JOURNAL 57655
OF THE tA
ARNOLD ARBORETUM
HARVARD UNIVERSITY
B. G. SCHUBERT
EDITOR
T. G. HARTLEY C. E. WOOD, JR.
D. A. POWELL
CIRCULATION
VOLUME 50
PUBLISHED BY
THE ARNOLD ARBORETUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS
1969
MissouRi BOTANICAL
DATES OF ISSUE
No. 1. (pp. 1-158) issued 15 January, 1969.
No. 2 (pp. 159-314) issued 15 April, 1969.
No. 3 (pp. 315-480) issued 15 July, 1969.
No. 4 (pp. 481-663) issued 17 October, 1969.
ie
.
TER, INDIANA
amy
alee
be Li
TABLE OF CONTENTS
CRETACEOUS ANGIOSPERM POLLEN OF THE ATLANTIC COASTAL
PLAIN AND ITS EVOLUTIONARY SIGNIFICANCE. James A. Doyle
COMPARATIVE ANATOMY AND RELATIONSHIPS OF COLUMELLIACEAE.
William L. Stern, George K. Brizicky, and Richard H. Eyde
“Notes on DIstTRIBUTION AND HasiTat oF COLUMELLIA. George
K. Brizicky and William L. Stern
THE Ecotocy oF AN EvFIN Forest IN Puerto Rico.
HILutTor AND Forest INFLUENCES ON THE MICROCLIMATE
oF Pico pet Orestr. Harold W. Baynton
4. TRANSPIRATION RATES AND TEMPERATURES OF LEAVES IN
Coot Humip ENVIRONMENT. David M. Gates ................
5. CHROMOSOME NUMBERS OF SOME FLOWERING PLANTS.
Lorin I. Nevling, Jr.
THE GENERA OF SENECIONEAE IN THE SOUTHEASTERN UNITED
States. Beryl Simpson Vuillewmier
ASPECTS OF THE CoMPLEX NopaL ANATOMY OF THE DIOSCOREA-
CEAE. Hdward S. Ayensu
ANATOMY OF THE PALM Ruapis EXCELSA, VII. Fiowers. N. W.
Uhl, L. O. Morrow, and H. E. Moore, Jr.
GLYCOSMIS PENTAPHYLLA (RUTACEAE) AND RELATED INDIAN
AxA. R. L. Mitra and K. Subramanyam
VASCULAR ANATOMY OF MONOCOTYLEDONS WITH SECONDARY
GrowTH — an IntTRopucTION. P. B. Tomlinson and M. H.
Zimmermann se
ASPECTS OF REPRODUCTION IN SauRAvUIA. Djaja D. Soejarto ....
THe Eco.ocy or AN ELFIN Forest IN Puerto Rico.
Attnraz,-Roors: A. My Galle .o.0....c.cccc cece
7. Som, Root, anp EarrHworm Rexationsuips. Walter
H. Lyford
Srupies or Stem GrowTH AND ForM AND oF LEAF StRUC-
TURE. Richard A. Howard
LEctToryPiricaTion or Cacauia L. (ComMPosITAE-SENECIONEAE).
Beryl S. Vuilleumier and C. E. Wood, Jr. ...........cc00cc
A REVISION oF THE MALESIAN AND Paciric RAINFOREST CONIFERS,
I. Popocarpacrar, IN PART. David J. de Laubenfels ............
A REVISION OF THE MALESIAN AND PaciFic RAINFOREST CONIFERS,
I. Popocarpacran, IN part (Concluded). David J. de Lau-
benfels .. i
THE Vascuar System In THE Axis oF DRACAENA FRAGRAN
(AGAVACEAE), 1. DistRIBUTION AND DEVELOPMENT OF PRI-
MARY Stranps. M. H. Zimmermann and P. B. Tomlinson ....
9
370
COMPARATIVE MorPHOLOGICAL STUDIES IN DILLENIACEAE, IV.
ANATOMY OF THE NODE AND VASCULARIZATION OF THE LEAF.
William C. Dickison
ANATOMY AND ONTOGENY OF THE CINCINNI AND FLOWERS IN
NANNORRHOPS RITCHIANA (PALMAE). Natalie W. UAL ........
Aspects OF MorpHoLoGy or A US FORMOSANA WITH A
OTE ON THE TAXONOMIC POSITION OF THE GENUS. Hsuan
Keng
A KaryoLocicaL Survey or Lonicera, II. Lily Riidenberg and
Peter S. Green
Nores on West Inp1an Orcuins, I. Leslie A. Garay ...........00000...:
POLLEN CHARACTERISTICS OF AFRICAN SPECIES OF VERNONIA. C.
Earle Smith, Jr.
A New Species or Ficus rrom Suriname. Gordon P. DeWolf, Jr.
A REVISION OF THE GENUS FLINDERSIA (RUTACEAE). Thomas G.
Hartley
A Stupy oF THE GENUS JOINVILLEA (FLAGELLARIACEAE). Thomas
K. Newell
THE Ecouocy or AN ELFIn Forest 1x Purrro Rico.
9. CHEMICAL STUDIES OF CoLorED Leaves. Richard J.
Wagner, Anstiss B. Wagner, and Richard A. Howard ....
THE GENERA OF PORTULACACEAE AND BASELLACEAE IN THE SOUTH-
EASTERN Unitrep States. A. Linn Bogle
STUDIES IN THE NortTH AMERICAN GENUS FOTHERGILLA (HAMAME-
LIDACEAE). Richard E. Weaver, Jr.
Tue Tripe MuvtTIsIEAE (COMPOSITAE) IN THE SOUTHEASTERN
Unirep States. Beryl Simpson Vuilleumier
A New Species oF ARENARIA FROM THE BHuTAN HIMALAYA.
N.C. Majumdar and C. R. Babu
THE Drrector’s REPORT
INDEX TO VOLUME 50
VoLuME 50 NuMBER 1
JOURNAL
OF THE
ARNOLD ARBORETUM
HARVARD UNIVERSITY
B. G. SCHUBERT
EDITOR
T. G. HARTLEY C. E. WOOD, JR.
D. A. POWELL
CIRCULATION
Be ER ISS
: PUBLISHED BY _~CO
THE Sees ARBORETUM OF HARVARD UNIVERSITY —
: _ CAMBRIDGE, MASSACHUSETTS
THE JOURNAL OF THE
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CONTENTS OF NUMBER 1
CRETACEOUS ANGIOSPERM POLLEN OF THE ATLANTIC COASTAL
PLAIN AND ITS EVOLUTIONARY SIGNIFICANCE. James A. Doyle
CoMPARATIVE A AND RELATIONSHIPS OF COL
COLUMELLIA-
CEAE. William, ‘® 1 tom George K. Brizicky, and —
H. Eyde
Nores on DistrisuTion aND HABITAT OF COLUMELLIA. Georg
K. Brizicky and William L. Stern
seu pre ese ELFIN gence: In Puerto Rico.
. Hixtor anp Forsst CES ON THE MICROCLIMATE
or Pico pEL OxsTE. Harold W. Baynton
4. TRANSPORTATION RATES AND TEMPERATURES OF
IN _— oS p ENVIRONMENT. David M. Gates ............
: CEAE. "Edward 8 Ayers
“Tock “Fe AE a ’
(Roracean) AND Retarep Brows
76
JOURNAL
OF THE
ARNOLD ARBORETUM
Vor. 50 JANUARY 1969 NUMBER 1
CRETACEOUS ANGIOSPERM POLLEN OF THE ATLANTIC
COASTAL PLAIN AND ITS EVOLUTIONARY SIGNIFICANCE
James A. DOYLE
ONE OF THE MAJor problems in the study of the evolution of higher
plants is the paucity of evidence from the fossil record on the origin and
evolution of the angiosperms. Because of the relatively sudden appearance
of angiosperms in the fossil record, the lack of recognized angiosperm pre-
cursors, and the lack of any striking peculiarities of the macroscopic remains
of Lower Cretaceous angiosperms (mostly leaves), almost all conclusions
on the origin of the group and the nature of its primitive members have
been based on comparative studies of its living representatives. Angiosperm
paleobotany has been primarily concerned with the geographic vegetational
and floristic changes in the Tertiary, which were due more to migration
and extinction than to evolution. The methods of Tertiary angiosperm
paleobotany, such as the procedure of identifying modern taxa for paleo-
ecological information, have been far less productive in the Cretaceous,
and the problematical nature of the results is undoubtedly largely respon-
sible for the present low level of activity in Cretaceous megafossil paleo-
botany.
In the past decade a new method has been applied in Cretaceous paleo-
botany which promises to shed light on the problems of angiosperm origin
and evolution. This is the study of fossil pollen and spores. In contrast
to the megafossil record, the palynological record shows clear-cut changes
in the morphology and diversity of the angiosperms from the time of their
first appearance. Most of the information on early angiosperm pollen has
been obtained for stratigraphic purposes (cf. Couper, 1964), and is, from
a botanical point of view, largely descriptive; the evolutionary implications
have been frequently mentioned but not discussed in detail.
The purpose of this paper is to review the characteristics of Lower and
early Upper Cretaceous angiosperm pollen floras and to discuss the evo-
lutionary and phylogenetic implications of the record. Much of this dis-
cussion is based on my own work on Cretaceous angiosperm pollen of the
Atlantic Coastal Plain, which appears to give a fairly representative picture
of the world flora. Comparisons will be made with correlative sequences,
which are now known in many other parts of the world. Much of the
2 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
detailed documentation of both the stratigraphic and systematic aspects is
in progress and will be presented later in more complete form, but the
general results seem clear enough to be summarized in this preliminary
paper.
In general, morphological terminology follows Erdtman (1952), though
some terms of Pflug (1953) are used in discussing triporate pollen.
GEOLOGICAL BACKGROUND
The Cretaceous is one of the longer geologic periods, covering some 72
million years between about 136 and 64 million years before the present
(Casey, 1964). The Cretaceous System is customarily divided into Lower
and Upper Cretaceous series, which are subdivided into six stages each
(TABLE 1). These stages were first recognized in western Europe and
subsequently extended around the world; they are now operationally
defined by ammonite zones of the Tethyan province.
Continental or near-shore marine sediments favorable for palynological
TABLE 1. Subdivisions of the Cretaceous
SERIES STAGES
Maestrichtian
=)
Campanian
Santonian Senonian (most common usage )
Upper CRETACEOUS <
Coniacian
Turonian
Cenomanian
Albian
Aptian
i
Barremian
LOWER CRETACEOUS < Re
Hauterivian :
Neocomian (most common usage)
Valanginian
| Berriasian
studies are fairly extensively developed in the Cretaceous, though few areas
have large parts of the system represented by continuous continental
deposition. For example, the uppermost Jurassic and much of the Lower
Cretaceous are well represented in the Purbeck and Wealden of southern
England, but late Lower Cretaceous rocks there are marine and only
marginally suitable for palynological study. The Upper Cretaceous consists
of the wholly unsuitable marine Chalk, and to extend the European
Cretaceous pollen record we must go to Central Europe.
TABLE 2, Presumed stratigraphic relations of ae Coastal Plain
nmarine Cretaceous formation
TIME-STRATIGRAPHIC UNITS ROCK-STRATIGRAPHIC UNITS
SERIES STAGES SOUTHERN AND CENTRAL MARYLAND AREA RARITAN Bay AREA, NEW JERSEY
?—?P—?
[6961
Cliffwood on
Magothy Formation Magothy Formation Morgan
Santonian Amboy nee Clay
—?—?—?—
Hiatus
Coniacian
—?—?—?—
Upper Hiatus Old Bridge Sand
Turonian South Amboy Fire Clay
Raritan Formation Sayreville Sand
es Bh Mek SS Woodbridge Clay
Farrington Sand
Raritan Fire Clay
Cenomanian
Subsurface Only
“Raritan” Formation
—?— —?—?—?— Subzone B
Patapsco Fm. —Zone II—
Albian Potomac Group Subzone A
Lower —?—?—?— Arundel Clay
Aptian a Zone I
Barremian? Patuxent Fm.
NATIOd WUAdSOIDNV SAOADVLAMO “ATAOG
: JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
An excellent section of the late Lower and early Upper Cretaceous for
pollen studies is found in the Atlantic Coastal Plain between Virginia and
New Jersey. Pre-Campanian deposition in this area took place mostly in
river flood plains and deltas. The result is a seaward-dipping wedge of
unconsolidated clays, sands, and gravels, often very rich in organic matter,
which is exposed as a wide northeast-trending band several hundred feet
thick at the landward margin of the Coastal Plain. The presumed strati-
graphic relations are shown in TABLE 2.
In Maryland and adjacent states the basal unit is the Potomac Group,
which until recently was defined as consisting of the Lower Cretaceous
Patuxent, Arundel, and Patapsco formations (cf. Clark e¢ al., 1911). The
Patuxent tends to be feldspar-rich and sandy or gravelly; the Arundel is
a huge lens of dark, organic-rich clay with siderite nodules, while the
Patapsco is rather heterogeneous, though its red and variegated clays are
most characteristic. As is often the case with continental sediments,
Potomac Group lithologies are highly variable, and there is much doubt
that the formations can be consistently separated in the field. The Arundel,
which was apparently deposited in a swamp belt, is the greatest exception
to this, and it is an important marker in dividing the Potomac Group;
however, it is definitely present only in the area between Washington, D.C.,
and Baltimore Co., Maryland.
To these three formations have recently been added higher beds tradi-
tionally designated Raritan Formation (Weaver et al., 1968), which appear
to be earliest Upper Cretaceous. These sediments are generally sandy and
lacking in fossils; the few samples that have been examined palynologically
(discussed below) indicate an age between the typical Patapsco and the
type New Jersey Raritan. In the absence of any distinctive lithological
similarity to the type Raritan, this Maryland “Raritan” should probably
be considered either part of the Patapsco or a new formation of the
Potomac Group.
In the Raritan Bay area of New Jersey, Coastal Plain deposits begin
with the Upper Cretaceous Raritan Formation, which appears to be largely
of deltaic origin (Owens et al., 1968). The Raritan has been divided into
six locally recognizable members, excluding the Amboy Stoneware Clay,
which on palynological and other grounds is better associated with the
overlying Magothy Formation (Wolfe & Pakiser, ms.). The palynologically
important Woodbridge Clay member is dark, massive, highly organic, and
siderite-bearing, like the much older Arundel Clay of Maryland, but the
South Amboy Fire Clay member is more varied lithologically, with lignitic
beds and sands as well as dark, often laminated clays. Most of the Raritan
Formation consists of deltaic sands.
The last nonmarine Cretaceous unit, the Magothy Formation, occurs in
both Maryland and New Jersey. It unconformably overlies both the
Potomac Group and the Raritan Formation; this unconformity represents
a considerable hiatus in deposition even in New Jersey, where older Upper
Cretaceous is present. The Magothy is lithologically distinctive and unlike
the lower units: it consists mostly of alternating sands and dark clays with
“eo
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 5
considerable lignitic material, and it shows much more continuity of
individual beds. It appears to be a deltaic deposit, with evidence of tidal
influence (Glaser, 1967); it is overlain by the often glauconitic offshore
shelf sediments of the Matawan and Monmouth groups.
Unfortunately, controls on the age of the Atlantic Coastal Plain con-
tinental units from marine fossils are poor. Except for an aberrant brackish
water fauna of uncertain age from a deep well on the Eastern Shore of
Maryland (Anderson, 1948), marine fossils are unknown in the Potomac
Group. Marine mollusks from the Woodbridge Clay in New Jersey,
recently restudied by Sohl (pers. comm.), date that unit as middle or late
Cenomanian, while a late Santonian ammonite was recently found in the
Magothy Formation of New Jersey (Owens & Sohl, pers. comm.). Bio-
stratigraphic correlations must therefore be based almost entirely on the
plant fossils, of which the pollen and spores are by far the most useful and
readily obtained. Palynological study reveals a sequence of biostratigraphic
zones which are consistent with the regional lithostratigraphy, and which
compare closely with better dated sequences in other parts of the world.
It is the angiosperms, which were apparently undergoing rapid evolutionary
diversification in the mid-Cretaceous, that are most useful in defining these
zones. Though questions might be raised on the exact correlation of the
angiosperm pollen assemblages in the absence of independent age control,
the relative times of appearance of major types are the same as elsewhere,
and correlations based on the angiosperms agree well with those made with
the spores and gymnosperm pollen alone.
PATUXENT AND ARUNDEL FORMATIONS
The pollen and spore flora of the lower two formations of the Potomac
Group, first described in detail by Brenner (1963), is representative of
mid-Lower Cretaceous floras of most of the world, just before the appear-
ance of typical angiosperm pollen. It is dominated by pteridophytes and
gymnosperms, notably: the fern families Cyatheaceae (or Dicksoniaceae),
Schizaeaceae (Cicatricosisporites, Appendicisporites, and possibly Trilobo-
sporites, Concavissimis porites, etc.), and Gleicheniaceae, as well as groups
of less certain affinities; conifers representing the living families Pinaceae,
Podocarpaceae, Cupressaceae (or Taxodiaceae), and Araucariaceae, and
extinct forms such as Classopollis (which was apparently produced by
plants known as the megafossil genera Cheirolepidium, Brachyphyllum,
and Pagiophyllum, Pocock & Jansonius, 1961), the possibly related Exest-
pollenites tumulus Balme, and the last of the seed ferns, the Caytoniales
(Vitreisporites). Smooth monosulcate grains probably represent the gym-
nospermous orders Cycadales, Bennettitales, and Ginkgoales, while the
Gnetales are represented by grains of the Ephedra type. The picture of
the flora obtained from the pollen and spores is in general agreement with
that provided by the megafossils, which are also predominantly ferns,
conifers, and cycadophytes (Fontaine, 1889; Berry, 1911).
The genus Eucommiidites, common throughout the Potomac Group, 1s
6 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
of special interest since it was first described by Erdtman (1948) as a dicot
from Jurassic rocks. Eucommiidites pollen is smooth and medium-sized,
with three furrows which initially suggest the tricolpate condition typical
of and restricted to dicots (Fics. 1a,b). However, one of the furrows is
wider and more cycad-like than the others, and the general shape of the
grains was shown by Couper (1958) to be more like that of monosulcate
gymnosperm than tricolpate angiosperm grains. Subsequently, Eucom-
miidites has been found in the micropyles of gymnospermous seeds in both
England (Hughes, 1961a) and Virginia (Brenner, aria It was pre-
sumably produced by an extinct group of gymnosper
The Patuxent-Arundel flora does include one oe Clavatipollenites
Couper, which has distinctive angiosperm characters. Clavatipollenites is
generally monosulcate, with the exine finely pilate (clavate), retipilate
(with free pila arranged in a reticulum), or reticulate (with the heads of
the pila fused to form a true reticulum). Couper (1958), in describing the
type species C. hughesii from the Barremian of England, pointed out that
while the monosulcate aperture condition is prevalent in gymnosperms,
pilate or retipilate sculpture is not known outside the angiosperms, and he
noted the similarity of the grains to those of Ascarina in the dicot family
Chloranthaceae. Clavatipollenites has been widely reported from the
middle and late Lower Cretaceous: the Barremian through Albian of
England (Hughes, 1958; Kemp, 1968), the Aptian and Albian of Portugal
(as Apiculatisporis vulgaris Groot & Groot, 1962), the Barremian through
Albian of West Africa and the Aptian and Albian of Central America
(Couper, 1964), the Albian of Australia (Kemp, 1966), presumed pre-
Albian rocks of southern Argentina (Archangelsky & Gamerro, 1967), and
the late (Norris, 1967) and middle (pers. obs.) Albian of the Canadian
Plains. A supposed latest Jurassic or earliest Cretaceous species, C. couperi
Pocock, from Canada (Pocock, 1962) and Egypt (Helal, 1966) is dissimilar
in its exine structure and is probably a cycadophyte (Pocock, pers. comm.;
cf. Kemp, 1968)
Clavatipollenites is so variable that it undoubtedly represents several
natural species. The coarseness of the sculpture varies greatly, and there
is every degree of fusion of the heads of the pila, up to a good reticulum
with large lumina. The grains usually have a simple sulcus, consisting of
a granulate or irregularly sculptured area in the pilate forms (Fics. Ic-e),
or a well-delimited unsculptured membrane in the reticulate ones (Fics.
1f,g). But especially in the overlying Patapsco Formation, the pilate grains
often have a more irregular, sometimes trichotomosulcate aperture (Fic.
1h), as figured by Groot and Groot (1962) as Apiculatisporis vulgaris from
Portugal, or they may be inaperturate or have several weak colpoid areas
(Fic. 1i).?
* Recently Hedlund and Norris (1968. Spores and pollen grains from Fredericks-
burgian (Albian) strata, Marshall County, Oklahoma. Pollen et Spores 10(1): 129-
hese grains o be essentially identical to the irregular-aperturate specimens
of Fg eae ney Hae the Potomac Group, but they show much more complete
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 7
The distinction between retipilate grains with an irregular sulcus and
reticulate grains with a clearly defined sulcus and a te ndency for the
reticulum to detach is important in Kemp’s (1968) separation of Clavati-
Fic. 1. Potomac Group gymnosperm and probable angiosperm pollen. Num-
bers in parentheses refer to slides. a and b, Eucommiuidites troedssonii, grain
sa main sulcus on upper side, two od levels (Aq 45-Ic: Patuxent
d, and e, Clavatipollenites sp., flattened pilate grain with sulcus on : lower
si (Ag 27-1g: Patuxent Fm., Zone I); f and g, C/ P aseaierigsa or se
culate grain, two focal levels (Aq eis. _Patu t Fm., ) a
Clixaipllentes sp., trichotomosulcate grain (B 27-Ic: pores a Subzone
B ); 1, Clavatipollenites sp., tetrac olpoidate grain 651-2 : Patapsco
Fm., ae B of Zone II); j and k, Peromonolites sp. (sensu pie, grain
with sulcus (?) on nie side, two focal levels (Aq 18-lc: Patuxent Fm., Zone
I). All figures * 1000.
intergradation from sulcoidate to colpoidate. This gives gaged support to the a
pothesis that zonaperturate (including tricolpate) pollen is derived from monosul-
cate through tri-, tetra-, or pentachotomosulcate Lamar
8 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
pollenites hughesii and her species C. rotundus. She also found size and
shape were reliable characters, but the retipilate forms in the Potomac
Group are much more variable in size and shape than C. hughesii in
England, overlapping considerably with C. rotundus, while the reticulate
forms often lack the characteristic infolding of the sulcus margin of C.
rotundus and sometimes have coarser sculpture. Brenner (1963) referred
the reticulate forms, which he reported only from the Patapsco, to Liliaci-
dites dividuus (Pierce) Brenner. However, the Liliacidites type inter-
grades with Clavatipollenites and does occur occasionally in the Patuxent-
Arundel, favoring Kemp’s treatment of both types as one genus. The status
of C. minutus Brenner, defined on the basis of smaller size, is doubtful,
since Kemp found it falls within the size variation of C. hughesii. Another
form which should be re-evaluated is the small, coarsely reticulate “Pero-
monolites” reticulatus Brenner, which, as Worrie (1967) suggested, may
be an angiosperm related to Clavatipollenites rather than a perinate spore
(Fics. 1j,k).
edit (1963) was skeptical about the angiospermous nature of Clava-
tipollenites, and he suggested that it represents an extinct group of gymno-
sperms. This possibility cannot be denied, but there is no concrete evidence
for it, and it loses its force because definite (tricolpate) angiosperm pollen
appears in the next formation, and all the morphological characters are
quite at home among the angiosperms. Much of the range of variation
(though not all the intermediates) may be found in the Chloranthaceae:
Ascarina pollen resembles the fine clavate-retipilate forms, Hedyosmum
the coarser clavate irregular-aperturate ones, while Sarcandra pollen is
reticulate and nearly inaperturate. Similar retipilate sculpture is seen in
the Myristicaceae and many dicots with tricolpate pollen, and variation in
the aperture from monosulcate to trichotomosulcate is common in several
monocot and “ranalean” families, e.g. Canellaceae (Wilson, 1964). In
general, Clavatipollenites has more in common with the “‘ranalean’’ dicots
than the monocots, which tend to have reticulate or tegillate rather than
pilate exines
If Clavatipollenites is tentatively regarded as of angiospermous origin,
it is the oldest definite pollen record of angiosperms (cf. Couper, 1964).
Older reports have gradually been rejected as more has been learned of
Mesozoic gymnosperms. Eucommiidites has been discussed; the alleged
nymphaeaceous pollen from the Scottish Middle Jurassic (Simpson, 1937)
appears to have been grains of the coniferous genus Zonalapollenites and
folded araucariaceous grains (Hughes & Couper, 1958), and Rouse’s (1959)
Upper Jurassic Pterocarya was a corroded Classopollis grain (Pocock &
Jansonius, 1961). Classopollis itself was originally misinterpreted by Pflug
(1953) as a tricolpate grain. A possible older occurrence of tricolporates,
in the Berriasian-Valanginian of the Netherlands (Burger, 1966), has not
yet been restudied.
The presence of primitive angiosperm pollen in the Patuxent-Arundel
is consistent with the megafossil record. Fontaine (1889) described dicot
leaves, Ficophyllum, Rogersia, and Proteaephyllum (in part), from the
(catia sitieneteeeniaienaneeeaen
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 9
Patuxent near Fredericksburg, Virginia. Palynological study of the matrix
(Harvard University Paleobotanical Collections: cf. Fontaine, 1889, p. 5)
shows that this locality is indeed of lower Potomac age. Berry (1911)
questioned that these leaves were dicots and suggested that they could be
Gnetum, but reinvestigation by Wolfe (pers. comm.) shows none of the
distinctive fine venation or fiber network characters of Gnetum, and instead
a series of presumed primitive angiosperm characters found in the living
Winteraceae. It should be noted that the distinctive permanent tetrads of
the Winteraceae are absent in the Potomac Group pollen flora, so a direct
affinity is questionable. Isolated entire margined dicot leaves are also
reported from the Aptian of the USSR (Vakhrameev, 1952).
No consistent way has been found to subdivide the Patuxent-Arundel
palynologically, and Brenner (1963) included both in one biostratigraphic
unit, Zone I. The age has not been defined more precisely than Bar-
remian, Aptian, or early Albian. Clavatipollenites and ephedraceous pollen
(Couper, 1964) and the schizaeaceous spore assemblage (Hughes, pers.
comm.) indicate a post-Hauterivian age. The general assemblage suggests
middle more than early Lower Cretaceous (cf. Pocock, 1962), and it is
very much like the flora described from undifferentiated Aptian-Albian
rocks of Portugal (Groot & Groot, 1962). Determinations of an early
Neocomian age based on the megafossils (Berry, 1911; Dorf, 1952) were
made when younger pre-Albian floras were practically unknown. The
upper limit on the age is defined by the absence of tricolpate angiosperm
pollen in the Patuxent-Arundel and its appearance at the base of the
overlying Patapsco Formation. The appearance of tricolpates, discussed
in the following section, is often taken to mark the Aptian-Albian boun-
dary, but the sporadic record of Lower Albian tricolpates leaves open the
possibility that the Arundel-Patapsco boundary lies within the Albian.
PATAPSCO FORMATION
Changes in the pteridophyte spore and gymnosperm pollen flora between
the Arundel and the Patapsco are rather minor: the entrance of a handful
of new species which Brenner (1963) used as index fossils for his Zone IT,
and the decline of some groups such as Classopollis and the Schizaeaceae
within the Patapsco. The most important event is the appearance of small
reticulate tricolpate grains. This pollen type is unlike the “pseudotricol-
pate” Eucommiidites type in having radial symmetry and a reticulate
exine, and it is at present restricted to the dicots. In the lower part of the
Patapsco (Brenner’s Subzone A) the tricolpates are very uniform and
present only in low percentages; in the upper part (Brenner’s Subzone B,
recognized by the entrance of several distinctive spores and gymnosperm
pollen) they become more diverse and abundant; on rare occasions they
constitute a majority of the pollen and spore flora. Their percentage
variation from sample to sample (Brenner in fact encountered Subzone B
assemblages with no tricolpates) suggests that they were restricted ecologi-
cally to certain habitats. Clavatipollenites also increases in abundance in
10 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
the iy ere forms with irregular aperture morphology are not uncommon
(Fics. 1h,i)
. 2. Patapsco — LanfOir All specimens from Patapsco Fm., Sub-
zone B of Zone II. a and b, colpopollenites cf. micromunus, two focal levels
(65—1-2a); c pee - Tricolpo pollenit es cf. minutus, two focal levels (65-S—3h) ;
e and f, Tricolpate type 1, pilate grain, two focal levels (65—-1-2a); g, h, and i,
Tricolpate type | 2. nearly Hsu srg ~ focal levels (65-0-2g); j and k,
V
m, Tricolpopollenites aff. crassimurus, pte ocal levels (65-2a-1b); n and 0,
i i t erc i
, Ip
(65-S—3i) ; s and t, Tricolporoidate type 2, oblate grain with subtriangular amb,
two focal levels (65-2a—1j). All figures 1000.
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 11
Patapsco tricolpate pollen shows some differentiation; at least in the
upper part of the formation perhaps a dozen form species might be recog-
nized. They are typically small (10-20), prolate or spheroidal in shape,
with a fairly thin retipilate or reticulate exine, and colpi without any
specialized margins or wide membranes. Many of the dominant forms
(Fics. 2a,b. Cf. Tricolpopollenites micromunus Groot & Penny, compared
by Brenner to pollen of Tetracentron, or Tricolpites albiensis Kemp) have
fine but well-defined retipilate or reticulate sculpture, without a continuous
tegillum, and sexine somewhat thicker than nexine (1.0-1.5, total exine
thickness). Tricolpopollenites minutus Brenner is very small (11, average
axial dimension), with a reticulum which may be missed without oil im-
mersion (Fics. 2c,d). Besides these and similar microreticulate species,
there are tricolpates with a Clavatipollenites-like exine, with free pila not
arranged into a reticulum (Fics. 2e,f), and at another extreme small forms
with a nearly continuous smooth tegillum which were not reported by
Brenner and are possibly restricted to the upper Patapsco (Fics. 2g-i).
Less common species are “Retitricolpites” vermimurus Brenner with a
loose vermiculate reticulum (Fics. 2j,k), and in the upper Patapsco large
prolate tegillate grains close to Tricolpopollenites crassimurus Groot &
Penny (Fics. 21,m), and a rather thick-walled spheroidal type with a coarse
reticulum in the mesocolpia and internal thickenings (costae) at the
margins of the operculate colpi (Fics. 2n,o).
A previously unmentioned but possibly significant morphological feature
of many of the Patapsco tricolpates, especially the Tricolpopollenites micro-
munus and T. minutus complexes, is a frequent buckling-out of the center
of the colpi, giving them a geniculate appearance and suggesting a rudi-
mentary os. This “‘tricolporoidate” tendency is prevalent in the upper
Patapsco (Fics. 2p-r). Another tendency, so far seen only in the upper
Patapsco and later, results in oblate grains sub-triangular in equatorial
outline, with the apertures at the protruding apices instead of sunken, as
is the rule in Patapsco forms. These grains show differentiation of the
nexine at the aperture and should probably be considered truly tricolporate
(Fics. 2s,t).
The appearance of tricolpate pollen seems to have been a major world-
wide event, and in all areas which have been carefully studied there is a
zone with small reticulate tricolpates but without triporates or typical tri-
colporates (cf. Krutzsch, 1963; Muller, 1968). This appearance generally
may be dated as early or middle Albian, but refinement is needed in most
areas. In England, the Patapsco-type Tricolpites albiensis Kemp appears
at the base of the Middle Albian, but Kemp (1968) found rare grains of
another tricolpate species in one Lower Albian sample. In western Canada
tricolpates are reported by Norris (1967) from the base of the Colorado
Group, which is considered basal Upper Albian (Norris) or late Middle
Albian (Jeletzky, 1968) on the basis of ammonites, while they are reported
to be absent from the underlying Mannville Group (Singh, 1964), which
is presumably Middle Albian at the top. However, mid-Albian tricolpates
cannot be ruled out here since the contact between the two groups is an
12 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 50
unconformity, and in fact rare tricolpates occur locally in the upper Mann-
ville (Pocock, pers. comm.; pers. obs.). The same relation holds in the
U.S. Western Interior: tricolpates are present in the Thermopolis and
Mowry shales of Colorado (lower Colorado Group equivalents; Tschudy
& Veach, 1965), and in the Fall River Formation (basal Colorado Group
equivalent) of the Black Hills, but they are absent from the underlying
Lakota Formation (Cahoon, 1968). In Portugal, they occur in undifferen-
tiated Albian but not in lower Aptian-Albian rocks (Groot & Groot, 1962);
there are other reports from Albian rocks in France (Taugourdeau-Lantz
& Jekhowsky, 1959), Germany (Krutzsch, 1963), and USSR (Zaklinskaya,
1961). In the central USSR Bolkhovitina (1953) reported tricolpates
from the Lower Albian on; Yedemskaya (1960) found them in the Albian
of the Caucasus, plus two isolated grains in the Aptian. In the Southern
Hemisphere, tricolpates occur in the Upper Albian or Cenomanian of New
Zealand (Couper, 1960), and in the Albian of northwestern Australia
(Kemp, 1966).
In view of theories of a tropical origin of angiosperms, it would not be
surprising to find earlier occurrences of angiosperm pollen or more pollen
types in the present tropical areas. However, it appears that here, too,
there is a zone with reticulate tricolpates and no triporates, and that the
tricolpates do not appear demonstrably earlier than in present temperate
areas. In North Borneo the Upper Albian or Cenomanian pollen flora
contains angiosperms only of the same tricolpate and tricolporoidate types
as in the Patapsco, associated with a very similar pteridophyte and
gymnosperm flora (but lacking Pinaceae) (Muller, 1968). In northeast
Brazil (Miller, 1966), the first angiosperm pollen is again reticulate
tricolpates; the age is early Albian or possibly late Aptian. In higher zones
(mid-Albian through Cenomanian, subdivision uncertain) these are joined
by Didymeles-type tricolpodiorates and polyporates, and later triporates.
In Upper Albian samples from Peru, Brenner (pers. comm.) found tri-
colpates and polyporates but no other angiosperm pollen. A very similar
sequence occurs in Africa: in Senegal and the Ivory Coast reticulate
tricolpates and tricolporates (tricolporoidates?) are the only angiosperms
in the Lower (?) through much of the upper Albian; polyporates and later
triporates enter higher in the Upper Albian-Lower Cenomanian interval
(Vachey & Jardiné, 1962; Jardiné & Magloire, 1965). The Albian and
Cenomanian of Gabon yield similar tricolpates, tricolpodiorates, triporates,
and polyporates (Boltenhagen, 1965). The most unusual elements are the
tricolpodiorates and polyporates. The latter are compared with the
Amaranthaceae, but it should be noted that similar pollen occurs in some
monocots (e.g. Alisma spp.). These pollen types do appear earlier in
South America and Africa: the tricolpodiorates in fact are unreported
elsewhere, but rare polyporates are known from the Cenomanian of Okla-
homa (Hedlund, 1966) and Bohemia (Pacltova, 1966). Still, the record
is consistent with a pre-Upper Albian interval with only tricolpate and
tricolporoidate angiosperms.
Brenner (1963) considered the Patapsco to be Albian, and the record
1969} DOYLE, CRETACEOUS ANGIOSPERM POLLEN 13
of the first tricolpates as reviewed here indicates that it is almost certainly
no older than Lower Albian and may in fact be younger. Considering the
record in England and the North American Western Interior, it is quite
possible, as suggested by Wolfe and Pakiser (ms.), that the underlying
Patuxent and Arundel are largely Lower Albian in age. More work on the
pollen flora near the Aptian-Albian boundary in well-dated sequences (e.g.,
in Texas) is clearly in order.
Aside from the Fredericksburg material mentioned above, the first
angiosperm leaves in the Atlantic Coastal Plain are found in the Patapsco
(Fontaine, 1889; Berry, 1911), where they are still a subordinate element.
Similar fossils occur in the (Middle?) Albian lower Blairmore flora of
western Canada (Bell, 1956), the Cheyenne Sandstone of Kansas (Berry,
1922), deposits in the Kolyma basin in eastern Siberia (Samylina, 1960),
Lower and Middle Albian rocks of Kazakhstan (Vakhrameev, 1952), and
the Albian of Portugal (Teixeira, 1948); Vakhrameev (1952) and Takh-
tajan (1960) have noted the characteristic small size of these Albian leaves
and suggested a relation to a still unperfected conductive system.
Brenner (1963) proposed an Upper Albian age for the upper Patapsco
(Subzone B) on the basis of a close specific similarity to the Upper Albian-
Lower Cenomanian angiosperm pollen flora of Portugal (Groot & Groot,
1962). It is generally in the Upper Albian that tricolpate pollen becomes
a characteristic though still subordinate element in the flora and attains
a certain low degree of diversity. Similar Upper Albian (-Lower Ceno-
manian?) floras are seen in the lower Colorado Group of western Canada
(Norris, 1967), the Thermopolis and Mowry shales of Colorado (Tschudy
& Veach, 1965), and Upper Albian-Lower Cenomanian strata in the USSR
(Bolkhovitina, 1953; Bolkhovitina et al., 1963). At the present time it is
impossible to rule out a Lower Cenomanian age for part of Subzone B,
considering the uncertain dating of the correlative deposits, the general
lack of well-dated Lower Cenomanian pollen for comparison, and the only
slightly more modernized floras of the Middle Cenomanian (cf. Hedlund,
1966, and below).
It is in the Upper Albian and Lower Cenomanian that we see the first
megafossil floras dominated by dicot leaves. Such floras are the Dakota
flora of Kansas (Lesquereux, 1892), long considered Upper Cretaceous
but now known to be in part correlative with the Upper Albian Mowry
Shale, the upper Blairmore flora of western Canada (Bell, 1956), and a
series of Upper Albian floras from Kazakhstan (Vakhrameev, 1952).
Characteristic for these floras are a variety of entire leaves and a large
number of lobate leaves with platanaceous venation which have been
referred to several unrelated modern genera (Aralia, Sassafras, Sterculia,
Liquidambar). A “‘Dakota” flora has not been described from the Patapsco,
but this may be because the early collections were made mostly near the
Potomac River, where the Patapsco appears to be pinching out; I have
seen upper Patapsco localities (65—2a, 65—S) rich in simple entire marginal
leaves of a type not described by Berry (1911).
14 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
RARITAN FORMATION
The basal Coastal Plain unit in New Jersey, the Raritan Formation, has
not been studied as comprehensively as the Potomac Group. The Wood-
bridge Clay member, near the base of the formation in the Raritan Bay
area, is best known palynologically (Groot, Penny, & Groot, 1961; Kimyai,
1966; Wolfe & Pakiser, ms.). This unit, dated as Middle or Upper
Cenomanian by marine fossils, is significantly younger than the typical
Patapsco of Maryland: the angiosperm pollen is much more diverse, with
several definite tricolporates (the dominant pollen type in modern dicots)
and low percentages of the first triporates, the genera Complexiopollis
Krutzsch and Atlantopollis Krutzsch of the Normapolles group of Pflug
(1953). Older beds which promise to close the gap between the Patapsco
and Raritan are becoming known to the south of Raritan Bay and in the
subsurface, as are younger beds of presumed Turonian age (the South
Amboy Fire Clay member) in the Raritan Bay area.
The Raritan is the first Coastal Plain unit in which angiosperms clearly
dominate the pollen flora, but gymnosperms and pteridophytes are still
important elements. These are mostly Pinaceae, Podocarpaceae (including
Phyllocladus-like forms and perhaps the bizarre genus Rugubivesiculites
Pierce, with a ruffled central body, which appears in the upper Patapsco
but is most typical of the North American Upper Cretaceous) , Taxodiaceae,
Cupressaceae, Araucariaceae, Cyatheaceae, and Gleicheniaceae; the Schi-
zaeaceae are in decline, and most of the extinct groups represented by
Classopollis, Eucommiidites, Vitreisporites, etc. are very rarely seen. The
angiosperm pollen of the Woodbridge includes reticulate tricolpates of the
Albian type, though many appear to be new species, and occasional mono-
sulcates (Clavatipollenites, Liliacidites). A larger portion is assumed by
small psilate tricolpates and tricolporates (Fics. 3a—d). Some of the most
characteristic of these continue a trend seen in the upper Patapsco: they
are oblate and triangular in equatorial outline, with apical apertures (Fics.
3e,f. Cf. Tricolporopollenites triangulus Groot, Penny, & Groot). Besides
these small, simple tricolpates and tricolporates, there are larger forms
with more complex exine structure (reticulate to completely tegillate) and
apertures (e.g., Fics. 3g-i). An unusual new pollen type is represented by
two forms with permanent tetrads: one is larger, with a smooth tegillum
supported by large pila, and with somewhat obscure colpi arranged accord-
ing to Fischer’s law (Fics. 3j,k); the other is smaller, psilate to retipilate,
with very irregular colpoid areas (Fics. 3l,m). Neither is like the familiar
tetrads of the Ericaceae; the smaller type is strikingly similar to pollen of
the monogeneric family Myrothamnaceae of South Africa and Madagascar.
The most striking new element is the first of the bizarre triporate Nor-
mapolles, which are dominant in the Upper Cretaceous and earliest Tertiary
of Europe. They are an extinct group, not directly comparable to any
living angiosperms, though if the complex protruding apertures of some
of the later form genera were reduced somewhat and the grains became
less triangular, they might approach pollen of some modern amentiferous
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 15
Fic. 3. Lower Raritan and Patapsco-Raritan transition zone ea oe rm a
All specimens except p-s from Woodbridge Clay member, Rari ‘m ind b,
Tricolpate type 4, geen hg ae two focal levels (68-10-1b); c — a
Tricolporate type 1, pro psilate grain, two focal levels (68 8-la); e and f,
Tricolporopollenites cf. pt 1 focal levels (68—-10~1b); g and h, Tricol-
porate type 2, reticulate grain with flat mesocolpia, two focal levels (NJ 2-1a);
i, Tricolporate type 3, reticulate grain with marginate colpi (68~8-1a); j es k,
Tetrad type 1, large, aie: grain, two focal levels (NJ 2-1a); land m trad
type 2, small, scar te grain, two focal levels (68—-10—-Ic); n, Compieaosélie
sp. (NJ 2-1b); Atlantopollis sp. (68—10—-1b); p and q, Tricolporate type 4
possible eens of mers aha group, oblique view, two focal levels
(TR(1551-3)-I1c: up atapsco-Raritan transition zone); T and s,
same, polar view, two focal pate (TR(1551-3)~Ic). All figures * 1000.
16 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
groups (Betulaceae, Casuarinaceae, Myricaceae, Rhoipteleaceae, Juglan-
daceae, Urticales). In fact, as was pointed out by Goéczan et al. (1967) in
their revision of the group, they cannot be rigidly separated from the
Tertiary “Postnormapolles” of Pflug (1953), which include many of the
modern “Amentiferae.”’ Complexiopollis and Atlantopollis are among the
oldest Normapolles in Europe. Atlantopollis differs from the psilate or
scabrate Complexiopollis in its coarsely reticulate or (in New Jersey)
verrucate sculpture. Apertures in both genera are very short colpi or
elongate pores, with the nexine differentiated into an endannular collar
just inside the pore (Fics. 3n,o). The multiple endannular rings seen in
European Turonian species and in the upper Raritan (Fic. 4a) are poorly
developed in lower Raritan forms.
The Normapolles and other pollen and spores provide an age determina-
tion consistent with that from the marine fauna of the Woodbridge. A
Normapolles assemblage with only Complexiopollis and its relatives was
first described from the Lower Turonian of Germany (Krutzsch, 1959),
but it has been extended an uncertain distance down into the Cenomanian.
The Cenomanian Peruc Formation of Bohemia (Pacltova, 1966; Pacltova
& Mazancova, 1966) is probably closest to the Woodbridge: very similar
Normapolles are present in very low proportions, while the rest of the
angiosperm flora contains reticulate and psilate tricolpates and tricolpo-
rates, including a triangular form of the Tricolporopollenites triangulus
type (but also polyporates unknown in the Raritan). Cenomanian deposits
of Portugal (Groot & Groot, 1962) also contain Complexiopollis and
Atlantopollis (as Latipollis). In North America, the Tuscaloosa Group of
Alabama, believed to be of late Cenomanian age, yields a flora with
Complexiopollis and Atlantopollis almost identical to that of the Wood-
bridge (Leopold & Pakiser, 1964). Complexiopollis (as Punctatricolpo-
rites) appears near the putative Cenomanian-Turonian boundary in the
Eagle Ford Shale of Texas (Brown & Pierce, 1962). Wolfe and Pakiser
(ms.) believe the Woodbridge is Upper Cenomanian, and considering the
low percentages of Normapolles this is probably correct, though the range
data permit a Lower Turonian age as well. It is probably younger than
most of the Middle Cenomanian, since Normapolles are not reported from
the Middle Cenomanian Woodbine Formation of Oklahoma (Hedlund,
1966), nor the “Dakota Group” of Minnesota (Pierce, 1959), though both
these floras have post-Patapsco elements such as psilate tricolporates and
similar conifers (diverse Phyllocladus-type and Rugubivesiculites) and
spores (common large Sphagnumsporites, Camarozonosporites, and Glet-
cheniidites ) .
Triporates other than Normapolles appear in other parts of the world
probably in the Cenomanian, though the age control is lamentably poor.
Thus in North Borneo grains with simple round pores appear in a zone
loosely dated as Cenomanian to Senonian (Muller, 1968); triporates com-
pared with Sapindaceae or Proteaceae appear near the end of an Upper
Albian to Lower Cenomanian interval in Senegal and the Ivory Coast
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 17
(Jardiné & Magloire, 1965) and late in the Upper Albian through Ceno-
manian interval in northeast Brazil (Miiller, 1966).
It is becoming clear that there was an interval in the Atlantic Coastal
Plain after the typical Patapsco and before the Woodbridge with angio-
sperm floras including tricolporates, many psilate and some triangular, but
without Normapolles. Extinct gymnosperms such as Classopollis are often
common in these floras. This Patapsco-Raritan transition zone is seen in
surface samples from Elk Neck in northern Maryland and near Trenton,
New Jersey (Wolfe & Pakiser, ms.), from the uppermost “Raritan” of
Bodkin Point, Maryland (pers. obs.), in the subsurface “Raritan” near
Delaware City, Delaware (Brenner, 1967), at the top of the “Raritan(?)-
Patapsco” in a well near Waldorf, Maryland, and in a well some thirty
miles downdip from the Raritan Bay outcrop area on the Toms River, New
Jersey (pers. obs.). The data of Wolfe and Pakiser and from the Toms
River well suggest that the first “Raritan”? elements to appear are the
more prolate psilate tricolporates and, soon after, the triangular forms;
other “Raritan” elements enter as Lower Cretaceous gymnosperms and
ferns decline, until the flora is very close to the Woodbridge except for
the absence of Normapolles.
In one of the uppermost pre-Woodbridge samples from the Toms River
well, an unusual tricolporate occurs with a shape and sculpture very much
like Complexiopollis but with longer vestigial colpi and no typical annulus
or endannulus. The apertures approach those of some of the more complex
Raritan triangular tricolporates, suggesting a link between less bizarre
tricolporates and the Normapolles (Fics. 3 ,
Wolfe and Pakiser (ms.) characterize the pollen flora of the South
Amboy Clay member as essentially the same as the Woodbridge flora
except for some new non-Normapolles triaperturates. However, samples
from four localities in the upper Raritan, including the classic Kreischerville
collections of Hollick (New York Botanical Garden Paleobotanical Collec-
tions: cf. Hollick & Jeffrey, 1909), yield floras which appear to be
significantly younger than the Woodbridge. Although many of the Nor-
mapolles might be considered advanced members of the Complexiopollis
group (Fic. 4a), most show characters of the mid-Turonian and younger
Plicapollis and Vacuopollis groups. Intergradations render generic separa-
tion difficult, but the genus Pseudoplicapollis Krutzsch, with rudimentary
endoplicae and a characteristic pore structure, is definitely present (Fic.
4b). In most of the grains the apertures tend to protrude less than in
Complexiopollis and consist of nearly round pores, with the nexinous collar
retracted or reduced to produce a true vestibulum (as in Plicapollis Pflug)
or atrium (as in Vacuopollis Pflug). Many show apparently structural
folds (Fics. 4c,d), though these are generally less regular than the “endo-
plicae” of typical (younger?) Plicapollis. Some of the subspheroical forms
approach the myricoid genus Triatriopollenites Thomson & Pflug (Fic.
4g). Also present are small psilate triporates perhaps unrelated to the
Normapolles (Fics. 4e,f), and large, oblate brevitricolporates (Porocol-
popollenites, sensu Leopold & Pakiser 1964: Fic. 4h). Other new tricol-
eT
18 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
porates appear to be forerunners of typical Magothy species (cf. Fics.
Sh-k). Most of these forms are reported from the McShan and Eutaw
formations of Alabama (Leopold & Pakiser, 1964), which are referred to
the later Turonian by Wolfe and Pakiser (ms.). Although strictly com-
parable floras have not been described from Europe, the range data of
Goéczan et al. (1967) suggest a Middle or Upper Turonian age.
MAGOTHY FORMATION
The Magothy Formation, which extends from Maryland through Long
Island, has a highly diversified angiosperm flora which has been described
by Stover (1964), Groot, Penny, and Groot (1961), and more completely
by Wolfe and Pakiser (ms.). As noted by Wolfe and Pakiser, the rich and
advanced Normapolles element indicates a sizable break in deposition
before Magothy time, though the presence of Turonian in the Raritan
may close some of the gap. The Raritan marks the end of the nearly
continuous mid-Cretaceous record; remarks on the Magothy flora will
hence be only of a general nature.
Normapolles are a dominant element, represented by at least a dozen
genera. The Plicapollis-Pseudoplicapollis group (with Y-shaped thicken-
ings and vestibula), the Vacuopollis group, now represented by typical
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 19
Vacuopollis (with large atria and thick annuli made up of minute inward
Si apo rods), and Trudopollis Pflug (with thick annuli and endannuli
ind a space or interloculum between inner and outer exine) are especially
common (Fics, 5a—d). There are many small Normapolles (Minor pollis,
etc.) and other simple triporates of the type seen in the upper Raritan
(Fic. 5e; cf. Fics. 4e,f). Such an assemblage must be at least as young
Fic. 5. Magothy angiosperm pollen. a, Plicapollis ‘7 (68-14-1a: Amboy
Stoneware Clay member); b, — sp. (68-14-la); ¢, Vacuo pollis
sp. (68-14 4-la); d, Trudopollis sp. (68-14-1a); e, eee type 3 (68-14-la);
a
Tricolporate type 6, two focal levels (68-16-1a); 1 and m, Monosulcate type ",
grain with sulcus on lower side, two focal levels (Ch-Bf 127 (441-2)-1b: Parte
Fm. undifferentiated). All figures < 1000.
