Evolution in Urginea altisima (L.) Baker .............

OYEWOLE, S. O. Cytotaxonomic Studies in the Genus Urginea Stein in West
Africa. III. The Case of Urginea indica (Roxb.) Kunth in Nigeria ......
OYEWOLE, S. O. Cytotaxonomic Studies in the Genus Urginea Stein in West
Africa. IV. Population Differentiation and Karyotype Variation in Ur-
ginea indica (Roxb.) Kunth

PorPENDIECK, HaNs-HELMUT. Monoecy and Sex Changes in Freycinetia

(Pandaceae)
PRANCE, GHILLEAN T. (See Scott A. Mori & Ghillean T. Prance) l.
RABAKONANDRIANINA, ELISABETH & GERALD D. CARR. Chromosome Numbers

of Madagascar Plants
RAVEN, PETER H. (See Hiroshi Tobe & Peter H. Raven)
RAVEN, PETER H. (See John E. Averett, Peter H. Raven & Elsa Zardini).
SCHUBERT, B. G. (See O. Téllez V. & B. G. Schubert)
SHIGENOBU, YouJI. (See Ahsan A. Vahidy, Gerrit Davidse & Youji Shige-

nobu)
SHMIDA, AVI. (See Michael Auerbach & Avi Shmida)
SMITH, Davip N. (See William C. D'Arcy & David N. Smith)...
SMITH, L. B. (See Robert Kral, L. B. Smith & Maria das Gracas de Lapa
Wanderley) ..
SODERSTROM, THOMAS R. & Emmet J. Jupziewicz. Systematics of the Amphi-
Atlantic Bambusoid Genus Streptogyna (Poaceae

871

STEIN, BRUCE A. Siphocampylus oscitans (Campanulaceae: Lobelioideae),
a New Name for Burmeistera weberbaueri from Peru

STEIN, BRUCE A. Synopsis of the Genus Burmeistera (Campanulaceae: Lo-
belioideae) in Peru

STEYERMARK, JULIAN A. Flora of the Venezuelan Guayana—ll 2...
STEYERMARK, JULIAN A. New Taxa of Rubiaceae from Venezuela .........
STEYERMARK, JULIAN A. Flora of the Venezuelan Guayana— III U.

STEYERMARK, JULIAN A. Yutajea, Another New Genus of Rubiaceae from
the Guayana Highland

STEYERMARK, J. (See A. H. Gentry & J. Steyermark)

STEYERMARK, JULIAN A. & RONALD LIESNER. Notes on Cornus (Cornaceae)
in South America

SYTSMA, KENNETH J. The Shrubby Gentian Genus Macrocarpaea in
Panama

TAYLOR, CHARLOTTE M. Reconsideration of the Generic Placement of Pal-
icourea domingensis (Rubiaceae: Psychotrieae)

TEBBS, MARGARET. A New Species of Piper (Piperaceae) from Central Amer-
ica

TÉLLEZ V., O. & B. G. SCHUBERT. Una Nueva Especie del Género Dioscorea
(Dioscoreaceae) del Estado de Queretaro, México

TETENYI, PETER. A Chemotaxonomic Classification of the Solanaceae .......
Tope, Hinosur. (See Porter P. Lowry II, Peter Goldblatt & Hiroshi
Tobe)

Tose, HiRosHi & PETER H. RAVEN. Systematic Embryology of the Aniso-
phylleaceae

Topzia, CanoL A. A New Variety of Hedyosmum (Chloranthaceae) from
Nicaragua

URQUIOLA, ARMANDO. (See Robert Kral & Armando Urquiola) U.

VaAHIDY, AHSAN A., GERRIT DAVIDSE & YoujJI SHIGENOBU. Chromosome
Counts of Missouri Asteraceae and Poaceae

WAGNER, WARREN L. (See Werner Dietrich & Warren L. Wagner)
WAGNER, WARREN L. (See Werner Dietrich & Warren L. Wagner) _...