20 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
as mid-Coniacian (cf. Géczan et al., 1967) and is probably younger, since
most of these genera become abundant only in the Santonian (Krutzsch,
1957). More specific evidence is provided by close relatives or new species
of the mid-Santonian and younger genera Praebasopollis Groot & Krutzsch
(with two endannular lips extending into the large vestibula: Fic. 5f) and
Pecakipollis Krutzsch & Pacltova (Plicapollis-like grains without clear
endoplicae and with some Trudopollis traits: Fic. 5g). A Santonian age
is verified by a late Santonian ammonite found in the upper Magothy of
New Jersey (Sohl, pers. comm.).
Tricolpates and tricolporates, many continuing from the upper Rari-
tan, are highly diverse in the Magothy. Upper Raritan and Magothy
pollen types commonly suggest modern families, but most species have
anomalous features or characters now found only in related families.
Thus the grain in Figures 5h and i has some nyssaceous and cornaceous
characters but would not be at home in either family, while the grain in
FIGURES 5j and k has rhamnaceous apertures but hippocrateaceous OF
celastraceous sculpture, and the common myricoid grains (cf. Fic. 4g)
have more of an endannulus than modern Myricaceae. Reticulate, psilate,
and spiny monosulcates very suggestive of modern palms are abundant
in some samples (Fics. 51m). Palm megafossils, among the oldest known,
are also found in the Magothy (Berry, 1916).
From the Turonian on, the world pollen flora is marked by provincial-
ism which contrasts strongly with the cosmopolitanism of the early Cre-
taceous. Zaklinskaya (1962) first pointed out the major provinces of the
Northern Hemisphere in the Senonian and Maestrichtian: the Aquilapol-
lenites province in Siberia and western North America and the Norma-
polles province in Europe and eastern North America (cf. Gdczan et al,
1967). Aquilapollenites Rouse is a peculiar extinct form with a prolate,
often heteropolar, central body and protruding arms bearing the apertures.
It is often associated with pollen of possible proteaceous affinities which
is common also in the Senonian of New Zealand, Australia, and Africa.
Aquilapollenites has been found in equatorial Africa (Jardiné & Magloire,
1965) and Borneo (Muller, 1968), but not in the Normapolles province
until the breakdown of provincialism in the Paleocene, when it occurs
briefly in the Gulf Coastal Plain (Tschudy, pers. comm.). Normapolles
are unknown in Africa and Borneo and very rare in Siberia. There are
also strong similarities between the pollen floras of Africa and Brazil in
the Upper Cretaceous, but these have not been studied as well (cf. Miiller,
1966). The peculiar distribution of the Northern Hemisphere provinces
is clearly a reflection of the epicontinental seas which extended from the
Gulf of Mexico to the Arctic Ocean and along the east side of the Urals
(cf. Tschudy, 1966).
The Magothy flora is a representative of the Normapolles province,
but it illustrates that the province should be divided into American and
European areas. Though many of the stratigraphically important genera
and groups of genera are common to Europe, Wolfe and Pakiser (ms.)
pa a at a
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 21
point out that many bizarre forms are restricted to Europe, and many
of the Magothy genera are new, being most nearly represented only by
relatives in Europe. They note that the Trudopollis group is absent from
the American Turonian, and that most of the Magothy dicots other than
Normapolles are still unreported from Europe. The Atlantic Ocean cer-
tainly acted as a barrier to migration in the Senonian, but it is still sur-
prising that it may have been less effective than the epicontinental sea
of the American interior.
GENERAL EVOLUTIONARY IMPLICATIONS
In the late Lower Cretaceous the angiosperms are a very subordinate,
undiversified element in the pollen flora; by the mid-Upper Cretaceous
they are dominant and highly differentiated, though far from modern in
total variation. The increase in diversity is regular, with new morpho-
logical types appearing not at random but in what can be read as series
that permit derivation of each type from an earlier one. Small etiilate
monosulcates are joined by small retipilate tricolpates; these pass into
tricolporoidates and then tricolporates of more diverse exine eichns
and these into the first triporates, which in turn diversify. The pollen
record by itself leads unambiguously to the conclusion that we are wit-
nessing a major adaptive radiation of a new group. Since we have the
time dimension, we can tell which way to read our series and hence de-
termine which character states are primitive (i.e. ancestral) and which
advanced (i.e. derived). The resulting trends of course apply directly
only to the plants of the time observed, and many of them have doubtless
been reversed in later evolution, but they are relevant to modern groups
insofar as the present major alliances are the result of this radiation and
much of the ancient range in morphology is retained today. Likewise, we
observe directly only evolution in pollen morphology, but this tells us
something about general phylogeny insofar as pollen morphology is useful
in recognizing taxa today, and as primitive or advanced characters in
different organs are loosely correlated as a result of lesser or greater
evolutionary rates in a given line (cf. Sporne, 1954).
The pollen record sets some limits on the time and place of origin
of the angiosperms. The group must have originated at the beginning of
the observed radiation (in the Barremian-Aptian) or earlier, though the
possibility that the earliest pollen with angiosperm characters (Clavati-
pollenites and the early Albian tricolpates ) was produced by plants which
had not reached the angiosperm level in other organs should not be ig-
nored, But at least in the Aptian, the typical dicotyledonous leaf morph-
ology had been attained as well.
So far, the pollen record provides no conclusive evidence on the
hypothesis that the angiosperms appeared and diversified first in the
tropics. The earliest tricolpates there are similar to those in the tem-
perate zones, though after the Middle Albian some pollen types seem to
have appeared earlier in the tropics. There is suggestion of a lag of
22 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
a third of a stage in invasion of the middle latitudes in reports of Lower
Albian and even Aptian tricolpates in Africa and South America on the
one hand and their poor record before the Middle Albian in England and
North America on the other, but the stratigraphy requires much refine-
ment before this can be considered established. In any case, the tropical
belt in the early Cretaceous undoubtedly covered areas which are now
temperate. The tree ferns and cycads in the Potomac Group suggest at
least an equable (warm temperate?) climate, though the lack of bisaccate
conifers in the tropics indicates some latitudinal differentiation.
One area that is ruled out as a center of angiosperm origin and evolu-
tion is the Arctic. Megafossil and microfossil floras from northern Siberia
and Alaska north of the Brooks Range rarely contain angiosperms until
well into the Cenomanian (Teslenko, 1958; Vasilievskaya, 1956; Smiley,
1966; Stanley, 1967; cf. Hughes, 1961b). Most of Seward’s Lower Cre-
taceous angiosperm leaves from the Kome flora of Greenland appear to
have come from the Upper Cretaceous, and the status of the one remain-
ing leaf is uncertain (Koch, 1964).
It is certainly possible that primitive, undiversified angiosperms existed
as a subordinate part of the flora long before the Barremian. We need
only compare the mammals, which originated in the late Triassic but did
not undergo major radiation until the Tertiary. Rare angiosperms with
cycad-like pollen, as in several “ranalean” families, might easily go un-
noticed in the Jurassic or Triassic. However, theories that postulate
that the angiosperms not only existed but diversified long before the
Cretaceous in isolated areas such as the tropical uplands (e.g. Axelrod,
1952) and simply migrated into other areas in the Cretaceous do become
implausible in the light of the progressive appearance of morphological
types. While we might expect a gradual increase in the number of types
as a result of such migration, we would expect a sequence of unrelated
derived types rather than convincing evolutionary series.
The record of early angiosperms is doubtless biased toward prolific
pollen shedders and wind-pollinated offshoots. Even so, if much more
highly evolved pollen was being produced by strictly insect-pollinated
plants, we would expect to see it occasionally. Isolated large, more highly
sculptured grains often found in the Potomac Group (e.g. ‘“Retitricol-
pites” geranioides (Couper) Brenner or the form in Fics. 2l],m) may in
fact represent such plants, but they are similar morphologically to their
smaller and more common associates. The simplest assumption, that the
pollen we see preserved is fairly representative of the morphological types
that existed at the time, is followed here
The concept of an evolutionary radiation of angiosperms beginning in
the early Cretaceous may appear to be in conflict with the megafossil
record. Lower Cretaceous leaves have been placed in such unrelated
modern genera as Populus and Sassafras and form genera (e.g. Ficophyl-
lum and Celastrophyllum) intended to suggest families as distant as
Moraceae and Celastraceae. To reconcile such diversity with the uni-
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 23
formity of the pollen flora by postulating that pollen evolution lagged
behind while other organs differentiated to nearly a modern level would
require an incredible amount of mosaic evolution in many lines. The
pattern of variation in modern angiosperms suggests rather that, in gen-
eral, pollen morphology has not behaved markedly unlike other character
complexes: families may be either very uniform or diverse palynologically.
What is needed is a re-evaluation of the leaf determinations, which were
important fossils. Pacltova (1961) found that cuticles of “Eucalyptus”
from the Cenomanian of Bohemia bore no specific relation to that genus;
the platanaceous venation of Dakota leaves placed in several unrelated
genera and Wolfe’s case for the winteraceous affinity of Ficophyllum have
been mentioned. A detailed study of the morphology of Cretaceous leaves
might prove of more evolutionary interest than attempts at identification
of taxa.
SPECIFIC TRENDS
The most striking evolutionary trends seen in early angiosperm pollen
are in the apertures and shape of the grains. Other more questionable
trends involve the exine sculpture and size. These trends are summarized
in TABLE 3 (p. 28).
It would appear that the monosulcate condition of Clavatipollenites,
the first convincing angiosperm pollen, is very primitive. Within this
group we see as later variants trichotomosulcates, inaperturates, and
grains with several ill-defined colpoid areas. There is a definite trend to
fusion of structural elements into a true reticulum, as in Liliacidites.
The record is consistent with derivation of the tricolpates, the next
most ancient major pollen type, from monosulcates of the Clavatipollenites
type. The similar retipilate exine structure in Clavatipollenites and the
earliest tricolpates favors this hypothesis over such alternatives as a
completely independent origin or derivation from the Eucommiidites type.
It is a general principle that in seeking ancestors for a group we should
consider its most ng (here earliest) members rather than advanced
(later) forms (Thorne, 1963). There are, unfortunately, no obvious in-
termediates between monosulcates and tricolpates in the Cretaceous record,
but the presence of trichotomosulcate apertures and irregular colpoids in
Clavatipollenites may be significant. A theory of the origin of the tri-
colpate condition by loss of the polar connection of the three arms In a
trichotomosulcate grain has been advanced by Wilson (1964). However,
intermediate forms with three colpi displaced toward the pole are lacking.
Another possibility is that the tricolpate condition represents the stabili-
zation of an irregular situation with several colpoids. In the Chloran-
thaceae a similar process may have produced the longitudinal colpi (usu-
ally six) in Chloranthus 2
* See footnote 1 on page 6.
24 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Within the triaperturate group trends are more readily documented,
and the wealth of intermediates leaves no need to invoke independent
origin of the more complex forms. Tricolporates may have originated from
tricolpates via tricolporoidates with only a slight weakening at the center
of the colpus membrane. In some lines this trend was associated with a
change in shape from prolate or spheroidal to oblate with a triangular
amb and apical apertures. A pervasive trend in all the triaperturate
classes (as well as the monosulcates), but most common in the tricol-
porates, was the fusion of exine structural elements into a complete
tegillum, often resulting in psilate grains. Permanent tetrads also appear
as a later offshoot in the triaperturate groups.
The small size of the early tricolpates suggests that this may be a
primitive character in the triaperturates. However, the occasional pres-
ence of large grains suggests size was an unstable trait from the beginning,
being subject to changes in pollination ecology. Small grains are often
associated with wind pollination, but the Albian forms are even smaller
than most amentiferous pollen. A comprehensive comparative study of
size-pollination relations in modern angiosperm pollen would be desirable.
The culmination of the trend toward apical apertures is evidently seen
in the triporate Normapolles, which may have been derived from con-
ventional dicots through triangular tricolporates in pre-Woodbridge time.
The Normapolles show various peculiar trends, such as the evolution of
atria, vestibula, and other elaborations of the pores, and the development
of endoplicae suggesting the arci of the Betulaceae, Rhoipteleaceae, and
Ulmaceae. Soon after the origin of the group, the shape trend was appar-
ently reversed to produce subspheroidal grains, as in most of the modern
“Amentiferae.” Other triporates, seen in the upper Raritan and parts of
the world where Normapolles are lacking, may be of independent origin;
they might originate by reduction of the colpus in a tricolporate grain or
perhaps by contraction of the colpus in a tricolpate.
Pollen with numerous scattered pores occurs in the Cretaceous of some
areas, apparently always after the entrance of tricolpates. Polyporates
are found today in both monocots and dicots: the fossil record is con-
sistent with derivation from either monosulcates or tricolpates and does
not indicate which alternative is correct. Occasional dicolpate and poly-
colpate variants of tricolpate Raritan species and tetraporate Normapolles
grains are within the normal variation of modern species; apparently such
variation did not lead to major trends in the Cretaceous.
The proposed evolutionary relationships among the major pollen types
are shown in their stratigraphic framework in Ficure 6. This scheme is
almost identical to the one proposed by Takhtajan (1959, 1966), based
on the comparison of the pollen of angiosperms which are believed to be
primitive or advanced in other characters. Many of the same trends are
implicit in the writings of Bailey and coworkers (e.g. Money, Bailey, &
Swamy, 1950) and of Wodehouse (1936). One of Takhtajan’s impor-
tant trends, from monosulcate to monoporate, is not shown since mono-
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 25
Turonian
(U. Raritan)
other triporates higher Normapolles
—_
oS !
(L. Raritan) (@ ? ? a
tetrads Complexiopollis
Cenomanian |
(Patapsco-
Raritan ; ,
Sranettien tricolporates tricolporates:
| triangular amb
ne nee
: ae
Altiien polyporates* tricolporoidates
(Patapsco) ?
2 aa
= ee tricolpates
Barremian-
Aptian * Africa, South
(Patuxent - monosulcates America only
Arundel) (Clavatipollenites type)
Fic. 6. Suggested evolutionary interrelationships of the major angiosperm
pollen types of the Potomac-Raritan interval (Barremian-Turonian).
porates are not reported from Turonian or older rocks, but it is suggested
by the record. Monoporates of a graminioid type are known from the
Maestrichtian of Africa (Jardiné & Magloire, 1965).
The trends proposed here are very different from those of Kuprianova
(1966), who lists as primitive a large number of characters found in such
groups as the Santalales, many amentiferous plants, and Upper Cretaceous
fossils including the Normapolles which suggest an ancestry with trilete
spores. This approach overlooks earlier Cretaceous fossils which point
toward simple tricolpate or monosulcate ancestral forms; the fossil sequence
clearly shows sporelike Upper Cretaceous forms and their modern analogs
are secondarily derived. As a rule, I would suggest that no exclusively
post-Middle Albian pollen types can be used directly to reconstruct primi-
tive conditions in angiosperms.
PHYLOGENETIC INFERENCES
Although the Cretaceous pollen record does not show us the origin of
26 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
the angiosperms, it does allow us to make more secure inferences about
the ancestors of the group. Thus the earliest monosulcate angiosperm
pollen points to a group of gymnosperms with monosulcate pollen. This
suggests an ancestor among the cycadopsids (i.e. the seed ferns and their
presumed derivatives) rather than the coniferopsids or pteridophytes. An
ultimate seed fern ancestor, making the angiosperms a parallel group to
the cycads, Bennettitales, and Caytoniales, is suggested by the compara-
tive morphology of the other plant organs (cf. Takhtajan, 1960; Cron-
quist, 1968). Clearly more must be known of Triassic, Jurassic, and, as
Hughes (1961b) emphasizes, Lower Cretaceous gymnosperms before a
more definite hypothesis may be presented.
Discussion of an ancestor of the angiosperms assumes the group is
monophyletic, at least in the loose sense of Simpson (cf. Cronquist, 1968).
This assumption is consistent with the record, which appears to show one
major radiation, with the more ancient representatives of putative lines
more instead of less similar to each other. Even the Normapolles may be
derived from earlier tricolporates. This argument holds only for the basi-
cally tricolpate groups and their immediate monosulcate ancestors (i.€.
the bulk of the dicots), and it does not mean all the characters we asso-
ciate with the angiosperm grade had evolved when the taxon originated.
In any case, it is quite likely that still more primitive groups with mono-
sulcate pollen reached the angiosperm level in several lines, resulting in
much of the heterogeneity of the living “Ranales
Speculation on the affinities of early sngioapern pollen might easily
lead to unwarranted conclusions on the age of modern taxa. It is not
difficult to find modern analogs of Albian pollen: we have seen that much
of the morphological variation in the Clavatipollenites type may be found
in the Chloranthaceae, while similar generalized tricolpates occur in the
Lardizabalaceae and Menispermaceae, or the Tetracentraceae, Hama-
melidaceae, Platanaceae, and related families of the Trochodendrales and
Hamamelidales of Cronquist (1968). However, the monosulcate and
tricolpate complexes were young and evolving rapidly in the Albian, and
their total diversity could probably be accommodated in two or three closely
related orders and perhaps five to ten families. In contrast, the modern
families mentioned are relictual and isolated from each other by specializa-
tion and extinction. It would probably be a mistake to believe the simi-
larities indicate any more than that such taxa have retained a primitive
pollen type, and hence perhaps other primitive characters.
Higher dicot groups (e.g. Salicaceae) may have reticulate tricolpate
and tricolporoidate pollen, but unlike the Ranunculales, Trochodendrales,
and Hamamelidales they are usually dominated by tricolporate pollen
(cf. Fagaceae, Elaeocarpaceae, Flacourtiaceae). In terms of the subclasses
of Takhtajan (1966) and Cronquist (1968), it is possible that Albian
angiosperms had not evolved beyond the level of the Magnoliidae and
lower Hamamelididae, and that the higher Hamamelididae (most of the
“Amentiferae”), Dilleniidae, Caryophyllidae, Rosidae, and Asteridae were
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 27
represented only by ancestors more primitive in pollen morphology and
many other characters than their present members. These taxa may have
differentiated in the radiation of basically tricolporate groups beginning
in the Cenomanian. Some Upper Cenomanian tricolporates already sug-
gest orders of the Rosidae such as the Cornales.
The fossil record indicates that extinct dicot alliances, represented by
the Normapolles and Aquilapollenites, flourished in the late Cretaceous
(cf. Krutzsch, 1963). The possibility of extinct major groups is largely
ignored in angiosperm phylogeny, but it is quite relevant, for example, in
the ““Amentiferae,” which may be in large part relics of the group repre-
sented by Normapolles pollen.
Since the basic monosulcate pollen type of monocots is common among
“ranalean” dicots, the fossil pollen record is ambiguous on the origin of
monocots. Though typical Clavatipollenites is most like the pollen of
some modern dicots, the usually younger reticulate Liliacidites type could
be either “ranalean” or monocotyledonous. Some Cenomanian pollen is
more convincingly monocotyledonous in origin.
The discussion of the last paragraphs shows the consistency of the rec-
ord with the systems of Takhtajan and Cronquist. In general, “ranalean”
theories of angiosperm phylogeny are favored, since the earliest angio-
sperm pollen is of types characteristic of or restricted to groups considered
primitive in such theories. On the other hand, systems which make the
wind-pollinated “Amentiferae” primitive become implausible. The Betu-
laceae, Casuarinaceae, Myricaceae, Rhoipteleaceae, Juglandaceae, and
Urticales all have basically triporate pollen (from the Normapolles?), a
definitely derived, though ancient, type. The Fagaceae, with generally
prolate tricolporate pollen, have a questionable status, but the unusual
complex protruding apertures in Trigonobalanus doichangensis (Camus)
Forman (Erdtman, 1967) suggest a relation to the Normapolles.
CONCLUSIONS
Angiosperm pollen types in the Cretaceous of the Atlantic Coastal
Plain appear in essentially the same sequence as in other areas, including
the tropics. In the Patuxent and Arundel formations (Barremian?-Lower
Albian?) the retipilate monosulcate genus Clavatipollenites, apparently
the oldest pollen with characters restricted to angiosperms, occurs In a
ora dominated by pteridophytes and gymnosperms. Clearly dicotyle-
donous reticulate tricolpate pollen appears at the base of the Patapsco
Formation (Lower-Middle Albian?); tricolpates increase in abundance
and diversity in the upper Patapsco (Upper Albian-Lower Cenomanian?),
where many show tricolporate tendencies. Definite tricolporates, often
psilate and with triangular amb, occur in beds transitional to the Raritan
Formation; in the lower Raritan (Upper Cenomanian?) they are joined
by the first triporates, Complexiopollis and Atlantopollis of the extinct
(pre-amentiferous?) Normapolles group. In the upper Raritan (Middle-
Upper Turonian?), these pass into more advanced Normapolles genera.
28 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The rich flora of the Magothy Formation (Santonian), which includes
some forms suggesting modern families, is representative of the Senonian
Normapolles province of Europe and eastern North America.
The expansion and diversification of angiosperm pollen in the Cretaceous
is believed to reflect the basic adaptive radiation of the group, within
which morphological series documenting evolutionary trends and the origin
of major types may be recognized. Though the angiosperms may have
originated well before the observed radiation, the idea that they were
highly differentiated at their first appearance in the fossil record conflicts
with the low diversity of Albian angiosperm pollen and the regular sequen-
tial appearance of morphological types. Trends such as monosulcate to
tricolpate, prolate tricolpate to tricolporoidate to oblate tricolporate to
triporate, and retipilate or reticulate to completely tegillate are in good
agreement with trends postulated on the basis of comparative morphology
and with systems in which the Magnoliidae and lower Hamamelididae
are considered primitive and the ‘““Amentiferae” advanced. Considering the
evidence for important evolution in pollen characters, it is hoped that
the megafossil record of early Cretaceous angiosperms will be re-examined
with modern techniques and a more evolutionary-morphological point of
view.
TABLE 3. Evolutionary trends in pollen morphology based on the
Cretaceous pollen record
GENERAL APERTURE TRENDS:
monosulcate, bilateral symmetry — tricolpate, radial symmetry
monosulcate or tricolpate — polyporate
simple colpi > complex apertures
MONOSULCATE GROUP
monosulcate > eiietishicisatitiade, inaperturate, or with several colpoids
pilate or retipilate — reticulate or completely tegillate
gpeenteris GROUP
colpate > iecokliee — tricolporate > triporate
aa — triporate:
prolate or subspheroidal — oblate, triangular amb, angulaperturate
retipilate or reticulate — psilate, completely tegillate
single grains — permanent tetrads
small size — large size?
NORMAPOLLES GROUP
pores nearly siete — pores with atria or vestibula
no endoplicae — endoplica
triangular amb —> circular amb
ACKNOWLEDGMENTS
I am deeply indebted to Professor Elso S. Barghoorn, whose sponsor-
ship made this study possible, for his continuing advice, encouragement,
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 29
and criticism. Dr, Alexandra Bartlett provided invaluable advice on
palynological techniques and interpretation of pollen morphology. I also
wish to thank the many persons acknowledged in the text for discussions
and unpublished information. E. T. Cleaves, J. D. Glaser, and H. J. Han-
sen, III, of the Maryland Geological Survey, J. P. Owens and C. F. With-
ington of the U. S. Geological Survey, and H. F. Becker of the New York
Botanical Garden supplied me with samples. I have received financial
support from the Committee on Evolutionary Biology (NSF grants
GB3167, GB7346; principal investigator, R. C. Rollins) and the Graduate
School of Arts and Sciences of Harvard University.
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APPENDIX
Loca.LiTies CITED IN TEXT AND FIGURES
PATUXENT FORMATION:
All specimens figured are from the lower Potomac Group exposed in construc-
tion of the Susquehanna Aqueduct between Baltimore and Aberdeen, Md.,
collected by E. T. Cleaves. Locality data are given in Cleaves (1968):
Aq 18 = Cleaves sample no. 18
q ? = ”? 9 be
27
Aq 44 = ” ” ” 44
Aq 45 = 7 7 7 45
Patapsco FORMATION:
65-1: exposure on E side of parking lot behind eget Center on 52nd St.,
ca. 0.2 mi. N of junction of Kenilworth Ave. (Md. 201) and Baltimore-
Washington Parkway, S of Bladensburg, Md. cee s priced 17). Gray clay
with cupressaceous twig compressions, 40-50’ above surface of parking lot
and Brenner’s sample, overlain and underlain by red and gray clay. Subzone B-1
of Zone II, vs. Subzone A of Zone II for Brenner’s sample.
65-2a: NW side of West Bros. Brick Co. pit on N side of Sheriff Rd., 1.0 mi.
E of Washington, D. C. city limit, ca. 0.7 mi. NW of Highland Park, Md.
(Brenner’s Seuttin 29). Thin gray clay lens with dicot leaf compressions near
34 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
top of predominantly red sit roughly same level as Brenner’s collection but
toward N side of pit. Subzone B of Zone II.
65-O: exposure on NE corner of Branch Ave. and O St. SE, Washington, D.C.
Gray clay lens with lignite bed, grading downward and laterally into red and
white clay, and overlain by cross-bedded sands with ironstone concretions
(“Raritan” Formation?). Subzone B of Zone II.
65-S: N side of Severn Clay Co. pit, 0.1 mi. N of road connecting Ritchie
Highway and Md. Rt. 648, ca. 0.5 mi. SE of Harundale, Md. (Brenner’s Station
11). Gray clay lens with dicot leaf compressions just above base of pit, overlain
by red clay. Subzone B of Zone II.
B-27: James D. Bethards No. 1 well (Socony-Vacuum Oil Co.), ca. 5 mi. SW
of Berlin, Md. (cf. Anderson, 1948). Gray clay core sample from 2735-2751’,
provided by Maryland Geological Survey. Subzone B of Zone II.
PATAPSCO-RARITAN TRANSITION ZONE:
68-65: exposure overlooking Chesapeake Bay, ca. 0.6 mi. S of Bodkin Point,
Anne Arundel Co., Md. Gray clay exposed just above beach level, passing
laterally into red clay, overlain by yellow-white sands with ferruginous ledges.
“Raritan” Fm.: Patapsco-Raritan transition zone.
Ch-Bf 127(536-7) and Ch-Bf 127(546-7): well ca. 1.5 mi. NE of Waldorf,
Md. (Ch-Bf 127: cf. Hansen, 1968). Medium gray clay core samples from
536-537’ and 546-547’, obtained from Maryland Geological Survey. Near top
of “Raritan(?) -Patapsco” Fm.: Patapsco-Raritan transition zone.
TR(1551-3): Toms River Chemical Co. Test Well No. 84, 39° 59’ 3” N
latitude, 74° 14’ 20” W longitude, Ocean Co., N.J. Gray clay core sample from
1551-1553’, obtained from H. E. Gill through J. P. Owens, U.S. Geological
Survey. Near top of Patapsco-Raritan transition zone: ecigey from 1369-1371’
and 1298-1300’ yield typical Woodbridge pollen and spore
RARITAN FORMATION:
NJ 2: “Woodbridge, N.J.” Light gray clay matrix from specimen of Magnolia
glaucoides Newberry, N.Y. Botanical Garden Paleobotanical Collections. Wood-
bridge Cla
68-8: S side of Sayre & Fisher Brick Co. pit, on S side of Main St., just NE
of Sayreville, N.J. Near top of massive dark gray clay exposure. Woodbridge
Cla
68-10: E side of same pit. Medium gray clayey sand at top of massive dark
aed Woodbridge
68-12: NE end of shies sand pit ca. 0.5 mi. NNE of Phoenix, N.J. Gray clay
capping thick cross-bedded sands. Old Bridge Sand?
68-23: W side of clay pit on N side of Washington Rd., ca. 0.5 mi. E of Parlin,
N.J. Near base of thin-bedded gray clay unit, underlain by light gray sand, at
low elevations in pit. South Amboy Fire Clay.
68-25: same pit. Near top of same clay unit, exposed just to W and 10-20’
higher. South Amboy Fire Clay.
8-26: same le Thin bed of laminated gray clay exposed near top of small
hill near NW corner of pit, underlain by white sand, and overlain by thin bed
of thinly eaaOG lignitic sand (68-27). South Amboy Fire Clay or Old Bridge
Sand.
68-27: see under 68-26.
68-28: S side of abandoned sand pit off E side of Hillside Ave., just N of high
voltage wires, Sayreville, N.J., above and to E of Sayre & Fisher pit (68-8,
1969 | DOYLE, CRETACEOUS ANGIOSPERM POLLEN 35
68-10). Gray clay lens in predominantly coarse-medium grained sand. Sayreville
Sand or South Amboy Fire Clay.
MacotHuy ForMATION:
68-14: SW corner of Madison Township dump, 0.3 mi. E of U.S. Rt. 9, ca.
0.9 mi. S of junction with Ernston Rd., and 1.5 mi. SSE of Ernston, N.J. Dark
gray clay overlain by thin-bedded alternating sands and clays of the Morgan
beds of the Magothy Fm., near lowest elevations in dump. Amboy Stoneware
Clay (J. P. Owens, pers. comm
68-16: bluff overlooking Raritan Bay NE of town of Cliffwood Beach, N.
Gray silty clay just below contact with wag Merchantville Fm. exposed
at top of bluff. Cliffwood beds of Magothy
Ch-Bf 127(441-2): same well as Ch-Bf 127 samples under Patapsco-Raritan
transition zone. Fine gray clayey sand core sample from 441-442’. Near top of
Magothy Fm.
Note: Brenner localities are those described in Brenner (1963). Unless other-
wise indicated, samples were collected by J. A. Doyle. All slides are located at
the Harvard University Paleobotanical Collections.
DEPARTMENT OF BIOLOGY
Harvarp UNIVERSITY
36 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
COMPARATIVE ANATOMY AND RELATIONSHIPS
OF COLUMELLIACEAE
WILLIAM L, STERN, GeorcE K. Brizicxy,'
Fs
AND RICHARD H. EypE
Género dedicado 4 Junio Moderato Columela, antiguo espaiol,
colocado por Linneo entre los padres de la Botanica, y que escribo
elegantemente en prosa y verso de Labranza y cultivo de Jardines
— Ruiz and Pavon 1794.
In 1961, Brizicky summarized information on the Andean genus Col-
umellia and presented a taxonomic synopsis of this puzzling group of plants.
The genus was described in 1794 by Ruiz and Pavén and David Don estab-
lished Columelliaceae in 1828. Eleven species have at one time or another
been ascribed to the genus and through his critical examination of all
available herbarium specimens, Brizicky reduced this number to four
more or less well-defined species. Evaluations of the taxonomic position
of Columellia and Columelliaceae have been set forth from the time of
A. L. de Jussieu and Ruiz and Pavén, but even the latest authors have
been unable to fix the relationships of these plants conclusively. ‘With
its peculiar combination of opposite, exstipulate leaves; bisexual, epigy-
nous flowers; somewhat irregular, sympetalous corollas; two stamens
with plicate and contorted anthers resembling those of some Cucurbitaceae;
two-carpellate, imperfectly two-locular ovaries; and imperfectly four-
locular capsular fruits, Columellia is indeed a unique genus’ (Brizicky
1961).
Although several positions have been proposed for Columellia and for
Columelliaceae, taxonomists agree that a plausible understanding of the
relationships of these plants requires comprehensive studies to clarify dis-
puted points and to complete our knowledge of their anatomy. It was
with this in mind that the present authors have examined the anatomy
of the flower and fruit, node, leaf, and secondary xylem.
Taxonomic position of Columellia
A. L. de Jussieu (1801) considered Columellia as a genus of Oleaceae
“hoc Genus ad Jasminearum ordinem pertinere.”’ Kunth (1818) placed
the genus in Scrophularinae, but noted, “‘An Gesnereis affinior?” At first
Reichenbach (1828) included the genus in Gesneriaceae (‘‘Gesnereae’’ as
a tribe of Bignoniaceae) but later (1837) he transferred it to Oleaceae
e K. Brizicky died in 15, 1968 in Cambridge, Massachusetts, during the
ie a of the preparation f this — It is to his memory that the sur-
yiving authors respectfully cook this pape
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 37
(“Jasmineae”). Bartling (1830) retained Columellia in Scrophulariaceae
among “Genera incertae sedis.” Sprengel (1830) supposed the affinity of
the genus to be with Gesneriaceae. In 1839, Endlicher placed Columellia
near Ebenaceae among “Genera Dubiae Affinitatis”; later (1841), he in-
cluded it in his classis (order) Petalanthae (Primulaceae, Myrsinaceae,
Sapotaceae, Ebenaceae, and Styracaceae) as a genus “‘Petalanthis affinis.”
Schnizlein (1843-1870) recommended an affinity with Saxifragaceae-
Escallonioideae (‘‘Escallonieen’”’), and particularly with the genera Argo-
phyllum J. R. & G. Forst., Brexia Nor. ex Thou., and Roussea Smith.
J. D. Hooker (1873, 1875) suggested referring the genus to Loganiaceae.
Baillon (1888) included Columellia in Gesneriaceae as a representative of
the monogeneric series Columellieae (between series Gesnereae and series
Cyrtandreae). Hallier at first (1901) placed Columellia in Rubiaceae as an
anomalous genus and later (1903) included it in Scrophulariaceae as ques-
tionably related to Veronica sect. HEBE Benth. of the tribe Leucophylleae.
Finally (1908, 1910) he transferred it to Saxifragaceae-Philadelpheae.
Herzog (1915) also regarded Columellia as a genus of Saxifragaceae.
Taxonomic position of Columelliaceae
David Don (1828), who founded the family Columelliaceae, considered
it allied to Oleaceae (“Oleinae” and “Jasmineae”) as well as to Styra-
caceae and Ebenaceae. Apparently following the suggestions of his brother,
George Don (1838) showed Columelliaceae (“Columellieae”) to contain
three genera: Columellia, Menodora Humb. & Bonpl., and Bolivaria
Cham. & Schlechtd. (= Menodora Humb. & Bonpl.). He placed the
family between Oleinae and Jasmineaceae. Grisebach (1839) presumed
a close relationship with Gentianaceae. Meisner (1836-1843) favored the
affinity of Columelliaceae with Oleaceae. De Candolle (1839) assumed
a close relationship with Gesneriaceae. Adrien de Jussieu (1848) placed
Columelliaceae in Rubiales between Caprifoliaceae and Valerianaceae.
Lindley (1835) put Columelliaceae in his alliance (order) Cinchonales
(Rubiales) between Vacciniaceae and Cinchonaceae (Rubiaceae) with
which families and Onagraceae he thought it related. He also presumed
an affinity of Columelliaceae with Caprifoliaceae. Agardh (1858) sug-
gested a close affinity of the family with Lythraceae (“Lawsoniae”).
Basing his conclusions on the contorted anthers in both Columelliaceae
and Cucurbitaceae, Clarke (1858) asserted that, “... if the nearest
affinity of this family [Columelliaceae] is not with Cucurbitaceae, yet
there is no other to which it more closely approaches. . . .” Following
de Candolle, Bentham and Hooker (1876), and several of the more recent
taxonomists — Fritsch 1894, Engler 1892 (unchanged in Melchior’s 1964
edition of Engler’s “Syllabus der Pflanzenfamilien”), Schlechter 1920,
Wettstein 1935, and Pulle 1952 —placed Columelliaceae near Gesneri-
aceae. Fritsch emphasized the similarity with Bellonia L. (Gesneriaceae).
Nevertheless, Wettstein stressed the continuing uncertainty of the sys-
tematic position of Columelliaceae. Warburg (1922) placed Columelliaceae
near Gesneriaceae also; however, he noted: “Am natiirlichsten diirfte die
38 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Stellung bei den Rubiaceen sein.” In 1959, Takhtajan allied Columelliaceae
closely to Gesneriaceae, particularly with the genus Ramonda Rich.
Here, and in his 1966 work, he stated that Columelliaceae is a derivative
of Gesneriaceae. Hutchinson (1959) placed Columelliaceae in Personales
with the families Scrophulariaceae, Acanthaceae, Gesneriaceae, Orobanch-
aceae, and Lentibulariaceae. Airy Shaw (in Willis 1966) stated: “Despite
the sympetaly, slight zygomorphy and curious anthers [in Columelliaceae],
probably related to Escalloniac. and Hydrangeac.; perhaps also to Loga-
niac.” In his recent conservative treatment of Saxifragaceae, Thorne
(1968) treated Columelliaceae as a subfamily adjacent to Escallonioideae
and Montinioideae. Columelliaceae is placed in Rosales by Cronquist
(1968) near the Pittosporaceae and Grossulariaceae.
Anatomists have examined the microscopic structure of Columelliaceae
in an attempt to establish its affinities with more certainty. Solereder
(1899) was able to study the structure of Columellia oblonga Ruiz &
Pavon ssp. serrata (Rusby) Brizicky (= C. serrata Rusby) and concluded
that the occurrence of scalariform perforation plates and fibrous elements
with conspicuous bordered pits in the secondary xylem precluded any
close affinity with Gesneriaceae. Rather, he thought, Columelliaceae
showed anatomical similarities to Saxifragaceae. Van Tieghem (1903),
having several species of Columellia at his disposal, confirmed Solereder’s
anatomical observations, thus establishing the homogeneity of secondary
xylem structure throughout the genus. However, van Tieghem believed
Columelliaceae to be best placed in his alliance Rubiales near Rubiaceae.
Metcalfe and Chalk (1950), having no further material at their disposal,
repeated Solereder’s findings. Erdtman (1952) stated that pollen mor-
phology of Columelliaceae does not give any positive indications of the
affinity of the family. He does remark, however, that “The following
families have been mentioned as possibly related [to Columelliaceae]:
Ebenaceae, Ericaceae, Gesneriaceae (the grains oF Bellonia {Gesneri-
aceae| are not similar to those of Columellia!). .. .
Columellia, or Columelliaceae, has been considered related to families
of both Sympetalae and Choripetalae, to families with superior ovaries
and to others with inferior ovaries. Some proposed relatives have stipules
and others are exstipulate; some proposed relatives have opposite leaves
and others have alternate leaves; some proposed related families are
largely herbaceous and others are mostly woody. Among the taxa sug-
gested as relatives, the following seem to predominate: The first proposals
indicated Oleaceae; later the Ericaceae-Vaccinioideae and Rubiaceae were
recommended; Scrophulariaceae appeared a few times in the literature
during the early 19th century; but Gesneriaceae seemed most strongly
defended in the late 19th and early 20th centuries. Although alliance with
Saxifragaceae was suggested in the mid-19th century, it was not until the
early 20th century and later that the proposal seemed to gain strength.
Several other families have been proposed, though not as often as the
foregoing: Ebenaceae, Styracaceae, Gentianaceae, Loganiaceae, Capri-
1969 } STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 39
foliaceae, and Onagraceae. Today, both the gesneriaceous and saxifra-
gaceous hypotheses of relationship seem to have equal standing among
plant taxonomists, although the most recent treatments favor alignment
with saxifragaceous taxa. It is clear, though, that the variety of families
proposed as relatives of Columellia (Columelliaceae) could not be much
more diverse.
MATERIALS AND METHODS
In drawing comparisons between Columelliaceae and other families,
it has been necessary for convenience and clarity to accept certain taxo-
nomic delineations and judgements. This is especially important in refer-
ring to the Saxifragaceae which has been treated in different ways by
different authors. Engler’s (1928) treatment is the most detailed to date
and his concept of the family is very broad. He divides Saxifragaceae
into several subfamilies, namely, Penthoroideae, Saxifragoideae, Lepuro-
petaloideae, Parnassioideae, Tetracarpaeoideae, Pterostemonoideae, Iteoi-
deae, Brexioideae, Kirengeshomoideae, Kanioideae, Baueroideae, Hydran-
geoideae, Escallonioideae, Montinioideae, and Phyllonomoideae. Thorne’s
(1968) outline is very reminiscent of Engler’s treatment. In our paper,
when ‘“‘Saxifragaceae, sensu lato,” is employed, it is used in this broad
Englerian sense.
Other taxonomists have chosen to disassemble the Englerian conglom-
erate into several smaller families; hence, Hutchinson (1967) treated
Engler’s subfamily Escallonioideae as the family Escalloniaceae and his
subfamily Hydrangeoideae as the family Hydrangeaceae. Engler’s tribe
Philadelpheae of Hydrangeoideae is considered as Philadelphaceae by
Hutchinson. The genus Rides L. is part of the subfamily Saxifragoideae
in Engler but Hutchinson treated it as the basis of the monogeneric fam-
ily, Grossulariaceae. Cronquist (1968), similarly, has dissected Engler’s
Saxifragaceae. Because our comparisons among the vegetative parts of
plants depend heavily on the information in Metcalfe and Chalk (1950),
we have used their taxonomic designations for the Englerian subfamilies.
The concept of Saxifragaceae employed by these two plant anatomists is
wholly herbaceous, and the woody taxa in Engler’s Saxifragaceae are
relegated to other families, e.g., Escalloniaceae, Grossulariaceae, and
Hydrangeaceae (including Hutchinson’s Philadelphaceae). “Saxifrag-
aceae, sensu stricto,’ as we have used it, refers to a strictly herbaceous
family conforming to the sense of Metcalfe and Chalk.
Terminology used in the descriptions of xylem anatomy follows that
prescribed by the Committee on Nomenclature of the International Asso-
ciation of Wood Anatomists (1957). Other terminology used in descrip-
tions of anatomical structures is that in current use and deviations from
common usage are explained where they occur.
TABLE | contains a detailed listing of specimens employed in the study
of the vegetative anatomy of Columellia; materials used for comparative
40 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
floral anatomy are cited in the text. Fluid-preserved material of about
30 flowers of C. oblonga ssp. oblonga was available from one of Tovar’s
collections (4033, USM). All study specimens of wood, stems, leaves, and
flowers (except for comparative floral material of Escallonia and Car-
podetus), are supported by herbarium vouchers and their place of deposit
is noted in TABLE 1 or in the text.
Methods of preparing specimens for study followed standard laboratory
techniques. Woods were boiled in water to hydrate and stored in 70 per-
cent ethanol prior to microtoming. Transverse, radial, and tangential
sections of wood were stained with Heidenhain’s iron-alum haematoxylin
and counter-stained with safranin. Macerations of wood were prepared
using Jeffrey’s fluid. Clearing of leaves was carried out using Arnott’s
(1959) method involving 5 percent NaOH followed by a saturated aqueous
solution of chloral hydrate. After washing in water, leaves were stained
in aqueous safranin to accentuate vascular detail, dehydrated, and mounted
on glass slides in Canada balsam. Transverse and paradermal sections of
leaves were also prepared after embedding in paraffin, These were stained
in Heidenhain’s iron-alum haematoxylin and safranin. Nodal and petiolar
anatomy were studied from hand-cut sections treated with phloroglucinol
and concentrated HCl to differentiate the lignified tissues. Observations
of floral anatomy were performed from serial microtome sections (trans-
verse and longitudinal), cleared thick sections, and cleared whole flowers
of Columellia oblonga ssp. oblonga. These preparations were made using
familiar microtechnical methods from flowers fixed in formalin-acetic acid-
alcohol.
ANATOMY
The flower
Transverse sections through the base of the Columellia gynoecium show
two locules separated by a thick septum (Fic. 1, d, d'). In successively
more distal sections the placentas appear first as single lobes on each side
of the septum (Fic. 1, e; Fic. 3), then as deeply two-lobed structures
bearing many unitegmic ovules (Fic. 1, f). In still more distal sections
there is an opening between the locules (Fic. 1, g, h), but the uppermost
level of the ovary may again be divided by a complete septum (Fic. 2)
through which the stylar canal enters the ovarian cavity.
If the stylar canal is followed distally its appearance in transverse sec-
tion changes from that of a single cavity to that of a pair of tracts filled
with pollen-transmitting tissue (Fic. 1, j, k). The pollen-transmitting
tracts expand greatly below the two-lobed stigmatic surface, producing
the unusual transectional effect shown in Fic. 4. The outer layers of
gynoecial tissue, from the stylar base to the corolla, constitute a nectary
of small cells with densely staining cytoplasm (Fic. 2).
Flowers of Columellia are devoid of unusual histologic features that
can be used as taxonomic markers. The hypanthium, like the foliage, is
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 41
Fic. 1. Columellia oblonga, flower. Camera lucida drawings of selected trans-
verse sections, arranged sequentially from pedicel (a) to upper part of flower (k).
D, dorsal carpel bundles; S, stamen supply.
Fics. 2 and 3. Columelia aban: “ey in transverse section. Fic. 2. “Upper
(free) part of gynoecium, showing nectary and upper ovarian septum, X 30.
Fic. 3. Lower part taf ower showing basal septum, arrangement of vascular
bundles (cf. Fic. le); arrows indicate bundles supplying the 2 stamens,
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 43
covered with simple, appressed trichomes. Floral tissues contain no con-
spicuous tannin cells or sclereids and no crystal inclusions except for a
few scattered druses. The anthers dehisce with the aid of the familiar
subepidermal banded layer (Fics. 6, 7); moreover, the sporogenous por-
tions, in spite of their peculiar external form, resemble in section the
corresponding parts of ordinary four-locular anthers. The anther sacs,
at least the young ones, are minutely glandular-hairy at the margins, the
glandular trichomes being more or less club-shaped. The gynoecium con-
tains a well-marked endocarp tissue, four to six cells deep on the dorsal
side of the locule, gradually decreasing in thickness in the vicinity of the
septum and the placentas. Cell walls of the endocarp are neither lignified
nor greatly thickened in newly opened flowers, and there is no anatomical
indication of a dehiscence line at this stage.
Floral vascular bundles, many of them amphicribral, diverge from a
continuous cylinder in the pedicel (Fic. 1, a, b). Well below the base of
the locules, the cylinder expands into the pattern shown in Fic. 1, c, with
an inner portion of the vascular tissue directed to the septum and the
placentas and an outer portion directed to other parts of the flower. A
few sections above this level, and still below the locules, the outer portion
separates into two series of traces, a gynoecial series and a series supply-
ing perianth and stamens. With additional branching at even higher
levels (Fic. 1, d, d?, e), the gynoecial series contains as many as 20
bundles per carpel, and the other series (now outermost) contains about
a dozen perianth traces plus two stamen traces. A stamen trace can be
united for part of its length with the basal extension of a sepal midvein
or it can be completely free of other bundles to the base of the flower. In
either case, the position of the stamen traces is the same; they occupy
roughly the same radius as the septum. The perianth traces, if followed
distally, become the major veins of sepals and corolla lobes. As in many
other kinds of flowers, there are lateral connections between these strands
at the level where the calyx and corolla become free of the ovary wall,
and minor strands diverge from the major ones within the perianth mem-
bers. The vascular tissue of the stamen broadens within the filament
(Fic. 1, k) and terminates in the connective with a great many short
branches.
The vascular supply to the placentas rises through the septum in a
massive and irregular column or plexus (Fic. 1, c-f). Branches to the
ovules diverge from the plexus all through the placental region, but this
Portion of the vascular system does not continue above the placentas.
The many outer gynoecial bundles, however, extend all the way to the
base of the style (Fic. 1, g-i). Although the dorsal bundle is not easily
distinguishable in sections through the lower half of the ovary, it is con-
spicuous in higher sections because of its proximity to the locule (Fic.
1, g,h). The dorsal bundle can be followed into the style, which it enters
as a single well-defined strand. About a third of the way up the style,
it divides into two or more strands, which subdivide further into many
[voL. 50
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4-7. Fic. 4. Columellia oblonga,
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STERN, BRIZICKY, & EYDE, COLUMELLIACEAE
ower. Fic. 8. Placental region, 25. Fic.
x ee below placenta to show basal septum and ventral bundles (arrows),
9. Another flower,
48 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
strands just below the stigma. The remaining bundles of the gynoecium
wall converge upon the dorsal at the base of the style (Fic. 1, i); how-
ever, they do not appear to merge with the dorsal, because it enters the
style with its cross-sectional shape and dimensions unchanged.
The leaf and the node
Hairs on leaves of Columellia are thick walled and simple, tapering to
the obtuse tip and slightly swollen or bulbous at the base (Fics. 10, 11).
Trichomes emanate from the center of saucer-shaped depressions in the
lower epidermis. These are formed from several radially oriented cells each
of which is thicker toward the periphery of the depression and thinner
toward the center where the hair arises (Fics. 10, 11).
The cuticle is thick and covers upper and lower epidermis. It is espe
cially pronounced toward leaf margins and in the trichome-base depres-
sions of the lower epidermis. It also covers all portions of hairs. The cuticle
is strongly modified in the stomatal region; it covers the exposed surfaces
of guard cells and it over-arches both the outer portion of the aperture
producing a front cavity and the inner portion producing a back cavity
(Fics. 11, 12).
Stomata are restricted to the lower epidermis. The stomatal apparatus 3
is anomocytic (sensu Metcalfe and Chalk, 1950), ie., the guard cells are
surrounded by cells of varying number which are indistinguishable 1
form or position from the remainder of the epidermal cells (Fic. 10).
Guard cell walls are thickened along the inner surface facing the spongy
mesophyll and on the outer surface (Fics. 11, 12). In paradermal view,
guard cells are elongate-reniform (Fic. 10). ;
The lower epidermis is uniseriate; the upper epidermis is biseriate
(Fics. 11, 13, 15). Since developmental studies could not be conducted,
it is not possible to determine if the inner layer is protodermal in origin
or if it arose from the ground tissue. Inner cells of the biseriate uppeT
epidermis are larger and conspicuously more rotund than those of the
outer layer (Fics. 13, 15). Leaves are dorsiventral and the mesophyll
is divided into a biseriate, upper palisade layer and a lower spongy lay =
In the thickish leaves of Columellia lucida and C. obovata, the transition
between palisade and spongy mesophyll is not sharp. Furthermore, 19
these two species, there is a tendency for a lower palisade layer to
formed and an isobilateral condition (Fic. 15). :
Leaves of all species of Columellia are glandular; in those species having
serrate leaves, the tips of the teeth and the apical point are glandular,
in species with entire leaves, the apex of the leaf may be glandular.
* Although Metcalfe and Chalk (1950), Fahn (1967), and Esau (1965) do not
agree, the first author would prefer to use the term stoma (Gr. a mouth) in its re-
stricted sense to mean the actual aperture or pore in the epidermis which is sur-
stoma, guard cells, and subsidiary (accessory) cells, if present. The maintenance of
separate terms for the aperture and guard cells seems meritorious in that it provides
for independent reference to each of these units and alleviates the possible redundant
implications of referring to the “aperture of a stoma.”
showing biseriate upper epidermis, biseriate palisade layer, and uniseriate lower
€Pidermis. The cuticle overarches the unevenly thickened guard cells externally
and internally to form front and back cavities. Bases of hairs are situated in
Saucer-like depressions of the lower epidermis.
12-15. Fic. 12. Columellia lucida, Friedberg 240, transverse section of
lower epidermis of leaf showing thickened cuticle and ae cuticular
modification in association with stomatal apparatus, X 600. Fic. 13. C. oblonga
ssp. oblonga, Tovdr 4033, transverse section through mid-vein of leaf showing
biseriate upper epidermis, uniseriate lower epidermis, bun sheath, and bundle
sheath extensions, * 210. Fic. 14. C. oblonga ne be Tovér 3785, cleared
whole mount of leaf showing a single glandular ration; ark bodies in glan
are fruiting structures of an aspergillous rea < 40. Fic. 15. C. lucida,
Friedberg 240, transverse section through mid-vein of leaf showing biseriate
u Ppe epi is, uniseriate lower epidermis, bundle sheath and bundle sheath
r epider
tensions, tendency to development of a —e palisade layer, and abundance of
thick. walled fibers in the vascular bundle, 180.
tte
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 51
Fics. 16 and 17. Sectional series through petioles of Columellia, (a) being
= (c) proximal, showing increasing distal development of sclerenchyma.
. C. oblonga ssp. sericea, Drew E-113. Fic. 17. C. oblonga ssp. oblonga,
leer & Gilbert 1749. (x) xylem, (p) phloem, (s) sclerenchy ma.
Glands are highly vascularized and massive (Fic. 14); proximally adjacent
to the secretory epithelium is a cupulate reticulum of vascular elements.
That the central portion of the gland contains a cavity is borne out by the
occurrence there of aspergillous fruiting bodies in some specimens. Apices
of glands are aperturate probably through schizogeny.
Vasculation of the petiole is characterized by a single collateral strand
of conducting tissue varying from crescentiform to cupulate to almost
semiterete in transverse section (Fics. 16-20). Xylem is adaxial and
phloem is abaxial. In all species examined, an abaxial sclerenchymatous
region develops progressively from the proximal to the distal portion of
the petiole (Fics. 16-18). In specimens of Columellia oblonga ssp. ob-
longa (Fic. 17) and C. lucida (Fic. 19), a well-developed lunate layer
completely subtends the phloem at the extreme distal end of the petiole;
in specimens of other species (Fics. 16, 18, 20) the sclerenchyma seems
not to develop into more than a series of widely-spaced rods at this point.
However, sections through the mid-vein of the lamina in C. oblonga ssp.
52 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Fics. 18-20. Fic. 18. Columellia oblonga ssp. serrata, Bang 1172, sectional
series through petiole, ~ bey * Lege (c) proximal, showing increasing dista
development of scleren C. lucida, Friedberg 240, distal section
of collie. showing one “cerenchynatos arc. Fic. 20, C. obovata, Vargas
7693, distal section of petiole wing sclerenchyma as an arc of rods at this
point. (x) xylem, (p) phloem, () sclerenchyma.
sericea (Drew E-113), which shows a series of sclerenchymatous rods at
the distal end of the petiole (Fic. 16, a), show a complete sclerenchyma-
tous layer subtending the phloem. It is likely, therefore, that in the
laminae of all species of Columellia, the mid-vein is supported by an
abaxial layer of sclerenchyma. The central vascular strand of the petiole
branches into a series of minor strands toward the base of the lamina
(Fics. 16-20).
In Columellia oblonga the mid-vein of the lamina is characterized by
secondary growth and several layers of secondary xylem and phloem are
produced (Fic. 13). In C. lucida and C. obovata, secondary growth is not
pronounced; furthermore, in these species most of the xylem in the mid-
rib and secondary veins consists of thick-walled fibers (Fic. 15). Bundle
sheaths surround secondary veins in all species. Bundle sheath extensions
(Wylie, 1952) reach upper and lower epidermises in C. oblonga (Fic. 13);
in C. lucida and C. obovata, there are no bundle sheath extensions asso-
ciated with the bundle sheaths of secondary veins.
The node in Columellia is unilacunar and a single trace emerges through
each of the two opposite gaps in the vascular cylinder (Fic. 21).
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 53
F Transverse section of stem illustrating the unilacunar node in Colu-
mellia: “a) xylem, (p) phloem, (It) leaf trace.
The secondary xylem
The wood of Columellia is generally without growth rings, although in
the immature specimens of C. obovata, represented by Weberbauer 5482
and Nunez 3309, more or less sharply defined rings occur. However, both
of these specimens show strong evidence of decay or disease and it is sus-
pected that the growth rings are related to these conditions. All woods
examined are diffuse-porous, the strictly solitary, uniformly-sized pores
being distributed evenly across the transverse surface (Fic. 23). Vessel
walls are thin and there are no tyloses. Pores are angular.
Data for measurements of vessel diameter, vessel element length, bars
per scalariform perforation plate, tracheid length, and heights of vascular
rays are presented in TABLE 2. Because both mature and immature wood
were examined, measurements for each are separated in the table to
provide a more meaningful basis for comparisons with xylem in other taxa.
Vessel elements are generally long and narrow although ligules as such
are short and sometimes lacking. End wall angle ranges from 10° to 45°.
Perforation plates are entirely scalariform (Fic. 27) and in some cases
bars are so profusely branched they give the appearance of pits. Openings
in scalariform perforation plates are completely bordered. Spiral thicken-
ings occur in the cell walls of ligules throughout all species, being more
prominent in some than in others. In specimens of Columellia oblonga
ssp. oblonga, vaguely outlined spirals are seen in the body segment of
vessel elements and they are strongly marked in the ligules; in C. oblonga
54 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
(
a
RPS
2 ee
RK Soars
se 8 aN | ae
Owe
oe Be ot
Fics. 22-25. Fic. 22. Escallonia myrtilloides, Rimbach 13, Yw 16920, trans-
verse section of xylem showing solitary distribution of angular pores, 100.
Fic. 23. Columellia oblonga ssp. sericea, Rimbach 122, transverse section of
xylem showing solitary distribution of angular pores, and scanty vasicentric and
diffuse axial parenchyma, * 100. Fic. 24. E. myrtilloides, pranayese section of
xylemswith biseriate vascular rays and spiral thickenings in tracheids and vessels,
X 100. Fic. 25. C. oblonga ssp. sericea, oo section of xylem showing
uniseriate vascular rays and tracheids, « 1
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 55
Fics. 26-28. Fic. 26. poet myrtilloides, Rimbach 13, Yw 16920, radial
section of xylem showing scalariform perforation plates, x 150. Fic. 27. Colu-
mellia oblonga ssp. sericea, Rimback 122, hi dial section of xylem showing
scalariform perforation plates, x 100. Fic. c. degegens W eberbauer 5482
longitudinal section of xylem showing spiral Sry toni in \ 500
56 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ssp. sericea, spirals are tenuous at best and appear only in ligular portions;
in C, cida spirals occur only in ligules; and in C. obovata ee are
peach throughout the lengths of vessel elements (Fic.
Intervascular pitting is generally absent owing to the nee nature
of vessels; however, a suggestion of intervascular pitting is sometimes
present in the overlapping ends of superposed vessel elements. In these
areas, the circular to elongate pits are sparse and irregularly scattered but
there is a tendency toward the alternate arrangement.
Imperforate tracheary elements are tracheids, the pits in these cells
being of the same order of magnitude as those which occur in the over-
lapping ligulate portions of vessel elements (Fic. 25). Pitting in tracheids
is ordinarily uniseriate; less commonly two rows of pits are present, stag-
gered alternately. Inner apertures of pits are elliptical, crossed in face
view, and included within the pit border. Tracheid walls vary from very
thin to thick.
Vascular rays are entirely uniseriate and comprise axially elongated or
upright cells only (Fic. 25). These rays are homocellular and the ray tis-
sue corresponds with Kribs’ (1935) Heterogeneous Type III.
TABLE 2.
Summary of Xylem Anatomical Measurements in Columelliaceae
Marovre * IMMATURE ”
VESSEL DIAMETER IN p
Average: 45 25
Range: 22-105 12-45
MFR *°: 30-70 15-36
VESSEL ELEMENT LENGTH IN pb
verage:
Range: 308-1100
MFR: 375-828
BARS PER SCALARIFORM PERFORATION PLATE
Average: 14 10
Range: 7-20 3-24
: 11-16 6-17
TRACHEID LENGTH IN p
Average: 866
Range: 378-1260
MFR: 625-1110
HEIGHT OF VASCULAR RAYs IN CELLS
Range: 1-6 1-47+
* Columellia oblonga : oblonga, Tovdr 4033, Wurdack 1732. C. oblonga ssp.
sericea, Rimbach 122 and 3
olumellia oblonga ii ski Psi & Pavén 1/52; Weberbauer 5584 and
7791. C. lucida, André K— 1444 and 4500. C. obovata, Weberbauer 5482, Herrera 3451.
Data from Nufez 3309, a d saat are not included here.
° MFR = Most frequent range.
1969} STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 57
Axial xylem parenchyma is largely scanty vasicentric, a few isolated
strands occurring about the vessels (Fic. 23). In addition a few strands
were seen embedded within the groundmass of tracheids (diffuse paren-
chyma).
DISCUSSION
In view of the sympetalous corolla of Columellia, it is not surprising
that taxonomists looked for its relationships among the sympetalous
families, and especially those with inferior ovaries and opposite leaves.
Among other features, the androecial peculiarities of Columellia, un-
matched in any other known taxon, persuaded David Don to establish
a separate family for this unusual group of plants. Time has shown him
to have been correct in his assessment of the individuality of Columellia.
Evidence from gross morphology
There are such sharp differences in floral structure between Columel-
ated on an axile placenta in each locule, can hardly be regarded as closely
related to Columelliaceae. The mostly herbaceous Gentianaceae-Gentian-
oideae show some similarities with Columelliaceae in their cymose in-
florescences, in the structure of ovaries and fruits (2-carpellate ovaries
with numerous unitegmic, tenuinucellate ovules on parietal intrusive to
axile placentas, septicidal capsules, small seeds, etc.), as well as in the
possession of opposite, exstipulate leaves. They are markedly different,
however, in their regular flowers; in the usually contorted aestivation of
corolla lobes; in their usually dorsifixed, introrse anthers; and in their
superior ovaries. Loganiaceae (excluding Desfontainea Ruiz & Pavon)
differ from Columelliaceae in their usually stipulate leaves, regular flow-
ers, and superior ovaries; in addition, in the subfamily Buddleioideae,
the presence of glandular and stellate hairs is widespread. Some relation-
ship with Scrophulariaceae and especially Gesneriaceae appears possible,
but both families have highly specialized, mostly hypogynous flowers
(only Gesnereae of Gesneriaceae, sensu Fritsch 1893, 1894, have semi-
inferior or inferior ovaries). Some genera of Rubiaceae agree with Col-
umelliaceae in floral structure (except for the non-reduced number of
stamens) and opposite leaves, but they differ in the presence of stipules.
A close relationship with Caprifoliaceae seems equally doubtful. The
only genera of this family which are perhaps comparable with Columel-
liaceae in possessing multi-ovulate, 2-carpellate ovaries, are Diervilla Mill.
and Weigela Thunb., genera apparently restricted to the temperate zones
of North America and eastern Asia. The gross-morphological similarities
between Lythraceae and Onagraceae and Columelliaceae are too scarce
even to suggest a relationship. Within the saxifragaceous families —
58 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Hydrangeaceae, Grossulariaceae, and Escalloniaceae— almost all the
gross-morphological characters in Columellia may be found: frutescent
and/or arborescent habit; opposite, ba os often glandular-dentate
leaves; 5-merous haplostemonous flowers (Escalloniaceae); sympetalous
corolla (Roussea of Escalloniaceae) ; a semi-inferior or inferior, 2-car-
pellate ovaries with parietal intruding placentas bearing numerous, ana-
tropous and apotropous, unitegmic, tenuinucellate ovules (Escalloni-
aceae and some genera of Hydrangeaceae). Septicidal capsules usually
have numerous small endosperm-containing seeds (Hydrangeaceae and
Escalloniaceae) with small embryos. Most of the features of Columellia
are represented in the family Escalloniaceae. Although alternate leaves
predominate in this family, opposite leaves are found in the genera
Grevea Baill., Roussea, and Polyosma Blume. Other genera, as Valdivia
Remy, have subopposite leaves.
Evidence from floral anatomy
It would not be practical, nor is it necessary, to attempt a detailed
anatomical comparison of the Columellia flower to flowers of all the plant
families with which Columellia has been allied. A brief commentary on
the Cucurbitaceae seems to be in order, however, because androecial
structure in that family has significance for the interpretation of the an-
droecium in Columellia.
Clarke (1858) considered the stamens of Columellia, because of their
contorted anthers, to be almost identical to those of many Cucurbitaceae.
He interpreted the androecia of certain cucurbits — those with three ap-
. pendages, one two-locular and two four-locular — as comprising two and
a half stamens, an opinion shared by some other 19th century botanists.
If this view were correct, the two-staminate androecium of Columellia
would not seem greatly different. In more recent times, however, an
alternative interpretation of such cucurbitaceous androecia has been con-
firmed again and again; that is, the two-sporangiate stamen is an entire
one, and the four- -sporangiate stamens are duplex appendages. Evidence
for the more modern view is now overwhelming. It is derived from on-
togeny; from vascular anatomy (the duplex stamens sometimes contain
two well-defined bundles that are derived from two different petal traces) ;
and from comparative studies of male, female, and bisexual flowers 0
many genera, some of them exhibiting transitional stages between the
five-staminate condition and the “two and a half’’-staminate condition.
Reviews of the evidence are given by Miller (1929) and McLean (1947)
and additional confirmation by Bhattacharjya (1954), Chakravarty
(1958), and Quang (1963).
Although Columellia stamens are superficially similar to the duplex
stamens of Cucurbitaceae, the vascular supply in Columellia is a solitary
bundle. In transverse sections through the filament or the connective, the
bundle is often very broad and may occasionally seem to have two xylem
patches, but its appearance within the inferior part of the flower gives no
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 59
hint of compound structure. Observing this, van Tieghem (1903) con-
cluded that the two members of the Columellia androecium are solitary
stamens, and most floral morphologists would accept his evidence. Thus,
it can be argued rather convincingly that the evolutionary modification
leading to the two-staminate condition in Columellia was a loss or “abor-
tion” of stamens rather than any sort of phylogenetic union of stamens.
The occasional occurrence of a third stamen in flowers of Columellia, re-
ported by Brizicky (1961), supports this argument.
This reasoning might be thought to favor the relationship of C olumellia
other gesneriads there are only two stamens. In the latter case, as in
Columellia, there are no staminodes. But the resemblance of Columellia
to the gesneriads is not so close as this information would suggest, for in
Gesneriaceae only genera with superior ovaries have the two-staminate
androecium (Fritsch 1893, 1894).
A satisfactory anatomical comparison of the Columellia flower with
gesneriaceous flowers is not yet possible because floral anatomy of the
Gesneriaceae has never been investigated to any great extent. Compara-
tive information is presently available only for flowers of a Kohleria hybrid,
K. amabilis & K. scladotydea (Teeri, 1968), and for those of Kohleria
elegans (Dcne.) Loes. (H. E. Moore 8190; US, BH). Serial sections of
the latter were prepared from fluid-preserved material especially for this
paper. Anatomically, flowers of the two gesneriads are much alike, and
they have several characters in common with Columellia. For instance,
the floral tissues are devoid of tannins, and general features of placentation
and vasculation do not differ greatly from those of Columellia. In addi-
tion, both gesneriads have two-lobed placentas and many gynoecial strands
(Fic. 8). On the other hand, there are differences in detail that may be
important. The two gesneriads have no well-developed endocarp tissue,
except for a single layer of transversely elongate cells adjoining the locule.
The style is hollow for all of its length, with a single canal (Fic. 5).
Floral trichomes are multicellular (but uniseriate). Vascular traces to the
stamens are united with sepal traces for part of their passage through the
inferior part of the flower, and the supply to the placentas is derived from
two large septal bundles (duplex bundles representing paired hetero-
carpous ventrals; Fic. 9) in the septum. Of course, a major floral dif-
ference is that the anthers of the gesneriads are not contorted. Perhaps
the most important difference aside from that is in the nectary: nectaries
of Gesneriaceae are usually very well developed and deeply lobed or even
divided into distinct appendages (Feldhofen 1933).
It is somewhat easier to compare flowers of Columellia with those of
Rubiaceae because a detailed survey of floral anatomy in Rubiaceae is
available (Rao, Ramarethinam, & Iyer 1964). Rubiaceae is a large fam-
ily, rather diverse in floral structure; therefore, it is almost to be expected
that some of the members would have characters in common with Col-
60 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
umellia. For instance, some Rubiaceae have separate vascular traces to
calyx, corolla, androecium, and gynoecium. And in some genera (e &
Guettarda L. ) there are a great many gynoecial bundles. Piece is
often similar to that of Columellia, and many genera have an epigynous
nectary resembling that of C olumellia. A difference that strikes one im-
mediately, when sectioned flowers of Columellia are compared with sec-
tions of rubiaceous flowers, is the absence of conspicuous tannins in the
former. Floral tannins are rarely lacking in Rubiaceae. Another differ-
ence is that a single stylar canal seems to be of universal occurrence in
the Rubiaceae. Furthermore, the peculiar androecial modification in
Columellia has no counterpart among the rubiads.
Floral anatomy of the more easily obtained members of Saxifragaceae,
sensu lato, is fairly well known through the investigations of many work-
ers, including Palmatier (1943), Morf Shee Dravitski (see Philipson,
1967), Gelius (1967), and Komar (1967). None of these studies has
produced evidence to support Hallier’s (1908, 1910) opinion that Col-
umellia belongs with the Philadelpheae. In fact, ontogenetic observations
on Philadelphus (Gelius, 1967) suggest that evolution has favored an
increase in stamen number in this group. In some other genera of Phila-
delpheae, a reduction in the number of ovules has led to forms that bear
little resemblance to Columellia (e.g., Jamesia Torr. & Gray, Whipplea
Torr.). Schnizlein (1843-1870) proposed Brexia and Roussea as close
allies of Columellia; however both Brexia and Roussea have superior ova-
ries with distinctly two-ranked ovules. Argophyllum, another genus men-
tioned by Schnizlein, is also very dissimilar to Columellia, for it has
peculiar corolline ligules and T-shaped trichomes like its ally Corokia A.
Cunn. (Eyde, 1966). If the relationships of Columellia are to be sought
among the escallonioids, attention should be given to genera other than
the aberrant Argophyllum and Corokia. Berenice Tul. can also be elim-
inated from consideration, because it has recently been transferred to
Campanulaceae on anatomical and palynological grounds (Erdtman &
Metcalfe, 1963). From the standpoint of floral anatomy, Escallonia Mutis
ex L. f. is not as close to Columellia as might be indicated by other evi-
dence. Tannins are abundant in floral tissues of Escallonia species and
the floral trichomes frequently are multicellular with globular terminal
portions; also, the gynoecial bundles are few and the ventral bundles
commonly accompany the dorsals into the style. Choristylis Harv. has
stamens united with corolla tube, but in other respects the flowers are
unlike those of Columellia. One difference is that the gynoecial bundles
are few; another is that the nectary is located on the lower part of the
corolla tube. The latter character may be sufficiently important to remove
Choristylis from its position adjoining Forgesia (Engler, 1928) and to
place it elsewhere in the Saxifragaceae, sensu lato. (Agababyan 1964,
links Choristylis with Itea on palynological evidence.) Flowers of F orgesia
have rather massive multicellular trichomes; otherwise they are anatomi-
cally similar to Columellia flowers. To judge from our one sectioned her-
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 61
barium flower, the gynoecial vasculature and the placentation approximate
those of Columellia. The nectary, if there is one (it is not easy to tell
from dried material), is part of the free portion of the gynoecium, and
the androecium shows indications of reduction (abortive locules in some
anthers). Forgesia, like many other Saxifragaceae, sensu lato, has free
styles that could be viewed as a precursor to the two-canal structure of
the Columellia style.*
In summarizing this section on floral anatomy, it must be conceded
that the cited points of similarity and dissimilarity do not tell us much
about the affinities of Columellia. The foregoing commentary includes no
strong evidence against the proposed relationship with Gesneriaceae, nor
does it include really firm evidence for such a relationship. The same can
be said of the possible alliance with Rubiaceae or with the escallonioid
group of Saxifragaceae, sensu lato. The reason for this is clear.
observed characters in the flowers of Columellia are widely distributed in
many plant families, except for the contorted anthers. Ironically, the
latter character does not help in placing Columellia because it has not
been found in any other group of plants, the resemblance to anthers of
certain cucurbits being demonstrably superficial.
Evidence from leaf anatomy
It does not appear possible to compare all features of the foliar anatomy
of Columelliaceae with those of families reputed to be allied to it, since
complete foliar surveys of these families are lacking from the literature.
An original study of leaves in all these families is surely outside the scope
of this investigation. Nevertheless, certain comparisons can be made.*
Leaves are dorsiventral in Gesneriaceae. Hairs are always multicellular
and they are often situated on a pedestal. They may be glandular or
non-glandular. A multiseriate hypodermis occurs in certain species. The
stomatal apparatus is often very large and anisocytic. Vascular bundles
in veins are not usually accompanied by sclerenchyma. Vasculation of
the petiole is various and many genera show a single leaf trace; Alloplec-
tus Mart., Besleria L., Episcia Mart., and others have three leaf traces
and Klugia notoniana A. DC. shows a large number of separate strands.
There is no “‘pericyclic’” sclerenchyma associated with the petiolar vascular
strand in Gesneriaceae. Gesneriaceous leaves differ markedly from those in
Columelliaceae in their multicellular and glandular hairs, anisocytic sto-
matal apparatus, and lack of sclerenchyma associated with vascular tissue.
* Observations on the floral anatomy of Escalloniaceae are based on serial sections
prepared especially for this paper. Material was obtained from the following sources:
Escallonia, fluid-preserved flowers from several cultivars growing in the Los Angeles
State and County Arboretum, not vouchered; Carpodetus serratus, fluid-preserved
owers from plants cultivated at the University of Auckland, New
vouchered ; Quintinia fawkneri, pressed flowers, Brass 4719, US; Choristylis
shirensis, pressed flowers, Swynnerton 607, US; Forgesia borbonica
de V'Isle 216, US. ? > g ca, pressed flowers,
amily circumscriptions follow those used by Metcalfe and Chalk (1950) for
convenience in making comparisons.
62 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Rubiaceous leaves are generally dorsiventral; centric and homogeneous -
leaf organization occur in a few species. Hairs may be unicellular, multi-
cellular and uniseriate, tufted, and rarely peltate. A hypodermis occurs in
many species. The stomatal apparatus is _paracytic (rubiaceous ) in most
species, as might be expected. The petiolar vascular strand is usually
shield shaped with more or less coe Se wings. There are also
variously shaped median vascular strands, nearly always associated with
smaller accessory bundles toward the wings. In such a large and anatomi-
cally diverse family as Rubiaceae, it is not surprising to find foliar re-
semblances to Columelliaceae. The only clear and consistent difference
is the more or less ubiquitous occurrence of the paracytic stomatal appa-
ratus in Rubiaceae.
Caprifoliaceae usually have dorsiventral leaves, but the palisade tissue
is poorly developed in species of Triosteum L. and Viburnum L. Hairs
may be glandular or non-glandular and unicellular, simple and multi-
seriate, tufted or stellate, and peltate. Glandular leaf teeth are present
in some species. Stomatal organization is frequently anomocytic, but
paracytic types occur in the same genera as anomocytic types. Except
for Diervilla, a single layer of palisade mesophyll occurs; in Sambucus
L. and Viburnum, cells of the palisade layer may have arms. The petiolar
vasculation shows a considerable range of structure from a solitary, slightly
crescentic bundle to an arc of 3—5 or more separate bundles to a closed
vascular cylinder. The anomocytic stomatal apparatus and solitary petl-
olar strand in Caprifoliaceae are similar to Columelliaceae, but the para-
cytic stomatal apparatus, single-layered palisade mesophyll, and multi-
strand and cylindrical vasculation of the petiole, which also occur in some
species of Caprifoliaceae, are very different from the situation in Columel-
liaceae.
Leaves in Saxifragaceae, sensu stricto, are dorsiventral and isobilateral.
Hairs are glandular and non-glandular and these may be simple, uniseriate
and multicellular; shaggy; and multiseriate. Stomatal organization 1S
anomocytic and sometimes subsidiary cells, smaller than neighboring
epidermal cells, are evident. The mesophyll in some species of Saxifraga
L. is undifferentiated, and in species where it is differentiated, the palisade
segment may range from 1 to 7 cells deep. Hydathodes are of common
occurrence. Petiolar vasculation is distinctive, especially in Saxifraga
where one concentric bundle or one hemi-concentric bundle may occur.
In other species of Saxifraga, there are three such bundles, each with its
own endodermis. Some Saxifragaceae have the usual collateral bundles.
but these may be scattered. The herbaceous Saxifragaceae resemble Col-
umelliaceae in the presence of an anomocytic stomatal apparatus, appar-
ently modified in some taxa; but the undifferentiated mesophyll in some
species of Saxifraga and multilayered palisade tissues in others, are very
different from the condition in Columellia. Petiolar vasculation in Saxi-
fragaceae bears little resemblance to that in Columelliaceae.
Leaves in Grossulariaceae are dorsiventral and bear unicellular and
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 63
also glandular hairs. Pairs of small, circular guard cells are characteristic.
The petiole is characterized by three separate vascular strands at the base
which fuse distally to produce a single crescentiform strand all supported
by sclerenchyma in the “pericyclic” region. The specialized, small circu-
lar guard cells vary from those in Columelliaceae but the abaxial scleren-
chyma associated with the petiolar bundle also occurs in Columelliaceae.
The proximally triple vascular strand differs from the condition in Col-
umelliaceae.
All leaves in Escalloniaceae are dorsiventral. In Escallonia, foliar hairs
are thick-walled and unicellular; in Adbrophyllum Hook.f., hairs are
glandular with unicellular heads; in some species of Escallonia hairs are
glandular-shaggy with multiseriate stalks; in Quintinia A. DC. peltate
hairs occur; and T-shaped hairs occur in Argophyllum. Stomatal organi-
zation is variable and pairs of nearly circular, small guard cells, resem-
bling those in Grossulariaceae, occur in Escallonia, Itea L., and other
genera; the stomatal apparatus in Quintinia is paracytic; and the stomatal
apparatus in Brexia is characterized by a double front cavity. A 1—3-
layered upper hypodermis occurs in species of Argophyllum, Carpodetus,
Escallonia, and other genera. A single-layered palisade mesophyll is pres-
ent in two genera, Three vascular bundles enter the base of the petiole in
Escallonia, but in E. macrantha Wedd. (= E. polifolia Hook.) and E.
rubra (Ruiz & Pavon) Pers., a single crescentiform petiolar bundle with
accessory strands is present. Apparently there is no abaxial sclerenchyma
present in Escallonia. Brexia appears unique, for besides the abaxial,
crescentiform vascular strand in the petiole, there is also a small cylinder
of xylem in the medullary region and two abaxial xylem cylinders. Certain
similarities between Columelliaceae and Escalloniaceae occur: unicellular,
thick-walled hairs; presence of a hypodermis; and a single petiolar vascu-
lar strand in at least two species of Escallonia. However, there are also
marked differences and Escalloniaceae show glandular and multicellular
hairs, grossulariaceous stomatal organization, and a triple vascular condi-
tion in petioles of most species of Escallonia.
Hydrangeaceous leaves are dorsiventra]l. Hairs are various with long,
unicellular trichomes in Jamesia; tufted trichomes in Broussaisia Gaudich.
and Pileostegia Hook. f. & Thoms.; and stellate, calcified, and unicellular
Deutzia, and Philadelphus. A hypodermis occurs in Broussaisia and in
species of Hydrangea, and the epidermis contains some _ horizontally
divided cells in Carpenteria Torr. The stomatal organization is paracytic
in species of Dichroa Lour. and Hydrangea L. and anomocytic in Phila-
delphus. Palisade mesophyll is uniseriate in Deutzia and Philadelphus.
The petiolar vascular strand differs throughout the family: It is single
and crescent-shaped in species of Deutzia, Jamesia, Philadelphus, Hy-
drangea, and Pileostegia; petioles of Decumaria sinensis Oliv., Dichroa
febrifuga Lour., and Hydrangea petiolaris Sieb. & Zucc. are characterized
by a main abaxial arc with several flat adaxial bundles between the ends.
64 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Additional strands are present in other species, including medullary
bundles. Although some foliar similarities exist between some taxa of
Hydrangeaceae and Columelliaceae — unicellular hairs, glandular leaf
teeth, hypodermis, anomocytic stomatal organization, and arcuate petiolar
vascular supply —the differences are equally clear. Multicellular and
tufted hairs, paracytic stomatal organization, and multistranded petiolar
vascular supply occur in Hydrangeaceae.
The remaining plant families which have at one time or another been
suggested as near relatives of Columelliaceae or Columellia — Scrophu-
lariaceae, Ebenaceae, Loganiaceae, Oleaceae, Lythraceae, Vacciniaceae,
Ericaceae, Gentianaceae, and Onagraceae — present a wide array of foliar
anatomical features, some similar and others different from Columelliaceae.
As should be apparent from the brief comparative summary above, no
family presents a consistent foliar pattern which is similar enough in
most respects to that in Columelliaceae to convince the critical botanist
that leaf anatomy is a key to understanding the relationships of the fam-
ily. To be sure, this is probably related to the lack of thorough anatomi-
cal investigation in those taxa reputedly related to Columelliaceae, but
as the situation stands now, foliar anatomy is at its best equivocal in
pointing the way to the relationships of Columelliaceae.
Evidence from nodal anatomy
According to Sinnott’s (1914) survey of the nodal condition among
seed plants, all members of the Tubiflorae, which include Scrophulari-
aceae and Gesneriaceae, are unilacunar. However, three or five gaps are
typical for Cyrtandra J. R. & G. Forst. (Gesneriaceae). Onagraceae,
Ericaceae, Ebenaceae, Oleaceae, Gentianaceae, Loganiaceae, and Rubi-
aceae, are also characterized by unilacunar nodes. In addition, some mem-
bers of Gentianaceae are multilacunar and some Rubiaceae are trilacunar.
Caprifoliaceae are generally tri- and sometimes pentalacunar. Cucurbit-
aceae are all trilacunar. Rosales, which include Saxifragaceae (treated in
the broad Englerian sense by Sinnott), are said to be mostly trilacunar
although five gaps occur in Brunelliaceae and in a few Saxifragaceae,
Rosaceae, and Leguminosae. Platanaceae exhibit seven
Plant orders are remarkably constant with respect to their nodal con-
ditions but Sinnott recognized that nodal anatomy is only one character,
that nodal structure is not always invariable, and that further study will
necessitate changes in his outline. In 1955, Marsden and Bailey presented
their penetrating analysis of the node and interpretation of the primitive
nodal condition. In contrast to Sinnott’s hypothesis that the trilacunar
condition is basic and primitive, Marsden and Bailey provided evidence to
indicate that the unilacunar, two-trace condition is ancestral and they
indicated possible means for deriving both the unilacunar, single-trace
condition and the trilacunar, triple-trace condition directly from it.
Furthermore, they hypothesized that the unilacunar node could give rise
to the trilacunar node through amplification, much as Sinnott derived
the multilacunar form from the trilacunar.
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 65
Takhtajan’s (1964) scheme of nodal evolution is similar to that of
Sinnott in that he accepted the primitiveness of the trilacunar node.
However, the median lacuna has a double trace: ‘Thus, from all of these
data one can conclude, it seems to me, that the node with three or more
lacunae (Fic. 9) is the primary type of node in angiosperms. At present,
it is impossible to determine more accurately the initial nodal type in
angiosperms.’
ecause of the studies of Marsden and Bailey, it is apparent that a
reassessment of the taxonomic value of nodal anatomy, as exemplified by
Sinnott’s treatment, is very much in order. The derivation of the uni-
lacunar, one-trace condition in Columelliaceae, rather than the condition
itself, is the key to taxonomic understanding. This is also true of the
largely characterized by trilacunar nodes, nor can we assign the relation-
ship of Columelliaceae to those families with unilacunar nodes, if we
agree with Marsden and Bailey that, ‘Structures which appear to be
similar at the nodal level may not be truly homologous, and conversely
differences which seem outstanding at the nodal level may acquire a dif-
ferent significance where comprehensive developmental studies at succes-
sive levels of the stem and leaf are made.’
Evidence from xylem anatomy *
A brief recapitulation of the salient features in the xylem anatomy of
Columelliaceae is in order here: perforation plates scalariform; pore dis-
tribution exclusively solitary; intervascular pitting usually absent, except
tending to alternate in regions of ligular overlap between superposed ves-
sel elements; axial parenchyma vasicentric scanty; vascular rays exclu-
sively uniseriate consisting solely of upright cells; spiral thickenings present
in walls of vessel elements; and imperforate tracheary elements are tra-
cheids.
Gesneriaceae all have simple perforations in vessel elements. However,
vestigial bars were noted in perforation plates of Solenophora calycosa
Donn. Smith. In all woods examined, pores are solitary, in radial multiples,
and in clusters except in Solenophora sp. (Yw 22822) where no clusters
were observed. Intervascular pitting is exclusively alternate, i in
Solenophora calycosa where transitional pitting was also seen. Axial par-
enchyma distribution is various; however, it is paratracheal nase for
um DC.,
and Solenophora calycosa, in which diffuse parenchyma occurs. In most
species the vasicentric parenchyma is scanty; vasicentric parenchyma is
abundant, however, in Columnea purpurata Hanst., Cyrtandra oenobar-
° Anatomical eat presented in this section are based on original observations in
Gesneriaceae, Gro riaceae, Hydrangeaceae, and Escalloniaceae. Microscope slides
examined were ae the Yale (Yw) and Say seapeon Sw) woed collections. Data
pe g comparisons,
onv
families are considered as circumscribed in Metcalfe oi 1 Chalk (19
66 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
bata H. Mann, C. spathacea A. C. Smith, and Gesneria sp. (Yw 16832).
In Drymonia sp. (Yw 17724), aliform and aliform-confluent parenchyma
occurs. Vascular rays are absent in Besleria spp. (Yw 12217, 12225).
In Columnea purpurata rays are 1 to 3 cells wide and in Solenophora
calycosa, rays are mostly uni- and biseriate. In all other species investi-
gated, rays are multiseriate. Rays are homocellular consisting solely of
upright cells in Drymonia spectabilis, Columnea purpurata, Rhytidophyl-
lum crenulatum, R. tomentosum (L.) Mart., and Rhytidophyllum sp.
(Yw 20017). Heterocellular rays occur in Cyrtandra oenobarbata, Gs
spathacea, Gesneria sp. (Yw 16832), Drymonia sp. (Yw 17724), Soleno-
phora calycosa, and Solenophora sp. There are no spiral thickenings 1n
vessels of Gesneriaceae. Imperforate tracheary elements are various:
septate elements occur in Besleria spp., Gesneria sp., Drymonia spectabilis,
Columnea purpurata, Rhytidophyllum crenulatum, R. tomentosum, Soleno-
phora calycosa, and Solenophora sp. Only Cyrtandra did not show septate
imperforate tracheary elements. Drymonia spectabilis exhibits only fiber-
tracheids and Gesneria sp., Drymonia sp., Rhytidophyllum crenulatum,
and Solenophora sp. show only libriform wood fibers. All other species
investigated show both fiber-tracheids and libriform wood fibers.
Except for the common occurrence of vasicentric scanty axial paren-
chyma in Gesneriaceae and Columelliaceae, the wood anatomy of these
two families is very different. Perforation plates in Columelliaceae are
scalariform; in Gesneriaceae they are simple. Pore distribution is strictly
solitary in Columelliaceae; in Gesneriaceae it is solitary and in radial
multiples and clusters in most of the species studied. Intervascular pitting
is virtually absent in Columelliaceae because of the independent distribu-
tion of vessels; in Gesneriaceae it is alternate. All species of Columelli-
aceae have vascular rays; in Gesneriaceae, Besleria lacks vascular rays.
Vascular rays are uniseriate in Columelliaceae; in Gesneriaceae all species
have vascular rays more than one cell wide. Vascular rays contain only
upright cells in Columelliaceae; in Gesneriaceae species may show both
heterocellular rays and homocellular rays with upright cells. Spiral thick-
enings are present in the vessels of Columelliaceae; in Gesneriaceae, veS-
sels lack spiral thickenings. In Columelliaceae all imperforate tracheary
elements are tracheids; in Gesneriaceae both fiber-tracheids and libriform
wood fibers occur, but no tracheids.
Grossulariaceae have scalariform perforations in vessel elements, but
some simple perforations were also observed. Pores are solitary, in radial
multiples, and in clusters. Growth rings are pronounced and the wood is
ring porous. Intervascular pitting is transitional and scalariform. Axial
parenchyma is absent. Vascular rays are multiseriate, broad, and hetero-
cellular. Sheath cells are of common occurrence in the rays. Spiral thick-
enings are absent from vessel walls. Imperforate elements are septate
tracheids and in Ribes viscosissimum Pursh, fiber-tracheids were also
recorded.
The presence of scalariform perforations and tracheids seems to pro-
1969] STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 67
vide the only common anatomical features between Grossulariaceae and
Columelliaceae. Solitary, radial multiple, and clustered pores; ring poros-
ity; broad, heterocellular vascular rays; septate imperforate tracheary
elements; and the absence of axial parenchyma in the wood of Grossu-
lariaceae are rather distinct anatomical characteristics which differ from
Columelliaceae
Perforation plates in vessel elements of Hydrangeaceae are scalari-
form.® Pores are exclusively solitary in Broussaisia arguta Gaudich., B.
pellucida Gaudich., Fendlera rupicola A. Gray, and Philadelphus sp. (Yw
11845). In Deutzia vilmorinae Lemoine & D. Bois, Hydrangea pana-
mensis Standley, and Philadelphus coronarius L., pores are solitary and
in radial multiples. Hydrangea bretschneideri Dipp. and Dichroa febri-
fuga show pores in solitary, radial multiple, and clustered dispositions.
Intervascular pitting is generally absent in Broussaisia arguta, B. pel-
lucida, and Fendlera rupicola. However, in the overlapping vessel ligules
of Broussaisia arguta, scalariform pitting was seen, whereas in this posi-
tion Fendlera rupicola shows a tendency to alternate intervascular pitting.
In Philadelphus coronarius, intervascular pitting is transitional; in Phila-
delphus sp., pitting is alternate with some opposite. Pitting in vessel
walls of Deutzia vilmorinae, Hydrangea bretschneideri, and H. panamensis
is scalariform. Vessel walls in Dichroa febrifuga show both transitional
These occur in conjunction with other heterocellular rays, two or more
cells wide. Rays up to 8-cells wide occur in Broussaisia pellucida. Deutzia
vilmorinae, Fendlera rupicola, and Hydrangea bretschneideri have only
uni- and biseriate rays. Sheath cells are common in species with wide
rays. In Deutzia vilmorinae, scalariformly perforated ray cells occur.
Tenuous spiral thickenings occur in the cell walls of vessels and tracheids
of Fendlera rupicola; in Philadelphus coronarius and Philadelphus sp.,
spirals occur in tracheids. Imperforate tracheary elements in Fendlera
rupicola, Hydrangea bretschneideri, Philadelphus coronarius, and Phila-
delphus sp., are exclusively tracheids. In Broussaisia arguta, B. pellucida,
and Dichroa febrifuga, both tracheids and fiber-tracheids appear. Deutzia
vilmorinae and Hydrangea panamensis show only fiber-tracheids. Imper-
forate tracheary elements are septate in Hydrangea panamensis and
Dichroa febrifuga.
There are several similarities between the woods of some species of
Hydrangeaceae and Columelliaceae: scalariform perforation plates, ex-
clusively solitary pores and concomitant absence of intervascular pitting,
a tendency to alternate intervascular pitting, and tracheids. Axial xylem
parenchyma is vasicentric scanty in Columelliaceae with some diffuse; in
® Metcalfe and Chalk (1950) report simple perforation plates in Deutzia glabrata
Kom. and in some species of Philadelphus.
68 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
all Hydrangeaceae studied, where axial xylem parenchyma was present, it
it vasicentric scanty and some strands were diffusely arranged. On the
other hand, there are also pronounced anatomical differences between
these families: Pores are exclusively solitary in Columelliaceae; in several
species of Hydrangeaceae pores are in both solitary and other arrange-
ments. Intervascular pitting tends toward alternate in Columelliaceae;
in Hydrangeaceae scalariform and transitional intervascular pitting occur
in several species. All species of Columelliaceae show axial parenchyma;
several species of Hydrangeaceae lack this tissue. In Columelliaceae, vas-
cular rays are all uniseriate and homocellular; all Hydrangeaceae have
uniseriate rays plus rays which are two or more cells wide and hetero-
cellular. Imperforate tracheary elements in Columelliaceae are tracheids;
some species of Hydrangeaceae show both tracheids and fiber-tracheids,
while other species have only fiber-tracheids.
Perforation plates in Escalloniaceae are exclusively scalariform except
in Brexia, where plates are mostly simple, and in Kania Schlechter,“ where
they are exclusively simple. All Escalloniaceae have solitary pores, 10
Escallonia floribunda H.B.K., E. fonkii Phil., and E. myrtilloides Lif,
pores are exclusively solitary. In E. pulverulenta (Ruiz & Pavon) Pers.,
E. revoluta (Ruiz & Pavoén) Pers., E. rubra (Ruiz & Pavon) Pers., and
E. tortuosa H.B.K., pores are also in radial multiples. Pores are solitary
and in radial multiples in Brexia madagascariensis Thou. ex Ker-Gawl.,
Itea sp. (Yw 20142), Ouintinia acutifolia T. Kirk, Q. serrata A. Cunn., and
Q. sieberi A. DC. In Quintinia, however, multiples are rare but tangentially
oriented groups of pores are conspicuous. Solitary, radial multiple, and
clustered dispositions are seen in Anopterus glandulosus Labill., Argo-
phyllum ellipticum Labill., and in all Polyosma species studied. Inter-
vascular pitting is sparse in Quintinia acutifolia and Q. serrata; pitting 1n
Q. sieberi is alternate with a tendency to opposite. In those species of
Escallonia with exclusively solitary pore distribution, the widely overlap-
ping vessel ligules provide areas of intervascular communication showing
alternate intervascular pitting. Species of Escallonia with radial pore
multiples show alternate intervascular pitting. Alternate intervascular
pitting also occurs in Anopterus macleayanus F. Muell., Argophyllum el-
lipticum, Brexia madagascariensis, Itea sp., and in all Polyosma species
studied except P. integrifolia Blume and P. serrulata Blume which have
exclusively opposite pitting. In addition to alternate intervascular pit-
ting, Anopterus macleayanus and Escallonia floribunda show transitional
pitting. Anopterus glandulosus only has transitional intervascular pitting.
In addition to alternate pitting, /tea sp. shows transitional and scalariform
pitting. All species of Polyosma with alternate intervascular pitting also
show opposite pitting. Escalloniaceae are characterized by apotracheal
axial parenchyma and all species studied show either a diffuse and/or
*Erdtman and Metcalfe (1963) have assigned this genus to Myrtaceae on anatomi-
cal and palynological grounds. Their evidence is so strong, that Kania will not be
considered further in this discussion.
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 69
diffuse-in-aggregates pattern. In Escallonia revoluta, E. rubra, all species
of Polyosma, and Quintinia sieberi, both diffuse and diffuse-in-aggregates
patterns occur. In Anopterus glandulosus, Escallonia myrtilloides, E. tor-
tuosa, Itea sp., Quintinia acutifolia, and Q. serrata, only diffuse axial
parenchyma was observed. Parenchyma in Brexia madagascariensis con-
sists of multiseriate bands. Short uniseriate bands occur in Escallonia
floribunda, in addition to the diffuse-in-aggregates pattern. Axial paren-
chyma is absent in Argophyllum ellipticum. All species of Escalloniaceae
have some uniseriate rays, although none was observed in Anopterus
glandulosus where rays are exclusively multiseriate. All species have some
heterocellular rays except for Brexia madagascariensis. The following
species has uni- and biseriate rays only: Anopterus macleayanus, Brexia
madagascariensis, Escallonia myrtilloides, and E. tortuosa. All other spe-
cies studied have both uniseriate rays and rays which are two or more
cells wide. Vascular rays are exclusively heterocellular in Anopterus glan-
dulosus, A. macleayanus, Argophyllum ellipticum, and Escallonia flori-
bunda. Rays in Brexia madagascariensis are homocellular and cells are
upright. In the following species, multiseriate and biseriate rays are
heterocellular and uniseriate rays are homocellular containing only upright
cells: Escallonia fonkii, E. myrtilloides, E. pulverulenta, E. revoluta, E.
rubra, E. tortuosa, Itea sp., and all species of Polyosma and Quintinia.
Species with wide multiseriate rays commonly exhibit sheath cells. Spiral
thickenings occur in walls of vessels in Escallonia floribunda, E. myrtil-
loides, E. rubra, and E. tortuosa. In E. myrtilloides and E. tortuosa, spiral
thickenings also occur in tracheid walls. Only tracheids occur in Anopterus
glandulosus, Escallonia floribunda, E. myrtilloides, E. revoluta, E. rubra,
E. tortuosa, Polyosma cunninghamii Benn., and Quintinia. Both tracheids
and fiber- tracheids occur in Anopterus macleayanus, Escallonia pulver-
ulenta, and Itea sp. Argophyllum ellipticum, Brexia madagascariensis,
Escallonia fonkii, Polyosma cambodiana Gagn. (?), P. ilicifolia Blume, P.
integrifolia, P. mutabilis Blume, and P. serrulata, exhibit only fiber-tra-
cheids. Septate fiber-tracheids appear in Argophyllum ellipticum.
The xylem anatomical similarities between species of Escalloniaceae
and Columelliaceae are striking: exclusively scalariform perforation plates
(except in Brexia), exclusively solitary pore distribution (in some species
of Escallonia and in Polyosma cunninghamii), spiral thickenings in vessels
(in some species of Escallonia), and exclusively tracheids (in Anopterus
glandulosus, in some species of Escallonia, Polyosma cunninghamii, and
Quintinia). The only major anatomical differences between these two
families are the presence of vascular rays which are two or more cells
wide and exclusively apotracheal axial parenchyma in all Escalloniaceae.
Among the species studied, the xylem anatomy of Escallonia myrtilloides
can hardly be distinguished from that of Columelliaceae, except for the
biseriate condition of some of the rays and exclusively diffuse axial paren-
chyma in the former (cf. Fics. 22 and 23, 24 and 25, 26 and 27).
Among the remaining families which have been suggested as close rela-
tives of Columelliaceae — Ebenaceae, Styracaceae, Gentianaceae, Logan-
70 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
iaceae, Caprifoliaceae, Rubiaceae, Onagraceae, Oleaceae, Vacciniaceae,
and Scrophulariaceae — xylem anatomy provides serious bases for compar!-
son only with Styracaceae, Caprifoliaceae, and Vacciniaceae. Styracaceae
typically show scalariform perforation plates and uniseriate, homocellular
vascular rays in some species; some Caprifoliaceae have scalariform and
simple perforation plates, spiral thickenings in vessels, and imperforate
tracheary elements with distinctly bordered pits; and most Vacciniaceae
have scalariform or scalariform and simple perforations and imperforate
tracheary elements with distinctly bordered pits. Ebenaceae, Loganiaceae,
Rubiaceae, Onagraceae, Oleaceae, and Scrophulariaceae, are characterized
by simple perforations. In addition, Gentianaceae-Gentianoideae univer-
sally possess internal phloem and medullary vascular bundles; Logania-
ceae-Loganioideae are characterized by included phloem; and Onagraceae
have internal phloem in the axis and a few genera show included phloem.
These dispositions of phloem are very specialized and are ordinarily in-
dicative of close relationship within specific taxa, sometimes on an ordinal
basis (e.g., internal phloem in families of Myrtales).
CONCLUSION
In reviewing the foregoing presentations of evidence and discussions, it
is clearly impossible to assemble an array of data from each form of evl-
dence presented — gross morphology, floral anatomy, foliar anatomy, nO-
dal anatomy, and xylem anatomy — which would affirm unequivocally
the relationships of Columelliaceae with any one of the several families
to which it has been allied. The similarities in gross morphology of flow-
ers and fruits among many families of various alliances probably indi-
cates parallel evolution rather than close genetic relationship. The evo-
lutionary development which has culminated in Columellia has proceeded
in such a manner that the complex of its characteristics is different from
any known taxon today. What baffles us now baffled our predecessors
and it is time to admit once and for all that Columelliaceae is a unique
plant family, probably with no really close living relatives. The clearest
line of evidence for the possible relationships of Columelliaceae is pro-
vided by xylem anatomy and it appears not too far from reality to assert
that this family belongs in the great saxifragaceous assemblage with the
Escalloniaceae, Hydrangeaceae, and Grossulariaceae. Data from gross
morphology, floral anatomy, palynology, etc., at least do not contradict
this probability. Perhaps its nearest relatives are in the Escalloniaceae.
If there was a common saxifragaceous ancestor, phylogenetic departure
must have occurred long ago, for transitional forms seem to have been
lost in the development of the modern plants of which this taxon is com-
posed. Evidence from xylem anatomy seems equally persuasive in negat-
ing an alliance with any other family or group of families. Unfortunately,
data from cytotaxonomy, embryology, and biochemistry, which might be
helpful in resolving our somewhat equivocal stand, are not available for
Columellia.