Wake, Davip B. Adaptive Radiation of Salamanders in Middle American
Cloud Forests

WANDERLEY, MARIA DAS GRACAS DE LaPa. (See Robert Kral, L. B. Smith
& Maria das Gragas de Lapa Wanderley)
Wess, C. J. & JAN LITTLETON. Flower Longevity and Protandry in Two
Species of Gentiana (Gentianaceae)

WEBSTER, GRADY L. A New Species of Jatropha (Euphorbiaceae) from
Nicaragua

WENDT, Tom & HENK VAN DER WERFF. A New Species of Ocotea (Lau-
raceae) from Southeastern Mexico

WERFF, HENK VAN DER. A Revision of Mezilaurus (Lauraceae) „u
WERFF, HENK VAN DER. Six New Species of Neotropical Lauraceae
WERFF, HENK VAN DER. (See Tom Wendt & Henk van der Werff)
WILLIAMS, Ion. (See Peter Goldblatt & Ion Williams)

Winc, Scorr L. Eocene and Oligocene Floras and Vegetation of the Rocky
Mountains

Wotre, Jack A. Contributions to a Symposium on the Evolution of the
Modern Flora of the Northern Rocky Mountains: Dedication
Wo re, Jack A. Memorial to Harry D. MacGinitie (1896-1987)
WOLFE, Jack A. An Overview of the Origins of the Modern Vegetation and
Flora of the Northern Rocky Mountains

ZARATE PEDROCHE, SERGIO. Clarification of the Taxonomy of Leucaena
salvadorensis Standley ex Britton & Rose

ZARDINI, Elsa. A New Combination in Lucilia (Compositae—Inuleae) .........
ZARDINI, Ersa. (See John E. Averett, Peter H. Raven & Elsa Zardini)

ZULOAGA, FERNANDO O. A Revision of Panicum Subgenus Panicum Section
Rudgeana (Poaceae: Paniceae)

413

Volume 74, No. 3, pp. 463-680 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN,

was published on December 22, 1987.

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CONTENTS

Contributions to a Symposium on the Evolution of the Modern Flora of the Northern Rocky
Mountains: Introductory Remarks Charles N. Miller, Jr.
Dedication Jack A. Wolfe
Memorial to Harry D. MacGinitie (1896-1987) Jack A. Wolfe
Memorial to Herman F. Becker (1907-1985) David Dilcher
Land Plants of the Northern Rocky Mountains before the Appearance of Flowering

Plants Charles N. Miller, Jr.
Angiosperms of the Northern Rocky Mountains: Albian to Campanian (Cretaceous)
Megafossil Floras David R. Crabtree
Eocene and Oligocene Floras and Vegetation of the Rocky Mountains Scott L.
Wing
An Overview of the Origins of the Modern Vegetation and Flora of the Northern Rocky
Mountains Jack A. Wolfe
Present-day Vegetation in the Northern Rocky Mountains James R. Habeck ....
Comparative Age of Grassland and Steppe East and West of the Northern Rocky
| Mountains E. B. Leopold & M. F. Denton AEN

Fossil Pollen of Sabicea (Rubiaceae) from the Lower Miocene Culebra Formation of

; Panama Alan Graham ee

‘Systematics of the Amphi-Atlantic Bambusoid Genus Streptogyna quiin Thomas
R. Soderstrom & Emmet J. Judziewiez

Wood Anatomy of Noteworthy Species of Ludwigia (Onagraceae) with Relation to Ecalogy
and Systematics . Sheruin Carlquist

The Origami of Botany: a Guide to Collecting and Mounting Specimens of
Cyclanthaceae Barry E. Hamme

| . Mesoamerican Sisyrinchium | (Iridaceae): New Species suit Records, ind Notes on "t

Typification Ms pue E. Henrich & Peter Caen.

681
683
684
689

692

707

William š.