1969] STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 71
ACKNOWLEDGMENTS
This study has been carried out under the sponsorship of the Yale
School of Forestry, the Smithsonian Institution, the Arnold Arboretum of
Harvard University, and the University of Maryland. We are grateful
to administrators of these institutions for the privilege of using their fa-
cilities. For their encouragement, suggestions, and critical advice, we
wish to acknowledge warmly Dr. Sherwin Carlquist, : rt F.
Thorne, Dr. Arthur Cronquist, and Dr. Hugh IItis. The peingiinarb
of various herbaria have been cooperative in allowing us to see specim
in situ, to borrow specimens for study, and to use bits of stems, ives
and flowers for microscopic observations: Royal Botanic Gardens, Kew;
Yale School of Forestry, New Haven; Herbario San Marcos, Museo de
Historia Natural, Lima; Arnold Arboretum and Gray Herbarium, Har-
vard University, Cambridge; Field Museum of Natural History, Chicago;
U.S. National Herbarium, Smithsonian Institution, Washington; and
New York Botanical Garden, Bronx. Microscope slides and woods for
sectioning of Columellia and other taxa were made available from the
Record Memorial Collection of the Yale School of Forestry and from
the collections of the Division of Plant Anatomy, Department of Botany,
Smithsonian Institution. We appreciate the generous cooperation of
botanists in these institutions. Dr. Oscar Tovar of the Herbario San Mar-
cos, Lima, provided the only fluid-preserved anatomical specimens of
Columellia available to us and we are especially thankful for his efforts
in our behalf. Dr. John J. Wurdack, Smithsonian Institution, kept our
needs in mind during a collecting trip to Peru and provided us with a fine
wood sample of C. oblonga ssp. oblonga. Mr. James Teeri, of the Univer-
sity of New Hampshire, and Miss Carolyn Bensel, who is working on the
floral anatomy of the Saxifragaceae, sensu lato, in Dr. Barbara Palser’s
laboratory at Rutgers University, were most kind to share their observa-
tions with us in advance of publication. We also extend our thanks to
Mr. Austin Griffiths, Jr., of the Los Angeles State and County Arboretum
for preserved flowers of Escallonia; to Miss Brenda Gee, of the University
of Auckland, for preserved flowers of Carpodetus; and Dr. Judy Morgan
for help with the sectioning and examination of floral material.
LITERATURE CITED
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fragales v svyazi s nekotorymi voprosami ikh sistematiki: filogenii. Izv.
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AGARDH, J G. 1858. emer systematis plantarum. xcvi + 404 pp. pls. 28.
C. W. K. Gleerup. Lun
ARNOTT, H. J. 1959. Leaf poate Turtox News 37: 192-1
BAILLON, H. 1888. Gesnériacées. Histoire des plantes. 10: ere Librairie
Hachette. Paris.
BARTLING, F. T. 1830. Ordines naturales plantarum. Dieterichianus. Gottingae.
72 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
BENTHAM, G., & J. D age 1876. Columelliaceae. Genera plantarum. 2:
989. L. Reeve & Co. Lon
BHATTACHARJYA, S. S. 1954. - Beitrag zur Morphologie des Androceums von
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Brizicky, G. K. 1961. A synopsis of the genus Columellia (Columelliaceae).
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CANDOLLE, A. P. pe. 1839. Columelliaceae. Prodromus systematis naturalis
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Cuaxkravarty, H. L. 1958. Morphology of the staminate flowers in the Cucur-
bitaceae with special reference to the evolution of the stamen. Lloydia 21:
49-87.
CxarkE, B. 1858. On the anthers of Columelliaceae and Cucurbitaceae. Ann.
Mag. Nat. Hist. London. III. 1: 109-113; pl. VJ.
CoMMITTEE ON NOMENCLATURE, INTERNATIONAL ASSOCIATION OF Woop ANATO-
MISTS. 1957. oe glossary of terms used in wood anatomy. Trop.
Woods 107: 1-3
Cronguist, A. See The evolution and classification of flowering plants. x +
396 pp. Houghton Mifflin. Boston.
Don, D. 1828. Descriptions of Columellia, Tovaria, and Francoa; with remarks
on their affinities. Edinburgh New Philos. Jour. 1828-1829: 46-53.
Don, G. 1838. Columellieae. A general nage of the dichlamydeous plants.
4: 57, 58. J. G. & F. Rivington. Lon
ENDLICHER, S. 1839. Columelliaceae. aie plantarum. 1839: 745. Fr. Beck.
Vindo bonae.
. 1841. Columelliaceae. Enchiridion botanicum. 366 pp. Guil. Engelmann.
Lipsiae-Viennae
ENGLER, A. 1892. ‘Syllabus der Vorlesungen iiber specielle und medicinisch-
pharmaceutische Botanik. xxiii + 184 pp. Gebriider Homtiaeeee: Berlin.
1928. Saxifragaceae. Nat. Pflanzenfam. ed. 2
ErpTMAN, G. 1952. Pollen morphology and plant taeonomy. Angiosperms.
xii + 539 pp.; frontis, Almquist & Wicksell. Stockholm.
C. R. Metcalfe. 1963. Affinities of certain genera incertae sedis sug-
gested by pollen morphology and vegetative anatomy. Kew Bull. 17: 249-
256; pl. 2.
Esau, K. 1965. Plant anatomy. ed. 2. xx + 767 pp. John Wiley & Sons. New
York.
Eype, R.H. 1966. Systematic anatomy of the flower and fruit of Corokia. Am.
Jour. Bot. 53: 833-847.
Faun, A. 1967. Plant anatomy — by SysiL Bromo-ALTMAN). vii + 534
pp. Pergamon Press. ork.
FELDHOFEN, E. 1933. Hee zur physiologischen Anatomie der nuptialen
Nektarien aus den Reihen der Dikotylen. Beih. Bot. Centralbl. 50: 459-634;
Taf. HI-XX XI.
FritscH, K. 1893, 1894. Gesneriaceae. Nat. Pflanzenfam. IV. 3b: 133-185
(133-144, 1893; 145-185, 1894).
———. 1894. Columelliaceae. Nat. Pflanzenfam. IV. 3b: 186-188.
Getius, L. 1967. Studien zur Entwicklungsgeschichte an Bliiten der Saxifragales
ssa lato mit besonderer Beriicksichtigung des Androeceums. Bot. Jahrb.
8 —303.
GriseBacH, A. H. R. 1839. Genera et ~— gentianearum. viii + 364 pp.
J. G. Cottae. Saran et Tubinga
1969 | STERN, BRIZICKY, & EYDE, COLUMELLIACEAE 73
Hauer, H. 1901. Uber die Ne der Tubifloren und
nalen. Abh. Naturw. Ver. Hamburg 16(2): 1
9 "1908. Ueber die Abgrenzung und Vervandtscat pe einzelnen Sippen
bei den Scrophularineen. Bull. Herb. Boiss. II. 3:
1908. Uber Juliania, eine Terebinthaceen- ee, ae ‘Cupula, und die
wahren Stammeltern der Katzchenbliitler. 210 pp. C. Heinrich. Dresden.
. 1910. Ueber Phanerogamen ie unsicherer oder unrichtiger Stellung.
Meded. Rijks Herb. Leiden 1: 1-4
Herzoc, T. 1915. Die von Dr. Th. piace auf seiner zweiten Reise durch
Bolivien in den Jahren 1910 und 1911 gesammelten Pflanzen. II Teil. Meded.
Rijks Herb. Leiden 27: 1-90;
Hooker, J. D. 1873. “Editor’s note.” ” In: E. LE Maout & J. apse A
general system of botany. (Transl. by Mrs. Hooker; edited .
Hooker.) xii + 1066 pp. Longmans, Green, and Co. London. Editor's
note, 594
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Fic A representation of the diurnal variation of intensity and frequency of
Bont occurrences.
night as in the day, whereas daytime rains were almost twice as heavy
as nighttime rains. Presumably daytime rains are predominantly convec-
tive, while nighttime rains are orographic.
84 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The effect of the forest on rainfall rate is illustrated in Ficure 4 where
the 1-minute accumulations of rainfall above and below the forest are
compared for a typical shower. During an 11-minute interval 0.54 inches
of rain fell above the forest. Within a minute the rain began below the
forest, continued for 12 minutes and totalled 0.38 inches. The forest acts
like a filter, delaying the onset of rain at ground level, lessening the peaks
and smoothing the variations of intensity, and prolonging the rain slightly
as it drips from the leaves.
T T T T T T T
> lOr ~ x————x ABOVE se
= o——-—-o BELOW
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TIME IN MINUTES
. A representation of the rainfall rate above and below the forest versws
‘eo near noon, 20 July 1966.
The rain above that is not caught by the lower rain gage has either
been intercepted by the trees or has reached the ground by means of the
trunks. Hydrologists define interception as that portion of precipitation
that never reaches the ground either as rain or trunk flow. This relation-
ship is expressed in the simple continuity equation:
Rain Below = Rain Above — Trunk Flow — Interception (1)
Wisler and Brater (1959) point out that after the initial wetting of the
leaves, branches, and trunks of trees, the interception rate becomes equal
to the evaporation rate from those surfaces. Since Pico del Oeste is usu-
ally shrouded in fog it follows that the interception rate is usually zero
and that trunk flow must account for almost all of the difference between
rain above and below the forest. This result contrasts with the rather large
1969 | BAYNTON, ELFIN FOREST, 3 85
interception and small trunk flow reported by Clegg (1963) in much
taller stands at lower elevations in the Luquillo Mountains.
Rainfall below the forest as a function of rainfall above was analyzed
by standard regression techniques. One hundred and twenty-four rains
were selected for the analysis. Each had the property that it was preceded
by a 6-hour drying period as shown by the hygrograph trace. Thus part
of each rain was used in wetting the foliage. The balance either ran
down the trunks or dripped through. Using the notation X = rain above
the forest in inches, Y = rain below the forest in inches, the analysis
yielded:
Y = 0.768X — 0.034
with a correlation coefficient of 0.99 between X and Y.
A correction was dictated by slight differences in the volume of water
required to tip the buckets in the two gages. The design value is 18.5 ml.
Calibration of the two gages in place gave 18.5 for the upper gage and 18.9
for the lower gage. Appropriate adjustment leads to the final result:
Y = 0.786X — 0.035
On rewriting the equation in the form:
Y = X — 0.214X — 0.035
and comparing it to equation (1) we can identify trunk flow as 21.4 per-
cent of the rain above and interception as 0.035 inches. Clegg cites other
investigators as setting trunk flow no higher than 10 percent of the total
rainfall. The explanation for large trunk flow on Pico del Oeste is un-
doubtedly found in the unusually high number of stems, a feature of this
forest that is well illustrated in Ficure 5.
The interception figure of 0.035 inches implies that the vegetation
over each square meter of ground is able to store 886 ml/m?. Studies of
the U.S. Forest Service reported by Wisler and Brater indicate storage
capacities of about 0.14 inch for hardwoods and 0.23 inch for pines in
North Carolina. Storage in the cloud forest of Pico del Oeste would be
expected to be very much less since it is only 10 to 12 feet high. It should
be noted that, for rains occurring when the forest is already thoroughly
wet, the relationship between rain below and above simplifies to
Y = 0.786X
Cloud Water. The difficulty with all cloud water studies is to relate
the observations of a collecting device to the amount of water that the
foliage itself extracts from the cloud. Although the same difficulty besets
the interpretation of the data collected on Pico del Oeste, different lines
of argument support the conclusion that cloud-water is of secondary im-
portance in this region of abundant rain. In the first place, cloud water
is not a means of sustaining the forest during drouth since cloud-free
periods coincide with rainless periods. Secondly, four distinct analyses,
each of which by itself is imprecise, give very similar results.
First ANALYsIs. The cloud-water sampler was in service for 258 days
during the year from June 1966 to May 1967. By extrapolating the data
86
JOURNAL OF THE ARNOLD ARBORETUM
a me
wh Pot oy
ex
*
RY ‘ Pt
% :
. -
[voL. 50
at number of stems in its composition.
A view of the elfin forest illustrating the gre
1969 | BAYNTON, ELFIN FOREST, 3 87
to a full year the annual total of cloud water is estimated at 325 liters/
square meters (1/m*). Since 1 mm. of rain is the same as 1 1/m?, the an-
nual rainfall total of 453 cm. may be expressed as 4530 1/m?. Although
the unit cross section is in a vertical plane for cloud water and in a
horizontal plane for rain, no adjustment is needed for trees such as those
and vertical planes. Moreover the wind speed through the thermometer
shelter housing the cloud-water collector was found to be nearly the same
as the wind in the forest halfway to the top, namely 17.6 and 16 percent,
respectively, of the 20-foot wind. Deferring for the moment any discus-
sion of differences in the collection efficiency of the aluminum shadescreen
and the foliage, and differences in the sampling period, the data imply
that cloud water is only 7.2 percent of rain water.
SECOND ANALYsIs. Another approach was based on a 1,000-hour period
from 18 July to 29 August 1966. Since cloud water intercepted by the
trees contributes to rain measured below the forest, the record of the
20-pen event-recorder was examined for occurrences of rain below the
forest without rain above. The event of interest, “rain below without rain
above,”” was defined as no rain above during the three hours including
the hours preceding and following an occurrence of rain below. Any run
of three hours without rain above the forest is a possible occurrence of
the event of interest. There were 324 possible occurrences and only 15
observed occurrences. During the 1,000 hours the total rainfall below the
forest was 15.83 inches. Of that total 0.15 inch occurred without rain
above and must therefore be attributed to cloud water. Additional cloud
water is also collected during rain but the exact amount cannot be deter-
mined. Again the implication is that cloud water is only a small fraction
of rain water.
Tuirp ANALysIs. The 15 cases were then examined in detail in an
attempt to relate the observed cloud water collection to the observed
rainfall below the forest. The approach was to count the number of tips
of the cloud-water collector during the time for 0.01 inch of rain to
accumulate in the below-canopy rain gage. The data are summarized in
TABLE 1. The entry for August 19 is suspect because of the long accumu-
lation time indicating that the foliage must have dried out and had to be
rewetted before the process of foliage drip could resume. The same might
be true for August 16. The analysis is imprecise because in many cases
some rain fell during the accumulation time. Omitting August 19 the
mean is 5.4 units of cloud water per 0.01 inch of rain below the forest.
Earlier it was shown that 78.6 percent of rainfall drips through when
the foliage is wet and the same will be true for cloud water. Thus the
collection of; < 0.01 inch! or 259 ml/m? of water below the canopy
18.9
i. : is the correction factor that accounts for 18.9 ml. being the actual volume
(18.
of water to tip the bucket rather than the design volume of 18.5 ml.
88 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
259
0.786
of cloud water by the foliage. During the same period this analysis shows
that 5.4 (50) = 270 ml/m? of water were collected by the cloud-water
detector. Therefore, the foliage is 1.2 (i.e. 330/270) times as efficient as
the cloud-water collector.
during a rainless period results from the collection of = 330 ml/m?
TABLE 1. For fifteen occurrences of 0.01 inch rain below the forest without rain
above, the amount of cloud water collected during the accumulation time,
the accumulation time.
Amount cloud water Accumulation time for
Date during accumulation time 0.01 inch rain below
July 29 8.1 units * 4 hours 11 min
30 9.6 3 37
Aug. 2 6.6 4 50
2 5.4 4 30
5 5.0 3 12
5 13 1 40
7 1.4 3 12
9 ye: 3 43
11 2.5 ! 9
13 42 5 14
13 4.0 4 54
14 55 4 7
16 10.0 11 8
19 16.5 24 39
22 5.0 3 10
* One unit, or bucket tip, equals 50 ml/m”.
Fourtu ANAtysis. The main shortcoming of the third analysis was
the truncation error associated with the collection of rain and cloud water
in discrete steps of a bucket. It appeared possible to sharpen the analysis
by replacing the tipping buckets with bottles and measuring exact vol-
umes of water collected by the two rain gages and the cloud-water col-
lector under personally observed weather conditions. The fourth analysis
summarizes the results of this approach carried out in December of 1967.
Five separate attempts were made to collect rain and cloud water under
known boundary conditions. When the data were analysed, errors in eXx-
perimental technique became evident. Generally there was doubt about
the boundary conditions. No interpretation of the first attempt was pos-
sible because it became apparent that the foliage was neither fully wet
nor fully dry. Another error in technique may be illustrated by the anal-
ysis of the data collected on 9 December.
The collecting of water began at 9:55 a.m., immediately after a mod-
1969] BAYNTON, ELFIN FOREST, 3 89
erate shower, and continued until 4 p.m. The error in technique was that
the sampling began so soon after the shower that its effect was still being
felt as drip from the foliage. The collected water was equivalent to 3227
ml/m? of rain above the canopy, 2746 ml/m? below the canopy, and
only 70 ml/m? of cloud water. Because the foliage was always fully wet
the relationship
Y = 0.786X
should apply where Y = 2746 ml/m*. But because of the faulty tech-
nique, we have
= Rain Above + (Cloud Water) E + A\R, with E being the ratio
between the collecting efficiency of the vegetation and the cloud-water
collector, and /\R being the unknown amount of rain above the canopy
immediately before 9:55 that is in the process of getting to the ground
via trunks and drip-through. Substituting both Y and X in the equation
gives:
2746 = 0.786 (3227 + 70E + AR)
whence:
E = 3.82 — 0.014 AR
We conclude therefore that E is less than 3.82 since /A\R is not zero,
but that is all we can sa
Two of the attempts were for periods that began with dry foliage, i.e.
no liquid water attached to plant surfaces. The collections in the three
gages should therefore be related by:
Y= 0.786X — 886
where the units of X and Y are ml/m? so that 886 replaces 0.035 in the
original equation because each hundredth of an inch of rain = 254 ml/m*.
Between 4:35 p.m. December 7 and 1 p.m. December 8 the amounts
collected were 3570 ml/m? of rain above, 2004 ml/m? below, and 16
ml/m* of cloud water. With so little cloud water the equation is too
sensitive to slight errors in the collected amounts above and below to per-
mit estimates of E, the relative collecting efficiency of the vegetation. We
can, however, get independent estimates of the Y-intercept, 886, by sub-
stituting E = 1.2, the value obtained earlier or E < 3.82,? the upper limit.
These choices of E yield a Y-intercept = 817 and <850.
Conditions were also dry on the peak at 11:45 a.m. December 3 when
collections were begun. By 3:45 p.m. December 4 the amounts were
1936 ml/m? of rain above, 906 ml/m? below, and only 22 ml/m? of
cloud water. Setting E = 1.2 and <3.82 gave values of 637 and <682
for the Y-intercept. The average of these two trials was 727 for E = 1.2
and <766 for E <3.82, providing fair confirmation of the value, 886,
obtained from the regression equation.
One attempt combined enough cloud water with light rain to permit
an estimate of E, the relative collecting efficiency of the forest. Between
? < is the symbol for “is less than.”
90
JOURNAL OF THE ARNOLD ARBORETUM
[voL. 50
4:30 p.m. on December 2 and 11:20 a.m. on December 3 the observed
collections were 2128 ml/m? of rain above, 310 ml/m? of cloud water,
and 1151 ml/m? of rain below. On December 2 the peak had been clear
for four hours during the day but had fogged in shortly before 4:30 p.m.
FREQUENCY IN PERCENT
nm w
i.e) oe)
D
WIND SPEED IN MI/ HOUR
oO
Ol
T
3JWiL GYVGNVLS 1V901
b2
! Lee |
NIVY JO AONINOSYS
J
8
Q33dS QNIM
Fic.
frequency
of rain.
6. A graphic comparison of
the diurnal variations of wind speed and
1969 | BAYNTON, ELFIN FOREST, 3 91
Presumably the forest was substantially, but not fully, dry. Substitution
in the regression equation gives:
1151 = 0.786 (2128 + 310 E) — 886,
from which E = 1.5 or slightly less since the constant, 886, is for fully
dry foliage.
While not providing the hoped for “acid test,”’ the fourth analysis con-
firms that the foliage is only slightly more efficient than the cloud-water
collector and that its storage capacity is substantially less than that re-
ported for other forests.
Correcting the first analysis for the greater collecting efficiency of the
foliage, we have annual cloud water of 1.2 (325) = 390 1/m?, which is
8.6 percent of the annual rainfall. Although this amount is relatively
unimportant to the water budget of Pico del Oeste it is equal to the nor-
mal annual precipitation for Denver, Colorado.
Wind Speed. The diurnal variation of wind speed above the forest is
shown in the upper half of Ficure 6. The data were for a month with little
daytime clearing, August, and a month with considerable daytime clearing,
October. Both months showed the same pattern and were therefore com-
bined. For comparison the diurnal variation of rainfall frequency is in-
cluded in Figure 6. The night maximum and day minimum show the influ-
ence of convection. During the daytime there is a downward flux of mo-
VERTICAL PROFILE OF WIND SPEED
or T T 1 T T q T Ly T
iB u
pb: 16E :
Z al | os :
2 ak TREE TOPS ve é .
ERQOL BOOAA
é Br Pe 7
. sar
sy,
tal / i
/
2+ ;
ral I l l i l | 1 i l l
a 10 20 30 00
WIND SPEED AS A PERCENT OF 20-FT. WIND
Fic. 7. A vertical profile of wind speed.
92 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
mentum below the mountain top and a consequent decrease in wind speed.
The same anomalous cycle has been reported on towers several hundred
feet above flat ground.
The almost identical cycle of rainfall frequency supports the interpre-
tation that the frequent nighttime rains are mainly orographic, and that
daytime rains are mainly convective.
The vertical profile of wind speed above and below the tree tops was
investigated by installing a sensitive Casella anemometer at various
heights on the tower. Wind speed averaged over an hour was expressed
as a percent of the 20-foot wind. The results are presented in FIGURE 7.
neem points below the tree tops might modify that portion of the
SUMMARY
Rainfall on Pico del Oeste, although twice as frequent by night, is only
half as intense as during the day. Rain is mainly orographic by night and
convective by da
Trunk flow accounts for 21 percent of the rainfall.
The canopy has a storage capacity equal to a depth of 0.035 inches or
886 ml/m?. :
On the average, water extracted from the clouds by the foliage 1s
slightly less than 10 per cent of rainfall.
Winds are strongest at night and weakest during the afternoon.
REFERENCES
ALAKA, M. A. Problems of tropical meteorology (A survey). Tech. Note no. 62,
p. 20. World Meteorological Organization, 1964.
Baynton, H. W. e ecology of an elfin forest in Puerto Rico, 2. The micro-
climate of See del Oeste. Jour. Arnold Arb. 49: 419-430. 1968.
Ctiece, A. G. infall interception in a tropical forest. Carib. Forest. 24(2):
75- 79. 1963. é
EKERN, P. C. Direct ger capes ee ane water on Lanaihale, Hawaii. Soil
Sci. Soc. Am. Proc. 28(3): 4 1964.
Howarp, R. A. The ecology of an ger forest in Puerto Rico, 1. Introduction
and composition studies. Jour. Arnold Arb. 49: 381-418. ;
Kraus, E. B. The diurnal precipitation change over the sea. Jour. Atmos. Sci.
20: 551-556. 1963.
Wister, C. O., & E. F. Brater. Hydrology, 2nd ed. p. 195. John Wiley & Sons,
New York, 1959.
NATIONAL CENTER FOR ATMOSPHERIC RESEARCH
BouLpEer, CoLorApo 80302
1969} GATES, ELFIN FOREST, 4 93
THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 4.
TRANSPIRATION RATES AND TEMPERATURES OF LEAVES
IN COOL HUMID ENVIRONMENT ?
Davip M. GATES
THE PURPOSE OF THE STUDIES reported here is to contribute some under-
standing of the adaptation, growth, and behavior of plants in the mist
forest at the top of Pico del Oeste, Luquillo Mountains, Puerto Rico.
The primary influence of climate on a plant is through the transfer of
energy. All physiological processes consume energy. Biochemical re-
actions are temperature dependent and some are light dependent. The
vitality of a plant depends on its temperature and its energy content. If
a plant is too warm, its vital processes slow down; and above certain
temperatures many physiological processes stop and denaturation of pro-
teins occurs. If a plant is too cool, its vital processes slow down. The
plant will not survive below certain temperatures. Most plants grow best
at an optimum temperature.
The energy content of a plant determines its temperature. Several
factors affect the energy exchanged between a plant and its surroundings.
The significant environmental factors are radiation, air temperature, wind
and humidity. In order for these factors to be translated into their effect
on the plant, they must be expressed as energy flow. The incident radia-
tion is a specific amount of energy. The air temperature and wind speed
are translated into energy flow by the concept of convection. The humid-
ity of the air affects the energy exchange for a leaf by means of the
transpirational cooling. The leaf temperature and transpiration rate are
dependent variables which are functions of the four independent variables:
radiation, air temperature, wind, and humidity. Therefore, it is seen that
one must deal with a six-dimensional problem. This is complicated, but
there is no choice. It is not valid to ask for the influence of air tempera-
ture on transpiration rate without specifying the values of all other vari-
ables simultaneously. It is this simultaneity of factors which makes eco-
logical problems complex.
ENERGY EXCHANGE
A leaf absorbs an amount of radiation which is designated Q,,, in cal
cm~-? min-!. The absorbed radiation is the sum of absorbed direct sun-
*Supported by grant No. AT(11-1)-1711 from the U.S. Atomic Energy Commis-
sion and the Center for the Biology of Natural Systems under PHS grant No. 1 P10
ES 00139-03 ERT. Field facilities and transportation were provided under the NSF
grant GB: 3975 to R. A. Howard.
94 JOURNAL OF THE ARNOLD ARBORETUM [vor. 50
light, scattered skylight, reflected light, and emitted thermal radiation
from ground, vegetation, and atmosphere. The leaf absorbs each incident
stream of radiation according to the absorptivity of its surface and the
leaf orientation. This is discussed by Gates (1968a) in detail. The leaf
consumes a very small fraction, maybe one or two percent, of the ab-
sorbed radiation in photosynthesis. The major portion of the absorbed
radiation is lost by radiation emitted from the leaf surface, by convection
and by transpiration. The energy budget for the leaf is given as follows:
1/2 od, (T1) — rh. sda (Te)
Que = co TY +k( ~) (T, — Ta) +L —
(1)
where ¢ is the emissivity of the leaf surface, o is the Stefan-Boltzmann
radiation constant, k is a constant, V is the wind speed in cm sec~’,
the width of the leaf in cm, T,; and T, the leaf and air temperatures respec-
tively, L the latent heat of vaporization of water (580 cal gm~' at 30°C),
the tribes. The hairs he calls ag :
hairs (Zwillingshaare) consist of two more or less isometric basal ce a ne
pulvi
Numerous elaborations and/or reductions of this basic type of double hair have
occurred in different species and genera of the family. (Cf. Senecio.)
110 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
hemispherical, composed of a single row [rarely more] of erect, usually
free, flat [keeled], green bracts; supernumerary bracts sometimes present;
receptacle slightly convex [flat], naked, foveolate. Florets dimorphic
(perfect and carpellate) or all tubular and perfect; pappus setose-capillary,
soft, white; ray florets, when present, carpellate, in a single outer series,®
the corolla with an irregularly toothed ligule, yellow; disc florets perfect,
the corolla tubular, shortly 5-fid, yellow to orange (rarely white or light
purple), the anthers with terminal appendages and truncate bases; pollen
spherical, more or less spiny, prominently tricolporate (cf. Greenman) ;
style branches of perfect florets truncate (to penicillate) or with a short,
pointed apex. Achenes subterete, 5—10-nerved, variously pubescent. LECc-
TOTYPE SPECIES: S. vulgaris L.; see Cassini, Dict. Sci. Nat. 48: 454. 1827.
(Name Latin, applied to groundsel, S. vulgaris; derived from senex, old
man, referring to the soft, white pappus which suggests the beard of an
old man). — RAGWoRT, GROUNDSEL.
An ubiquitous genus, possibly the largest of the flowering plants, esti-
mates varying from 900 (Bentham & Hooker) to 3000 (Cabrera) species.
There has been no treatment of the entire genus since that of De Candolle
(1838) in which he divided the genus into several “series” based on geo-
graphical distribution. Most authors have arbitrarily accepted this treat-
ment as a basis and have worked within one geographical area (e..,
Muschler, Africa; Cufodontis, northern Eurasia; Cabrera, Chile; Green-
man, North and Central America). Greenman placed the North and
Central American species in 21 sections; five sections (some of which are
not very distinct) with eleven species occur in the Southeast and four
introduced and 15 native species are known from the eastern United
States as a whole
The European section Senecio ($ Annui Hoffm.), composed of weedy
annual herbs, is represented in our area only by the now almost cosmopoli-
tan S. vulgaris L., 2n = 40, which differs from our other species in its
annual habit and uniformly discoid heads of yellow flowers. Two species
of sect. SANGUISORBOIDEI Greenman, characterized by the perennial habit
of its species and the more or less glabrous, once or more pinnately parted
leaves, occur in the Southeast. Senecio glabellus Poir., 2n = 46, is wide
ranging in wet habitats from Mexico eastward to Florida and north to
Oklahoma, Kansas, Missouri, Illinois, Indiana, Kentucky, Tennessee, and
North Carolina, whereas S. Millefolium Torr. & Gray (including S. Mem-
mingeri Britton), with basal leaves two or three times pinnate, is restricted
to rocks and cliffs in a few counties in the mountains of southwestern Vir-
ginia, western North and South Carolina, and northernmost Georgia. The
remainder of the species of this pence are found predominantly in the
uplands of Mexico and Central Amer
Most of our species fall into sect. ja Rydb., which is composed of
* The presence of ray florets in normally radiate species is not an absolute character.
The number of ray florets is also subject to great variation, mainly according to the
number of involucral bracts
1969 | VUILLEUMIER, GENERA OF SENECIONEAE 111
perennial, usually glabrous herbs with petiolate simple or lyrately parted
basal leaves and cauline leaves reduced upward. Six of the 22 species of
this group reach our area: S. aureus L. (including S. gracilis Pursh),
2n = 44; S. Robbinsii Oakes ex Rusby, 2m = 46; S. obovatus Muhl.
(including S. rotundus (Britton) Small), 2n = 40; S. Smallii Britton, 2n =
44; S. pauperculus Michx. var. Crawfordii (Britton) T. M. Barkley; and
S. plattensis Nutt. Senecio aureus, S. Robbinsii, and S. pauperculus var.
Crawfordii all frequent moist to wet meadows and bogs, S. obovatus and
S. plattensis prefer drier areas, and S. Smallii grows primarily in fields,
roadsides, and open woods. Senecio Robbinsii occurs in the Southeast
only as a remarkably disjunct population on Roan Mountain (Tennessee-
North Carolina border) with the principal populations located far to the
north in the mountains of New York and New England and in adjacent
Canada.
Many of these species commonly hybridize where their ranges and
habitats overlap, which often makes identification of intermediate plants
difficult. However, hybrids are usually restricted to “hybrid’”’ habitats
and do not seem to swamp out the parental species. The species of sect.
AurEI, their ecology, natural history, and evolution, have been thoroughly
discussed by Barkle
Members of sect. TomeNtos1 Rydb. differ from those of sect. AUREI
primarily in a tendency toward being permanently tomentose. The major-
ity of the species are centered in the Rocky Mountains, but the range of
the type species, Senecio tomentosus Michx. (including the glabrous-leaved
f. alabamensis (Britton) Fern.; S. alabamensis Britton), 2n =
stretches across the country in weedy areas from Arkansas and Texas
to Florida and north to southern New Jersey. Hybrids between S. tomen-
tosus and S. aureus are fairly common (Barkley), and the hybrid of
S. tomentosus and S. Smallii also occurs. The latter has been studied cyto-
logically (in the first meiotic division 2m = 21-22 bivalents and 1-3
univalents). This evidence and other studies of Barkley have shown that
some of the sections used by Greenman are artificial and should possibly
be abandoned. In the southwestern United States (Colorado, New Mex-
ico, Texas) hybrids are formed between S. mutabilis Greenm. (sect.
TOMENTOsI) and S. neomexicanus Gray (sect. ToMENTOsI), S. mutabilis
and S. multilobatus Torr. & Gray (sect. Lopatt Rydb.), and between
S. mutabilis and S. tridenticulus Rydb. (sect. AUREI). An earlier study
had already indicated that the species of the S. multilobatus group be-
longed in an integrated complex with species formerly considered to
belong to sects. BoLANDERANI Greenm., LosaTI, and AUREI. (See also
the cytological data of Ornduff et al., 1967. )
The last section, RUGELIA (Shuttlew. ex Chapm.) Greenm., contains
only the unique Senecio Rugelia Gray (Rugelia nudicaulis Shuttlew. ex
Chapm.), winterwell, 2n = 56, a perennial herb with alternate, undivided
leaves and large, nodding discoid heads of white or light purple florets
in a simple corymbose raceme. This species grows in partial shade in
112 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
cool woods (usually of Picea rubens Sarg. and/or Abies Fraseri (Pursh)
Lindl.) at high elevations (ca. 1300 m.) in the Smoky Mountains of
western North Carolina and eastern Tennessee. On the basis of both
the morphology and the chromosome number, Ornduff e¢ al. have reiterated
that this species should be removed from Senecio.
Considering the size of the genus, relatively few studies have been
made on its embryology, cytology, and anatomy. Palmblad (1965) and
Ornduff et al. (1967) have recently added much new cytological infor-
mation and discussed some of the possible significance of chromosome
numbers within the genus, but too many species have still not been
counted to allow decisions concerning the genus as a whole. Gustafsson
reported no apomixis in the species of Senecio he examined, a finding
corroborated on other species by Afzelius and Haskell. The breeding
system in the few cases studied appears to be one of facultative out-
breeding with occasional inbreeding (Knuth, Haskell). Hauman postu-
lated that the arborescent senecios (sect. ARBOREI Hoffm.) of Africa are
all obligate inbreeders.
Anatomical studies have been made most extensively on the African
arborescent species of Senecio (cf. Hare). Recent c comparative work by
apparently because in the Senecioneae stem anatomy is easily modified
under different environmental conditions. Yet within Senecio itself, Carl-
quist found that clustering of species on the basis of wood anatomy was,
in some cases, consistent with groupings based on other morphological
criteria. Hare and Carlquist concur that the woody members of the
Senecioneae are derived from herbaceous ancestors and that the stem
structure of Senecio is advanced in comparison with other genera.
A recent study by Drury & Watson on some of the Eurasian sections
of Senecio has revealed that the leaf and achenial hairs, pappus types,
and the kinds of ovarian crystals — when carefully and critically examined
—provide useful taxonomic characters. They call for a reassessment of
many characters usually superficially examined in species of the Com-
positae and imply that the use of these characters might help in pro-
ducing a more natural classification of such troublesome genera as Senecio.
The specialized anatomy of the achenial double hairs of Senecio vul-
garis (see footnote 5 under Arnica) has been described by Macloskie and
J. Small and that of several other species of Senecio by Hess. The basal
cells, as in the double hairs of most Compositae, act as pulvini sensitive
to moisture. In several species of this genus, the two hair cells are further
specialized and are filled with a spiral tongue of a mucilaginous substance
which is extruded when pressure due to water absorption forces the hair
cells to separate. Apparently, the seer sticks the achenes to soil
particles and thus helps to insure germina
Alkaloids reported in at least 75 species (cf. ‘Willaman & Schubert) un-
doubtedly account for the medicinal use of various species of Senecio.
1969} VUILLEUMIER, GENERA OF SENECIONEAE 113
In the United States, only S. aureus was extensively used, the leaves first
being dried, then steeped in water, and the liquid used as a stimulant,
diuretic, and uterine tonic. The last use of this brew by North American
Indian women led to the common name of squaw-weed. In other parts
of the world, shoots and leaves of several species are eaten raw or cooked.
Some species (especially those of Cineraria L., if this genus is merged with
Senecio) are cultivated as ornamentals.
Much attention has been directed toward Senecio and the Senecioneae
because of the writings of James Small, who attempted to prove in an
elaborate series of papers (1917-1919) that Senecio was the ancestral
genus of the Compositae. His theory has, however, been dismissed by
most workers with only a partial explanation. It thus seems worth noting
here that four general concepts, now considered to be erroneous, lay at
the base of his argument: (1) derivation of the Compositae from the
Campanulaceae subfam. Lobelioideae; (2) acceptance of the now refuted
Age and Area hypothesis of Willis; (3) the uplift of the Andes in the
early Cretaceous; (4) belief in the doctrine of evolution by saltation. As
a consequence of these tenets, Small proposed that the ancestral pre-
Composite had a woody habit, a zygomorphic bilabiate corolla, and united
anthers (free anthers are now considered primitive); * that Senecio, the
largest and most widespread genus of the family was naturally the oldest;
that the uplift of the Andes in the Cretaceous (rather than in the Pliocene-
Pleistocene as is now accepted) gave the genus ample time to spread
around the world; and finally, that evolution by saltation, combined with
this (presumed) early Andean uplift created a situation in which the
lobelioid pre-Composite evolved and radiated as the Andes rose and
thereby created a plexus of species able to migrate throughout the world.
Small also had a number of ideas concerning morphology which re-
inforced his conviction that Senecio was the ancestral Composite:
1. The pappus was developed from a structure that was morphologically
a hair. Consequently, a fine capillary pappus (as is found in Senecio)
should be primitive. The pappus now considered by most taxonomists to
be the ancestral type is composed of broad, flat bristles resembling the
a of the calyx, from which it is thought to be derived.
. The inflorescence of the pre-Composite was an umbel with all of
the pedicellar bracts except the outermost series already suppressed. Fur-
ther reduction would have resulted in a head with a flat or convex naked
receptacle and, correspondingly, a uniseriate involucre. Additional series
of receptacular or involucral bracts would be produced by the abortion
of florets in the head. Although it is still debated whether the primitive
Composite possessed an umbel or a panicle, most authors now accept
entham’s view that receptacular bracts and a multiseriate involucre are
unspecialized. A uniseriate involucre and a naked receptacle, as in Senecio,
are now considered to be advanced reductions.
“See Cronquist (1955) for a discussion of the evolution of the Compositae and
an enumeration of characters considered to be unspecialized in the family.
114 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
3. Through a series of drawings, Small showed how all the types of
style branches now found in the Compositae could be derived from the
flat, truncated style arms of Senecio. Similarly, he derived all the anther
types from the senecionid type with its terminal appendage and tailless
base. Yet, since this kind of hypothetical derivation from a selected
prototype can be made using almost any form (except for the obviously
highly modified ones) as a starting point, it really has little biological
meaning
4. Several corolla characters in Senecio were also suggested by Small as
primitive. Yellow, for example, was considered the ‘‘unspecialized” flower
color. Bilabiate corollas found in the ray florets of some species of Senecio
were deemed unspecialized because they were like those of Lobelia. Al-
though yellow may be a basic flower color in the Compositae, the tubular
corolla is now believed to be primitive and to have given rise to both the
bilabiate and the ligulate corolla (cf. Koch).
5. The chromosomal evidence available to Small suggested that five was
the base number for Senecio. More recent evidence, however, indicates
that ten is actually the base number for the genus (Ornduff et al.).
In spite of these mistaken ideas about Senecio, Small’s studies provide
one of the most complete comparative morphological surveys ever made
on the Compositae. Even without the bibliographies and summaries of
previous work, his research is an indispensable reference on Senecio and
the Compositae i in general.
REFERENCES:
All references listed under subtribe Senecioninae are pertinent.
AFzELIUs, K. Embryologische und zytologische Studien in Senecio und verwand-
ten Gattungen. Acta Horti Berg. 8: 123-219. 1924.
ALEXANDER, E. J. Senecio Rugelia. a 20: 29, 30. pl. 655. 1937. Sene-
cio Millefolium. Ibid. 31, 32, pl. 6
Bark_ey, T. M. A revision of — aureus Linn. and allied species. Trans.
Kansas Acad. Sci. 65: 318-408. 1962. icooeay treatment of Senecio
aureus fa ap group with comments on evolut
intergradation of Senecio plattensis ey Sélacie pauperculus in Wis-
consin. _Rhodors 65: 65-67. 1963.
. Taxonomy of Senecio see arrecg and its allies. Brittonia 20: 267—
284. 1968, "Tncludes S. Millefolium, 2
. Intergradation of Senecio sections se Tomentosi and Lobati through
Senecio mutabilis Greenm. (Compositae). Southwest. Nat. 13: 102-115.
1968.
re saci A. L. El género Senecio en Chile. Lilloa 15: 27-501. 1949. [In-
cludes 208 spp., numerous illustrations, little of generic or specific rela-
tionships. |
CLutTe, W. N. The meaning of plant names. XLVII. Senecios and others. Am.
Bot. 37: 105-109. 1931.
Cotron, A. D. The megaphytic habit in the tree Senecios and other genera.
Proc. Linn. Soc. Bot. London 156: 158-168. 1944.
1969] VUILLEUMIER, GENERA OF SENECIONEAE 115
Cronguist, A. Phylogeny and taxonomy of the Compositae. Am. Midl, Nat, 53:
478-511. 1955. [A general work on evolution of family with list of charac-
ters considered primitive and key to tribes. ]
Curopontis, G. Kritische Revision von Senecio sectio Tephroseris. Repert. Sp.
Nov. Beih. 70: 1-266. pls. 1-5. 1933.
Drury, D. G., & L. Watson. Anatomy and taxonomic comes of gross
vegetative morphology in Senecio. New Phytol. 64: 307- 65.
& . A bizarre pappus form in Senecio. Taxon - enn 311. 1966.
Cie pappus. ]
GREENMAN, J. M. Monographie der nord- und centralamerikanischen Arten der
Gattis Senecio. Bot. Jahrb. 32: 1-33. 1902. [A general treatment for
this area, with discussion of anatomy, morphology, relationships, evolution,
and key to sections. |
- Monograph of the North and Central American species of the genus
Senecio — Part II. Ann. Missouri Bot. Gard. 2: 573-626. pls. 17-20. 1915;
3: 85-194. pls. 3-5. 1916; 4: 15-36. pl. 4. 1917; 5: 37-107. pls. 4-6. 1918.
[The basic treatment. ]
Gustarsson, A. Apomixis in higher plants. Lunds Univ. Arsskr. II. Sect. 2.
“2: 1-68. 1946; 43: 69-372. Vs
Hare, C. L. The arborescent senecios of reece a study in ecological
anatomy, Trans. Roy. Soc. Edinb. 60: 355-371. 1940.
HASKELL, G. Adaptation and the breeding coating in groundsel. Genetica 26:
468-484. 1953. [S. vulgaris. ]
Hauman, L. Les “Senecio” arborescents du Congo. Etude morphologique,
phytogéographique et systématique. Revue Zool. Bot. Afr. 28: 1-76. pls.
1-11. 1935. [Also includes photographs and comparisons with the paramos
of S. Am.
Kocu, M. F. — in the anatomy and morphology of the Composite flower
II. The corollas of the Heliantheae and Mutisieae. Am. Jour. Bot. 17:
995-1010. pls. 51, 52. 1930.
Mactoskie, G. Achenial hairs and fibers of Compositae. Am. Nat. 17: 31-36.
883. [A short preliminary attempt, but interesting. For more detail, see
Hess under subtribal references. | d
MuscHter, R. Systematische und pflanzengeographische Gliederung der afri-
kanischen Senecio-Arten. Bot. Jahrb. 43: 1-74. 1909
Ornourr, R. Evolutionary pathways of the Senecio lautus alliance in New
Zealand and Australia. Evolution 18: 349-360. 1964.
PALMBLAD, I. G. Chromosome numbers in Senecio (Compositae). I. ——_
Jour. Bot. 43: 715-721. 1965. [65 collections representing 30 spp.; in-
cludes 5S. aaron Britton, 2n = 46, endemic to shale barrens of
Virginia, W. Virginia, and Penn. to the north of our area. |
Smatt, J. The a aa: ‘mechanism in the Compositae. Ann. Bot.
29: 457-470. 1 oe
Wittaman, J. J., _ G. Scuusert. Alkaloid-bearing plants and their con-
tained alkalcids, S. Dep. Agr. Tech. Bull. 1234: 1-287. 1961. [Senecio,
71-75.]
3. Cacalia Linnaeus, Sp. Pl. 2: 834. 1753; Gen. Pl. ed. 5. 362. 1754.
Tall caulescent herbs arising from a rosette of alternate, petiolate,
Spathulate, ovate, reniform, or hastate, entire, undulate, crenate, or
116 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
toothed [or lobed] basal leaves; stem leaves petiolate or sessile, decreas-
ing in size toward the inflorescence. Inflorescence a compound cyme of
numerous campanulate discoid heads with cylindrical or campanulate
involucres composed of a single series of herbaceous, lanceolate, winged
or flat bracts (sometimes with an outer series of supernumerary brac-
teoles); receptacle flat, naked, or with a fleshy projection in the center.
Florets monomorphic, perfect; pappus capillary, white; corolla deeply
5-cleft, white, cream, or pinkish; anthers with terminal appendages and
obtuse bases; style branches truncate or with short conical [or elongate
in some Mexican species] appendages. Achenes fusiform to cylindrical
with a variable number of ribs, smooth. (Synosma Raf. ex Britton &
Brown, 1898 [Hasteola Raf. ex Pojark., 1960, nom. superfluum]; includ-
ing Arnoglossum Raf., 1817 [Mesadenia Raf., 1838, nom. superfluum].)
Lectotype SPEctEs: C. hastata L., largely typified by the removal of the
original species to other genera; see Miller, Gard. Dict. Abr. ed. 4. 1754;
De Candolle, Prodr. 6: 327. 1838; Kitamura, Mem. Coll. Sci. Kyoto
Univ. B. 16: 170. 1942; Shinners, Field Lab. 18: 79. 1950; Pojarkova,
Fl. URSS 26: 684. 1961.8 (Name from Greek, kakalia, a name given
by Dioscorides to a plant believed to be a Tussilago.) — INDIAN PLAN-
TAIN.
A genus of perhaps 40 species distributed from eastern Europe to eastern
Asia, in eastern North America, Mexico, and Central America, and in
South America along the Andes southward to Bolivia. The Mexican
species were referred to Psacalium Cass., Odontotrichium Zucc., and
Pericalia Rydb. by Rydberg (1924) and recently to these plus Digitacalia
Pippen by Pippen (1968). Some Asiatic species have been removed to
Syneilesis Maxim. and Méiricacalia Kitamura. Nine species distinctly
divisible into two sections occur in the southeastern United States. Sec-
tion Cacaria (treated as the genus Synosma by Small and by Britton &
Brown, ed. 2) is represented only by Cacalia suaveolens L., 2n = 40,
which occurs in moist woods from Massachusetts to Minnesota, south-
ward to Missouri, Tennessee, and western North Carolina. Morphologi-
cally, it is distinct from all the other eastern North American species in
having hastate leaves with pinnate venation, large heads with 12-15
involucral bracts (plus a ring of bracteoles), a naked receptacle, and
numerous florets.® Its affinities lie with the Eurasian C. hastata L., 2n =
“The typification of Cacalia will be discussed in a subsequent paper. Rydbe
(1924), Cuatrecasas (1960), and Pippen (1968), contrary to the course followed spin
have maintained that Cacalia should be typified by C. alpina L., which was remove
from Cacalia as the type species of Adenostyles Cass. This choice restricts the name
to a genus of four or five species of Central Europe
*T have seen one atavistic specimen of Cacalia suaveolens (Moore, Rosendahl &
1969 | VUILLEUMIER, GENERA OF SENECIONEAE 117
60, and its Asiatic relatives rather than with any other North American
ecies
The seven other species in the Southeast form a closely knit distinctive
group (cf. Pippen) which constitutes sect. ConopHoRA DC. (Arnoglossum
Raf., Mesadenia Raf.). All are morphologically similar in having palmately
nerved leaves, five involucral bracts, a fleshy projection in the center of
the receptacle, and five florets.’ Cacalia Muhlenbergii (Sch.-Bip.) Fern.
(C. reniformis Muhl.), 2n = 50, occurs in woodlands from New Jersey
and Pennsylvania, west to Minnesota and south to Missouri, Alabama,
and Georgia. Cacalia lanceolata Nutt. var. lanceolata, 2n = 56, occurs
in moist to wet habitats from eastern Texas and Louisiana to Florida,
northward into southeastern North Carolina, and C. lanceolata var.
Eliottii (Harper) Kral & Godfrey (M. Eliottii Harper, C. Elliottii (Har-
per) Shinners) '° occurs from peninsular Florida northward into South
Carolina. Cacalia diversifolia Torr. & Gray also occurs in swampy areas
of southern Georgia and northern oe westward to Louisiana, and
C. floridana Gray is endemic to the dry, sandy oak and pine woods of
central and northern Florida. Cacalia striplicifolia LL, 20 = 50, 52, 54,
56, and C, apa rape (probably including C. plantaginea (Raf.) Shin-
ners), 2n = 54, wide ranging: the former in dry woodlands from
New York to helio and Nebraska, south to Oklahoma, and east to
Mississippi, Alabama, and Georgia, and the latter in damp prairies from
Ontario to Minnesota, south to Oklahoma and Texas. The last species
of this section, C. sulcata Fern., is restricted to sandy bogs in southern
Georgia and western Florida
Within the North American species, evidence supports the distinctive-
ness of the two sections, but the inclusion of both into one genus. The
morphological differences between the two groups of species are reinforced
y differences in germination. Three species of sect. CoNopHORA which
have been investigated (Cacalia tuberosa, C. atriplicifolia, and C. Muhlen-
bergii) have achenes which need four days to two weeks for germination
and cotyledons which are strongly curved upon emergence. In contrast,
C. suaveolens (sect. CacALIA) needs only 48 hours for germination and the
cotyledons are only slightly curved when they emerge. Afzelius reported
the infrequent occurrence of two embryo sacs in ovules of C. suaveolens,
each formed from a separate megaspore mother-cell. In one sac, the
€gg apparatus was invariably crushed but the antipodals were normal,
* The placement of Cacalia ovata Walt. is at present uncertain. This taxon, pre-
treatment is followed, C. cocoa var. Elliottii becomes C. ovata var. ovata, and
- lanceolata var. lanceolata will req under C. ovata a new combination based on
the oldest legitimate varietal cpithet “Gf available).
118 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
while in the other, the egg apparatus appeared normal, but the antipodals
were crushed. Greene later reported that good seeds of C. suaveolens
were difficult to find, and Wadmond stated that it was impossible to locate
viable seed of C. Muhlenbergii. These two observations suggest a similar
sort of meiotic irregularity in C. Muhlenbergii.
Another feature which links the Southeastern American species (sect.
ConopHora) and the Asiatic species (sect. CAcALIA) is the presence of
the same type of asexual reproduction. Kral & Godfrey reported, as a
general phenomenon in the Florida species, the production of lateral
rosettes which become disconnected from the parent plant by disintegra-
tion of the connecting stolons. Liubarsky described the same phenomenon
in greater detail for several Russian species. Two Japanese species,
Cacalia auriculata DC. var. bulbifera Koidz. and C. farfarifolia Sieb. &
Zucc., produce bulbils in the leaf axils (cf. Ohwi).
Within sect. ConopHora, the species are very similar morphologically,
seemingly closely related, and apparently rather removed from other
species in the genus. Hybrids within this section appear to be rare, how-
ever. The only natural hybrid reported, that between Cacalia atriplicifolia
and C. Muhlenbergii (also produced artificially by Coleman), was ex-
ceedingly sterile (2—9 per cent pollen staining and no seed set).
Generically, Cacalia is ill defined from Senecio L. and its satellite genera.
Originally, Linnaeus included in Cacalia the herbaceous perennials treated
here (and other species) and a group of shrubby African plants now con-
sidered to constitute either the genus Kleinia Mill. or Senecio subg.
Kleinia (Mill.) Hoffm. Bentham placed both the herbaceous and the
shrubby groups in Senecio, while Hoffmann separated the two, retaining
the herbaceous species as Cacalia and referring the species of Kleinia to
Senecio. In opposition to Hoffmann’s treatment, however, large and soli-
tary crystals of calcium oxalate (rather rare in the Compositae according
to Metcalfe & Chalk) have been found in both Cacalia and Senecio subg.
Kleinia.
Chromosome numbers of the 27 species reported as Cacalia are 2n = 40,
50, 52, 54, 56, 58, 60, 70, and 120. The 16 counts recently reported by
Pippen as species of the segregate genera Digiticalia, Odontotrichum,
Pericalia, and Psacalium are all 2n = 60, with the exception of that for
O. Palmeri which is 2n = ca. 50. Ornduff et al. (1967) suggest that the
basic chromosome numbers in Cacalia are 20 and 30 and that other num-
bers have been derived by aneuploid reduction from m = 30. More
chromosome counts and further study of the generic limits of Cacalia
on a worldwide basis undoubtedly are needed.
It seems possible, especially in view of the morphological continuity
with Senecio in Africa, that the genus is of Old World origin, but Liu-
barsky’s postulation of an origin in the region of the upper Amur River
is highly questionable.
,
PS rate
1969] VUILLEUMIER, GENERA OF SENECIONEAE 119
REFERENCES:
Under scepin references see BENTHAM, on & Hooker, De CANn-
DOLLE, HOFFMAN, ORNDUFF et al., and RICKE
AFZELIUS, K. aeames und aay Studien in Senecio und verwand-
ten Gattungen. Acta Horti Berg. 8: 123-219. 1924. [C. suaveolens, 162.]
ARANO, H. Cytological studies in subfamily Carduoideae (Compositae) of Japan
XVIL. The karyotype analysis in Cacalia and open is. Bot. Mag. To-
kyo 77: 86-97. 1964. [Gives counts for 9 spp. Cacalia
CoLeMaN, J. R. Natural and artificial cag of Gade atriplicifolia and C,
M uhlenber ii. Rhodora 67: 55-58.
Cuatrecasas, J. Studies on Andean acne ikea Brittonia 12: 182-195.
1960. [Includes typification of Cacalia
GREENE, E. L. Studies in a Compositae. eT Part 3. The genus Mesadenia.
Pittonia 3: 180-183.
GREENE, H. C. Differences in Senge characters and germination in some species
of Cacalia L. Am. Midl. Nat. 39: 758-760. 1948. [C. atriplicifolia, C.
Muhlenbergii, C. suaveolens, C. tuberosa. |
Harper, R. M. Mesadenia lanceolata and its allies. Torreya 5: 182-185. 1905.
1905.
Kitamura, S. Recognition of the genus Syneilesis Maxim. (In Japanese.) Jour.
Jap. Bot. 10: 699-703. 1934.
: — mia du Japon. (In Japanese.) Acta Phytotax. Geobot. 7: 236-
251:
AL, R. ‘ = K. Goprrey. Synopsis of the Florida species of Cacalia (Com-
positae). Quart. Jour. Florida Acad. Sci. 21: 193-206. 1958
Lruparsky, E. L. Notes on the genus Cacalia in the southern par of the Mari-
. 1961
Metcatre, C. R., & L. CuHatk. Compositae. Anat. Dicot. 2: 782-804. 1950.
[Cacalia and Kleinia, 786.]|
Ouw1, J. Flora of Japan (in English). F. G. Meyer & E. H. WALKER, eds.
ix + 1067 pp. Wash. D.C. 1965. [Miricacalia, 882; 13 spp. of Cacalia, 882-
884; Syneilesis, 887, 888.
Pippen, R. W. Mevicna * ‘cacalioid” ge allied to Senecio (Compositae).
Contr. U.S. Natl. Herb. 34: 363-448.
Poyarkova, A. Notae criticae de genere aes de fb Lee Russian.) Not.
Syst. Leningrad 20: 370-391. 1960. [Lectotype sp. = C. atriplicifolia;
adopts Hasteola Raf. (ex Pojarkova) for C. hastata, C. suaveolens, an
relatives. |
. Cacalia. Fl. URSS 26: 683-697. 1961. [Lectotype sp. = C. hastata L.]
Rypgerc, P. A. Some senecioid genera—I. Bull. Torrey Bot. Club 51: 369-
378. 1924. [Includes incorrect typification of Cacalia. |
Suinners, L. H. The Texas species of Cacalia (Compositae). Field Lab. 18:
79-83. 1950. [C. plantaginea and C. lanceolata, with comments on the
application of the name Cacalia
TaKEsHiITA, M. Cytological studies on Cacalia and its related genera. I. The
chromosome number of three species and one variety of acalia and one
variety of Miricacalia. (In Japanese.) Jap. ie Genet. 36: 217-220.
1961.*
Wapmonp, S. C. The Indian plantain. Asa Gray Bull. 6: 52. 1898. [C. reni-
formis = C. Muhlenbergii.]
120 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
4. Erechtites Rafinesque, Fl. Ludov. 65. 1817.
Robust annual [perennial] caulescent herbs with fibrous roots and alter-
nate, toothed or parted, glabrous or pubescent leaves. Inflorescence a
“panicle” or cyme of numerous heads, each with a basally swollen in-
volucre composed of a single series of narrow lanceolate scabrous bracts
often surrounded by a series of supernumerary bracteoles; receptacle
flat, alveolate or fimbrillate. Florets monomorphic and perfect [or in
some cases florets dimorphic, the ray florets then carpellate with filiform
4-5-parted corollas]; pappus thin, white, soft, copious; corolla tubular,
regular, 5-toothed, whitish or yellowish; anthers with obtuse bases; style
branches of perfect florets with a terminal appendage of fused papillose
hairs surrounded at the base by a semicircular crown of collecting hairs
(cf. Belcher). Achenes oblong to linear in outline, more or less 10[—20]-
ribbed, glabrous. (Erechtites sensu Bentham & Hooker and Hoffmann,
in part.) Type species: EF. praealta Raf. = E. hieracifolia (L.) Raf.
ex DC. (Name from Greek, Erechthites, a name given by Dioscorides
to a species of Senecio.) — FIREWEED, PILEWORT.
Two sections with five species native to the Americas (sensu Belcher)
and adventive in the Pacific region, Asia, and Europe. One wide-ranging
species, Erechtites hieracifolia (L.) Raf. ex DC., of sect. ERECHTITES
(§ Hieracifoliae Belcher), occurs in weedy habitats in the southeastern
United States.
Erechtites usually has been distinguished from Senecio and its allies
by its two to several series of outer carpellate florets with filiform, eligu-
late corollas. Belcher, however, narrowed the genus to include only
New World species which possess what he considers the diagnostic feature
of the genus: style branches with an appendage of fused papillose hairs
with a semicircular crown of collecting hairs at its base. He returned
several Australian and Indonesian species traditionally included in Erech-
tites to Senecio and placed five New Guinean species (one population in
New South Wales) in Arrhenechthites Mattfeld. The characters used to
separate Arrhenechthites from Erechtites include the presence of func-
tionally staminate disc florets and reduced, astigmatic (and thereby with-
out the critical character of Belcher’s Evechtites) style branches. In
view of the obvious correlation between sterile (abortive) ovaries and
reduced stigmas, the question arises as to whether these species are not
simply inbreeders derived from the outbreeding American species. Also,
the chromosome number of Senecio (Erechtites) minimus Poir., the only
Old World species counted, has a diploid count of 2m = 60, less like the
most common number in Senecio, 2n = 40, than E. hieracifolia with 2n =
40. Regardless of which circumscription is used, the taxonomy of our
species is unchanged.
Our species, Erechtites hieracifolia, has three varieties, two of which
are now widely distributed weeds. Varietas hieracifolia (including vars.
intermedia Fern. and praealta (Raf.) Fern.) occurs naturally from Can-
1969 | VUILLEUMIER, GENERA OF SENECIONEAE 121
ada southward through the Greater Antilles. It is distinguishable from
the other varieties by its short bracteoles less than 1/4 the length of
the involucre and by its 10-ribbed achene. It has been introduced into
Hawaii and Europe. Varietas megalocarpa (Fern.) Crong. (E. megalo-
carpa Fern.), separable from var. Aieracifolia in its much larger receptacle
(twice as wide) and its 16—20-ribbed achene, is endemic to sandy coastal
habitats from southeastern Massachusetts to New Jersey. Belcher’s sug-
gestion that the plants may be tetraploids derived from var. hieracifolia
seems not to have been tested. The third variety, var. cacalioides (Fisch.
ex Spreng.) Griseb., is found in Central America, the Lesser Antilles, and
South America to Argentina, and is now established as a weed in Asia.
It differs from the other two varieties in its longer bracteoles with multi-
cellular hairs (instead of being glabrous or bearing unicellular hairs).
Intermediates between this and var. hieracifolia occur in the West Indies.
The success with which Erechtites hieracifolia has managed to colon-
ize new areas is apparently due to its adaptability to new environmental
conditions and to its easily dispersed achenes. Ridley listed it as one
of the first plants to recolonize Krakatau after the volcanic eruption of
1883.
Although this species is usually an annual herb, Carlquist has exam-
ined a Hawaiian specimen over six feet tall which had secondary xylem.
The change from the annual habit in the Hawaiian population has
occurred, Carlquist postulated, because plants on oceanic islands are
“released” from the selection pressures of a cyclical climate. However,
there must be some positive selection for woodiness on islands (see also
Carlquist, 1965).
In Brazil, the leaves of Erechtites hieracifolia (and of E. valerianifolia
(Wolf) DC.) are cooked with palm oil (Corréa), and Ochse reports that
in the East Indies the upper leaves, called “lalab,” are eaten raw or steamed
with rice and are rumored to be beneficial for nursing mothers.
REFERENCES:
Under subtribal peau see BENTHAM, BENTHAM & Hooker, HOFFMANN,
OrnvurF et al., and RICKET
BELCHER, R. O. A revision of the genus Erechtites paacaany” wan inquiries
into Senecio and Arrhenechthites. Ann. Missouri Bot. Gard. 43: 1-85. 1956.
Carxouist, S. Wood anatomy of Senecioneae (Compositae). - 5: 123-146.
1962. [Comparative anatomy with conclusions as to relationships within
tribe and correlations with ecology. ]
Island life. 451 pp. New York. 1965. [Chapter 8. Some remodeled
plants, ]
Cooper, G. O. Cyt tological investigations of Erechtites hieracifolia. Bot. Gaz.
98: 348-355. 1936. [Describes both micro- and mega sporagenesis.
Corréa, “~ P. Diccionario das plantas uteis do Brasil e das exoticas cultiva-
vols. Ministerio da Agricultura, Rio de Janeiro. 1926, 1931. [Erech-
tites, * 96. 1931.]
122 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
FERNALD, M. L. The genus Erechtites in temperate North America. Rhodora
19: 24-27. 1917. [E. hieracifolia, its varieties, and E. megalocarpa. |
OcuseE, J. J. Vegetables of the Dutch East Indies. 1006 pp. Buitenzorg, Java.
1931. [Erechtites, 132-134. ]
Riptey, H. N. The dispersal of plants throughout the world. xx + 744 pp. pls.
1-22. oy Kent, England. 1930. [Erechtites, 133, 160, 656.
SANForD, S. N. Erechtites megalocarpa in Rhode Island. Rhodora 28: 111.
1926. [See on H. K. Svenson, Rhodora 41: 256. 1939. |
5. Emilia Cassini, Bull. Sci. Soc. Philom. Paris III. 1817: 68. 1817.
Annual or perennial caulescent herbs arising from a rosette of lyrate-
pinnatifid or spathulate, dentate [entire], glabrous or glaucous leaves.
Stem leaves alternate, dentate or lyrately lobed, decreasing in size toward
the lax corymbose inflorescence [plants sometimes monocephalous] of
discoid heads. Involucre tubular, often swollen at the base, composed of
a single row of lanceolate slightly scarious bracts; receptacle flat, naked.
Florets perfect; pappus setose, soft, white or purplish; corollas tubular,
shortly 5-fid, lavendar [white] or red; anthers truncate at the base; style
branches terete with penicillate appendages surrounded by a ring of hairs.
Achenes 5-angled, truncate at both ends. Typr species: Cacalia sagit-
tata Vahl, nom. illegit. = Emilia javanica (Burm.) C. B. Robinson.
(Derivation of name not explained but apparently from the French proper
name Emilie.) — Cupip’s PAINTBRUSH.
A genus of about five species native to the Tropics of Africa and the
Far East. Two weedy species, Emilia sonchifolia (L.) DC. ex Wight,
2n = 10, and E. javanica (Burm.) C. B. Robinson (E. coccinea (Sims)
G. Don in Sweet, E. sagittata (Vahl) DC., E. flammea Cass.), 2n = 20,
are naturalized in disturbed and weedy habitats in the warmer parts of
peninsular Florida, where the latter is more frequently encountered. Both
are also rather widely naturalized in the West Indies, Central America,
northern South America, and Brazil.
The application of the names of these two species has been the source
of much confusion. There has never been any doubt that there are two
entities: one a species with lyrate-pinnatifid lower leaves, small heads,
lavender (rarely white) corollas which barely exceed the involucre, laven-
der anthers and styles, and white pollen; the other with ovate-spathulate
dentate leaves, carmine corollas which extend conspicuously beyond the
involucre, orange anthers and styles, and bright yellow pollen. The first
species is Emilia sonchifolia, and the other long has been known as E.
sagittata in the Old World and FE. coccinea in the New. When Cassini
described Emilia he listed Cacalia sagittata Willd. as type species. Will-
denow, in turn, referred to Cacalia sagittata Vahl, excluding the synonym
Hieracium javanicum Burman. Because Willdenow excluded the synonym
(Vahl’s inclusion of this earlier legitimate name as a synonym made the
name Cacalia sagittata superfluous) and because the type specimens of
1969] VUILLEUMIER, GENERA OF SENECIONEAE 123
Vahl’s and Burman’s names had not been examined, it has been assumed
that two different taxa were involved. However, Mattfeld (1929) estab-
lished that Vahl’s type belongs to the large-headed, red-flowered species,
and Fosberg (1966) finally located the Burman type and reported that
it, also, was this species. Since Burman’s is the oldest legitimate name
available, it must replace the other names now used. That Cassini had
the showy red-flowered species in mind when he described the genus
Emilia is evidenced by his changing the name of the type species to EF.
flammea (nom. illegit.).
The two species apparently do not interbreed in nature, and attempts
to produce artificial hybrids (cf. Lee) have failed, indicating that a
sterility barrier (as well as the difference in chromosome number) is
involved. The two species are not only incompatible, but also have dif-
ferent mechanisms of reproduction (at least in Jamaica): Emilia sonchi-
folia is an obligate inbreeder, while E. javanica is outbreeding.
e genus seems to be a natural group of species closely related to, but
distinct from, Senecio L. Three of the five species counted have a chromo-
some number of 2” = 10, and the other two have 2m = 20. The species
with five pairs of chromosomes are annuals apparently derived from less
specialized species with 2n = 20 (cf. Ornduff et al.).
The genus has no true commercial value, although Emilia javanica is
sometimes used in tropical areas as an ornamental. Baldwin mentioned
that the plants were eaten in the Far East, but not in the New World.
REFERENCES:
Under subtribal references see BENTHAM, BENTHAM & Hooker, DE CANDOLLE,
HOFFMANN, and RICKETT
BaLpwin, J. T., Jr. Cytogeosraphy of Emilia Cass. in the Americas. Bull.
Torrey Bot. ‘Club 73: 18-23 6.
———.. Cytogeography of mais in Wes Africa. Ibid. 76: 346-351. 1949.
FosBerc, F. R. Miscellaneous notes on Hawaiian plants. Occas. Pap. Bishop Mus.
23: 129-138. 1966. [Typification of Emilia sagittata Vahl and Hieracium
javanicum Burm.
GaRABEDIAN, S. A revision of Emilia. Bull. Misc. Inf. Kew 1924: 137-144. 1924.
Goopine, E. G. B., A. R. Lovetess, & G. R. Proctor. Flora of Barbados. 486
pp. London. 1965. [E. sonchifolia, 437, fig. 27.]
Koster, J. Notes on Malay Compositae III. Blumea 7: 288-291. 1952. [Dis-
cusses application of names E. sagittata and E. javanica; but see Fosberg. ]
Ler, B. Jamaican species of Emilia. Sci. Notes sie Jamaica 2: 14, 15. 1966.
[Breeding systems of E. sagittata and E. javanica. |
MarttFELp, J. Die Compositen von Papuasien. a Jahrb. 62: 386-451. 1929.
[Emilia, 443-447, nomenclature. |
Sims, J. Cacalia coccinea. Scarlet-flowered Cacalia. Bot. Mag. 16: pl. 564. 1802.
(E. javanica. |
THE ARNOLD ARBORETUM Present address: THE GRAY HERBARIUM
OF
OF
HARVARD UNIVERSITY HARVARD UNIVERSITY
124 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ASPECTS OF THE COMPLEX NODAL ANATOMY
OF THE DIOSCOREACEAE ?
EDWARD S. AYENSU
THIS PAPER IS AN ATTEMPT to explain how the vascular tissue of two
successive internodes maintains continuity in the complex nodal structure
between them in stems of the Dioscoreaceae, especially in the genera
Dioscorea and Tamus. Because of the economic importance of this family
early emphasis (Mason, 1926) was placed on the relation between struc-
ture and function, This led physiologists to take a look at the anatomy
before they had a full knowledge of how food substances are translocated
in the plant.
The Dioscoreaceae is a monocotyledonous family which is distributed
throughout the tropics and subtropics of the world. It is, by all standards,
one of the most economically important foodstuffs in the diet of most
tropical peoples (cf. Coursey, 1967). Attention has recently been focused
on this family, especially the genus Dioscorea, because a precursor of
cortisone and other related steroidal drugs is derived from the tubers of
some species.
The unique anatomy of the nodes of the stems of the Dioscoreaceae
was brought to attention by Mason (1926) when he studied the rate of
sugar transport in Dioscorea alata L. Earlier, Falkenberg (1876) had
called the glomerulus of the node an imperfect knot in his study of D.
villosa L. Mason noted that the phloem was of a markedly abnormal type.
He further observed that the sieve tubes of the successive internodes did
not join with each other directly but through a glomerulus which was
composed of a great number of oblong thin-walled parenchymatous cells,
each with a distinct nucleus, running fairly parallel with each other.
Behnke (1965a) questioned the presence of nuclei in the glomerulus cells.
Present studies show that nuclei occur at certain stages in the ontogeny
of these cells (Fic. 2).
In his study of the ontogeny of the stem of Tamus communis L., Bur-
kill (1949) disproved Mason’s claim that glomeruli were absent from
the nodes of T. communis. Present studies reveal that glomeruli are cer-
tainly present in the nodes of Tamus (Fic. 12) and, although they cannot
be easily overlooked, it should be emphasized that the glomeruli in this
genus are not so pronounced as those of most species of Dioscorea (Fics.
4-11).
A full account of the vegetative anatomy of the Dioscoreaceae will be includ
in Bs Anatomy of the Monocotyledons. Dioscoreales, ed. C. R. Metcalfe, Oxford
University Press.
1969 | AYENSU, ANATOMY OF DIOSCOREACEAE 125
Happ (1950) wrote his thesis on the nodes of the Dioscoreaceae but a
copy is not available to me. However, a comment on it appeared in
Braun’s (1957) work. Essentially, Happ investigated by means of serial
sections the interlacing of the xylem-phloem glomeruli in the vascular
system of the node.
Brouwer (1953) published his account of the arrangement of the vas-
cular bundles in the nodes of Dioscoreaceae and presented a diagram of
the elements of the node. Brouwer concluded that the sieve tubes of two
successive internodes were connected in the following manner: sieve tubes,
funiculus cells, bast tubulus cells, glomerulus cells, bast tubulus cells,
funiculus cells, and sieve tubes. Brouwer, following Mason (1926), con-
cluded that the phloem-glomerulus cells were (a) densely filled with cyto-
plasm; (b) with a persistent nucleus with nucleolus; and (c) without
sieve areas.
A comprehensive study of the nodal anatomy of Dioscorea batatas
Decne. and Tamus communis was conducted by Braun (1957). He con-
cluded that (a) the xylem-glomerulus consists of very numerous short
tracheids of various sizes, the orientation of which is difficult to trace;
(b) the phloem-glomerulus, which is divided into several partial glomeruli,
is composed of a new type of translocatory cell, called phloem-glomerulus
cells; and (c) the phloem-glomerulus cells possess thin walls without sieve
pores and without visible pitting; they are distinguished from parenchyma
cells by their lack of starch. Behnke’s (1965c) electron microscopic
studies show that sieve areas are, in fact, present in the phloem-glomerulus
cells.
The present study, involving more species than were available to earlier
investigators, essentially supports and extends their conclusions.
MATERIALS AND METHODS
My observations are based on 180 specimens of 112 species. A com-
plete list and citations are given elsewhere (Ayensu, 1966).
Most of the specimens examined were fluid-preserved in formalin acetic
alcohol. Microscopic details were studied in serial sections at 10u, and
those produced on a sliding microtome usually at 162. Depending upon
the nature of some specimens, sections were cut up to 90u. The sections
were stained in safranin and counterstained with Delafield’s haematoxylin
followed by conventional differentiation, dehydration, clearing in xylene,
and mounting in Canada balsam.
NODAL ANATOMY
As pointed out in an earlier paper (Ayensu, 1965), the vascular strands
between the petiole and the stem at the nodes of many species of Dios-
coreaceae are highly distinctive and are believed to be unique in the family.
Longitudinal serial sections of the node reveal two groups of interlacing
vascular elements, each forming a plexus close to the petiole insertion.
126 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
VT
)
|
)
f
00
)
\
i
Fic. la (LEFT). Schematic diagram illustrating the arrangement of the ele-
ments of xylem-glomerulus in the nodal region of stems of Dioscorea and
amus.
Fic. 1b (RIGHT). Vessel-like tracheid showing a reticulate perforation plate
(lower) and bordered pits (upper).
1969 | AYENSU, ANATOMY OF DIOSCOREACEAE 127
Xylem-glomerulus. Serial sections and macerations reveal that the
mature xylem glomerulus is mainly composed of short tracheids of vari-
able shape closely fitted together, thus resembling the distinct parts of a
composite jig-saw puzzle. These peculiar tracheids are confined to the
node and have large bordered pits. Presumably in the internodes water
moves freely from vessel element to vessel element through the scalariform
perforation plates. Exactly how materials are translocated through the
nodal region is not clearly understood.
The phyllotaxy determines the width of the glomerulus in the nodes.
In the species having simple, alternate leaves, a single glomerulus oc-
cupies about one-third of the area of the node. In an opposite (or decus-
sate) arrangement, the glomerulus occupies about two-thirds of the nodal
area. In species that exhibit a whorled arrangement, the glomerulus oc-
cupies almost all the nodal area.
The tracheids vary in width and length within species. The widths
varying from 40» to 110u, and lengths from 80, to 260 have been re-
corded for different species. These tracheids are closely fitted together,
and have numerous pit-pairs on their common walls. The exact pathway
of the contiguous tracheids between successive internodes is very com-
plicated and variable within a species. (See Fics. 4-12.) Longitudinal
| ©,
Fic. 2 (LEFT). Schematic diagram of the phloem glomerulus. : :
Fic. 3 (RIGHT). Schematic representation of the stem showing the relationship
of and the position of the xylem and phloem glomeruli in the region of the leaf
Insertion.
128 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
sections and macerations of the node give a partial elucidation of the
complicated sequence of the tissue structures. As Braun (1957) inter-
preted D. batatas and Tamus communis, a vessel just about to enter a
node is attached to 1, 2, or 3 cells which Braun referred to as “‘vessel-like
tracheids.” The end wall of the vessel-like tracheid (VT) facing the vessel
(v) has a reticulate perforation plate, while the other end wall has
bordered pits (Fics. 1a, b). The elements that constitute the bulk of the
xylem-glomerulus lie between the vessel-like tracheids. The tracheids of
the first group (Tj) are closely fitted to those of the second group (T2)
and to other successive tracheid groups, thus establishing the normal com-
munication between them. The lengths of the tracheids vary from one
node to the other within a species. In this respect variation in tracheid
length does not have any taxonomic value. Those of the first few groups
(T; —Ts3) are shorter than those of T, and T;. It is also observed that
the tracheid groups increase in number from T; to T;, presumably for
enlarging the water conducting tissues in the node. The surface area of
the water conducting tissues is further increased by the complex arrange-
ment of many xylem-glomeruli at a node. Each glomerulus is S-shaped
and longitudinally orientated. A xylem-glomerulus diagram (Fic. 1) is
presented for the sake of simplicity, but the full complexity of it is demon-
strated by Fics. 4-12.
Phloem-glomerulus. The construction of the phloem-glomerulus
(Fic. 2) follows essentially the scheme presented for the xylem-glomerulus
(Fic. 1). The phloem-glomerulus is made up of what Braun (1957)
named ‘‘glomerulus sieve-tubes” (GS). Earlier, the same tissues had been
called “funiculus cells” by Brouwer (1953) and “funnel-cells” by Mason
(1926). Recently, Behnke (1965a) has called the same tissues “connect-
ing sieve-tubes.”’ Essentially, these tissues are composed of somewhat
funnel-shaped, thin-walled cells having numerous small simple pits at the
end walls adjoining the PH. They differ from ordinary sieve tubes in the
presence of sieve plates only at the end adjoining the sieve tubes. The
glomerulus sieve-tubes adjoin the cells that make up the bulk of the
phloem glomerulus. These cells were designated “phloem-glomerulus cells”
of the first (PH), second (PH), and third (PH;) orders by Braun
(1957). Similar cells had earlier been called “bast tubulus” and ‘“‘glomer-
ulus cells” for PH, and PHg orders, respectively, by Brouwer (1953).
PH, and PH: had also been called “Nodal sieve-tubes” and “Nodal
sieve-elements” respectively by Behnke (1965a). The PH; of Braun may
actually be the over-lapping ends of PH» and PHoel.
The phloem-glomerulus cells vary in length, and as was observed in the
case of the xylem-glomerulus, some of the cells of the phloem groups are
shorter than others. In this case, the cells of the PH, order vary from
20p to 60u in length, while those of PH» vary from 60, to 140,.
The cells of PH; and PHg have thin walls (about 1p thick) with simple
pits that can hardly be seen with a light microscope. Whether the walls
are interconnected by cytoplasmic threads (plasmodesmata) or by any
1969 | AYENSU, ANATOMY OF DIOSCOREACEAE 129
other mechanism has not been demonstrated with the light microscope.
Dr. Behnke of Bonn University informed me that his electron microscope
studies show that plasmodesmata are indeed present in the cells of PH;
and PH». His recent publications (Behnke, 1965a, b, c) support his find-
ings. It is, however, certain that these phloem cells are specialized and
differ from sieve tubes and sieve cells of ordinary phloem tissue. Micro-
chemical tests reveal the absence of starch-grains from the phloem-glo-
merulus cells; the surrounding parenchyma cells possess starch. The his-
tochemistry of the phloem will have to await critical studies.
Cleared and stained portions of young and old stems reveal that at the
node (Fic. 3) three major vascular bundles (LT) enter the petiole from the
stem through the node without joining other vascular bundles, coming
through the underlying internode as peripheral vascular bundles. These
leaf-trace bundles are V-shape
The vascular bundles of the stem axis lying in front of the point of
entry of the leaf traces, and those of the inner and outer circles become
enlarged and join to form the xylem and phloem glomeruli (X, PH).
These glomeruli lie obliquely above each leaf insertion at the same height
as the axillary bud (AB). Opposite the outer circle of the vascular
bundles in the internode they appear somewhat towards the outside and
project into the base of the axillary bud or the lateral shoot. Five cauline
vascular bundles leave a glomerulus into the internode above (CB), but
only two enter it from below (GB). The latter are the characteristic large
vascular bundles which are arranged in the gaps between the three leaf-
trace bundles, which lie on the inside of the stem furrows. Hence the five
cauline vascular bundles forming the circle are made up of the two vascu-
lar bundles from the glomerulus (GB) and the three vascular bundles of
the leaf-trace (LT).
The vascular bundles of the axillary buds come from the glomerulus
directly. Just after they leave the glomerulus, each divides into two (an
upper xylem branch and a lower phloem branch), which come from the
upper and lower regions of the glomerulus respectively. Occasionally the
lower phloem branch subdivides into two with one establishing itself above
the xylem branch.
DISCUSSION
The structure of the xylem and phloem glomeruli in the nodes of the
Dioscoreaceae seems to be unique amongst the monocotyledons. Futher-
more, the presence of tracheids and the distinct type of sieve elements in
the node has considerable implications regarding the evolutionary history
of these tissues in the angiosperms.
The anatomical studies of the xylem by Bailey and Tupper (1918)
showed that the most logical phylogenetic sequence is the derivation of
vessels from tracheids in the angiosperms. Cheadle (1943) working with
the xylem of monocotyledons confirmed Bailey’s work. In the light of
the above theory it is interesting to examine the developmental aspects
130 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
of the tracheal elements in the node of the Dioscoreaceae. The bulk of
the xylem glomerulus is made up of tracheids which are considered primi-
tive in the phylogenetic sense. Similarly, the cells of the phloem glo-
merulus are considered to be of a relatively primitive type (cf. Braun,
1957, and the papers he quotes).
It is significant that such a difference can occur in a stem with primitive
structures in the nodes and more advanced structures in the internodes.
Bailey (1956) stated that “It is now clearly demonstrated that evolu-
tionary modification of the xylem of stems and roots is not necessarily
closely synchronized with phylogenetic trends in the specialization of the
angiospermic flower. Either trend of evolution may be accelerated or
retarded in relation to the other.” The above can be extended with a
statement that vessel development in an individual part of an organ can
be delayed or advanced within that particular part as demonstrated in
the node and internode of the Dioscoreaceae respectively.
This study demonstrates that in the midst of the complex nodal vascu-
lar system lies an orderly and systematic mechanism that permits the
transport of assimilatory materials through the stems of the Dioscoreaceae.
However, any attempt to gain full understanding of the exact pathway,
and therefore, the movement of material through the phloem glomerulus
must first confirm the present observations which are based on a recon-
struction from serial microtome sections. A more reliable understanding
of the pathway will hopefully be gained when the writer is able to study
the vascular system of the Dioscoreaceae using the motion-picture analy-
sis technique employed by Zimmermann and Tomlinson (1965, 1967).
The complexity of the phloem glomerulus in the Dioscoreaceae raises
some fundamental questions about the current hypotheses on transport
mechanisms in plants. Esau, Currier and Cheadle (1957) summarized the
hypotheses as (a) mass or pressure flow; (b) mass flow together with
activities of parenchyma cells associated with the phloem that account for
the turgor gradients necessary for mass flow; (c) transport of solutes in
the sieve tube along protoplasmic interfaces; (d) accelerated solute move-
ments in sieve tubes resulting apparently from some special kind of
cytoplasmic movement or flow; (e) independent solute movement result-
ing from one or more as yet unknown active transfer processes that occur
in the sieve element cytoplasm.
The unique anatomical characteristics of the phloem glomerulus in this
family seem to suggest that perhaps more than one of the above methods
is responsible for the movement of assimilatory substances in the Dio-
scoreaceae. Arisz’s (1952) suggestion that every substance moves its own
way, and that different mechanisms may be involved in translocation
should be considered in the light of the anatomical variation in the phloem
of this family.
Although I have no proof as to the exact function of the phloem
glomerulus, it seems likely that rapid translocation is achieved by the
numerous cells that form the bulk of the nodal region.
1969] AYENSU, ANATOMY OF DIOSCOREACEAE 131
SUMMARY
The complex nodal anatomy which is unique and basically uniform in
the Dioscoreaceae, especially in Dioscorea and Tamus, is described. The
width of the two masses of tissues referred to as glomeruli is correlated
with the phyllotaxy in each species. The xylem-glomerulus is composed
of numerous short tracheids of various sizes and shapes which are closely
fitted together. The phloem glomerulus, whose construction is essentially
that of the xylem-glomerulus, consists of thin-walled cells without visible
pitting and sieve areas. Because of the presence of primitive xylem and
phloem structures in the nodes in contrast to more advanced structures
in the internodes, it is postulated that vessel development in an individual
part of an organ can be delayed or advanced within that particular part
as shown in the node and internode of the Dioscoreaceae respectively. The
peculiar nature of the vascular bundle glomeruli is presumed to have some
effect on the rate of fluid transport in the stem. It is suggested that
another technique, such as the motion-picture analysis method, should
be employed to study further the nodal structure and its relation to trans-
location.
ACKNOWLEDGMENTS
I am very grateful to Drs. P. B. Tomlinson, R. H. Eyde, and H. Robin-
son for reading the manuscript.
LITERATURE CITED
Arisz, W. H. 1952. Transport of organic compounds. Ann. Rev. Pl. Physiol.
3: 109-130.
AYENSU, E. S. 1965. Notes on the anatomy of the Dioscoreaceae. Ghana Jour.
Sci. 5(1): 19-23.
1966. Vegetative anatomy and taxonomy of the Dioscoreaceae. Ph.D.
Thes esis. University of London, pp. 383 [unpublished].
gee I. W. 1956. Nodal anatomy in retrospect. Jour. Arnold Arb. 37: 269-
. Tupper. 1918. Size variation in tracheary cells. I. A com-
Parison between the secondary xylems of vascular cryptogams, gymno-
BEHNKE, H.-D. 1965a. Uber das phloem der Dioscoreaceen unter besonderer
i. “thick Phloembecken. I. Zeitschr. Pflanzenphys. 53(2):
97-12
3 ed Uber den Feinbau “gitterartig” aufgebauter Plasmaeinschlusse
in den Siebelementen von Dioscorea reticulata. Planta [ Berl.| 66: 106-112.
mit besonderer Beriicksichtigung eines neuartigen Typs assimilateleitender
Zellen. Ber. Deutsch. Bot. Ges 322.
Brouwer, R. 1953. The arrangement of the vascular bundles in the nodes of
the Diowcsireaceec: Acta Bot. Neerl. 2: 66-73.
132 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
BurkitL, I. H. 1949. The ontogeny of the stem of the Common Bryony,
Tamus communis Linn. Jour. Linn. Soc. Bot. 53: 313-382.
CuHeapie, V. I. 1943. The origin and certain trends of ir apse of the
vessel in the si ea a Am. Jour. Bot. 30:
Coursey, D. G. 1967. Yams. Longmans, London, pp. 230.
Esau, K., H. B. canes. & V. I, CHEADLE. 1957. Physiology of phloem. Ann.
Rev. Pl. Physiol. 8: 349-374.
FALKENBERG, P. 1876. Vergleichende Untersuchungen iiber den Bau der Veg-
etationsorgane der Monocotyledonen. Stuttgart. p. 202.
Harr, H. 1950. Die Dioscoreaceen-Knoten. Staatsexamensarbeit, Miinchen
[unpublished].
Mason, T. G. 1926. Preliminary note on the physiological aspects of certain
undescribed structures in the phloem of the great yam Dioscorea alata
Linn. Proc. Roy. Dublin Soc. 18: 195-198.
ZIMMERMANN, M. H., & P. B. ToMLINson. 1965. Anatomy of the see Rhapis
excelsa, I. Mature vegetative axis. Jour. Arnold Arb. 46: 160-178.
& . Anatomy of the palm Rhapis excelsa, W. Vascular
development in apex of vegetative aérial axis and rhizome. /bid. 48: 122-
Hela oF BoTANY
SMITHSONIAN INSTITUTION
WasHINGTON, D.C. 20560
EXPLANATION OF PLATES
4-12. Longitudinal sections of the stem ee region illustrating the
copaetec of the xylem and phloem glomeruli, « 8
PLATE I
Fic. 4. Dioscorea hirtifora Benth., showing an example of the meeting se
between the phloe Map tale a cells (PH.) of the second order, and a trans-
verse section a a gece tu
scorea discolor Kunt h, interlacing of xylem glomerulus cells.
Arrow points es a transverse section of a vessel (V) just entering the node.
PLA sen ne
Fie: '6. Rrmcaage? cies gh ana Hochst., exhi oh faa orientation of
xylem and phloem glomeruli. Vessel Gstieal (vi glomerulus of the
first (PH,) and ee (PH.) orders. Xylem Aare galls (XG), phloem
glomerulus cells (PHG).
FI Dioscorea multiflora Mart., showing a vessel element (V) and xylem
glomerulus cells (XG).
PLATE III
Fic. 8. Dioscorea —— Schauer, showing transverse sections of phloem
glomerulus cells (PH
Fic. 9. De composi ita Hemsl. (D. tepinapensis Uline ex Knuth).
Arrows pointing to vessel (V), vessel-tracheid (VT) and xylem anmeraiun
in transverse section (XG).
Fro. 1
a ome elem
Dioscorea dregeana (Kunth) Th. Dur. & Schinz, end plate of vessel-
fdea (VT) and phloem glomerulus cells (PHG).
PLAT
so pentaphylla L., she hbo of a vessel-tracheid (VT) and
PLATE V
12. Tamus communis L., exhibiting the presence of xylem (XG) and
pret (PHG) glomeruli.
Jour. ARNOLD Ars. VoL, 50
AYENSU, DIOSCOREACEAE
Jour. ARNOLD ARB. VOL. 50 PriateE II
AYENSU, DIOSCOREACEAE
Jour. ARNOLD ARB. VOL. 50 PLaTE II
AYENSU, DIOSCOREACEAE
Jour. ARNOLD Ars. VoL. 50 PuaTe IV
opt
bgt
ba tom -
ie
CY ff he
y s& ~ et
Fg {*
OY Like
F
ie
AYENSU, DIOSCOREACEAE
PLATE V
Jour. ARNOLD Ars. VoL. 50
AYENSU, DIOSCOREACEAE
138 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ANATOMY OF THE PALM RHAPIS EXCELSA, VII. FLOWERS *
N. W. UHL, L. O. Morrow, anv H. E. Moore, Jr.
Tur GENus Rhapis is one of a group of six genera in the subfamily
Coryphoideae centered in the southeastern United States (Rhapidophyl-
lum) and southeastern Asia (Liberbaileya, Maxburretia, Trachycarpus,
Rhapis) with a Mediterranean outlier (Chamaerops). These genera are
notable for complete apocarpy coupled with an apparently specialized in-
florescence (relative to the subfamily as a whole), polygamy or dioecism,
and slight to marked morphological distinction between staminate and
pistillate flowers. Among them, Rhapis appears to be most highly special-
ized in having greater dissimilarity between staminate and _pistillate
flowers and a gamophyllous corolla.
That perfect flowers may sometimes occur is suggested by the forma-
tion of apparently normal seed on an isolated pistillate plant at Cornell
University and further by the comments of Tomlinson and Zimmermann
(1968). In young stages of pistillate flowers, anthers appear normal but
in the mature flowers they are small and do not normally contain pollen.
There is a basic similarity between staminate and pistillate flowers in
young stages. Differences — functional versus abortive carpels and anthers
and elongate staminate corolla tube—are obvious only in later and
mature stages. The nature of most palms makes morphogenetic experi-
a pot plant in greenhouses. When the present anatomical series is com-
plete, it may prove an excellent subject for studies of development and
morphogenetic experiments on different aspects of flowering. The purpose
of this paper is to describe the anatomy of the two morphological types
of flowers in order to continue the anatomical series on Rhapis, to add to
a survey of floral anatomy in palms, and to provide the basis for further
WOrkK.
MATERIAL AND METHODS
A partial description of the floral anatomy of Rhapis was previously
prepared by one of us (Morrow, 1965) from collections vouchered by
* This study was undertaken at the request of and in collaboration with Drs.
P. B. Tomlinson and M. H. Zimmermann. We would like to thank them for this
appeared in Jour. Arnold Arb. volumes 46 (1965), 47 (1966), 48 (1967), and 49
(1968).
1969} UHL, MORROW, & MOORE, RHAPIS EXCELSA 139
Read 701 and 774. Further study of this material and of new collections
(Moore & Uhl 9561, 9562) has resulted in the more complete description
presented in this paper. Staminate and pistillate flowers (Moore & Uhl
9561, 9562) at anthesis were cleared and sectioned as described previously
(Uhl, 1966). Serial sections were studied in polarized light and by cine-
matography (Zimmermann & Tomlinson, 1965), a technique we are find-
ing most useful for analyzing flowers where many bundles are present.
RACHILLAE
As described in a previous paper of this series (Tomlinson & Zimmer-
mann, 1968), both staminate and pistillate inflorescences of Rhapis are
small panicles with up to three orders of branching. Bract to branch
relationship, although somewhat obscured by adnation, reveals a simple
monopodial system similar to that described for Nannorrhops ritchiana
(Tomlinson & Moore, 1968). Flowers are inserted in irregular spirals
(Fics. 1, 7) on branches of the first, second or third orders and on the
terminal part of the main axis, these axes being rachillae as defined by
Tomlinson and Moore (1968).
2
Fics. 1-17. Fic. 1, portion of staminate rachilla, x 2; Fic. 2, staminate
flower in vertical section, x 8; Fic. 3, stamin ah flower, x 4; as as — swt
Calyx, X 4; Fic. 5, staminate flower, calyx ved, X 4 6, stam
flower expanded, X 4; FiG. 7, portion of pistillate ictal, x - om 8, Distillate
flower, x 4; Fic. 9, pistillate calyx, exterior view, X 4; Fic ‘10, pstillate —
foteeiae view, x 4: Fic. 11, pistillate flower, calyx remov FI
Pistillate flower, vertical section, < 8; Fic. 13, gynoecium, x 8:
Synoecium, vertical section, X 8; Fic. 15, carpels in transection, X 8; nh fas
ie bead. dorsal view, X 8; Fic. 17, one ports ventral view, X 8. DETAILS:
r, bractl
140 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Pistillate inflorescences (Fic. 7) have fewer, commonly shorter branches
than staminate, with flowers more widely spaced (1-2 mm. apart). In
staminate inflorescences (Fic. 1), third order branches are more common,
often longer, and flowers are more crowded (0.5—1 mm. apart), sometimes
opposite, or in pairs. Many more flowers are produced in a staminate
than in a pistillate inflorescence (TABLE 1).
Anatomy. Anatomically all rachillae are similar. Epidermal cells are
small with rounded to slightly papillose outer walls. A thin cuticle is
present. The cortex is moderately wide and of unspecialized parenchyma
cells which increase in diameter centripetally. Some of the cells contain
tannins. The vascular complement consists of both large and small
bundles. Each larger bundle has one or two large vessels, a single phloem
strand, and a fibrous sheath four to five cells wide next to the phloem
and two to three cells wide next to the xylem. Smaller bundles have fewer
vascular elements; a few may contain only a phloem strand or be com-
pletely fibrous. In general these axes differ from the main axis in having
less lignified ground tissue and fewer cortical fibrous strands.
There is a definite arrangement and orientation of axial bundles in
rachillae of many palms. In Rhapis, a transection of a rachilla at any
level shows some large central bundles, one or two peripheral groups of
smaller bundles, and some scattered fibrous or very small vascular bundles
in the inner cortex. This configuration is easily explained in terms of
origin of bundles to the flowers.
Slightly below and opposite a floral insertion, six to ten axis bundles
branch (Fic. 18, fls) to form the bundles supplying the flower. A single
axial strand may produce one to four small branches in close vertical
succession or in a horizontal plane. Commonly the vertical derivative
continues as an axial bundle; however all branches of a bundle may be-
come floral traces. The peripheral clusters of small bundles are traces
to higher flowers; consequently, the number of small bundles varies de-
pending on the proximity to a floral insertion.
One or two axis bundles as well as branches from many others extend
directly into each flower. The total number of bundles in a rachilla is thus
progressively reduced distally (TABLE 1). Bundles in the axis branch fre-
quently, providing the numerous traces to flowers. Absolute numbers of
bundles are difficult to determine because bundles branch frequently, the
levels at which bundles are counted cannot be considered perfectly com-
parable, and fibrous sheaths of main strands and branches are often con-
fluent. Mere vigor or order of the branch may also affect the number
of bundles in a rachilla. However, the number of bundles in floral stalks
and organs seems to vary within definite limits. Approximately 6 to 9
bundles are present in staminate floral stalks below the abscission zone,
and a larger number (20-25) in pistillate floral stalks.
Rachillae are not terminated by flowers. In pistillate branches 4
rounded or pointed projection of the axis extends beyond the flower; some
14 to 16 vascular bundles are present in this reduced tip. Staminate
1969]
triangular group of bun
stalk;
UHL, MORROW, & MOORE, RHAPIS EXCELSA
abs, abscission zone;
oh
wn
pe, petal traces; se, gee ra, rachilla.
fis,
142 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
rachillae usually end less abruptly, one to seven abortive flowers being
present. Bracts subtending these abortive flowers are more prominent
than bracts of normal flowers, which are often obscured as the axis and
flower increase in size. A difference in growth patterns is suggested in the
two types of inflorescences. More branches and more flowers per branch
are formed in staminate inflorescences, suggesting that factors affecting
branch and floral initiation are more active and that cessation of growth
is less abrupt.
TABLE I. Flowers and Bundles per cm. of Length in Rachillae
, Pistillate rachilla Staminate rachilla
1 cm. intervals,
base to apex bundles/cm. flowers/cm. bundles/cm. flowers/cm.
base 50 42
1 43 4 36 3
2 40 3 3D 7
Bi 40 + 35 10
- 39 4 Sl 9
5 ot 3 28 15
6 16 1 28 12
7 16 26 13
9 19 14
10 12 13
11 5 7 abortive
PISTILLATE FLOWER
Bracts. Each pistillate flower is subtended by a bractlet (Fic. 7, br).
Bractlets subtending basal flowers on rachillae may be larger than bract-
lets of distal flowers which are usually small, crowded between the flower
and the axis, and apparent only when flowers are detached (Fic. 7). A
small trace, originating as a branch of an axis bundle, is usually present
in the bractlet. One or more floral traces may originate from the same
stelar bundle from which the trace to the bractlet diverged at a lower level.
Morphology (Fics. 7-17). Although considerable connation and ad-
nation are present in floral organs, a 3-3—6-3 floral plan is obvious both
morphologically and anatomically. Sepals of pistillate flowers (Fics. 8,
9, 10) are connate forming a shallow parenchymatous cup about 1 mm.
high with three pointed lobes 1-1.5 mm. long. The three petals (Fic. 11)
are also connate for approximately 3 mm., above which the free lobes
are briefly imbricate and then valvate reaching an additional length of
1-2 mm. The staminodes (Fic. 12) resemble the stamens in staminate
flowers but are smaller. The filaments are linear, adnate to the petal tube
for 1 mm., and free above that for about 0.5 mm. In the material studied
the reduced anthers did not produce pollen.
1969 | UHL, MORROW, & MOORE, RHAPIS EXCELSA 143
The three separate carpels (Fic. 13) are wedge-shaped with flat ventral
sides and rounded and grooved dorsal sides (Fics. 13, 16). Each carpel
has a distinct stalk which is fused with the petal-staminode tube for a
very short distance basally (Fic. 24). A locule with a single basal ovule
occupies the lower half of the carpel (Fics. 12, 14). Distally the style
is wide; the upper part is distended abaxially and converges abruptly
toward the ventrally situated, conduplicate, tube-shaped stigma (Fic. 28).
Thus the styles of these carpels are enlarged and are also histologically
specialized, as described below.
A single, hemianatropous ovule with a large funicular aril is attached
basally in the ventral angle of the locule (Fic. 27). There are two integu-
ments which are free for about 1/3 the length of the ovule. The outer
integument is six to seven cells wide and increases to about nine cell
layers around the micropyle. The inner integument consists of two cell
layers and is widened to three to four cells around the micropyle to form
a short beak. The inner layer of the inner integument is specialized as
an integumentary tapetum.
Anatomy (Fics. 18-32). Pistillate flowers appear to be sessile (Fic.
7). Anatomically, however, a very short stalk with a distinct group of
floral traces, can be recognized (Fic. 19). As explained above, the major-
ity of the bundles of the floral axis originate as branches from strands
in the rachilla, one or two of which also extend directly into the flower
without branching. The number of bundles supplying the pistillate flower
(Fic. 19) is about 23
An abscission zone forms a characteristic feature of floral stalks of both
staminate and pistillate flowers (Fic. 18, abs). This zone is distinguished
by the absence of fibrous bundle sheaths and by smaller ground paren-
chyma cells (Fic. 20) through which bundles can be followed.
Generally, in palm flowers, even when organs are connate, the origin
of their traces indicates a spiral insertion. This is not apparent in the
Sepals or petals of Rhapis. Directly above the abscission zone, most
bundles of the floral axis branch at about the same level (Fic. 21) to
form about 30 sepal traces. The origin and horizontal divergence of so
many bundles at one level results in a collar-like complex in which inner
bundles extend radially between outer strands and some lateral fusion of
bundles occurs (Fic. 21). Individual bundles may be followed through
this complex. Ficure 38 is a radial plot of a single major bundle of the
floral axis. The sepal trace (se 1) originating from this bundle branches
to form three other sepal traces (Fic. 39, se 2, se 3, se 4) and these
bundles in turn branch forming the continuing vertical bundles VB 2,
VB 3, and VB 4. Smaller (minor) bundles of the floral axis may produce
only a single sepal trace or extend directly into the sepal.
Above the sepal complex about 30 vertical bundles form a central group
(Fic. 22). Some 30 to 40 petal traces diverge at an acute upward angle
(30° to 40°) from these as opposed to a near 90° angle of divergence
for sepal traces (Fic. 18). Smaller vertical bundles (Fic. 39, VB 2 and
144 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
VB 3) may extend directly into a petal without branching. Most petal
traces, at the level of their origin, contain phloem only and fibrous sheaths
of main bundles and branches are often confluent (Fic. 23, pe). At higher
levels where traces are separate, a few scalariform xylem elements are
present. Sclerenchymatic sheaths of petal traces are thinner walled than
those of sepal traces. As in the sepals, a few lateral bundles may branch
and a median and two lateral veins extend into each petal tip.
38. 24-27. Pistillate flower, continued. Fic. 24, transection through stalks
of the three carpels, outer ring of bundles are petal traces, inner six large
bundles supply staminodes, all bundles in carpel bases are provascular, X 18;
Fic. 25, transection through petal-staminode tube and three carpels at level of
funicular attachments, X 18; Fic. 26, transection showing petal-staminode tube
and expanded styles of carpels, x 18; Fic. 27, longitudinal section of one carpel,
35. DETAILS: ar, aril; cs, carpel stipe: pe, petal trace; stm, staminode trace.
1969 | UHL, MORROW, & MOORE, RHAPIS EXCELSA 145
Fic. 28. Three dimensional drawing of one carpel to show vascular supply.
Dorsal and ventral bundles labeled, remaining are lateral bundles. Seven lateral
bundles are not completed for clarity, ca. 50. For details see Fics. 29-32.
About 20 relatively large receptacular bundles, each with a complete
fibrous sheath (Fic. 23, central bundles) are present above the origin of
the petal traces. Just above this level, considerable reorientation and
146 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
branching of strands takes place. Traces to staminodes (Fic. 24, stm)
are formed, often as a central branch of a trifurcating receptacular bundle.
Traces to antisepalous staminodes diverge at a slightly lower level than
those to the antipetalous ones. The remaining bundles become oriented
into three groups, one group representing the supply to each carpel.
Fibrous bundle sheaths extend as far as the stalk of each carpel but
are absent in the carpel base where all bundles are procambial. Three
or four of the bundles in each carpel base are Jarger than the remainder
and possess xylem elements which are birefringent. The central of these
larger strands extends across the carpel base and distally around the
locule to the base of the stigma (Fic. 28, db). Two of the other larger
strands remain in ventral positions (Fics. 28, 29-32, vb). Thus a dorsal
Fics. 29-32. Series of transections through the base of one carpel drawn with
Wild M20 research microscope and drawing tube, to show origin of ovular
supply. Ovule traces shaded, bundles with birefringent xylem shown divided,
> 60. Derarts: db, dorsal bundle; Ib, lateral bundle; Ic, locular canal; vb,
ventral bundle.
1969 | UHL, MORROW, & MOORE, RHAPIS EXCELSA 147
and two ventral bundles can be recognized by size, position, and maturity
(Fics, 29-32). Remaining strands form the 20 to 24 lateral bundles
present in an irregular ring in the carpel wall (Fics. 31, 32). Four of
these are larger and more mature (Fics. 31, 32, lb). Some lateral bundles
fuse with others near their upper limits, the major ones extending toward
the locular canal (Fic. 28). The ventrals extend slightly higher and the
dorsal ends just below the stigma (Fic. 28).
The origin of the vascular supply to ovules varies in palms (Uhl, un-
published). In Rhapis, a branch from the dorsal bundle, a branch from
each of four or five lateral bundles, and a branch from one ventral bundle
form a group of strands (Fics. 29-32) which extends into the funiculus
(Fic. 28). These strands fuse near the chalaza and the resulting large
with traces derived from dorsal and ventral carpellary bundles are not
frequent. Such taxa occur in groups usually considered to be primitive,
as Magnoliaceae (Canright, 1960) and Nymphaeaceae (Moseley, 1961).
STAMINATE FLOWER
Morphology (Fics. 1-6). A comparison of staminate and pistillate
flowers shows both differences and similarities. Bractlets are alike in both
types of inflorescence. Sepals in the two flowers (Fics. 3, 4) are also
Similar in shape and size; those of staminate flowers are perhaps slightly
less fleshy. Petals are about the same length (4-5 mm.) and are 2/3 to
3/4 connate (Fics. 3, 5). In staminate flowers, however, the petal tube
is definitely obovoid or clavate and much less fleshy than that in pistillate
flowers (cf. Fics. 8 and 11 with 3 and 5). The diameter of the petal
tube immediately below the free lobes is approximately 2 mm. in staminate
flowers and 3 mm, in pistillate flowers. Staminate petals are valvate and
often incompletely connate, a groove of varying depths showing the limits
of each petal. In pistillate flowers, however, the petal-tube is smooth
and free lobes are briefly imbricate. Filaments (Fic. 6) are wider in
Stamens than in staminodes and bear well-developed, latrorse anthers with
dark, tannin-containing connectives (Fics. 33, 37). Three very tiny
vestigial carpels are present (Fics. 33, 35, vc).
148 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Fics. 33-37. Staminate flower. Fic. 33, longitudinal section, X 18; Fic. 34
cleared staminate flower, levels of riGuRES 35~37 indicated by one Op sbe beo
5* Fic. 36,
numbers, * 10; Fic. 35. transection through base of flower, 3
transection through petal-stamen tube, * 18; Fic. 37, tr cota ES of ae part
of flowe <18. DeETAILs: pe, petal traces; se, sepal; st, stamen trace;
vestigial t carpel.
1969 | UHL, MORROW, & MOORE, RHAPIS EXCELSA 149
14404
1080-
720+
360-
in microns
Length
60
40 240 360 720
Distance from center of axis in microns
Fic. 38. Diagram of the radial path of a major betyrs ¥ hime floral axis.
woe c, carpel traces; pe, petal trace; se, sepal trace; stm, staminode trace
VB 1, ontinuing vertical bundle. Dotted lines indicate any Ghrou bundle
sheaths ; are conflue
bundles. About three strands remain in the floral receptacle above the
origin of the stamen traces. These disappear just below the vestigial
carpels.
Histology. Histological features in floral organs are sometimes diag-
nostic in palms (Uhl, unpublished). In R/apis, tannins are present ran-
150 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
domly in sepals and filaments, near the adaxial surfaces of petals and in
all cells of connectives. Fibrous bundle caps are lacking in receptacular,
lower petal, and stamen bundles of staminate flowers; in carpels; and in
abscission zones of both flowers. It is perhaps significant that there are
few, if any, crystals in fleshy sepal bases and petal tubes. The abaxially
distended styles are histologically the most specialized parts of the flowers,
containing raphides, tannin cells in radial rows, and distal, cap-like layers
of sclereids (Fic. 26).
DISCUSSION
Comparison with vegetative organs. Emphasis in this series on
Rhapis has been on vascular pathways throughout the plant. Careful
analysis of the flowers shows continuity of bundles from those of the
se |
Fic. 39. Part of a transection, drawn with the Wild M20 research microscope
and drawing tube, to show the continuation of the sepal trace (se 1) diagrammed
in Fic. 38; se 2, se 3, and se 4 are sepal traces derived as branches of se 1.
Each of these branches to form a continuing vertical bundle (VB 2, VB 3, and
VB 4) as indicated, 125.
1969] UHL, MORROW, & MOORE, RHAPIS EXCELSA 151
rachilla to the ovule or stamen. The pattern of origin is a simple one.
Bundles of the floral axis, derived as branches of rachilla bundles, branch
in turn at appropriate levels to provide traces to sepals, petals, staminodes
or stamens, and carpels.
This vascular continuity throughout RAapis, which is now completed
in the description of floral vasculature, shows a similar pattern throughout
every kind of axis on the plant (e.g. seedling, aérial axis, rhizome, inflores-
cence axis, rachilla, and pedicel). The same principle of vascular organ-
ization is expressed in the flower, but it is somewhat more difficult to
recognize here than in the vegetative organs because the floral axis is
condensed and the lateral organs are small. Nevertheless we may say that
the divergence of traces to sepals, petals, and staminodes or stamens,
involving axial continuity, is comparable to the departure of leaf traces in
rhizome and aérial stem (Tomlinson & Zimmermann, 1966; Zimmermann
& Tomlinson, 1965). This is most obvious when an individual bundle is
followed through the floral axis. The radial path resulting from such an
analysis is presented diagrammatically in Ficure 38. In addition very
short bundles, which may be interpreted as bridges (Zimmermann &
Tomlinson, 1965), often link diverging traces with bundles of the re-
ceptacular system. More detailed comparison of floral and vegetative
vascular pathways must await a more complete understanding of mono-
cotyledonous vascular development.
Comparison with other palms. Among coryphoid palms, Rhapis
may be considered intermediate in specialization. The connation in sepals
and petals and corresponding derivation of sepal and petal traces in whorls
are evidences of specialization, as is also the adnation of stamens and
staminodes to the petal tube. Several features of the carpel are note-
worthy. Completely free, stipitate, spirally inserted carpels are con-
sidered primitive in palms and angiosperms. However, the large dorsally
extended styles and completely closed ventral sutures of Rhapis indicate
specialization. The orientation of the ovule is intermediate between the
primitive anatropous and the most advanced orthotropous position. The
multiple derivation of traces to the ovule from the dorsal, several laterals,
and a ventral carpellary bundle suggests laminar placentation (Eames,
1961) and may be a basic pattern in palms. In a preceding paper of this
series it was stated that Rhapis has a relatively specialized inflorescence
(Tomlinson & Zimmermann, 1968). Similarly it may be said that among
the Coryphoideae the flowers are relatively specialized.
LITERATURE CITED
Canricut, J. E. 1960. The comparative ey a relationships of the
Magnoliaceae. III. Carpels. Am. Jour. Bot. 47: —155.
Eames, A. J. 1961. Morphology of the ey aa -Hill. N.Y.
Morrow, L. O. 1965. Floral morphology and anatomy of certain Coryphoideae
(Palmae). Ph.D. Thesis. Cornell Univ. [Unpublished].
152 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Mose ey, M. F. 1961. Morphological studies of the Nymphaeaceae. II. The
flower of eae Bot. Gaz. 122: 233-259.
Tomitnson, P. B. & H. E. Moore, Jr. 1968. Inflorescence in Nannorrhops
ritchiana fies Jour. Arnold Arb. 49: 16-34
Tom.inson, P. B. & M. H. ZIMMERMANN. 1966. Anatomy of the palm Rhapis
excelsa, II. Rhizome. Jour. Arnold Arb. 47: 248-261.
& . 1968. Anatomy of the palm Rhapis excelsa, V. Inflorescence.
Ibid. 49: 291-306.
Unt, N. W. 1966. Morphology and anatomy of the inflorescence axis and
flowers of a new palm, Aristeyera spicata. Jour. Arnold Arb. 47: 9-22.
ZIMMERMANN, M. H. & P. B. TomLinson. 1965. Anatomy of the palm Rhapis
excelsa, I. Mature vegetative axis. Jour. Arnold Arb. 46: 160-180.
L. H. Battey Hortortum
CORNELL UNIVERSITY
IrHaca, NEw York 14850
(Uhl and Moore)
A
ND
RANDOLPH MACON COLLEGE
ASHLAND, VIRGINIA
(Morrow
7 1969 | MITRA & SUBRAMANYAM, GLYCOSMIS 153
GLYCOSMIS PENTAPHYLLA (RUTACEAE) AND
: RELATED INDIAN TAXA
| R. L. Mirra AND K. SUBRAMANYAM
THE PUBLICATION of a new series, Limonia arborea, by Roxburgh (PI.
Coromandel. 1: 60. ¢. 85. 1798) and his providing the plant which he
believed to be “Limonia pentaphylla Retz.” (Roxb. loc. cit. t. 84) with
a detailed description and illustration, as well as the subsequent discovery
of the authentic type specimen of Limonia pentaphylla Retz. by Tanaka
(Bot. Not. 1928: 156-160. 1928), has led to some controversy in the
, nomenclature of these two species now included in the genus Glycosmis.
In the interest of clarity, relevant parts of the earlier works are reviewed
in brief.
Tanaka (loc. cit.) pointed out that Limonia pentaphylla Retz. is con-
specific with Limonia arborea Roxb. and is entirely different from the
plant treated by Roxburgh as “Limonia pentaphylla Retz.” He therefore
treated Glycosmis arborea (Roxb.) Correa (= Limonia arborea Roxb.)
as a synonym of Glycosmis pentaphylla (Retz.) Correa (= Limonia
pentaphylla Retz.), and in Botaniska Notiser (1928: 159. 1928) proposed
Glycosmis mauritiana (Lam.) Tanaka (= Limonia mauritiana Lam.) for
the plant erroneously treated by Roxburgh as “Limonia pentaphylla
Retz.”
, Narayanswami (Rec. Bot. Surv. India 14(2): 26. 1941) did not agree
; with Tanaka’s view and maintained Limonia pentaphylla Retz. and
Limonia arborea Roxb. as distinct from each other; accordingly the cor-
rect names in the genus Glycosmis should be G. pentaphylla (Retz.)
Correa and G. arborea (Roxb.) Correa, respectively. Brizicky (Jour.
Arnold Arb. 43: 88. 1962) upheld Tanaka’s view on the conspecificity of
Limonia pentaphylla Retz. and Limonia arborea Roxb. and remarked,
“Narayanswami (1941), apparently having overlooked Tanaka’s article on
the type of Retzius’ species, came to the conclusion . . . that Tanaka’s
interpretation of L. pentaphylla was entirely incorrect. . ..” Brizicky
also pointed out that De Candolle (Prodr. 1: 538. 1824), instead of Correa
(Ann. Mus. Hist. Nat. Paris 6: 386. 1805), should be assigned the author-
ship of these two binomials, G. pentaphylla and G. arborea, since De Can-
: dolle made these combinations for the first time in the sense of the Code.
However, Brizicky’ s conclusion on their nomenclature is untenable,
not being in accordance with the existing Code. Brizicky (Joc. cit.,
P. 87) is of the opinion that “...... Glycosmis pentaphylla DC.
was based on the plant identified and illustrated by Roxburgh as
‘Limonia pentaphylla Retzius’ and only questionably on Retzius’ species
154 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ea eee, . . . Limonia pentaphylla Retz. obs. 5. p. 24? Roxb.
84.’).” Brizicky (loc. cit., p. 89) further argues, Then Gly-
cosmis pass: DC., based on Roxburgh’ s plant, not on that of Retzi-
us, must be regarded not as a new combination, but as a new name in
combination G. mauritiana (Lam.) Tanaka... Since G. pentaphylla
DC. cannot be applied to Retzius’ Limonia pentaphylla, the next available
name for the latter species is Glycosmis arborea (Roxb.) DC.” In treating
Glycosmis pentaphylla as a new name and not as a new combination
Brizicky was probably applying the provisions of Art. 72. However, this
article is not operative in this case; it is clear from Roxburgh’s treatment
of “Limonia pentaphylla Retz.” that he was not describing a new species
under a homonymous name, but was only misidentifying Retzius’ plant.
hus, the question of De Candolle’s basing the binomial G. pentaphylla
on “Roxburgh’s plant — Limonia pentaphylla’ does not arise. Moreover,
De Candolle, in making the transfer (Prodr. 1: 538. 1824), gave a direct
reference to Retzius’ plant, though with a question mark, “.... - -
Limonia pentaphylla Retz. obs. 5. p. 24? Roxb. cor. |. t. 84.” It is evident
from above that De Candolle was not certain about the identity of the
two plants involved in the confusion. Hence, Brizicky’s argument for
bag
menclature (ed. 1966) clearly states, “When, on transference to another
genus, the specific epithet has been applied erroneously in its new position
to a different species, the new combination must be retained for the species
*Dr. Brizicky, who died on June 15, 1968, saw an earlier but hardly different
version of this paper and, on May 4, 1968, set down the comments which follow.
These comments were duly communicated to the authors of this paper, who are
pee not agreeable to the arguments placed by Dr. Brizicky.
e authors of this paper believe that my view of Glycosmis pentaphylla DC. as
a new name, rather than combination, is untenable from the standpo int of the Code,
making transfer, gave a direct reference to Retzius’ plant, “Limonia pentaphylla Retz.
. 5. p. 24?.. .”’ Curiously, though, applying the Code mechanically [Art. 557],
the authors disregard the question mark which follows the complete citation of
a ri
sumed to be possessed by a taxonomist who publishes a new combination, new status,
1969] MITRA & SUBRAMANYAM, GLYCOSMIS 155
to which the epithet was originally applied, and must be attributed to the
author who first published it.” Glycosmis pentaphylla (Retz.) DC., there-
fore, must be retained as a new combination based on L. pentaphylla Retz.
The nomenclature of the relevant taxa follows:
Glycosmis pentaphylla (Retz.) DC. Prodr. 1: 538. 1824, quoad
basionym; Tanaka, Jour. Indian Bot. Soc. 16: 229. 1937.
ie pentaphylla Retz. Obs. Bot. 5: 24. 1789.
Limonia arborea Roxb. Pl. Coromandel. 1: 60. t. 85. 1798.
G. pe ee (Roxb.) DC. Prodr. 1: 538. 1824; Narayanswami, Rec. Bot. Surv.
India 14(2): 20. 1941; Brizicky, Jour. Arnold Arb. 43: 90, 1962
oe pieces es var. linearifoliola Tanaka, Jour, Indian Bot.
: 230. 1937, “linearifoliolis.”
G. arborea var. linearifoliola (Tanaka) Narayanswami, Rec. Bot. Surv. India
14(2): 26. 1941, “linearifoliolata.’
Glycosmis mauritiana (Lam.) Tanaka, Bot. Not. 1928: 159. [4 Apr. ]
1928; Bull,
, A 2a
excl. syn. G. june DC
Limonia mauritiana Lam. Encycl. Méthod. Bot. 3: 517. 1792.
Limonia pentaphylla auct. non Retz.: Roxb. Pl. Coromandel. 1: 60. ¢. 84.
1798.
G. Rnlaresicaty sensu Narayanswami, Rec. Bot. Surv. India 14(2): 12. 1941,
xcl. syn. L. pentaphylla Retz.
Tanaka proposed this combination in a manuscript sent to the Bulletin
de la Société Botanique de France on January 13, 1928. Ina subsequent
paper sent to Botaniska Notiser on February 18, 1928, he referred to his
etc. There certainly have been cases when because of unavailability of the ig pes and/or
ned o
of misinterpretation of some taxa the new combinations tur ave bee
made xa different from those for whi ey were intende os ceaare, in all
these cases the authors of the combinations sincerely believed that their taxa and
those the epithets of which were taken as io were identical (conspecific, con-
varietal, etc.) parently, there are so ene if any, combinations which :
on the epithets of doubtfully conspecific a that there has been no necessity to
mention them in the Code, and the ee ee those cases has been left to the good
t ‘
“Thus, since De Ca ndolle himself indicated that Limonia pentaphylla Retz. cries
t
as basionym, there is no reason to r d Glycosmis pentaphyl new combina-
tion based on Limonia pentaphylla Retz, On the contrary, pestis pentaphylla,
gesting a combination was not that, but a new name. Thus, A. Gray took Malvastrum,
the name of De Candolle’s section of Malva L., as the name of his genus without
having based the genus on De Candolle’s section. yeaa & Ree
156 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
proposed combination “Glycosmis mauritiana (Lamk.) Tanaka in Bull.
Soc. Bot. Fra with basionym and a few more synonyms; but this
latter paper appeared in print first. Tanaka’s awareness of this changed
situation is evident from his subsequent citation in the Journal of Botany
(68: 226. 1930).
Glycosmis mauritiana var. andamanensis (Narayanswami) Mitra &
Subr., comb. nov.
G. pentaphylla var. andamanensis Narayanswami, Rec. Bot. Surv. India 14(2):
16. 1941.
Publication of Glycosmis mauritiana var. andamanensis Tanaka (Jour.
Indian Bot. Soc. 16: 229. 1937) was not valid, for a Latin description was
not given.
Glycosmis mauritiana var. insularis (Kurz) Tanaka, Jour. Indian Bot.
Soc. 16: 229. 1937
G. arborea var. insularis Kurz, Jour. Bot. 14: 38. 1876, pro parte. ‘
G. pentaphylla var. insularis (Kurz) Narayanswami, Rec. Bot. Surv. India
14(1): 20. 1941.
Glycosmis mauritiana var. fuscescens (Kurz) Mitra & Subr., comb.
nov.
G. trifoliata var. fuscescens Kurz, Jour. Bot. II. 5; 37. 1876.
G. pentaphylla var. fuscescens (Kurz) Narayanswami, Rec. Bot. Surv. India
14(2): 20. 1941.
We could examine only one sheet present at CAL (King’s Collector
1884, acc. no. 74984) from which the diagram given by Narayanswaml
(Joc. cit. page 21, fig. 5) was drawn. We agree with Narayanswami in
treating this as a distinct variety.
Noses cymosa var. linearifoliola (Tanaka) Mitra & Subr., comb.
G. cyanocarpa var. aft ea Tanaka, Jour. Indian Bot. Soc. 16: 229.
1937, “linearifoliol
G. cymosa var. ecestbiia Narayanswami, Rec. Bot. Surv. India 14(2): 32.
1941.
DOUBTFUL TAXON
Glycosmis pentaphylla var. latifolia (Kurz) Narayanswami, Rec.
Bot. Surv. India 14(2): 20. 1941
We are doubtful whether this plant needs a distinct rank. The lone
sheet examined at CAL (Helfer 525, acc. no. 74981) has only two
twigs mounted on it, the inflorescence being altogether lost. The leaf
1969 | MITRA & SUBRAMANYAM, GLYCOSMIS 157
character of the right-hand specimen appears to be that of the Glycosmis
mauritiana var. andamanensis while the left-hand one is closer to G.
mauritiana var. insularis.
BOTANICAL SURVEY OF INDIA
76 ACHARYA JAGADISH Bose Roap
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CONTENTS OF NUMBER 2
Vascutark ANATOMY OF MOoNOCOTYLEDONS WITH SECONDARY
GrowTH — An Intropuction. P. B. Tomlinson and M. H.
Zimmermann ... 159
teeeeer
ASPECTs OF REPRODUCTION IN SaurautA. Djaja D. Soejarto ........
ipsa dled AN Erin Forest 1x Purrto Rico.
6. Aéptat Roots. A. M. Gill . 197
ae is: Root, AND Ranerwonm Reuationsuies. Walter H.
- Lyford ee
8, lg Gives as as ts a Se
TURE. Richard A. Howard 225°
JOURNAL
OF THE
ARNOLD ARBORETUM
VoL. 50 Aprit 1969 NUMBER 2
VASCULAR ANATOMY OF MONOCOTYLEDONS
WITH SECONDARY GROWTH — AN INTRODUCTION
P. B. TomMLINSON AND M. H. ZIMMERMANN
ARBORESCENT PLANTS with secondary growth from a vascular cambium
represent a small minority of monocotyledons. Nevertheless their sig-
nificance is out of proportion to their abundance because they exhibit a
growth habit comparable to that of familiar dicotyledonous and gymno-
spermous trees. Monocotyledonous secondary vascular tissue, however,
unlike that of other trees, includes discrete vascular bundles.
Earlier botanists in their study of palms and other arborescent monocot-
yledons devoted considerable attention to those few forms with secondary
vascular tissues and it is largely on the efforts of these nineteenth-century
anatomists that our present knowledge is based. It was therefore inevita-
ble that in our own studies of palms we should follow this earlier tradition
and turn our attention to monocotyledons with secondary growth, espe-
cially as we had found that early work on the vascular system of monocot-
yledons had been incomplete and furthermore had become misrepresented
in modern textbooks (Tomlinson & Zimmermann, 1966).
In the present article we review the vascular anatomy of monocotyledons
with secondary growth in order to provide the background for our observa-
tions which will be published separately. Such a lengthy and independent
introduction is justifiable because few botanists have first-hand familiarity
with these plants and the literature about them is old and not readily
available. Nevertheless our introductory review is necessarily very selec-
tive, because the early literature is extensive and some of it is no longer
very informative. We have largely cited those articles which contributed
significantly to knowledge about these plants. Reviews of the subject
do already exist. Those by Cordemoy (1894) and Cheadle (1937) are
lengthy. Cheadle’s paper is relatively recent and accessible, but it is re-
stricted to a study of small tissue samples. General organization, growth
habits, and overall distribution of vascular tissues are not discussed in it.
This is somewhat in contrast to the approach adopted by earlier genera-
tions, as indicated by Cordemoy’s review, and reflects the way in which
anatomists have lost sight of the plant as a functioning whole in their
160 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
preoccupation with histological detail. It is with the object of re-instating
the approaches of botanists concerned with overall aspects of growth and
construction that our own studies have been undertaken.
TAXONOMIC DISTRIBUTION
The occurrence of aborescent genera in certain families of monocoty-
ledons which possess secondary vascular tissues is indicated in TABLE 1. We
treat these genera in a very broad sense but appreciate that some, notably
Yucca (e.g. Trelease, 1902) have been subdivided. Two systems of classi-
fication are compared, that of various authors in Engler and Prantl (1930)
and that of Hutchinson (1959) in order to emphasize that the taxonomic
distribution of these plants is far from certain. We do not intend to dis-
cuss the evolutionary relationship between arborescent and herbaceous
monocotyledons, but we suggest that anatomical investigation is likely to
contribute significantly to a resolution of these problems. Of the listed
genera, Beaucarnea, Cordyline, Dasylirion, Dracaena, and Yucca consist
of species ranging in size from trees to shrubs and rhizomatous herbs. The
genus Aloé is predominantly herbaceous, but includes a few aborescent
TABLE 1. Taxonomic distribution of larger monocotyledons with
secondary vascular tissues.
ENGLER and PRANTL REPRESENTATIVE GENERA HUTCHINSON
AMARYLLIDACEAE AGAVACEAE
subfamily Agavoideae Agave, Furcraea tribe Agaveae
LILIACE
subfamily reais
tribe ceae Yucca tribe Yucceae
tribe Nolineae ron alte (Nolina) tribe Nolineae
Dasylirion
tribe Dracaeneae c oii, Dracaena tribe Dracaeneae
(Pleomele
subfamily Asphodeloideae LILIACEAE
tribe Aloineae 0é tribe Aloineae
Pen Ya RS Sec esi eNO
tribe Lomandreae Lomandra, Xanthorrhoea XANTHORRHOEACEAE
tribe Calectasieae Kingia
PSUR SSH Date INN Notte me eee ne No
TRIDACEAE IRIDACEAE
subtribe Aristeinae Aristea (Nivenia), Kiattia, tribe Aristeae
Witsenia
1969] TOMLINSON & ZIMMERMANN, MONOCOTYLEDONS 161
species, notably A. bainesti and A. dichotoma, Agave and Furcraea do not
really fit a strict definition of a tree although some species achieve mas-
sive proportions. The same seems true of the Xanthorrhoeaceae although
there is little information about their size, growth habit, and the extent
of secondary tissue except in the work of Floresta (1902). The iridaceous
genera are listed, although they are little more than shrubs, because sec-
ondary tissue is extensive and has been well described (e.g. Adamson,
1926; Scott & Brebner, 1893). On the other hand, we have omitted many
monocotyledons which possess a limited amount of secondary growth but
are otherwise essentially herbaceous. These include a number of genera
in the Liliaceae, like Aphyllanthes, Veratrum and others in Hutchinson’s
Agavaceae. Fleshy rhizomes with secondary tissues, as in the Dioscorea-
ceae, are also disregarded. Vascular tissue which is by definition secondary
may be quite common in other, unrelated, herbaceous monocotyledonous
families [e.g. Bromeliaceae, Krauss (1948); Musaceae, Skutch (1932);
Zingiberaceae, Chakraverti (1939) |] where it seems to be associated with
root insertion. However, before any major evolutionary significance can
be attached to secondary cambial activity, we must attempt to under-
stand it from a developmental point of view.
MORPHOLOGY
Growth habits (Fics. 1-11). Growth form is quite diverse although
it can be seen to depend on a common pattern of development. Leaves
are linear, usually rigid, often thick and fleshy. They are rarely distinctly
petiolate as in some smaller species of Cordyline and Dracaena. Axes are
made up of short, often very congested internodes. In slow-growing plants
this results in the characteristic terminal tufts of leaves or, if the main
axis is very much shortened, in the basal rosette which characterizes the
Agave-habit (Fic. 7). Branching is usually sparse; the reason for this
is discussed below. In Agave and Furcraea the vegetative axis may be
unbranched so that the plant is monocarpic. Otherwise the rosette in
these plants is propagated by basal and usually stoloniferous suckers.
Stoloniferous shoots are not usually present in other genera but they are
common in herbaceous relatives (e.g. Sansevieria). The habit of most
arborescent monocotyledons is quite tree-like, and some may even be
mistaken for a dicotyledonous tree by a superficial observer, as noted by
Wright (1901). However, some species of Dracaena, especially those in
its segregate genus Pleomele, look more like shrubs with their much-
branched crown and fine twigs (Fic. 8).
It is evident that shoot diameter on a single plant is not entirely
dependent on the amount of secondary growth. In smaller and much
branched species variation in crown diameter is considerable and seems
related to the vigor of the shoot. Basal, erect shoots are thickest and most
vigorous; distal horizontal shoots are narrow and least vigorous. We have
noted a range in primary shoot diameter of 6 to 30 mm. in Pleomele.
Some of the simpler growth forms can be looked upon as juvenile stages
162 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
fff ( Ml i
fi My
| SIO
P| WA Da aa i i} d AN a rs —
(N ) Yj ie fh t
: Nb
1 |
. /
WY ww, NY Y/
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VWNAN Ww si a Ree eee ee
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Growth habits in arborescent er eee with secondary
1-4. Beaucarnea recurvata.
Fics. 1-11.
thickening (all to approximately same scale). Fic
1969] TOMLINSON & ZIMMERMANN, MONOCOTYLEDONS 163
in the development of the larger forms, which are fixed permanently. De-
velopment of a large Beaucarnea (Fics. 1-4), for example, begins with a
rapidly growing main axis which remains unbranched for several years.
Leaves may be long persistent so that they clothe the axis of quite tall
specimens. Many species of Yucca do not develop much beyond this stage
(Fic. 10). A link between the specialized rosette of Agave and this juve-
nile habit is provided by a number of species of both Agave and Furcraea
with relatively tall stems (e.g. F. longaeva illustrated in Engler & Prantl
(1930) p. 419). Otherwise, normal development of the tree form con-
tinues with branching, the loss of leaves from the older stem parts, thick-
ening of the base of the stem, and development of a fissured bark. The
evolutionary relation between ontogeny and phylogeny is suggested by
Cordyline in New Zealand. Cordyline indivisa can be equated with the
unbranched juvenile stage of C. australis (Fic. 9) and in turn the low
rosette of C. pumilio with a younger stage still.
A disproportionate thickening of the base of the stem characterizes
mature plants (Fics. 4, 5, 13) and has probably led to some exaggerated
statements about their longevity. Speculations about possible great age
have particularly centered around Dracaena draco. The early literature
about this is summarized in the paper of Wossidlo (1868). Perhaps the
most famous individual tree in this respect was the specimen of Dracaena
draco of Orotava on Teneriffe, described by Alexander von Humboldt
(1850). Its historical record goes back to the fifteenth century. But
estimates that it dated back to the period of the building of the pyramids
(4,500 years) are probably exaggerations, especially in view of the known
rate of growth of Dracaena reflexa (Wright, 1901). In 1799 the famous
tree of Orotava had reached a height of about 70 feet and a circumference
of 48 feet at the base of the trunk. A hurricane destroyed it in 1821.
There is no certain method of telling the age of a specimen in the absence
of planting data. A more meaningful time scale is given by a specimen
of Beaucarnea recurvata (Fic, 13) in Fairchild Tropical Garden which
is 25 feet high, 19 feet in circumference at a height of 2 feet and yet is
known to be not more than 50 years old. Rates of growth otherwise
appear not to have been determined for any of these plants.
Inflorescences are always terminal. On unbranched axes they are large
and very conspicuous as in Yucca and Xanthorrhoeaceae (Fic. 11) and
€ven on young specimens of Beaucarnea. They reach massive proportions
in Agave and Furcraea. In the much-branched forms flowering is usually
simultaneous on all or most distal shoots and renders the tree very
conspicuous. In temperate species flowering is seasonal as in Cordyline
Plants of successive ages to show development of massive trunk. 1, Unbranched
sapling 5-6 years old. 2-3, Early wahiarkonaue of branch system in older stages.
4, Mature specimen in flower. Flow begins in saplings of the size a
in Fig. 1 hte it is ei branching 5, Aloé dichotoma. 6, Cordy
australis. sp. 8, mele (Drac aena) reflexa. 9, Cordyline indivisa
7, Aga
10, Yucca aloifolia. tt. ssshieohned quadrangulata.
164 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
IG. 12. Specimens of _Cordyline australis, growing in their natural habitat
near ana New Z
Beaucarnea recurvata, ay Dies ma
Nee he ‘ety aise “|
a pie < : ? ope,
ae 9 te \
*
A etl oe 4"
a ®
®
-
*
é =
J
~
os tone
S a'78e @e cep
o2® @@s ve Otic
©. 8's ae8 8 Ge. ess
s
ay .
8s
atta “en
at Ys
G23 ¥ ucca aloifolia, big aies eae of stem eaeety low ae pe Nap
38 show 7“ primary y ( sue. Growth rings ar
evident in the secondary tissue particularly if nae illu ustri oe is viewed pee
a distance C. Leaf siLT)t sige Do radially.
Fics. 24, 25. Dracaena fragrans, pete se otis of ro t 33. A small
amount of secondary tiss ssue is present. [Note the position of endodermis show-
ing that cambial activity began in places inside it, in other places outside of it.]
7
ss
i=)
Q
a8
er
always thickest in the region of insertion of lateral roots, and suggested
that secondary thickening is initiated in this region.
It is quite obvious that the somewhat conflicting observations of differ-
ent workers have a rational explanation in terms of growth and the factors
which influence cambial development and activity. The problem has to
be studied by following the origin and subsequent growth of adventitious
roots in seedlings of different age, as Wright (1901) suggested. Wright
174 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
also made the observation that the cambium originates in the pericycle
of the very short hypocotyl, thence spreading upward into the stem and
downward into the root. Further observations of this kind are needed to
establish a clear understanding of secondary growth in roots of Dracaena.
THE RELATION OF PRIMARY TO SECONDARY GROWTH
The earlier studies on growth and development of these monocotyledons
were carried out at a time when the understanding of plant growth in
general was at a very primitive stage. It was also inevitable that theories
of plant growth were dominated by concepts derived from studies of
dicotyledons, and some of the earliest interpretations of monocotyledonous
growth made unfortunate comparisons between monocotyledons and dicot-
yledons. To this early period belong a series of studies concerned with
the relation between the secondary “thickening ring” and the meristematic
tissues of the shoot apex proper as in the investigations of Karsten (1847),
Schacht (1852), Nageli (1858) and Sanio (1863). It seems that these
studies were based on examination of single sections cut in transverse and
longitudinal planes and that no attempt was made to trace the distribution
of developing vascular bundles and the 3-dimensional relation between
primary and secondary growth. We will show in a later article that this
kind of investigation is crucial to the understanding of this relation.
One of the features of arborescent monocotyledons which captured the
interest of earlier workers was the apparent continuity between the
secondary meristem and the meristematic tissues of the crown. Some
authors considered these two meristems to be discontinuous (e.g. Scott &
Brebner, 1893). This discontinuity is also implied by Millardet (1865)
who gave measurements of the distance below the apex at which the
secondary meristem could be first recognized. This varied from as little
as 3 mm. in Yucca aloifolia to as much as 22 cm. in Dracaena marginata.
On the other hand many authors regarded the two meristems as continuous
(e.g. Wossidlo, 1868; Lindinger, 1908). Hausmann (1908) reviewed the
extensive literature on this topic and himself supported the latter point of
view, concluding in fact that the distinction between the two meristems was
rather artificial. In a developmental sense this is true, because establish-
ment and activity of secondary tissue is dependent upon growth of the
primary meristem. Nevertheless, earlier authors have often adopted a
very dogmatic point of view, largely in an effort to establish whether the
secondary meristem originated in tissue which had completed its matura-
tion or not, and was therefore, by definition, truly “secondary.”
A similar dogmatic preoccupation which is also largely a semantic one,
was with the level, in a radial direction, at which divisions which initiated
the secondary meristem occurred. The problem was to decide whether
there was a region in the monocotyledonous stem, to which the term
“pericycle” could be given. This is entirely an artificial concept, since in
most monocotyledonous stems, cortex and central cylinder each ends where
the other begins. A true understanding of the development of that region
1969] TOMLINSON & ZIMMERMANN, MONOCOTYLEDONS 175
of the stem in dicotyledons for which the older term “pericycle’’ was
devised has been forthcoming only in recent years (Blyth, 1958). The
term pericycle has no application in monocotyledonous stems (Carano,
1910).
In terms of the overall distribution of the monocotyledonous cambium,
one factual error does deserve comment. Réseler (1889) and apparently
some earlier authors stated that the cambium does not extend into the
leafy zone of the shoot. This is manifestly so untrue a generalization,
whatever may have been the situation in the material on which it was
based, that it is not surprising that it was soon corrected (e.g. by Corde-
moy, 1894). The presence of functioning leaves, the traces of which must
cross the cambium and secondary tissues, does raise interesting physiologi-
cal and developmental questions to which we will return in a later article.
One reason for the conflicting reports on these topics which appears in
the literature was that many authors failed to appreciate the variability
in the time of appearance of the cambium and its vigor, which in turn
seems largely to depend on the vigor of the shoot. We have already com-
mented upon the variation in vigor expressed in the different diameters
of shoots in one plant. This variation extends to the secondary cambium
and may depend largely on the type of shoot. Seedling axes, for example,
initially produce secondary tissue very actively. This activity declines on
distal branches. Newly released buds, either below inflorescences or
decapitated shoots, are dependent on an active production of secondary
tissue in the early stages of growth in order to establish vascular continuity
with the parent axis. In view of this variation it is not surprising that
reports by early authors conflict, since they are probably based on com-
parison of shoots in different positions and of differing vigor.
COMPARATIVE INVESTIGATIONS
A few authors have been concerned with the relation between those
monocotyledons with secondary growth and those without. Notable are
Mangin (1882) and Petersen (1893). Chouard (1936) was concerned
with the same topic, but his interpretations of monocotyledonous growth
are not easy to comprehend. Petersen studied a number of monocotyledons
which together represented a wide variety of families and growth forms.
He came to the conclusion that in the group as a whole there was a con-
tinuous series with all intermediate steps, from those, like the orchids
with no trace of a secondary meristem, via those in which one is briefly
active, as in the Bromeliaceae, to the continually active cambium of
Dracaena which permits unlimited growth.
Mangin (1882), on the other hand, was concerned with the way in
which adventitious roots develop and establish vascular continuity with
the conducting tissues of the parent axis. Adventitious roots arise in a
meristematic region (couche dictyogéne) between cortex and central
cylinder. This meristem also gives rise to a plexus of vascular tissue
(réseau radicifére) which connects conducting tissues of root and stem.
176 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The extent of this plexus varies in different kinds of monocotyledons.
Mangin considered that in some arborescent monocotyledons, like Agave,
this meristematic region remains active throughout the life of the plant.
In others, like Dracaena and Yucca, the root meristem is replaced by the
secondary meristem. When more is known about the factors which
stimulate and maintain an active cambium in monocotyledons it will be
possible to approach the topic on a comparative base. Nevertheless Man-
gin’s contribution to anatomical literature remains a notable one.
CONCLUSIONS
It is obvious from the previous pages that a reappraisal of this subject
from first principles is needed. We hope to present in future articles the
results of studies which to a large part resolve much of the conflicting
literature. In particular we will describe the course and developmental
pattern of the primary vascular bundles, the constructional relation be-
tween primary and secondary vascular bundles and demonstrate how the
initiation and activity of the secondary meristem is dependent upon shoot
growth. These will be related to growth of the shoot system as a whole.
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ihr Verhiltniss zum Dickenwachsthum desselben. Bot. Zeit. 16: 185-190,
193-198
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PETERSEN, O. G. 1893. Bemaerkninger om den gr ONS staengels Tyk-
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412.
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1969] TOMLINSON & ZIMMERMANN, MONOCOTYLEDONS 179
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[PBT] [M.H.Z. |
FAIRCHILD TROPICAL GARDEN HARVARD UNIVERSITY
10901 OL_p CUTLER Roap CaBoT FOUNDATION
MriamI, Fiorwa 33156 PETERSHAM
MASSACHUSETTS 01366
180 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ASPECTS OF REPRODUCTION IN SAURAUIA
Dyaja D. Soryarto !
THE GENUS Saurauia is a widespread tropical member of the Actinidia-
ceae with representatives in both the Old and the New World. The
American range of distribution extends from Central Mexico to southern
Bolivia, through Andean South America. According to a recent study by
Hunter (1966), 22 species occur in Mexico and Central America, and m
present study indicates that 49 species are represented in South America.
The genus is not represented in the West Indies, and there are no records
of its occurrence in the Guianas or Brazil.
During the course of field work in southern Colombia in 1965, I ob-
served that some individuals of Saurauia tomentosa (H.B.K.) Sprengel
have flowers with sessile stigmas, in contrast to the flowers with long styles
(5-7 mm.) of individuals commonly held to be characteristic of the spe-
cies. Later herbarium studies indicated that several other South Ameri-
can species are similar to S. tomentosa in this respect.
To be certain that such a phenomenon had not previously been described
in Saurauia, I have searched the literature and found that nothing con-
clusive has ever been published. There are several references, however,
to the reproductive system of Saurauia. Gilg (1895) and Gilg and Werder-
mann (1925) described the flowers of Saurauia as hermaphroditic to
polygamo-dioecious. Brown (1935), who observed the flowering pattern
of S. subspinosa Anthony, an Asiatic species, noted that the ovary de-
velopment in this species lags behind the development of the anthers by
about five days, suggesting that cross-pollination may be dominant.
Hunter (1966) mentioned that some species in Mexico and Central
America have flowers with “aborted” pistils. Killip (Jour. Wash. Acad.
Sci. 16: 570. 1926) referred to the flowers of S. micayensis Killip as uni-
sexual, while Benoist ( Bull. Soc. Bot. France 80: 334. 1933) described the
flowers of his S. Aypomaila as staminate.
A few field workers have noted the existence of “male” and “female”
plants in some species of Saurauia. Lorenzo Uribe Uribe, for example,
noted the peculiarity in S. isoxanthotricha Busc. (L. Uribe U.’s collection
number 4802): “Este pie, que crecia cerca a mi No. 4801, no tenia sino
flores femeninas.” (This tree, which grew close to my No. 4801, had only
female flowers. )
The flowers of Saurauia are borne in a thyrsiform inflorescence, con-
sisting of a peduncle, rachis, and axillary scorpioid cymes arising in 4
spiral pattern. Each cyme is borne in the axil of a bract. The flowers are
*The author is currently engaged in the revision of the South American species of
Saurauia.
1969 | SOEJARTO, REPRODUCTION IN SAURAUIA 181
actinomorphic, pedicellate, each subtended by a bract and two lateral
bracteoles; basically, the flowers are pentamerous and are usually de-
scribed as bisexual or hermaphroditic. To the best of my knowledge,
there is no true “male” or “female” plant; in other words, there is no
true sexual dioecism in Saurauia.
The observations discussed in this paper were made to obtain more
conclusive evidence about the reproductive system and its operations, and
to suggest the implications for evolution in the South American species
of Saurauia. This paper is the basis for more detailed studies on the breed-
ing systems of the group which are in progress.
MATERIALS AND METHODS
The present investigation has been based primarily upon data obtained
from herbarium specimens. Initially, the work consisted simply of sort-
ing specimens with reproductive parts into long- and short-(obsolete-)
Styled groups. The next step was examination of the pollen grains (their
morphology, size, and fertility) of individuals in each of the two groups.
Pollen fertility count was obtained either from open flowers or from ma-
ture flower buds. The best results were obtained by boiling the flowers
or flower buds (sufficiently mature) to obtain the anthers for maceration.
Boiling restores the dried material to a natural texture, which makes dis-
section and measurement of the floral parts more accurate. The pollen
grains were mounted in glycerine jelly and stained with cotton blue dis-
solved in water. All pollen fertility counts reported here were obtained
by using a Wild M20 phase contrast microscope, with bright-field il-
lumination with or without a green filter. Percentage numbers were
based upon a count of between 100 and 500 pollen grains on a single
preparation. From two to five samples were prepared from one individual.
Stamen counts were made for taxonomic purposes. More important to
this study, however, was to ascertain whether or not stamen number has
any significant relation to floral dimorphism. All counts were made by
boiling the (mature) flower buds, since counts based upon open flowers
may be inaccurate, as some stamens may have aborted or others may have
been broken and fallen during the process of drying and handling.
Measurements of style length were made mainly from open flowers
and/or fruits, since the styles are persistent in Saurauia, Style length is
not reduced much by drying, so boiling was only occasionally necessary.
When neither open flowers nor fruits were available, measurement was
made from the mature flower buds. This is a valid and reliable substitute,
as will be obvious from the following discussions.
OBSERVATIONS
Analysis of data. I have examined all species from South America
for my taxonomic revision, but; due to lack of data, only species with
sufficient representation are included here for discussion. These are Sau-
182 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Individuals Scored
Individuals
Scored
Distribution of
bistribution of
? N
2 Stamen Number
i Stamen Number jai
; RW
°%50 100 160 200 : Q 0 To 20 #30 40
3
15 Ou
S Y
BE >
pat rt 2
Y Yy . Yy 2B! = ie
vs i £E
“ts & 0 i
104 sgt a a* Be
3 re aS Bink
z rat a3 ADO
4 of on 1 0
we > 9 : Yi; & © f Style
par 5 ao Li rs Length
a ao (ces )
oe 2% .
2 »
5} 7 -)
4 or 2 oO
mo fp
Se Bis
a3 va Z
| a & & y/ 4 Individuals Scored
tyle
Ld Ld canat :
' 2 3 5 6 71
ees 2 Distribution of
oo Stamen Number
pod abage bao Fag
Stamen Num
300 40
<%
wo
P
od
Lan
A
+e
ial
Uj, ws.
i=
®
g nl
a
o
De
Fruit not recorded
N
e
Cf
-
al
et
A
»
é
S
o
a
3
a,
Pollen Fertility 0%
Fruit recorded
Fruit not recorded
.
:
Ficures 1-4, Graphic representation of the distribution of long- and short-
s aed son fepiciented by herbarium collections, together with histogram dis-
tribution o e stamen numb
. Sample examined for each species consists ‘of short-
or long-styled forms in more or less equal numbers.
rauia bullosa Wawra, S. brachybotrys Turczaninow, S. excelsa Willdenow,
S. Humboldtiana Buscalioni, S. tomentosa (H.B.K.) Sprengel, S. omich-
lophila R. E. Schultes, S. putumayonis R. E. Schultes, and S. ursina
Triana & Planchon. Data for each species, such as presented in TABLE 1
for S. bullosa, and in Taste 2 for S. omichlophila, have been converted
into graphs, Fics. 1-8. Of the eight species, seven show a definite correla-
tion between low pollen fertility (or absolute pollen sterility and a long-
Table 1.
Saurauia bullosa Wawra
POLLE STYLE
COLLECTOR i 2, je . FERTILITY LENGTH he veesond FRUIT oS ALTITUDE
(%) (mm.) m.
Soejarto 496 _ — §.5 _ + Aug. 2900
Soejarto 1504 — 6 — + Aug. 3000
Soejarto 1533 - — 6.5 _~ + Aug. 2700
Soejarto 1472 + 0 5 160 — Aug. 3100
Soejarto 1336 - - 7 — + Aug. 2900
Soejarto 495 + 0 6 115 + Aug. 2900
Soejarto 1015 + 0 5 127 + July 3200
Soejarto 1478 + 0 5.5 220 ~ Aug. 2900
Soejarto 1595 + 0 7 160 Sept. 3000
Soejarto 500 + 0 5 125 ~ Aug. 2900
Cuatrecasas 20805 + 0 6 100 + Apr. 3100
Cuatrecasas 20414 + 1 4.5 85 - March 3200
Cuatrecasas 23316 + 2 5 70 Nov. 3000
Jorge Castro 78 + 0 5 140 - Apr. 3400
. Uribe Uribe 5278 + 0 55 100 + July 3000
nes Mexia 7599 + 10 5 140 - Aug. 3000
Soejarto 1496 on 75 0 175 Aug. 3100
Soejarto 1435 + 80 0 225 _ Aug. 3100
Soejarto 1484 + 85 0 150 _ Aug. 3000
Soejarto 1491 a 93 0 164 - Aug. 3000
Soejarto 1045 + 80 0 150 - July 3600
Soejarto 1508 + 92 0 127 —- Aug. 3000
Soejarto 1474 + 88 0 149 - Aug. 3000
Soejarto 1335 + 40 0 240 _ Aug. 3200
Soejarto 1473 + 96 0 145 a Aug. 3000
Fajardo G. 81 + 96 0 125 -— Apr. 3400
I. F. Holton 23 + 97 0 a _ Jan. 3000
Cuatrecasas 20997 + 98 1 145 — Apr. 3000
Hitchcock 20888 + -- 1.5 150 - Aug. 3400
L. Uribe Uribe 5328 + 90 0 160 — July 3100
VINVUAVS NI NOILONGOUdAY ‘OLUV[AOS [696T
est
184 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Individuals
Scored
Distribution of
10: Stamen Number
20)
0'$0 100 150 200 250
Individuals Scored
20.
Distribution of
10 Stamen Number
i]
o
i
050 100 156 200 250
5
g
3 of 3 nd
- Sng 3
>
2% a5 4%
Yt © 5 oO iy
Se & > or
a Pear ners aq
i ef: tS
Styl
Yj, 6 Wa ran
te} 2 3 4 6 6 a (mn. )
Distribution of
Stamen Number
Distribution of
Stamen Number
Pollen Fertility (25-)65-97 %
Fruit not recorded
Pollen Fertility 0-10(-70) %
Fruit recorded
Style Style
3 . Lengt 8 ps Length
. (mm. ) oe
2 3 4 5 6 7 (mm. )
Ficures 5~8. Graphic representation of the distribution of long- and short-
styled forms represented by herbarium collections, together with histogram
distribution of the stamen numbers. Fic. 5, Saurauia
brachybotrys Turcz.; Fic. 7, S. Humboldtiana Busc.; Fic. 8, S.
Sample examined for each species consists of short- and long-styled forms in
more or less equal numbers.
styled condition, and between high pollen fertility and a short-(obsolete-)
styled condition. This correlation breaks down in S. omichlophila, where
both long- and short-styled plants have high pollen fertility. Plants of
each type occur in approximately equal numbers within a sampling col-
lection of each species, which may reflect the distribution in the natural -
populations. Another significant correlation is that specimens bearing
fruits have been recorded only from plants with long styles. This is cer-
1969 | SOEJARTO, REPRODUCTION IN SAURAUIA 185
tainly not mere coincidence, since all eight species discussed here (and
many others for which statistics are not included) show this condition
throughout. Morphological examinations from free hand sections of ad-
vanced ovaries in short-styled flowers show that these are aborted and
they simply “do not grow” (PLaTeE II, Fic. 19). In long-styled flowers
fruiting is accompanied by good seed set, except in several individuals
where seed set is poor, notably in Soejarto 1043 (S. tomentosa).
Short or long condition of the styles is not in any way correlated with
high or low number of the stamens. As is obvious from Fics. 1-8, and
from TaBLes 1 and 2, the distribution of the stamen number is continuous
throughout the population, regardless of the style length. From the mea-
surements of flower parts (data not included here), it also appears that
a short- or long-styled condition is not correlated with the size of the
flowers.
From field observations and from herbarium records, there is no indi-
cation of any particular flowering and fruiting season among the South
American species of Saurauia. Flowering is usually associated with the
wet months of the year. In most species, however, flowering and fruiting
are continuous throughout the year, although fruiting is recorded in the
herbarium collections (at least, in the eight species under discussion)
only from February through October.
Pollen grains (PraTe I). All eight species have 3-colporate pollen,
which is oblate spheroidal (cf. also Erdtman, 1952). Several individuals
of S. excelsa have 3-colpate, prolate pollen grains. Most of the South
American species that I have examined have oblate spheroidal pollen
grains, although occasional prolate pollen is also present. No single
pollen shape is restricted to a particular species. This condition applies
only to fertile pollen grains, where the cell content stains uniformly with
cotton blue and appears light to dark blue with bright-field illumination.
The cell wall is smooth, with no observable wall sculpturing (at least
with the present processing technique). The (fertile) pollen grains are
binucleate at the time of anthesis (PLATE I, Fic. 14; cf. also Brewbaker,
1967); the generative cell is ellipsoidal or spindle-shaped, and the vege-
tative cell is roundish. The vegetative cell usually does not take acetocar-
mine stain so well as the generative cell. The binucleate condition of
the grains may be seen (with cotton blue stain) in a sufficiently mature
flower bud, prior to anthesis, and it is assumed that this condition indt-
cates that the pollen is fertile.
The sterile pollen grains, on the other hand, have no fixed shape or
any orientation. They may be lenticular, roundish or simply irregular in
shape, but, lacking contents, do not stain. The cell wall usually is shriv-
elled. Some roundish pollen grains have minute dark granules within.
The size of the pollen varies, and no serious attempt has been made
to measure size variation species by species. I am convinced, however,
that pollen size is not taxonomically significant. Pollen size variation is
always present in any preparation from a single plant, and size variation
186 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
between species is very slight. According to Erdtman (loc. cit.) the pollen
size of S. Prainiana Busc. from Pert is 18.5 20 microns (oblate
spheroidal), and that of a species from Bolivia identified as S. brachy-
botrys Turez. (probably a misidentification, since S. brachybotrys occurs
only in southwestern Colombia) is 19 % 15 microns (subprolate). Ac-
cording to my rather crude measurements, fertile pollen grains vary in
pollen grains from a short-styled plant and those from a long-styled
plant; nor between sterile pollen grains from a long-styled and from a
a eis plant.
Androecium and gynoecium (Prater II). There is no pollen dimor-
phism in Saurauia, in the sense of two types of pollen grains differing
morphologically and correlated with floral dimorphism. The correlation
in most cases is straightforward: long styles and low pollen fertility (ab-
solute sterility) vs. short styles and high pollen fertility. The term long
used here is relative, depending on the individual species involved. Spe-
cies with large flowers (3-5 cm. in diameter), such as S. bullosa and S.
tomentosa, have long styles 5-7 mm. long, and short styles 1-2.5 mm.
long, whereas species with smaller flowers (0.5—1 cm. in diameter), such as
S. pseudoleucocarpa Busc. and S. micayensis Killip, have long styles 3-5
mm. long, and short styles 0.5-1 mm. long. Styles less than 0.5 mm. long
are considered to be obsolete.
The ovary of the American species of Saurauia is mostly five-carpellate,
but in some species, e.g. S. yasicae Loes., S. peruviana Busc., and S. leuco-
carpa Schlecht. may be three- to five-carpellate or, in Saurauia sp. (a new
species from Bolivia to be described by me), five- to seven-carpellate.
Each style of the long-styled flower is surmounted by a capitate stigma.
The stigmatic surface is either roundish or cordate (1-2 mm. broad in
S. bullosa), covered by minute papillae. The size of the stigma — and
for that matter of all other floral parts — varies with the size of the flower.
At the time of anthesis, the stigmas turn dark brown and become sticky.
This condition lasts, in S. bullosa, for four to seven days. On the other
hand, the styles of a short-styled flower are tipped by simple stigmas
which are non-papillate, and according to my field observations, there
is no change in color or stickiness during anthesis.
Pollination. It appears from field observations that pollination in
Saurauia is promiscuous. Most flowers have persistent green sepals an
white petals (free for most of their length, but coherent at the base, falling
as a unit with the stamens’). Occasionally, some species (e.g. S. #50-
xanthotricha Busc.) have both white and pink flowers, but I have never
seen species with only pink flowers.* The stamens, with ‘white filament and
* As a result, flowers examined after anthesis are often described as “unisexu
al
*S. Conzattii Busc. from Mexico has red, beautiful flowers (Schultes, personal
comm.).
1969 | SOEJARTO, REPRODUCTION IN SAURAUIA 187
yellow anthers, characteristically form a yellow clump at the center of
the corolla. The anthers consist of two thecae, versatile and extrorse at
the time of anthesis; the point of attachment of the filament is at the
junction of the two thecae, which fork in most cases about two-thirds the
_ distance from the (embryonic) base. The versatile anthers and the pale,
morphologically unspecialized flowers represent, in a way, an adaptation
for wind pollination. There is no definite nectary present in the flower,
but nectar-secreting tissue is found inside at the base of the corolla, partly
hidden by the stamens (cf. Brown, 1935); also, most flowers have a faint,
Sweet scent, which in some species, especially S. omichlophila, is moder-
of S. peduncularis, S. omichlophila, S. brachybotrys, and S. chiliantha R. E.
Schultes.®
Fruit and seed dispersal. The fruit of Sawrauia is a berry filled with
numerous small seeds embedded in a mucilaginous pulp. The color of the
fruit is green, even when mature, although sometimes there is a purple
to purple-red tinge on the green, glabrous pericarp. The sepals are per-
sistent, as are the styles. Maturity of the fruit is indicated by an abundance
of mucilage, which is rather sticky, clear, sweet and edible. Dehiscence
of the fruit is septicidal along the longitudinal sutures, the septa often
being membranaceous; the central column and the septa remain intact after
dehiscence. In S. budlosa, dehiscence may occur from one to three days
after a ripe fruit is detached from the tree (faster when conditions are
wet) with little mechanical stimulation. The dehiscence lines start at the
apex of the fruit and run gradually towards the base, at the same time
discharging, or more precisely, exuding the mucilage which includes the
seeds. This is, I believe, the way that the seeds are dispersed naturally,
aided by the rain wash. Dispersal by birds certainly is not uncommon.
Birds have been seen frequently feeding on Saurauia fruits (common
name: moquillo or dulumoco, referring to the mucilage of the fruit). How-
ever, the effectiveness of bird dispersal must be further investigated. In
all probability, diaspores may not be transported great distances in Sau-
rauia; survival is insured by an abundant production of the seeds.’
“Pollen of wind- pollinated plants is sioner characterized by simplicity of structure,
and by the small size of the grains (betwee a microns, cf. P. Echlin, Sci. Amer.,
Apr., old such is the case in Saurauia ane
matica R. E. Schultes and S. eeceaenes R. E. Schultes were given their
enithets tai of the strong and heavy scent of their flowers
. Uribe Uribe (no, 2888) has observed numerous bees visiting the flowers of S.
Py mee
urauia are usually minute, areolate, dark brown; the testa is
fragile (cf Pee I, FIG. 21). That Saurauia seeds are viable for relatively long periods
is evident from the following notes, S. kegeliana Schlecht. (1836) was “described
from living plants at Halle, Germany, that grew from seeds in soil found about the
Toots of plants imported from Guatemala” (note by Standley & Steyermark, Field-
iana 24(6): 431. 1949). S. spectabilis Hook. (Bot. Mag. 69: pl. 3982. 1842) was de-
Scribed from a “plant raised by Mr. Knight, of the Exotic Nursery, King’s Road,
188 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Geographical distribution. The center of distribution of Saurauia
excelsa lies in the Venezuelan Cordillera de Mérida, while that of S. Hum-
boldtiana is found in the Cundinamarca region, Cordillera Oriental of the
Colombian Andes. There is an overlapping of range between these two
species in the Santander region. S. ursina is centered in Antioquia, along
the Cordillera Central, and its range overlaps that of S. Humboldtiana and,
perhaps, that of S. excelsa as well. S. brachybotrys centers in the Cauca-
Valle region, between the Cordillera Central and Occidental, but its range
extends north to Antioquia, and south to the Narifio-Putumayo region.
The Narifio-Putumayo area is located near the Colombian-Ecuadorian
frontier, where the species concentration of the genus is highest. S. budlosa,
S. tomentosa and S. omichlophila have their centers of distribution in
this region also. S. bullosa and S. tomentosa have the broadest ranges of
the South American species. S. putumayonis occurs in the Putumayo re-
gion, along the Cordillera of Portachuelo.
The four species, Saurauia excelsa, S. Humboldtiana, S. ursina, and
S. brachybotrys are not effectively isolated from one another geographically
or altitudinally. Although S. bullosa and S. tomentosa overlap geo-
graphically with other species, they are effectively isolated from the others
altitudinally and are themselves frequently sympatric in their distribution,
geographically, ecologically, and altitudinally. S. omichlophila and S. putu-
mayonis are effectively isolated from the others, particularly ecologically,
and they are spatially allopatric.
Cytology. I have examined the meiotic chromosomes of seven of the
eight species under discussion: Saurauia bullosa, S. brachybotrys, S.
Humboldtiana, S. tomentosa, S. omichlophila, S. putumayonis, and S.
ursina (Soejarto, 1969). Chromosome behavior at meiosis in these species
appears to be normal, and the chromosome size and morphology are re-
markably stable. The haploid chromosome number of all seven species is
n = 30. Cytokinesis is of a simultaneous type, and the tetrad arrange-
ment is tetrahedral. Chromosome counts were all made from pollen mother
cells.
DISCUSSION
In Saurauia, at least among the species from South America, two kinds
of flowers can be distinguished. The differences lie in the size and
morphology of the styles, and in the degree of pollen fertility. Anther
height and pollen size and morphology appear to be fixed. It is for this
reason, perhaps, that the existence of floral dimorphism in Saurauia has
passed unnoticed for so long. Most workers on this group considered
a short-styled condition to be peculiar to a particular individual or species,
Chelsea, England, from seeds imported from the Republic of Bolivia, in 1838.” How-
ever, I have attempted several times to germinate Saurauia seeds for cytological
studies without success.
1969 | SOEJARTO, REPRODUCTION IN SAURAUIA 189
or a sign of immaturity, and apparently did not appreciate the biological
significance of their observations. The present study shows that floral
dimorphism does exist in Saurauia, but that this type of dimorphism is
not distyly or heterostyly in the true physiological sense of the word,
since it appears (at least now) that no incompatibility system is involved.
Low pollen fertility (to complete sterility) in a plant with long-styled
flowers, and high pollen fertility in a plant with short-styled flowers is
a mechanism that promotes outcrossing. In this respect, the flower of
Saurauia must be described as functionally dioecious. The short-styled
form with high pollen fertility may be considered a functionally staminate
plant (the pistil being nonfunctional), while the long-styled form with
low pollen fertility (to complete sterility) is a functionally carpellate
plant (the stamens being nonfunctional). For those individuals, partic-
ularly populations of S. omichlophila, which are truly hermaphroditic
(Taste 2; hermaphrodites are characterized by a long-styled flower with
pollen fertility 80% or more) within the dimorphic populations, further
investigation is needed to demonstrate whether any self-compatibility be-
tween the pollen and the stigma of the same flower exists. From the pollen
size and morphology, there seems to be no reason why it should not occur.
If this is the case, species like S. omichlophila must be described as an-
drodioecious.
The widespread occurrence of functional dioecy, an outbreeding sys-
6 6 °O
Fic 23A-F. Anther orientation in the flower of Saurauia Humboldtiana
Buse. "Se pals and petals removed to show details. A, short rt-styled form, bud
stage; B, cee styled form, at anthesis; C, mcd form, bud stage; D, long-
styled fo orm, at anthesis; E, anther, bud stage; F, anther, at anthesis. As a re-
190 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
tem, is further enforced by the peculiar anther orientation during an-
thesis in the American species of Saurauia (Fic. 23, A-F). The end of
the anther (the embryonic base) is directed away from the center of the
flower as it opens, and the anther rotates about 130° on the filament
so that the pollen discharge is directed away from the stigmas. Pore
openings and dehiscence of the anther start at the embryonic (morpho-
logical) base and “zip” ventrally about two-thirds the length of the
thecae. My field observations of this dehiscence and reorientation of the
anther during anthesis are confirmed by Hunter’s (1966) interpretation,
from histological observations of the vascular trace of the stamen (Hunter
interprets the reorientation of the anther at anthesis as 180°, with which
I cannot agree). Therefore, in individuals which are truly hermaphroditic,
like those in S. omichlophila, self-pollination is averted as much as possible.
Prevention of self-pollination among the hermaphrodites is indicated by
the position of the stigmas which is well above the surface of the androe-
cium. Nevertheless, the chances of self-pollination are rather good.
From an evolutionary point of view, the immediate consequence of
outbreeding is its capacity for genetic recombination to produce variability
or the action of selection and other external forces which direct the
evolution of populations (Stebbins, 1950). The greater part of the geno-
typic variation within a cross-breeding population is due to segregation
and recombination of genic differences which have existed in it for many
generations. As a result, in a comparable environment, the outbreeders
may show great, more or less continuous, morphological variation, which
is an expression of genetic variability from plant to plant. Most species
populations of Saurauia, those which are functionally dioecious, are
characterized by this type of morphological and environmental continuity.
Because of a low selective pressure, variability within a population tends
to obscure any clear-cut distinction between populations. The situation
is further confounded by a more or less free gene flow between species
populations, due to an incomplete isolating mechanism: spatial, ecological,
ethological or, perhaps, genetical; this last mechanism must be further in-
vestigated. The lack of a complete genetic barrier is demonstrated by
the frequent occurrence of natural hybridizations where two or more species
populations are in contact or where they are sympatrically distributed. I
have collected several natural hybrids of Sauwrauia from southwestern
Colombia, where the greatest concentration of species is located, and the
hybrid status of at least four of these plants has been confirmed by meiotic
irregularities of the chromosomes (Soejarto, unpubl.). Although altitudinal
isolation is usually effective, nevertheless some population contact is al-
ways present. There seems to be no effective barrier against interspecific
pollination in most cases, which is reflected in the relatively uniform
floral morphology. Only size variation of the flower, which is conspicuous,
exists within as well as between species. Correspondingly, selective pres-
sures are relatively weak at the stage of flowering and also at the fruiting,
or dispersal stage. On the chromosome level, the differences between
species populations appear to be even less significant; that is, as far as
Table 2. Saurauia omichlophila R. E. Schultes
FL. BUDS POLLEN STYLE STAMEN COLLECTING
COLLECTOR OR FLs. FERTILITY LENGTH NUMBER FRUIT DATE ALTITUDE
(%) (mm. ) (m.)
Soejarto 1493 + 90 5 18 + Aug. 2900
Core 1019 + 95 5 30 + July 2700
Soejarto 1176 + 90 4.5 28 + July 2500
Soejarto 1046 ao 96 4 14 + July 3200
Soejarto 977 + — 4.5 _ + July 3000
Soejarto 1511 - — 4 — + Aug. 2800
Garcia-Barriga 13023 + 85 4 21 — July 2800
Uribe Uribe 3876 + 98 4 25 Sept. 3000
Soejarto 1501 + 90 0 23 Aug. 3000
Soejarto 1509 + 88 1,5 20 Aug. 3100
Schultes 3236 + 95 0 26 - Feb. 3200
Schultes 7560 + 90 0.5 26 March 2900
101 4 96 0.5 37 s July 2700
Schultes 7560A + 88 0.5 ay — May 2900
Soejarto 1502 + 85 0.5 29 - Aug. 3000
Hernandez 79 + 95 0 22 - -- 3000
Soejarto 1598 + 95 0 20 — Sept. 3000
Schultes 20098 + 92 0 21 — June 2800
Schultes 7550 +- 85 0 21 - June 3000
Schultes 7771 + 90 0 26 _ June 3000
Soejarto 914 + 0-80 0 21 _ July 3000
[6961
TI NOLLONGOWdAY ‘OLUV[AOS
VIAVANVS
T6T
192 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
my present investigations on the cytology of the South American Saurauia
show. The inevitable consequence of all this is the difficulty of drawing
clear-cut boundaries between species populations, and, consequently, in
the delimitation of the species within the genus. It is unfortunate that
species of Saurauia are unfavorable subjects for garden experiments be-
cause of climatic intolerance, poor seed germination, the large size of the
plants and the length of time before they reach the flowering stage. These
drawbacks, however, should not discourage workers on the group from
continuing their efforts. There are several other things that can and must
be done; one of these is more vigorous field work and collecting of her-
barium material. The more herbarium collections accumulate, the better
we can evaluate and analyze the limits of variation within and between
species. Considering the relatively young age of the group, Tertiary
(Eocene?; cf. Langeron, 1900, Hollick, 1936), it appears to me that
evolutionary differentiation is proceeding in the genus Saurauia.
Finally, the realization that functional dioecy is prevalent in Saurauia
may further confirm our opinion with regard to the phylogenetic relation-
ship of the genus with the closely allied, predominantly dioecious Actinidia.
SUMMARY
The reproductive system(s) of the following eight South American
species have been described and discussed: Saurauia bullosa, S. brachy-
botrys, S. excelsa, S. Humboldtiana, S. tomentosa, S. omichlophila, S. putu-
mayonis, and S. ursina. As far as the present data show, seven of the eight
species appear to be functionally dioecious, and one, S. omichlophila, is
androdioecious. The flowers of these plants are dimorphic: a long-styled
form with high pollen sterility (functionally carpellate) vs. a short-
styled form with high pollen fertility (functionally staminate). Anther
height is fixed, and pollen dimorphism related to style dimorphism has
not been seen.
Although data have been compiled exclusively from herbarium examina-
tions (presented here in graphic form, Fics. 1-8), the following observa-
tions, based upon field and laboratory studies, have also been briefly de-
scribed: pollen grain, androecium vs. gynoecium, pollination, fruit and
seed dispersal, geographical distribution, and cytology.
The widespread occurrence of functional dioecy may be a useful guide
in confirming the phylogenetic relationship between Saurauia and the
closely allied, predominantly dioecious genus Actinidia. It is further
suggested in the discussion, that the extensive morphological variability
is the result of the outbreeding nature of the group, because the immediate
consequence of outbreeding is its capacity for genetic recombination to
produce variability in the action of selection and other external forces
which direct the evolution of populations.
1969 | SOEJARTO, REPRODUCTION IN SAURAUIA 193
ACKNOWLEDGMENTS
I am deeply indebted to Professor Reed C. Rollins for his generous
suggestions and painstaking criticism of the manuscript. Dr. Richard E.
Schultes has very kindly helped me edit the English of the text, for which
I wish to express my thanks. I also wish to thank Dr. Carroll E. Wood,
Jr. and Dr. Beryl Vuilleumier for several discussions on heterostyly. I
have received financial support from the Committee on Evolutionary
Biology (NSF grants GB3167, GB7346; principal investigator, R. C.
Rollins), Harvard University. Finally, I want to thank my wife, Mariela,
for her constant and cheerful encouragement, and for typing the first
draft of the manuscript. Any errors or misinterpretations found in this
paper, however, are my sole responsibility.
LITERATURE CITED
BREWBAKER, J. L. 1967. The distribution and phylogenetic significance of
binucleate and trinucleate pollen grains in the angiosperms. Am. Jour.
Bot. 54: 1069-1083.
Brown, E. G. S. 1935. The floral mechanism of Saurauia subspinosa Anth.
re
Crowe, L. K. 1964. The evolution of outbreeding in plants, I. The angio-
sperms. Heredity 19: 435-457.
Erptman, G. 1952. Pollen morphology and plant taxonomy. 539 pp. Alm-
: : jae
Gite, E. 1895. Dilleniaceae — Actintoiinan: Actinidieae and ariel Sau-
rauieae. Jn: Engler & Prantl, Nat. Pflanzenfam. III. 6:
& E. WERDERMANN. 1925. Actinidiaceae. Nat. era ed. 2. 21:
7.
Hoxiicx, A. 1936. The tertiary floras of Alaska. U.S. Geol. Surv. Prof. Paper
1
Hunter, G. E. 1966. Revision of the Mexican and Central American Saurauia
(Dilleniaceae), Ann. Missouri Bot. Gard. 53: 47-89.
Bie ie M. 1900. Contribution a Vetude de la flore fossile de Sézanne. Bull.
c. Hist. Nat. Autun 13: 333-3
Sorsatte, D. D. 1969. Saurauia species and their chromosomes. Rhodora [in
pre: . .
Sreteties G. L., Jk. 1950, Variation and evolution in plants. 643 pp. Columbia
Univ. Press, New York & London.
DEPARTMENT OF BIOLOGY Present address: DEPARTAMENTO DE BIOLOGIA
Harvarp UNIVERSITY UNIVERSIDAD DE ANTIOQUIA
A
CAMBRIDGE, MASSACHUSETTS MEDELLiIN, Cotompta, S.A.
194 JOURNAL OF THE ARNOLD ARBORETUM {voL. 50
EXPLANATION OF PLATES
PLATE I
Ficures 9-16. Pollen grains in Saurauia. Fics. 9, 11, 14, 15, 16, fertile pollen;
9,
prolate pores As ins. except FIG. 14 prepared from herbarium samples,
stained in cotton blue; Fic. 14 prepared na anthers fixed in Carnoy’s solution
and stained in ig cio only a generative cell clearly visible). Fics. 9 and
10 approx. < 400, the others aA xX 1600.
PLATE II
big view of the stigmas at anthesis. Fic. 21, seeds. All to the same scale .
FIG. 19, and all from S. bullosa. Fic. 19 photographed from dried flowers
(boiled he eee all others from material fixed in Carnoy’s solution.
Scale in FIG. 19 is
Jour. ARNOLD Ars. VoL. 50 PLaTE I
SoEJARTO, REPRODUCTION IN SAURAUIA
Jour. ARNOLD ArB. VOL. 50 Prate II
SOEJARTO, REPRODUCTION IN SAURAUIA
1969 | GILL, ELFIN FOREST, 6 197
THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 6
AERIAL ROOTS !
A. M. GILi
IN TEMPERATE REGIONS aérial roots are rare and although they may be
found on a few vines they are absent from the trees and shrubs. In the
moist elfin forest of Puerto Rico, however, many of the trees, shrubs, vines,
and herbs form aérial roots. The tree fern Cyathea and the lowly Selagi-
nella also form aérial roots in this environment.
Many of the aérial roots hanging freely from the plants are very
characteristic of the species while some other species are difficult to dis-
tinguish by the characters of their aérial roots alone. In this study some
of the distinctive characters of the roots are described and the frequency
of aérial root formation on Pico del Oeste is documented.
OBSERVATIONS ON THE DISTRIBUTION OF ROOTS IN TOTO
The roots in the study area are found in four general habitats: in the
soil; immediately above the soil beneath a layer of cryptogams and/or
leaf litter; appressed to the trunks and branches of the trees and shrubs;
and hanging freely in the air.
All the roots in the last three habitats named may be considered
“aérial.” Those in the second category occur in a gaseous environment
immediately below the forest floor and above the soil. A mat of roots
up to five centimeters thick may be formed (Fic. 1) which appears to have
arisen not merely by erosion of soil but by the growth of roots out of the
soil and over its surface. On steep slopes roots of sufficient rigidity may
even grow through the forest floor into the atmosphere. On gentle slopes
this achievement has been attained by growth along tree trunks and
fallen branches beneath a layer of cryptogams and thence out to the atmos-
phere.
The roots of many of the vines and of the bromeliad Vriesea are found
closely attached to rigid organic surfaces. They are often found beneath
a mantle of cryptogams but are also found where such a covering is
lacking. This latter type of root may also be considered “aérial’” but
the affinity of the roots to their supports distinguishes them from the final
group which is the main subject of this paper.
The aérial roots to be considered here are those found hanging freely
in the atmosphere. They arise above ground and are not closely appressed
‘The first two papers in this series were published in Jour. Arnold Arb. 49: 1968.
See: R. A. Howarp, The Ecology of an elfin forest in Puerto Rico, 1. Introduction and
composition studies, 381-418; and H. W. BaynTon, 2. The Microclimate of Pico del
Oeste, 419-430.
198
cryptogam layer but
Fic. 1. Mat of roots immediately below the litter and
immediate} ly above the soil.
to any surface. They may become anchored in the substrate and undergo
considerabie secondary thickening and in such cases have been termed
rop” or “stilt” roots by other authors. The aérial portions of such
anchored roots may exhibit phenomena different from roots of the same
species in the freely-hanging stage — those to be considered here.
AERIAL ROOTS OF THE TREES AND SHRUBS
Many of the data Sew. to the aérial roots of the trees and shrubs
of the area are shown in TAB
rigin. Aérial roots ee arise from the undersides of branches and
from the main axis of the plant. They are often associated with the for-
mation of sprouts (probably arising from dormant buds) and in such
cases are found at the base of the sprout where it joins the main stem.
This condition was observed in Ocotea, Ilex, Miconia pachyphylla, Calyp
1969 | GILL, ELFIN FOREST, 6 199
TABLE 1. The aérial roots of the trees and shrubs
Tip PROPERTIES AT ROOT ORIGIN
Max
incre- Min.
ment Max. Lateral Min. oe ——
Max. before replace- Rigidity roots stem i
diam. none ment and without diam. bane hee
SPECIES (mm.) (cm.) tips Color alignment injury* (mm.) (cm. ) ne
Prestoea 17 19 4 pale orange stiff & + 58 45 —
montana to pale pink brittle,
simple
curves
Hedyosmum 3.5 80 3 white apex, unbent, — 4 5 +
arborescens then lemon, flexible
then green
nd
cotea 36 4 pink to unbent, ~ 12 30 _
spathulata brown flexible
Trichilia (1.2) 9 1 creamy brown’ unbent, _ 15. HS ?
pallida to pink flexible
ex 0.7 5 3 white to unbent, -- 13.5 110 ?
sintenisii flexible
Torralbasia 0:6: 12 5 orange unbent, a 3 30 +
neifolia flexible
za 89 2 white apex, unbent, - 13.5 SO +
grisebachiana yellow and flexible
brown behind
Calyptranthes 1.2 14 1 white to unbent, - 2S) Ws +
krugii red-brown flexible
Eugenia 24 1 white to unbent, — “HO 2m. +
borinquensis red-brown flexible
Calycogonium 1.55 14 3 bright unbent, — = 2
squamulosum pink flexible
Mecranium 9 2. white to unbent, = 2 2 Ss
amygdalinum pink flexible
iconia pink — 2 3 ?
foveolata
Miconia 10 2 white to unbent, ae 3s ob +
bachyphylla pink flexible
Grammadenia 2.0 9 2 white to unbent, = .) oe
Sintenisii light brown flexib]l
Wallenia 10 10 4 white to unbent, 5 5 1 +
yunquensis pink xible
Micropholis 27 kT ? white unbent, _ 47 ~ &50 ?
garciniaefolia flexible
Symplocos 0.8 4 6 white unbent, = — 55 6 +
micrantha flex. t
200 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
TABLE 1 — continued
Tie PROPERTIES AT ROOT ORIGIN
incre- Min.
ment Max. Lateral Min. distance Second.
Max. before replace- Rigidity roots stem to __ thick.
diam. laterals ment and without diam. _ leaves before |
SPECIES (mm m.) tips Color alignment injury* (mm.) (cm.) ground
Haenianthus 5° 23 6 cream ochre unbent to — 8 15 +
salicifolius to brown hang in
cluster
Tabebuia 2 10 5 creamy lime unbent, = 1 ae
rigida to weak flexible
yellow
Gesneria 0.5 4 7 white to unbent, — 3 3 ?
sintentsti tan flexible
Psychotria 0.7 7 2 beetroot unbent = 4 11 ?
berteriana to pale flexible
white
Lobelia white to unbent, ~ vf 0 ?
portoricensis pale green flexible
* + represents presence, — represents absence.
tranthes, Grammadenia and Torralbasia. The same thing may occur
below the ground with some species of trees in temperate areas. In Massa-
chusetts it has been seen in Fraxinus americana: when a tree is cut down
and new shoots arise at the base from beneath the soil-surface, new roots
may be formed at the junction of the new shoot and the parent stem.
No root was found within the leafy zone (distal to the most proximal
leaf and with or without a few leafless nodes included) of the trees and
shrubs except in Lobelia. In Tasie 1 the minimum distance of an aéria
root from the leaf zone has been noted and also the minimum stem diam-
eter on which an aérial root has been found. Most of the species have
aérial roots very close to leaf zones but not within them. Miconia pachy-
phylla was recorded with roots at the junction of the leaf zone and for
most of the species aérial roots have been found within 50 centimeters of
the leaf zone.
The aérial roots are usually found too far away from the leafy zone
to determine if there is any association between the root origin and the
nodes of the stem. With Miconia pachyphylla (Fic. 2) one root was found
at a node and no roots were observed where a definite lack of such an
association could be seen. No anatomical observation of the origin of the
aérial roots was made but association with a node would suggest develop-
ment of a preformed primordium giving rise to the aérial root.
Aérial roots may also arise laterally from other aérial roots, a condition
discussed in a later section.
Tip properties. The maximum diameters of the root tips vary widely
between species. Maximum values were taken since they are more dis-
1969 | GILL, ELFIN FOREST, 6 201
2. Roots of Miconia oat ta with droplets of water at the apices.
fae eesti of Dr. R.A
tinctive than average values. Tip size may decrease with the order of
branching, with distance from the point of origin, and in some species
served with smaller tips than those on large trees). Prestoea has tips up
to 17 mm. in diameter (Fic. 3) while the maximum recorded for Torral-
basia was 0.6 mm. The root tips of the other species were of diameters
intermediate between these values. The significance of tip size in between-
species comparisons is not known, but in the roots of the trees in central
Massachusetts at least, tip size within a given root system is an important
parameter, and other properties of the root are associated with it, e.g.,
lateral frecmency: number of protoxylem poles, and other anatomical
features, as well as the probability of secondary thickening.
Color variations in the tips may be distinctive. Torralbasia roots are
often orange in color, those of Miconia spp. usually a bright pink, and those
202 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
rial roots of the palm, Prestoea aioe 8 inhibited mage
Fic Aé
root te At in the absence of
aérial roots of Hedyosmum arborescens. Fic. ve Brox 90m - like cluster of nérial
— of Tabebuia rigida, developed as a response to deg oeey iauey. ” Secondaty
thickening is bag evident near the point of attachment. Fic. 6. Distally anchored
agvial roots of Clusza a showing devel mae of root tips and con-
siderable oe thicken
of Hedyosmum most often lemon. However, the color of the roots may
be considerably muted under some conditions and for many of the species
darker environments may cause all color to be lost from the root tips.
1969 | GILL, ELFIN FOREST, 6 203
Most of the newly emerged tips are straight and flexible. The palm
root tips, however, may be stiff and rather brittle and they may curve
down towards the soil. When the roots are longer their alignment and
rigidity may change. The roots of Clusia become long and rubbery and
the roots of Tabebuia and Torralbasia tend to hang in clusters. The roots
of Ocotea, however, usually maintain the initial direction of growth with
some bias downward.
The living roots of Hedyosmum arborescens are usually bathed in a
gelatinous fluid.* This may hang down from the apex of the tip for two
millimeters as part of a drop around and below the apex (Fic. 4). Shortly
above the apex the material becomes much thinner and is very thin 4 or 5
centimeters from the apex. Two roots with a considerable drop on their
ig ebe apices were stripped of their coating, using two fingers to remove
. The volume from both was approximately 2 milliliters.
aye experiment was initiated in an attempt to discover how quickly
the material covering the root was replaced. After a week there was par-
tial replacement of the material. After this period approximately 1 milli-
liter of fluid was removed from the three stripped roots. It should be
noted however, that similar iy ia (with care not to exert pressure)
eventually resulted in root death (R. A. Howard, personal communication).
In addition it may be noted that the material does not occur on dead
roots nor those with brown, apparently inactive, tips.
The gelatinous substance is common on the healthy roots of Hedyosmum
but has been found only rarely on the aérial roots of other species in this
area. It has been found only twice on the aérial roots of Calycogonium
and once on a root of Miconia pachyphyila. Many aérial roots of the
latter two species were observed, but in only these cited cases was the
gelatinous material seen. On these specimens the material was less gelat-
inous and more readily removed than that on roots of Hedyosmum. How-
ever, it was not dislodged as easily as a drop of water might be and it was
jelly-like in texture.
A cut in the aérial roots of three species caused drops of milky exudate
to be formed. Such material is common in the leaves and stems of the
three families represented and this observation indicates that the lactif-
erous system does extend into the aérial roots. The plants concerned are
Clusia grisebachiana, Lobelia portoricensis and Micropholis garciniaefolia,
members of the families Guttiferae, Campanulaceae, and Sapotaceae, re-
spectively.
Lateral root formation. Patterns of lateral root development con-
tribute to the specific character of many of the aérial roots. The rope-like
Clusia roots and the broom-like Tabebuia roots (Fic. 5) are very distinc-
tive for this reason. In many cases the lateral root formation from the
freely hanging aérial roots appears to be entirely dependent upon injury.
With such a stimulus, one to seven replacement tips may arise behind the
*This fluid has a high content of algae including diatoms and desmids. Some six
Carbon sugars were found in the material but no higher sugars (R.A.H.).
204 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
injured portion (TABLE 1). These tips may arise not only close to the
injury but also several centimeters behind it (e.g. Tabebuia and Torral-
basia).
With the aérial roots of the palm Prestoea lateral roots develop regu-
larly without injury (Fic. 3) but have very limited growth. The laterals
are short (up to 5 mm.) and pear-shaped. Their bases are narrow but
their diameter increases markedly (to 2 mm.) within a short distance and
then tapers to the apex. They arise in three to eight regular rows depend-
ing on parent root diameter.
Some roots may achieve great lengths before any lateral is formed.
The length attained is a reflection of the growth rate and the interval
between injuries, which in Clusia and Hedyosmum may amount to
2.5 to 3 meters. However, the maximum length without laterals for most
of the aérial roots is usually between 4 and 40 centimeters in this area.
When a freely hanging aérial root becomes anchored in the substrate,
prolific lateral root formation may occur in the subterranean portion. In
addition, however, new laterals may arise on the aérial portion with no
apparent stimulus from injury (Fic. 6). This contrasts with the develop-
ment of lateral roots before anchorage and is evident in the aérial por-
tions of anchored Clusia and Ocotea roots.
Growth rate. The growth rate appears to be very variable through
time as single uninjured roots are found to have variations in diameter
and color suggesting growth pulses. Injury, of course, prevents growth
and a decrease in length may result. Injury may be environmentally in-
duced and desiccation is a probable agent.
The growth in length of the aérial roots of ten species was measured
as it occurred during a 7 to 11 day period in December, 1967. The growth
rates varied from 0 to 2 millimeters per day. This may be contrasted
with the rate of growth of the aérial roots of Rhizophora mangle (the red
mangrove) in Miami, Florida, which grew up to 7 millimeters in length
per day during the months of April and May, 1968 (P. B. Tomlinson and
author). In the soils of New England, roots may achieve growth rates of
12 millimeters per day during summer (W. H. Lyford and author). Thus
the growth rates of aérial roots in the equable climate of the elfin forest
may be regarded as low.
Secondary thickening. Many of the aérial roots of trees and shrubs
may commence secondary thickening well before they reach the substrate.
Examples of this may be found among the roots of Ocotea, Clusia, Miconia
pachyphylla, Tabebuia, Haenianthus and Eugenia. In FiIGURE 5 thick-
ening of the root of Tabebuia near the point of attachment to the branch
may be observed readily.
The aérial roots of Ocotea near the base of the plant may become an-
chored and considerably thickened. In cross section these roots are oval
with the larger axis vertical and the morphological center in the lower
half. In Clusia however, similar roots are more nearly circular in cross
section and arise from up to 3 meters above ground.
1969 | GILL, ELFIN FOREST, 6 205
AERIAL Roots OF THE VINES
Origin. The aérial roots of vines are often found within the leafy zone
of the plant (in contrast to those of trees and shrubs). Not all roots are
found within the leafy zone but this is a common occurrence. In the
species under study here all the aérial roots were associated with a node
in one way or another. This is not always the case with vines: in some
vine species the roots are apparently formed at random along the stem as
in the ornamental Hydrangea anomala ssp. petiolaris and in the native
Rhus radicans in Massachusetts.
The associations with the node were varied. Marcgravia sintenisii has
up to four aérial roots produced in a row parallel to the axis of the stem
and running proximally from the leaf base. Gonocalyx, an ericaceous
vine, has a similar arrangement of aérial roots but on the distal side of
the leaf base. The genus Mikania of the family Compositae has roots
formed between the leaf bases or in an axillary position. Psychotria
guadalupensis (Rubiaceae) has aérial roots formed just distal to and be-
tween the nodes. Such specific positions of origin suggest a regular for-
mation of root primordia in these positions as the shoot grows.
In Marcgravia at least, it appears that new roots may be formed on
TABLE 2. The aérial roots of the vines
Trp PROPERTIES
Roots Max.
oots assoc. number Max. Rigidity Laterals
in leaf with per diameter _and without
SPECIES zone? nodes ? node (mm.) Color alignment injury?
Rajania yes yes 1 0.3. + white flexible, fat
cordata wrinkled
r yes yes 1 0.1 white nt, os
emarginella flexible
delicate
Marcgravia yes yes 4 0.6 cream unbent, =
Sintenisii (2.0) tend to
be rigid
Gonocalyx yes yes 5 0.3 white flexible, +
bortoricensis crinkled
Hornemannia yes yes 1 0.3 white to crinkled, +
racemosa pale pink flexible
to brown
Ipomoea yes yes 2 0.5 white weak ian
repanda flexibility,
curves
Psychotria yes yes 4 0.5 cream to flexible, +
guadalupensis light crinkly
green
Mikania yes yes 3 0.5 white to flexible,
pachyphylla pale crinkled —
206 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
old parts of the vines where they may be 2 or 3 centimeters in diameter.
In this case the tips produced may have different properties from those
formed within the leaf zone. In Taste 2 (summarizing the data collected
for the vines) the value for tip diameter recorded on older wood is noted
in parentheses. Whether these new roots develop from latent primordia
formed in association with the leaves is not known.
Tip properties and lateral root formation. The tips of the aérial
roots formed within or close to the leaf zone are usually very fine. Ap-
proximately 0.1 mm. to 0.6 mm. diameter is the range encountered. The
larger value in this range was recorded for Marcgravia, which has rapidly
tapering aérial roots — from 0.8 mm. to 0.4 mm. over one centimeter of
length. In this species aérial roots with a length greater than a few centi-
meters have not been found free-hanging in or near the leafy zone.
Some of the species were observed to have lateral roots formed appar-
ently without injury to the parent. The species in which this was ob-
served are recorded in TABLE 2.
The aérial roots of vines appear rather fragile in comparison with those
of trees and shrubs, both because of their small diameter and the fact that
they are often irregularly bent.
AERIAL Roots oF THE HERBS
Origin. The aérial roots of herbs may be found within the leafy zone
in most species. In most cases also the roots are formed at well defined
morphological positions. TABLE 3 presents a summary of the data. Selagi-
nella and Dilomilis both form roots at the branch junctions. Most of the
other species root only at the nodes but internodal roots have been ob-
served in Pilea yunquensis.
Tip properties and lateral root formation. The roots of the Sela-
ginella are green as are the apices of the aérial roots of Dilomilis. In the
latter species the region of the tip behind the apex was creamy in color
and green only at the apex.
A few of the species were observed to have lateral roots formed in the
absence of injury and these are recorded in TaBLe 3.
DISCUSSION
Frequency of formation of aérial roots in species. Some species
such as Miconia pachyphylla are usually found with aérial roots but other
species have been found to have no aérial roots. The reasons may be that
too few specimens have been examined or that they do not in fact ever
form them under the conditions experienced in this area. Woody plants
such as Cleyera and Ardisia were not seen with aérial roots, but these
species are not common on the site.
Some of the epiphytes, such as the bromeliad Vriesea, had readily
visible roots but these were not observed hanging free of the host. Simi-
larly no root of the vine Peperomia hernandiifolia was seen hanging free.
1969 | GILL, ELFIN FOREST, 6 207
TABLE 3. The aérial roots of the herbs
Min. Tip PROPERTIES
distance Roots
leaf Max. aterals
. zone at diameter Rigidity and without
SPECIES (cm.) nodes? (mm.) Color alignment injury
Selaginella QO branch 04 ~~ green flexible -
li junctions but wiry,
unbent
Tsachne yes 0.6 white straight & —
angustifolia to pale flexible
green
Dilomilis 2 branch 1.8 green corrugated, —
montana junctions apex, flexible
cream
behind
ilea QO yes 0.3. white to curled,
obtusata pink flexible
Pilea 0 no 0.2 reddish curled, +
yunquensis brown flexible
Sauvagesia QO yes 0.2 pale straight & —
erecta cream flexible
Begonia 1 yes 0.4 white to straight & _
decandra flexible
One large Cecropia peltata was examined and found to have secondarily
thickened “prop” roots, but no tip was seen above ground and in this
Case it appeared that the roots had been exposed by erosion. Some of the
grasses and carices had a preponderance of leafy tissue above ground, and
very little stem tissue and no aérial root was observed. The scrambling
grass [sachne has many stems above ground and in the most humid situa-
tions aérial roots are found. The tree fern Cyathea forms aérial roots in
some cases but these were not studied. Other ferns were also omitted
tom the investigation.
The environment and aérial root formation. Two stages may be
distinguished in aérial root formation. The first is the production of a
Primordium either in association with normal growth and development
of the shoot or formed de novo under certain conditions and in older
In Populus nigra for example, root primordia are present in the aérial
stem but no aérial root is formed. Under the proper conditions of mois-
ture and darkness however, aérial roots may be forced into active growth
(Shapiro, 1962). However, the “proper conditions” for the appearance
208 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
of aérial roots in various species differ. The aérial roots of Rhizophora
mangle in Florida, for example, often show great development in environ-
ments where there is always a high light intensity. In the elfin forest
light intensities are low and roots often arise beneath a mass of crypto-
gams, but the importance of the light factor can only be surmised at present.
Mechanical tissue and/or lack of injury may be important in some
species as roots are often associated with new shoots, which may cause
wounding of the parent shoot as they grow and which are composed of
relatively soft tissue. The humid environment may be essential to out-
growth of roots as desiccation seems to be an important cause of injury to
apices.
Lateral root formation. The freely hanging aérial roots of the plants
in the study area rarely produced laterals in the absence of injury. How-
ever, when these roots enter the soil they branch immediately. Thus there
are two types of control to the lateral root formation, an external (environ-
ment) and an internal (through injury). In the external environment
of the aérial root the high humidity appears to be incapable of inducing
lateral root development in many of the species, and some other environ-
mental factors such as light intensity and nutrient-environment may be
involved.
Some plants with aérial roots fail to develop laterals in the absence of
injury, although many herbs and vines do not. The major member of the
latter group is the palm Prestoea, but its lateral roots are inhibited.
One cause of injury seems to be desiccation. The dead apices of the
roots sometimes show no signs of physical injury but the rare periods of
desiccation seem a likely cause of death. One case of physical injury was
observed on a root of Clusia, which appeared to have been chewed.
Function of the aérial roots. Aérial roots may enable a plant to
spread vegetatively to the surface of another plant or to the soil away
from the base of the parent plant. If the roots from the shoot system
reach the ground the path that nutrients have to travel will be shortened
and this may be an advantage. Vegetative spread from detached portions
of a tree is possible as broken branches with no connection to a parent
tree or the soil have been seen with new roots and shoots. In an area where
trees and shrubs may be pulled over by vines and/or the weight of epi-
phytes and water, or toppled on the steep slopes after a little soil erosion,
the ability to form aérial roots may constitute a valuable property for
survival.
The presence of copious quantities of a gelatinous material on the apices
of the aérial roots of the Hedyosmum is remarkable. Samtsevitch (1965)
has noticed relatively small gel-like caps on the roots of some plants such
as Zea mays in soil and artificial media, and considers that they have
several important functions including protection of the root apex from
mechanical injury, improvement of the root penetration of soil, and pro-
motion of root hair growth. In the area of study it seems that protection
from desiccation is the most likely function of the material, as death of
tips follows its removal (R. A. Howard, personal communication).
1969 | GILL, ELFIN FOREST, 6 209
The anchored roots of Clusia and Ocotea certainly provide support
to their parent trunks. In their absence however, subterranean roots may
provide the plants with the same stability. Thus the adaptive value of
these roots for support is questionable.
SUMMARY
In the humid conditions of the Puerto Rican elfin forest many freely
hanging aérial roots are found on the trees, shrubs, vines, and herbs. Those
of the trees and shrubs are not found in the leafy zone of the shoot system
and lateral root development in the absence of injury is rare. In contrast
the aérial roots of the vines and herbs arise in definite morphological posi-
tions within the leafy zone of the shoot system, and more commonly de-
velop laterals in the absence of injury. Patterns of lateral root develop-
ment may be distinctive, but other properties of the root tips such as
color, rigidity, alignment, diameter, and the presence of secretions, may
also contribute to the character of the aérial roots of the various species.
ACKNOWLEDGMENTS
This study was carried out over a period of 15 days at Pico del Oeste in
the Luquillo mountains of Puerto Rico. The trip was made possible by
a grant to Dr. R. A. Howard, director of the Arnold Arboretum of Har-
vard University, by the National Science Foundation (Grant # GB-3975).
Excellent housing facilities close to the site were kindly provided by Mr.
J. B. Martinson. Mrs. R. J. Wagner helped with the identification and
checking of plant specimens.
REFERENCES CITED
SAMTSEvITCH, S. A. 1965. Active excretions from plant roots and their signifi-
cance. Fiziol. Rast. 12: 837-846. [ Biol. Abstr. 48: 1968
SHAPIRO, S. 1962. The role of light in the growth of root primordia i in 7“ stem
of the lombardy poplar. In: The Physiology of Forest Trees, ed. K. V.
THIMANN, Ronald Press. pp. 445-465.
Wesster, T. R., & T. A. SrEEvES. 1967. Developmental morphology of the
roots of Selaginella martensii Spring. Canad. Jour. Bot. 45: 395-4
Casot FounpATION, HARVARD UNIVERSITY
VARD ForEST, PETERSHAM
MASSACHUSETTS 01366
210 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 7
SOIL, ROOT, AND EARTHWORM RELATIONSHIPS
WALTER H. Lyrorp !
ECOLOGICAL STUDIES Of a small area of elfin woodland on the narrow
ridge top of Pico del Oeste, a 1000 meter-high mountain on the extreme
eastern tip of Puerto Rico where rainfall is about 453 centimeters per
year (Baynton, 1968), were made over a period of two years by several
investigators. The background for the overall study has been given by
Howard (1968).
This report points out some soil and vegetation relationships and the
possible importance that earthworms and other fauna have on soil genesis.
PROCEDURE
Soils were examined and described in three trenches, 6 to 15 meters
long and about 1 meter deep, dug across the 10 to 15 meter-wide ridge of
Pico del Oeste (Fic. 1) at representative sites. Soil horizons were mapped
at a scale of 1:12 and a few soil samples collected for approximate deter-
Fic. 1. The broad western side of Pico del Oeste. The soil study was along
the narrow spine and extended from a location near the right hand side of the
photo to the summit.
* Field work was carried out during March 1-13, 1967, as a part of National Sci-
ence ieee Grant GB:3975 to R. A. Howard, whom I thank for making the
study possible. L. Theobald, W. E. Gensel and R. W. and Mrs. Wagner gon
laboratory and ns assistance, and A. R. Gill, a photograph, for all of which I a
very grateful.
1969 | LYFORD, ELFIN FOREST, 7 211
minations of ignition loss, pH, and moisture content. Weights of forest
floor and the epiphyte-soil blanket on tree stems were obtained. Earth-
worms were collected and their influence on the soil studied.
RESULTS
Soil characteristics. On the narrow ridge of the mountain the soil,
developed on residuum from fine grained volcanic rock, is wet and has a
muck-like surface 25—30 centimeters thick. This mucky surface thins out
and disappears as the soil becomes steep. Under the mucky surface there
is a gray gleyed horizon mottled distinctly with browns, yellows, and reds.
Under this, in turn, lies reddish yellow, massive, plastic, nonsticky clay,
in places with a noticeable content of soft, weathered rock fragments. With
greater depth the soil becomes redder and more fragments of weathered
rock are present. The relationship between the horizons is shown in the
scale diagram of the three trenches (Fic. 2).
Following is a detailed description of the soil. The terms are those in
common use in the United States (Soil Survey Staff, 1951).
O1 Forest floor: a continuous cover of recently oo leaves and twigs
horizon with up to 25 percent of the surface covered with living green bry-
3—0 cm. ophytes (mostly liverworts) and algae; most ‘sae are fragmented
0 : 1 i _»__§ werens
Fic. 2. Scale ees of the soil horizons in three trenches dug across the
Narrow spine of the mountain.
02
horizon
0-30 cm.
Al
horizon
0-30 cm
A2g
horizon
30-60 cm.
B21, B22
horizons
60-90 cm.
JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
and lie directly on the surface of the underlying soil or on roots; in
ee places under the leaf and twig cover there is a 2—2.5 cm. thick
oot floor” consisting of clean coarse and fine roots.
Dark brown or very dark brown (7.5YR 3/2 or 10 YR 3/2 wet)
mucklike material so well decomposed the original plants cannot be
identified, about 40-60 percent organic matter as judged from igni-
tion loss; massive in place and well permeated by fine grass-like roots
and some woody roots; nonplastic, nonsticky; very large earth-
on.
be identified from the remains in the soil takes the material out of
the peat class.
Reddish brown (5YR 4/3 wet) or dark reddish brown (SYR 4/2
with many medium, cre mottles of dark brown (7.5YR 3/2)
and olive gray (SY 5 , in some places with mottles of strong
brown (7.5YR 5/6) ae yellowish red (5YR 5/6); “silty clay
i friable, plastic, nonsticky; many roots; distinct
1 mm. red borders around many of the dead roots; large earth-
worms present and many earthworm tunnels; no or few coarse
fragments of strongly weathered rock.
This horizon is adjacent to the wetter A2g horizon and is at the
top of the very steep slopes. As a whole this horizon has a brown
color in contrast to the overall gray color of the A2g. The texture
feels like a silty clay loam of the northeastern United States but the
soil probably is mostly clay
Dominantly olive gray (5Y 5/2 wet) with 10-20 percent dark
brown (7.5YR 3/2) or reddish brown (5YR 4/3) distinct moderate
size mottles ; ; massive; firm in place, plastic, non-
sticky; many grass-like roots, many large earthworm tunnels filled
with dark brown organic matter.
In places this horizon can be divided into a somewhat browner
upper portion in which the colors are dark brown (7.5YR 3/2) and
light olive gray (SY 6/2) in about a 60-35 proportion with the re-
mainder made up of red and black fine mottles. The gray mottling
in this upper portion is distinct and as a whole this portion appears
to be gray, but somewhat less gray than the lower part.
Reddish yellow (7.5YR 6/8, 7/8 wet) with reddish yellow (SYR
6/8), yellowish red (SYR 5/8) and red (2.5YR 5/8) distinct mot-
tles; “silty clay loam”; massive; firm to friable and digs out readily
with a shovel, plastic, nonsticky, dense and nonporous; in places
with up to 5 percent very pale brown (10YR 7/3) or nearly white
fine scattered mottles; few 5-15 cm. pieces of saprolite agi
with a noticeable yellow or red rind, hard but can be broken with
the edge of the trowel; organic matter-filled earthworm tacks are
common but fewer than in the A2g horizon.
The B21 and B22 horizons are generally yellowish in the upper
part, becoming redder with depth. Mottles are distinct and not
1969]
B23
horizon
and deeper
LYFORD, ELFIN FOREST, 7 213
noticeably in a reticulate (network) pattern. The B21 horizon
shown in Fic. 2 has more conspicuous mottling than the B22 and
was designated as a gleyed horizon (B21g) in the field. It does not,
Weak red (10YR 4/4) and strong brown (7.5YR 5/6) “silty clay
loam”; massive; firm, dense, plastic, nonsticky; contains up to 60
percent angular, hard but thoroughly weathered rock fragments that
break readily by a blow from the edge of a trowel.
The overall color of this horizon is reddish and this contrasts
strongly with the yellowish colors of the B21 and B22 horizons.
Below the B23 horizon at depths ranging from one to several me-
ters is red, yellowish red, and reddish yellow thoroughly weathered
rock (saprolite) that retains its original stratification but is par-
tially fragmented and the fragments can be broken readily. The
bedrock probably is a fine-grained volcanic.
Analyses. The ignition loss, moisture and pH analyses of soil samples
collected from the trenches are listed below. These analyses are approxi-
mate and are presented as preliminary data to provide a rough idea of
the range of some of the soil characteristics.
Moisture to Ignition loss p
nearest 5 to nearest 5 Glass
percent percent electrode
Horizon (Oven dry basis) = (Oven dry basis) 1:1 water
02 (mucky) 270 50 4.4
320 40 4.3
sie 65 mies
Al (edge of slope) 90 20 4.7
den 20 es
A2g (grey, gleyed) 100 25 4.5
80 20 4.9
— 10 a
= 15 a
_ 10 ae
B22 (yellowish) — 15 =
—_— > —
B23 (reddish) 60 5
SS 10 4.7
— 5
= 5 oe
Although the mucky horizon (02), by feel, seems to be mostly organic
matter, the ignition loss analyses indicate that about half is mineral matter.
The gray, gleyed A2g horizon, on the other hand, seems by feel to be
highly mineral but actually has 10 to 20 percent organic matter content.
214 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The soil is very strongly acid throughout as would be expected for any
soil developed from strongly weathered rock under extremely high rainfall.
By feel, the soil in the mineral horizons seems to be sandy, but it is
probably quite high in clay-sized particles. Clay, in many soils of tropical
regions, is aggregated into silt or sand-sized particles and these are not
readily dispersed between the fingers by rubbing, or even by the use of
dispersing agents in the laboratory. Determination of moisture charac-
teristics, however, reveals that a high proportion of the soil is of clay size.
This property is indicated in many of the soils sampled in connection
with soil surveys currently being made in Puerto Rico. (Soil Survey In-
vestigations Report No. 12, August, 1967. U.S. Dept. of Agri. Soil Con-
servation Service in cooperation with Puerto Rico Agri. Exp. Station.)
Analysis of a soil (S58PR—11~-1) collected about two miles away from
the Pico del Oeste site shows that on the basis of moisture characteristics
some of the “sand” and “‘silt” particles have clay-like properties.
The soil on Pico del Oeste seems to have the requisite properties for
an oxic horizon and it qualifies for an Oxisol in the 1965 USDA Soil
Classification (Soil Survey Staff, 1960, 1967). It qualifies as a Latosol
in the older classification scheme.
Water in the soil. The peaty soil of the narrow ridge top is continu-
ously wet judging from observations made during the course of the study.
The path that bisects the area is always muddy and water-proof foot-
wear is required if the feet are to be kept dry. There is a more or less
continuous drip of water from the epiphyte-blanketed trees even when the
sun is shining.
When the trenches were being dug water moved into the excavations pri-
marily by seepage rather than by overland flow. This seepage water
came into the trenches from all horizons suggesting that the whole soil
is waterlogged and not just the upper highly organic portions. For ex-
ample, when the trenches were being dug, earthworm tunnels in the B2
horizons were full of water and under a hydrostatic head, and they
drained suddenly and conspicuously when exposed by the shovel.
Moisture content of the mucky horizon is high and so is that of the
mineral horizons. This high content of moisture is evident if a clod of
freshly collected soil is left in the sun; almost a whole day of good dry-
ing conditions is necessary before the surface of the clod dries.
A few observation wells 20 to 30 centimeters deep were made to ob-
tain some idea about the fluctuation of the water tables in the mucky
surface horizon. In these simple unlined wells, depth to the water sur-
face varied considerably from day to day during the period March 1 to
April 17, 1967, and there were only a few days when water did not stand
within these shallow wells. During the period April 17 to June 15, some
water generally was in the wells but the fluctuation from day to day was
less.
Forest floor characteristics. A forest floor horizon, (01 horizon)
about 2 to 4 centimeters thick lies on the mucky horizon (02 horizon).
1969 | LYFORD, ELFIN FOREST, 7 zA3
i
9 MARET-6
West Peak
PR
3. Components of the forest floor from an area 30 X 30 cm. square.
ABO : Tree leaves in various stages of disintegration are on the right, living
plants (mostly liverworts) in the middle, roots on the left. BELow: Entire and
fragmented tree leaves
216 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
This forest floor is readily separated into three components; namely, liv-
ing plants growing on the fallen leaves (mostly liverworts and algae),
fallen tree leaves and twigs, and roots (Fic. 3). Samples were collected
from six areas 30 30 centimeters square and the average oven dry
weight was determined for each component.
Living plants 314 g/m? (2800 pounds /acre)
Fallen tree leaves 78 g/m? ( 700 pounds/acre)
ts 577 g/m? (5200 pounds/acre)
Bryophytes (mostly liverworts) growing on the fallen leaves occupy per-
haps 10-25 percent of the area. Some are growing in place; others ap-
parently were growing on the leaves while the leaves were still attached
to the tree and continued growth after the leaves fell. In addition to
the bryophytes many of the fallen leaves are about half covered by a thin
layer of green algae.
Fallen leaves and twigs completely cover the surface of the soil and
no bare areas can be seen. Apparently leaves of these evergreen trees fall
one by one throughout the year. Most are entire when they fall but
after being on the surface of the soil for a short while show some signs
of disintegration: parenchyma is removed and the leaves are broken into
fragments (Fic. 3). The amount of organic material (other than roots)
above the mucky 02 horizon at any one time is about equivalent to the
amount that falls yearly in most deciduous forests in the northeastern
United States.
Clean woody roots lie in a layer between the fallen leaves and the mucky
02 horizon. This “root floor” is about 2 to 2.5 centimeters in thickness
and there are many places where the roots arch away from the surface
leaving open spaces beneath,
Roots in the soil. Both aérial roots and roots in the soil are common.
The characteristics of aérial roots on the study area have been described
by Gill (1969),
Many of the roots in the soil grow between the litter and the mucky
surface and constitute the previously described “root floor.” The roots
in this layer are essentially free of adhering soil and are as clean as
though washed with water (Fic. 4). Possibly some of these roots are
clean because they have never grown in the soil beneath. Gill (1969)
observed some roots growing above the soil surface; in fact, some were
even exposed to the atmosphere.
Woody roots on or just under the soil surface, extend for at least 7 to
8 meters laterally and are well exposed in the path (Fic. 4). On the basis
of observations made while digging the trenches it is estimated that 80—90
percent of the woody roots are either just under the fallen leaves in the
“root floor” or are within the upper 2 to 10 centimeters of the surface of
the soil. These woody tree roots seem to be much the same in overall
growth habits as those of forest trees at the Harvard Forest in central
Massachusetts (Lyford & Wilson, 1966).
ok clue, =
< ia me “geet
Ore gait
i a
P = A ‘.
1 NIATA GYOAAT
»
“LSAUYO
/
Fic. 4. Woody roots in the soil. Lerr: Layer of clean roots that commonly exists just under the leaf litter. RIGHT:
Woody roots in the upper portion of the soil that have become exposed in the path,
218 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Vertically descending grass-like roots are numerous in the gray gleyed
A2g horizon and give the soil material a sod-like character. Some of the
dead roots in this horizon are bordered by a thin layer of red soil and
these vertical “pipes” are noticeable when the soil is examined. These
pipes are well known features in many wet soils.
Amount of epiphytes and soil on tree stems. A rather large mass
of epiphytes and roots grows on the stems and branches of the trees. In
addition there is brown soil-like material adhering to the bark and inter-
mingled with the roots and green plants that blanket the stem. This
brown mucky material is essentially identical in appearance to the brown
mucky material that makes up the surface layer of the soil. To obtain
some idea of the amount of this soil-like material on the trees all material
around a portion (20 centimeters long) of the stem of each of six trees
was removed. This was subdivided into portions comparable to those
used for the forest floor, namely, living plants (mostly liverworts), roots,
and soil-like material. Subdivision was made under water to enable com-
plete separation of the soil-like material. Following are the results ex-
pressed as grams of oven dry material per square meter of stem area.
Living plants 265 g/m?
Roots 125 g/m?
Soil-like material 112 g/m?
Weight of green material on the stems is 265 grams per square meter ~
compared with 314 grams for the leaves on the forest floor. Together green
material and roots on the stem total 390 grams per square meter as com-
pared with 392 grams for the combined leaves and bryophytes on the for-
est floor. In other words, the amount of organic matter (other than that
which makes up the soil-like material) on tree stems is not far different
from that on the surface of the soil if compared area for area.
A few determinations of ignition loss were made on the soil-like ma-
terial on tree stems and branches. Most of the material has a 90 to 95 per-
cent loss on ignition so its origin probably is from the decomposition
of plant material in place. The one sample with an ignition loss of 64
percent may have had a different origin.
Probably the soil-like material on stems and branches is largely the
result of the decay of plant tissue in place. Some of the material however,
is granular and definitely coprogenic. Fauna of various kinds are common
under, and in, the plant material which clothes the stem. Millipedes,
centipedes, enchytriads, beetle larvae, tiny red ants and sow bug-like in-
sects are numerous and a black earthworm 10 to 15 centimeters long, like
those in the soil, was found on one tree stem at a height of about 1 meter.
This amount of faunal activity raises a question about the origin of the
soil-like material. Conceivably a large part of it could originate in the
soil and be carried into the trees within the bodies of fauna. In such a
case, however, the casts would probably have a rather large content
of mineral matter; larger than that indicated by the analyses of the
1969 | LYFORD, ELFIN FOREST, 7 219
Fic. 5. Soil occurs in some trees. Lert: Termite nest in a tree by a roadside
within sight of the study area. RicHT: Termite tunnels on a tree stem. W. L.
Theobald is used for scale.
soil-like material observed on the study area. At lower elevations termites
have large nests high in the trees (Fic. 5). They carry a good deal of
material into the trees so the process leading to the presence of soil-like
material in trees is not unusual.
Earthworms and their significance. Earthworms are common in
the soil of the study area and were collected from each of the three trenches
(Fic. 6). Two species were collected but have not yet been identified.
One species is black, about 15 to 20 centimeters long and weighs about
5 grams. It seems to be most common in the upper part of the soil and,
indeed, a single specimen was observed on a tree stem about 1 meter above
the soil surface where it was well protected by the wet epiphyte blanket.
The second species is very large and has a dark gray or dark olive color.
It is up to 60 centimeters in length and 10 millimeters in diameter. Each of
the earthworms weighs about 30 grams. These large earthworms are com-
mon to depths of 50 centimeters and are in all the upper horizons. Their
tunnels are conspicuous when filled with dark-colored soil material and
especially so in the gray gleyed A2g and the yellowish B2 horizons (Fic.
7)
Roughly 1 to 5 percent of the soil mass in the upper 50 centimeters
is occupied by earthworm tunnels. Tunnels in the mineral portion of the
soil are made by ingestion of soil material rather than by simple pushing
aside, and some of the ingested mineral material later is expelled on the
surface. Organic matter also is ingested in large amounts. Leaves on the
surface of the soil serve as food and some of the disintegration of leaves
220 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ste tee BDRM:
Fic. 6. Two species of earthworms from the study area. Apove: Close-up
of one of the large olive specimens. BELow: The four large olive pg
shown in the upper part of the photo. are up to 60 cm. long and on in
diameter. The smaller black earthworms shown in the bottom part of ne atts
are up to 20 cm. long
shown in Fic. 3 is the direct result of earthworm action. Earthworms
probably ingest some of the soil material near the root floor and, in fact,
these roots may have their clean appearance because of the earthworms.
Puddled masses of earthworm casts 5 to 10 centimeters in diameter are on
roots at intervals of about 40 to 50 centimeters, showing that earthworms
are active on the surface; and it is possible the open spaces beneath the
root floor are the result of earthworm action.
1969] LYFORD, ELFIN FOREST, 7 221
Fic. 7. Dark colored areas in the A2g clods are earthworm tunnels filled with
soil of high organic matter content. Scale in the photo. above is in centimeters.
Organic matter ingestion is also evident because of the dark color and
the high ignition loss of the casts in the tunnels. In the A2g horizon the
ignition loss for two samples of earthworm casts from the dark-colored
222 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
tunnels was 65 and 75 percent, whereas that of the soil immediately adja-
cent was 10 and 15 percent. A single sample of earthworm casts from a
tunnel in the B22 horizon had an ignition loss of 50 percent; that of the
soil immediately adjacent was 5 percent.
Thus there is considerable mixing of the organic and mineral matter of
the soil within the bodies of the large earthworms, and a rather great
amount of transportation from one place to another. This mixing and
transportation is not rapid enough, however, to cause the soil horizons
to lose their identity.
In addition to the mixing and transportation there is some segregation
of the larger particles because they are too large to be ingested.
DISCUSSION
The climate and vegetation of the study area very likely is much the
same now as it has been for centuries because of the particular location
of the Pico del Oeste in respect to the ocean and the constantly blowing
easterly trade winds. The soil, therefore, probably represents a kind that
has developed and been maintained under continuously wet tropical con-
ditions. Soil development processes may have been modified from time
to time by addition of volcanic ash, but no known additions of this kind
have been made recentl
Continuously wet conditions have permitted the build-up of a mucky
mineralization of the organic matter. A considerable amount of organic
matter is returned to the surface of the soil each year not only from the
dead leaves of trees but also from epiphytes that grow on the leaves.
The B horizon in this soil seems to qualify as an oxic horizon. This in-
dicates that the minerals of the original bedrock from which the present
soil materials have been derived, are thoroughly weathered. The remain-
ing materials are probably high in kaolinitic clay and the amount of iron
and aluminum compounds present are considerably higher than in the un-
weathered bedrock. In general, this would mean a lack of available nu-
trients, but where the soil has a highly organic surface this is probably
not the case, because organic matter has a high cation exchange capacity
and holds the nutrients so they are not readily leached out.
Action of large earthworms in soil development seems to be significant.
Their large tunnels allow rapid movement of water from place to place in
the soil. Their movement and mixing of mineral material and organic
matter by ingestion is estimated to occur actively in at least 5 percent of
the soil volume. They do not seem to use the same tunnels all the time
but make new ones, so much of the soil material in the upper foot or two
has been passed through their bodies at one time or another.
Earthworm casts are numerous at the surface and are readily observed
when the forest floor is removed. The rate of this deposition on the surface
suggests that there is a minimum of a centimeter or two of soil added to
1969 | LYFORD, ELFIN FOREST, 7 223
the surface every 100 i.e This estimation is based on similar studies
with ants (Lyford, 1964). But if there is this much action there is a
question how the various fone are able to preserve their identity.
Possibly, the earthworms we see now are recent colonizers or perhaps they
are now experiencing an unusual population surge. The large earthworms
are known to occur elsewhere in the island (Luis Maldonado, pers. comm.),
but probably are confined to certain restricted habitats.
On steep slopes, the action of earthworms in returning soil to the surface
is considerable. Their action was observed not only on the steep slopes
of the study area but also in two other places in the Luquillo Mountain
area, on steep slopes. This constant return of earthworm casts to the
surface may have some rather important geomorphological implications
because fresh soil material is always available on the surface for trans-
port by running water or gravity.
Earthworms not only return soil to the surface, they also consume the
leaves that fall on the surface of the soil. They do not, however, consume
all the leaves but allow enough to remain to form a complete cover of
the soil, thus, their activities are not completely detrimental and, perhaps
they have achieved a desirable ecological balance.
Presence of a considerable amount of soil, or at least soil-like material,
on the stems and branches of trees suggests that some epiphytes may have
their roots in soil even though the entire plant is on a tree. “Soil” in trees
raises a problem for the soil scientist because he is not in the habit of
finding it here, and indeed, this makes some revision of the definition of
soil in order. Yet termite mounds several meters in height are common in
tropical regions. These insects not only return soil material to the surface
of the soil, they raised it several meters above the original surface. It is
not too much of a mental jump then to think of soils in trees, particularly
when it is readily apparent that some kinds of termites, ants, and other
soil-moving fauna make their nests in trees.
SUMMARY
Soil on the steep narrow ridge of Pico del Oeste is wet and the mucky
surface, 25 to 30 centimeters thick, has about 50 percent organic matter.
Fallen tree leaves, on which grow liverworts and algae, lie on a layer of
clean roots and these in turn are on and in the soil. Tree stems and
branches are blanketed with liverworts, algae, and other epiphytes, and
the amount of organic matter on the stems per unit area approximates
that on the surface of the soil. Soil-like material occurs on the stems and
branches of trees. Some may be carried up from the soil by fauna. Large
earthworms up to 60 centimeters long and 1 centimeter in diameter are
common in the soil and ingest and move large amounts of it.
LITERATURE CITED
Baynton, H. W. 1968. The ecology of an elfin forest in Puerto Rico, 2. The
microclimate 7 Pico del Oeste. Jour. Arnold Arb. 49: 419-430.
224 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Gitt, A. M. 1969. The ecology of an elfin forest in Puerto Rico. 5. Aérial roots.
Jour. Arnold Arb. 50: 197-209
Howarp, R. A. 1968. The ecology of an elfin forest in Puerto Rico, 1. Intro-
duction and Composition Studies. Jour. Arnold Arb. 49: 381-418.
Lyrorp, W. H. 1963. Importance of ants to Brown Podzolic aoe genesis. Har-
vard Forest Paper 7. Harvard Univ., Petersham, Mass.
& B. F. WItson. 1966. Controlled growth of forest a roots: technique
and a. Harvard Forest Paper 16. Harvard Univ., Petersham,
Mass. 0
SOIL camel penn 1951. Soil Survey Manual. U.S. Dept. Agriculture Hand-
book 18.
. 1960. Soil Classification. A Comprehensive System. 7th apg
265 pp. (Supplement 1967. 207 pp.) Soil Conservation Service, U.S. Dep
Agriculture.
HARVARD FOREST
HARVARD UNIVERSITY
PETERSHAM, MASSACHUSETTS 01366
1969 | HOWARD, ELFIN FOREST, 8 225
THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 8. STUDIES
OF STEM GROWTH AND FORM AND OF LEAF STRUCTURE
Ricuarp A. Howarp*
IN THE FIRST PAPER of this study (Howard 1968), the species compris-
ing the elfin forest on the summit of Pico del Oeste were listed in systematic
order, classified by growth form, and indicated in their frequency through
transect and plot studies. It was shown that 14 taxa of monocotyledons
and 40 taxa of dicotyledons were the common components, and that these
could be distinguished as 25 taxa of woody trees or shrubs, 19 taxa as
herbs or herbaceous vines, 4 taxa as woody climbers, and 6 taxa as
epiphytes.
Various suggestions to explain the reduced stature of such a forest were
also listed. These suggestions involved the poor aeration of saturated soils,
the reduced light reaching the forest through a heavy persistent cloud
cover, the reduced transpiration suggested by the high humidity of the
atmosphere and frequent and high precipitation, and the restrictive effect
on growth of the high wind velocities. The frequent occurrence of trees
which had blown over or had toppled due to the subsurface erosion were
Suggested as factors increasing the density of the woody vegetation
The present study will consider the nature of the growth of the indi-
vidual and component species as factors in the form of the forest and will
document the presence or absence of characteristics of plant structure
which have been observed in comparable forests elsewhere by other
workers.
GROWTH AND FORM
The short stature of the elfin forest on Pico del Oeste is in part due to
the nature of the growth and of the mature form of the individual species.
The growth and form of a particular plant may be the expression of a
normal genetic factor exhibited at the specific, generic, or familial level
as the herbaceous, climbing, woody, rhizomatous, or rosette-forming habit.
Unusual or sbnortasl variation from the basic pattern may be an indi-
vidual characteristic due to biotic factors of the environment or to muta-
This study was possible only with the financial support of a grant from the Na~-
on ae Foundation (GB:3975) for which I am deeply indebted. The present
rs in tabular form data which represent much detailed work of counting
ig dn measuring. I repeat my gratitude to my wife Elizabeth Howard and our children
Jean, Barbara, and Bruce for their efforts in collecting, counting, and measuring leaves.
The work of drying foliage material, of weighing and obtaining water contents, and
of a the pH values of leaves involved the cooperation of sad ger Wagner
and his wife, Anstiss Wagner. Mrs. Helen Roca-Garcia prepared m f the slides,
patiently d on the leaf section and stomatal measurements, and eaten: ‘the: illustrations.
226 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
tions. We did find on rare individuals a fasciation of branches resulting
from insect damage to an apical meristem. Witches’ brooms or comparable
fasciations due to fungi or mutations did not appear within the forest
under study. A few shrubs, e.g., Cleyera and Symplocos, had scrambling
branches when intermediate in height. Aberrant leaf forms were encoun-
tered and these were the result of abnormal laminal development following
insect damage of primordia or very young leaves.
The genetic-based habit of the plant can often be a growth pattern
associated with the production of flowers. The production of a terminal
inflorescence may impede the normal vegetative extension of a stem and
result in a branch dichotomy. Vegetative development may be slowed by
the development of a resting or short shoot area. Die-back or self-pruning
of a limited amount of shoot development occurs in some species to restrict
ultimate growth. By contrast axillary or cauliflorous inflorescences can be
produced below the apical meristem of the branch or in the axils of sub-
terminal leaves and have no effect on the ultimate habit of the plant.
MONOPODIAL GROWTH
Growth can be described as continuous if a dormant terminal bud is
not formed and if leaves are produced in a regular sequence throughout
the year. Such a condition is often attributed to tropical forests. The
terminal growth of a branch can be interrupted by a resting period, whether
a bud is formed or not, and new growth can be by a flush of new leaves
with the subsequent lengthening of the internodal zones. The last leaves
formed in a flush of growth may be closely associated, indicating that
internodal elongation was reduced. The subsequent new apical growth
may also have reduced basal internodes, and in fact, the first leaves formed
may be aberrant in size and shape and be termed scales or cataphylls. In-
ternodal areas in or above the zone of scales or cataphylls may be longer
than average and the flush may again terminate in an apparent rosette
of leaves. Such a growth pattern is basically a monopodial production of
a long shoot terminated by an area of a short shoot. Jlex sintenisit ex-
hibits this growth pattern on Pico del Oeste.
SYMPODIAL GROWTH
Occasionally the long shoot is produced not from the terminus of the
short shoot, but sympodially from an axillary bud of a lower leaf. When
the growth of a single leader extends vertically, the lateral offset of the
new shoot may be noticeable for only a short period of time. It is evident,
however, that the leader was terminated by a short shoot development
and although the sympodial offshoot continued growth in the same direc-
tion, a restriction was imposed in the rate of apical elongation. Such a
growth pattern was found in Torralbasia.
Within the Pico del Oeste forest nine genera exhibited a restriction in
vertical growth by the production of long shoots and short shoots and
1969 | HOWARD, ELFIN FOREST, 8 227
>
= ‘i
Va NN SS
: if
iy
WW)
Ficure 1. Wallenia yunquensis. Plant grown from seed in greenhouses of the
Arnold Arboretum. Short shoots, or terminal rosettes of leaves, and the single
unit sympodial branching are all shown.
were further affected by the development of lateral branches. Wallenia
yunquensis commonly exhibits a vertical shoot producing a single lateral
flush of growth as a branch (Fic. 1). The erect main shoot of Wallenia
228 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
may be interrupted by short shoot areas separated by areas of long shoot
development. From one or more leaf axils in the short shoot zone a single
fast-growing lateral branch may develop which appears naked at the base
but does in fact have widely separated cataphylls of very short duration.
The naked shoots are terminated with a short shoot zone possessing an
aggregate of leaves. The lateral branch may also originate from the area
of cataphylls. In Wallenia the lateral flush shoots never branched or con-
tinued growth beyond the initial flush.
An example of repeated sympodial lateral branching is readily seen in
Ocotea spathulata. The development of flush shoots appeared to be from
a short shoot zone in all cases, but there developed additional and com-
parable lateral shoots from the terminal short shoot zone of the lateral
branch. This growth pattern results in a sympodial development of lateral
branches in a flat plane. The principal branches are tiered in appearance,
the tiers being separated by an unbranched, seemingly naked stem. This
growth form has been described as candelabra-branching, Terminalia-
branching, or as pagoda trees.
Corner (1952, p. 32) described this growth pattern for Terminalia as
follows: “The leader-shoot rather suddenly lengthens into a long vertical
fea clothed with a lax spiral of leaves . . . its growth slackens. . .
. another terminal rosette is produced. From the base of this
ashe several twigs grow out to form the next tier of branches . . . The
positions of the branches in successive tiers usually alternate so that only
those of every other tier are superimposed.”
Lateral branches from the terminal short shoot area may also grow
vigorously, producing scales or cataphylls before developing normal leaves
and, ultimately, each its own terminal shoot. Corner (1952, p. 31) de-
scribed the lateral growth as follows: “Each twig which grows from the
leader-shoot of the tree does so rapidly and at a wide angle from it; then,
as its growth slackens, it turns up at the end and from its lower side, just
at the bend, a branch arises to grow out as another twig which will follow
the same course by turning up at the end and branching in its turn. - -
In the first horizontal part of such a twig the internodes are lengthened;
the leaves, or their scars, are widely spaced on the slender stem; and the
growth has been rapid so that the new shoot has quickly been thrust be-
yond the parent rosette of leaves. In the second, vertical or upturned,
part of the twig the internodes are very short or absent and the leaves,
or their scars, are very crowded on a stout stem so that, while many more
leaves are being produced than in the previous stage of the twig’s develop-
growing, withers and falls off. When such a limb .. . is growing out
from the trunk of the tree, it diverges from its neighbours and begins to
branch sideways: this it does by producing every now and again not one
twig but a pair of twigs, or even three, which grow out from each other
at a wide angle; and thus the limb develops into a fan-shaped leafy spray.”
The Terminalia-type of branching was particularly conspicuous in Ocoted
spathulata (Lauraceae), one of the dominant trees of the elfin forest.
1969 | HOWARD, ELFIN FOREST. 8 229
Ib
W 3 Z
6
he
3b
3a
Ss
le
2b
3b
4a
Tc
2b “
Ib 2a
. 4b
? 3a
IGURE 2. Diagram of growth pattern of Ocotea spathulata. A, side view
showing tiers of repeated sympodial branching; B, view from above showing
the relationship of the tiered branches. Drawing by Pamela Bruns.
a
230 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Ficure 2A shows, diagrammatically, a plant of Ocotea of average stature
in the forest with four tiers of sympodially developed lateral branches. It
is evident that the terminal growth is impeded while the lateral sympodial
growth proceeds. Lateral branches at 4a or 4b show four additional
flushes of growth while only 3 periods of reactivation of the terminal shoot
are evident. The lateral branches, la and Ic, have each had one reactiva-
tion of growth while the terminal remains static. The relationship of the
lateral branches is illustrated in Ficure 2B and shows a conflict with
Corner’s observation that the branches of alternate vertical flushes are
superimposed.
Specimens of Ocotea were found within the elfin forest with 27 flushes
of sympodial lateral growth on the oldest branch. Only the last 13 of these
sympodial flushes retained any foliage demonstrating a die-back of the
upturned short shoot after a considerable period of continued lateral
sympodial growth.
Terminalia-branching was also observed in Ardisia and Grammadenia
(Myrsinaceae), in Torralbasia (Celastraceae), and in Calycogonium
(Melastomataceae). Terminalia-branching is conspicuous in Hedyosmum
arborescens and is due to the naked areas of elongation and the terminal
production of leaf pairs or of an inflorescence. Continued growth of the
sympodial branches in this species was restricted in many examples by the
production of an inflorescence.
The sympodially branched lateral shoots can develop vertical extensions
as well as horizontal branches. Vertically developed shoots as elongate
leaders of vigorous growth have been observed on the lateral branches
of specimens of Ocotea and Calycogonium, but in all cases additional
sympodial branching also occurred from the same upturned short shoot
and beyond it.
Attempts were made to induce either vertical elongation of the up-
turned shoot or the production of new or additional sympodial branching
by pruning the leader shoot. During the three year time interval of the
study all of the branches which had been pruned of lateral growth failed
to respond by any new development from the areas of the upturned shoot.
Likewise, vertical leaders when partially or completely decapitated by
pruning failed to develop any sympodial lateral branches.
Although many branches were marked along the trail to record growth
phenomena, we were unable to draw conclusions on the frequency with
which sympodial branching occurred normally. In no case where sympodial
branching was noted in early stages of development (and the branch
tagged for observation) was there any further sympodial branching. We
could only conclude that the sympodial branches were not produced an-
nually on any branch we had marked for observation.
Gill has reported the occurrence of adventitious or aérial roots on the
species within this forest. Although adventitious roots were observed on
the horizontal branches of the species exhibiting sympodial lateral branch-
ing, the roots did not appear to be associated with the short shoot area
or the curved portion of the lateral branch. Ocotea, which had the most
“ _“-s-----
1969 | HOWARD, ELFIN FOREST, 8 231
conspicuous Terminalia-branching, rarely produced adventitious roots
from the horizontal branches but did develop “prop” roots from the base
of the stem.
The development of sympodial branching or of vertically continuous
long shoot-short shoot growth patterns was not associated with flowering
in Ocotea, Grammadenia, Torralbasia, Calycogonium or Wallenia. Ardisia,
however, did develop a terminal inflorescence, and following the maturity
of the fruit and the fall of the inflorescence axis, a lateral but vertical
continuation of the stem developed as a sympodial flush of growth.
DICHOTOMOUS BRANCHING
The dichotomous branching of upright shoots was observed in a number
of the components of the elfin forest. Dichotomous growth and branching
was most conspicuous in Calyptranthes where it occurred, on the average,
every three internodes. The new shoots developed in pairs and normally
two pairs of leaves developed in each flush before elongation stopped. At
the apex of each shoot there was a terminal aborted primordium. Subse-
quent branching occurred lateral to the terminal aborted primordia but
remained consistently in one plane. Calyptranthes appeared to be a col-
lection of upright fans of dichotomous branches.
he three species of Miconia always developed upright dichotomous
shoots when growth was terminated by the production of a terminal in-
florescence. Two lateral buds continued the upright vegetative growth
after the inflorescence had matured fruit and had fallen. Subsequent
growth consisted of but one or two pairs of leaves per flush. In mature
plants flowering followed the maturation of each flush of growth on an
annual basis.
In Eugenia borinquensis one or two pairs of leaves formed each flush
of growth. Flowering occurred only on mature stems and terminated the
branch or was formed in an axillary position on the old wood.
Both species of Psychotria produced a terminal inflorescence and there
was no further vegetative growth on that shoot while the inflorescence
persisted. With the maturity and desiccation of the inflorescence, how-
ever, two basal axillary buds developed in Psychotria berteriana, produc-
ing a dichotomous growth pattern. In Psychotria guadalupensis, however,
only a single bud developed at the base of the inflorescence and the result-
ing growth was falsely monopodial.
Among the herbaceous vines Jpomoea repanda, with alternate leaves,
produced a terminal inflorescence of many flowers which matured over
an extended period of time. Axillary vegetative shoots often developed
while the inflorescence was only partially mature. In Mikania pachyphylla,
with opposite leaves, a terminal inflorescence appeared to restrict apical
growth while the terminal inflorescence matured, but then a dichotomous
growth pattern developed through activity of two axillary buds.
The heteroblastic growth of Marcgravia sintenisii also showed an asso-
Ciation with the production of a_ terminal inflorescence. Subsequent
oa2 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
growth was by the development of axillary buds below the inflorescence.
In all cases observed, leaves of the initial production on this axillary
shoot were of the juvenile form whether or not the branch was in contact
with a trunk or branch
Plants of Tabebuia rigida formed the second major component of the
forest and these plants produced flowers throughout the year mostly from
new growth. The terminal flush of growth consisted of an average of
branches. Most of the plants of Tabebuia which were observed in the
canopy of the forest also showed a significant die-back of the flush growth
during the winter season. Subsequent apical growth, therefore, came from
adventitious buds in the axils of lower leaves or from opposite buds of such
a node
DIE-BACK
Regular die-back of terminal and, less frequently, of lateral or sym-
podial shoots was observed in Jlex sintenisii, Cleyera albopunctata, Eugenia
borinquensis, Hornemannia racemosa, Ardisia luquillensis, Micropholis
garciniaefolia, Alloplectus ambiguus, Gesneria sintenisii and Lobelia porto-
ricensis. Although regular die-back has been described for plants of
temperate areas, its occurrence as a factor in the size of a plant in tropical
areas has not been recognized previously (Garrison & Wetmore 1961).
Relatively long flushes of shoot development were observed in Gono-
calyx and Hornemannia consisting of 5~10 leaves or internodes per flush.
The young leaves were brightly colored and soft until full extension of the
shoot, or the full development of the leaf size, was completed. Both
species were climbers and the soft shoot development was often injured
mechanically and the entire flush of growth abscissed.
The height of the forest may be affected by the environmental factors
previously suggested, but clearly the low stature of the component woody
species may also be due to genetic factors expressed as long shoot-short
shoot development, dichotomous branching associated with a terminal
inflorescence, the abortion of the shoot tip, and the die-back of seasonal
flushes of growth.
Continuous production of single leaves or leaf pairs occurred in all of
the herbaceous species in the elfin forest. Continuous production of single
leaves or leaf pairs appeared to occur in both juvenile and adult shoots of
Marcgravia, in Symplocos, Cleyera, Ilex, Gesneria, Clusia, and Micro-
pholis. The herbaceous vines Rajania, Ipomoea ani Mikania also ap-
peared to produce leaves continuously unless affected by flowering
The rosette and epiphytic habit of the two members of the ee
found within the elfin forest can be regarded as a family genetic character.
Following flowering, however, the two species continued growth in differ-
ent patterns. Rosettes of both species died following flowering, but plants
of Guzmania produced one or, rarely, two basal vegetative rhizomes which
1969 | HOWARD, ELFIN FOREST, 8 233
developed laterally before terminating into a rosette or crown of leaves.
This growth pattern caused the plants to form a ring around the host tree
and Guzmania was most commonly found on the large trunks of Prestoea
montana. Vriesea sintenisii by contrast, produced-a single basal rhizome
which tended to grow upward immediately and formed a new crown in
close competition with the parent rosette. The new growth could be to
the right or the left of the parent plant but always extended upward.
Plants with 7 generations of rosette-rhizome vertical development were
found. When Vriesea sintenisii occurred on a branch extending horizontal-
ly, the plants persisted for only 2 or 3 growth generations before being
extended slightly off center and, seemingly top heavy, falling over to
break free and drop to the ground.
BUD PROTECTION
Richards (1952, p. 77) notes that “buds of rain-forest trees and shrubs,
as might be expected, are less well protected than those of trees in other
climates.” An examination of the terminal foliage buds or the shoot apex
in resting condition revealed that the leaf primordia are better protected
in the plants of the elfin forest than might be expected from Richards’
statement. (PLATE I).
Protective stipules are present in Hillia and Psychotria of the Rubiaceae
and in Calyptranthes of the Myrtaceae. In Hillia (PLATE Ib) the stipules
form a sheath around the young leaves which is compressed at the apex.
The developing young leaves force an opening in the apex of the sheath.
Psychotria species have smaller stipules consisting of an ochrea-like base
with four short free apices. The apices tend to be closely associated in
very young buds but their protective function would be of short duration.
Calyptranthes possesses a peculiar type of stipule protection for which
we have not found a description elsewhere (PLATE Ig). In fact Berg, in
a monograph, reports the family to be estipulate, as have subsequent
the original description of Calyptranthes krugii, Kiaerskou notes, “Quaque
innovatio e duobus internodiis constat, quarum alterum breve duo cata-
phylla opposita cito decidua, alterum longum duo euphylla fert.” In a
footnote he equates “cataphylla” with “Niederblatter Germanorum.” In
our observations of Calyptranthes krugii on Pico del Oeste the vegetative
shoot increase is by production of 1 or 2 pairs of leaves in a flush. The
apical meristem aborts although an inflorescence of one, rarely two,
flowers may be produced in one or both terminal leaf axils. Subsequently,
after flowering or resting, two axillary shoots develop and in each the apex
is covered with a pair of laterally folded bud scales. The young leaves in-
crease in size uniformly and are appressed by their ventral or adaxial
surfaces. As the leaves increase in size, the bud covering is forced apart
or torn free at the base, and the two halves separate as conduplicate folded
sheaths. The bud scales are a light yellow or cream color in contrast to
234 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
the green shoots. When separate they dry white, then brown and shrivel
before falling from the shoot. Although these bud scales are rarely found
on herbarium sheets, they were conspicuous in the living plants of Calyp-
tranthes krugii in the study area and were also found on a population of
Calyptranthes which may represent a different species in the Cerro de
Punta area. The term cataphyll although broadly inclusive for the early
leaves of a plant or shoot as cotyledons, bud scales, etc. (Jackson) is
scarcely descriptive of the folded protective scales of the young leaves
of Calyptranthes.
Apical buds may be protected by leaf bases as in Clusia which has
opposite leaves or by the cluster of leaves in the several genera which
produce terminal or lateral long shoots where the terminal apical elonga-
tion is reduced. In Clusia (PLATE If) the mature leaves conceal the young
buds as the leaf bases of opposite leaves of a pair are tightly appressed.
In Wallenia, Ocotea, Grammadenia, Torralbasia, Ilex (PLATE Ic), and
others, the vegetative shoot in resting condition is terminated by a dense
cluster of small leaves or primordia. In subsequent development of the
shoot represented by these primordia, the outer ones enlarge only slightly,
frequently failing to develop a leaf blade even though the petiole may
elongate. Such scales or cataphylls are found at the base of the long shoot
and the internodes between them may or may not have elongated. Clearly
these cataphylls have served a function of protection for the inner leaves
and the apical meristem.
The buds or apical meristem of shoots of Hedyosmum are enclosed
within the sheathing stipular base of the leaves (PLaTe Ie). Bud protec-
tion here is evident in the enclosure of the primordia in the sheathing leaf
base.
The apex of the stem of Micropholis and Symplocos have the young
leaf primordia tightly invested in a protective covering of brown trichomes.
As the leaves expand these trichomes are separated and in many cases
break off. In Tabebuia rigida the young leaves or primordia are tightly
and completely encased in a shield of brown peltate scales. Again with
leaf enlargement the scales are separated and often persist in isolated posi-
tions on the mature leaves.
The species Gesneria sintenisii appears to have a large naked meristem
where the leaf primordia are separated and evident from an early age
(PiateE Id). The leaf primordia and the apex of the stem have a dense
resinous covering. As the leaf starts to expand the resinous covering is
cracked and usually flakes off although sections of the covering may per-
sist even on the mature leaf blade and the petioles.
The young leaves of Marcgravia (Pate Ia) and Cleyera are convolute
in bud and appear to unroll in development. The apical meristem is en-
closed within this pointed bud and the youngest leaf primordia do receive
some protection.
It is clear from these examples that the young leaves are not without
protection in the majority of the species that comprise the woody com-
ponents of the elfin forest on Pico del Oeste.
1969 | HOWARD, ELFIN FOREST, 8 235
LEAF SIZE AND MORPHOLOGY
Plants of tropical forests have been grouped on the basis of leaf size:
dimensions and areas. Raunkiaer proposed a classification of life forms
on leaf-size classes which has been used for comparison and description by
many authors. Leaves have been termed leptophylls if their area does
not exceed 0.25 cm.*; nanophylls if their area is between 0.25 and 2.25
cm.*; microphylls if the area is 2.25-20 cm.?; mesophylls if the leaf area
is 20-182 cm.”; and macrophylls if the area is 182-1640 cm.” Cain et al.,
found, in a Brazilian rain forest, that the phanerophytes are strongly
mesophyllous and reported a tendency for the small leaf size classes to
have a higher percentage in taller strata than in lower ones
Brown, in his study of the mossy forest on Mount Maquiling in the
Philippines, found only the leaf-size classes of microphyll and mesophyll
represented in approximately the same numbers.
The elfin forest on Pico del Oeste had a single species (Peperomia
emarginella) of a size class smaller than the nanophyll classification and
the majority of plants were of the microphyll size class. The total classifi-
cation in numbers of taxa and percentage of the totals is the following:
leptophylls 1 1.9%
nanophylls 6 11.5%
microphylls 30 57.6%
mesophylls 13 25.0%
macrophylls 2 3.8%
Compound leaf types were represented only by Trichilia pallida, a
species clearly only surviving and not reproducing in the elfin forest zone.
At lower elevations Trichilia pallida becomes a small tree while most of
the plants found on Pico del Oeste were weak saplings, dependent for
support on the surrounding vegetation and nearly scrambling through
the elfin forest. A single plant of Weinmannia pinnata (Cunoniaceae) with
compound leaves was found on the peak but was not encountered in the
transects. Brown did not have a compound-leaved plant in the Philippine
study area and Lebrun notes such plants are less than 15% in African
elfin forests.
The largest leaves, macrophylls, were those of Prestoea montana, a
palm, restricted to the leeward erosion valleys and Anthurium dominicense,
an epiphytic member of the Araceae.
When grouped according to habit the following leaf-size classification
was obtained:
LEAF SIZE VINES-SCRAMBLERS HERBS EPIPHYTES TREES & SHRUBS
leptophylls 0 1 0 0
nanophylls 1 2 2 l
microphylls 3 10 2 a
mesophylls 1 2 2 8
macrophylls 0 0 1 1
Brown added data on leaf dimensions and leaf margins to his study of
the Mt. Maquiling forest in the Philippines. He found the leaves were
236 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
0-10 cm. long in 11 species or 70%, and 10-20 cm. long in 5 species or
30% of the plants. The leaves were 0-5 cm. wide in 12 species or 75%,
and 5-10 cm. wide in 4 species or 25%.
In the elfin forest of Pico del Oeste the leaves were 0-10 cm. long in
7 species of monocotyledons and 31 species of dicotyledons or 70% of the
total flora; and 10-20 cm. long in 7 species of monocotyledons and 9
species of dicotyledons or 30% of the total. The leaves were 0-5 cm. wide
in 10 species of monocots and 33 species of dicotyledons or 79%, and 5—10
cm. wide in 4 species of monocotyledons and 7 species of dicotyledons or
21%. The leaves selected for these measurements were taken from the
mature growth and were averaged for the plant. Variation in leaf size
within a given plant ranged from the cataphylls and reduced leaves of
initial growth of long shoots to the larger leaves of vegetative shoots,
when compared with those of flowering branches. Heterophylly was found
in dimorphic pairs of leaves in Pilea krugii, Pilea yunquensis (Urticaceae)
and in Alloplectus ambiguus (Gesneriaceae). Heteroblastic growth was
found only in Marcgravia sintenisii with appressed smaller leaves on
juvenile and climbing shoots and larger leaves on the free arching branches.
Heterophylly with age was observed in Ocotea spathulata and Symplocos
micrantha where the leaves of seedling plants appeared to be quite dif-
ferent in size and shape from those of adult plants. Although Cleyera
albopunctata appeared to have larger than average leaves on some vigor-
ous growing branches this could not be documented with measurement of
samples. However, the leaves of sterile or vegetative branches of Clusia
grisebachiana did possess larger leaves than were found on shoots which
were mature or produced inflorescences. Macrophylly on adventitious
shoots was not encountered within this forest.
Much attention has been given in existing studies of tropical forests
to the shape of the leaf, the nature of the margin, apex and base of the
blade, and to the presence of a cuticular layer in relation to the retention
of water or the presence of epiphyllous organisms.
Brown noted in his study of the mossy forest at 1000 meters in the
Philippines that as the altitude increases there is a marked increase in
the percentage of small leaves and a decrease in the percentage of leaves
with entire margins. It has been suggested that the presence of marginal
teeth aids the runoff of water from the leaf surface, and Brown found
entire leaves in eleven species of Philippines plants in the study area and
five species in which the margin was not entire. In Puerto Rico on Pico
del Oeste 29 taxa had entire leaves while eleven taxa of dicotyledonous
plants had leaf margins with coarse or blunt teeth or with marginal un-
dulation.
The extended leaf tip, often called a drip-tip, has a popular association
with wet tropical forests. The conclusion of Junger has been cited re-
peatedly that the function of the pointed leaf tip was to hasten the run-
off of water from the leaf, and thus help prevent insects and lower plants
from attacking them. Baker recorded 37 of 41 species of plants belonging
to 20 families with leaves drawn out into a tip, in a forest in Ceylon, and
1969 | HOWARD, ELFIN FOREST, 8 237
Richards observed, “Pointed tips to leaves are characteristic of plants of
wet regions and especially of tropical rain-forests, but I doubt whether
any rain forest can show the phenomenon more markedly than the Sin-
haraka.” Richards observed that drip tips are common and better devel-
oped in the lower than in the upper strata of the forest and in juvenile than
in mature leaves of tall trees. Cain e¢ al. reported that 70.6% of the
leaves in the Brazilian forest they studied had acuminate tips and 28%
of the 150 species studied had rather abruptly long tips of the drip point
type.
By contrast, Shreve found drip tips uncommon in the montane rain
forest of Jamaica, and Vaughan and Wiehe reported a similar observation
for upland climax forest of Mauritius.
In the Pico del Oeste forest 13 of the 14 taxa of monocotyledons had
the leaves acuminate at the apex and the other taxon had leaves acute.
Among the dicotyledonous plants the apex could be classified as acuminate
in 15 taxa of which 6 would qualify as drip tips; 14 taxa had the leaves
acute at the apex and 11 had the leaves obtuse, blunt, or emarginate.
Although previous authors have not considered the leaf base, it seems
that if leaf shape is important for drainage in one direction, it is equally
so in the other. Only 5 of the 14 taxa of monocotyledons have petioles
and of those, Prestoea montana, the mountain palm, has lacerate or com-
pound leaves; Rajania cordata and Anthurium dominicense have the basal
lobes extended and Renealmia antillarum and Brachionidium parvum have
the leaf base obtuse. Of the dicotyledonous plants 24 taxa had the leaf
base blunt, acuminate, or decurrent on the petiole while 16 are best de-
scribed as cordate to peltate at the base.
All of the leaves which were cordate, hastate, or peltate at the base had
an acuminate apex or a drip tip. All leaves which were blunt at the apex
or rounded or emarginate had acute or decurrent leaf bases except for
Micropholis garciniaefolia and Eugenia borinquensis. In these two taxa
the attitude of the leaves to the stem tended to be either upright or droop-
ing and in this manner adapted to the runoff of water. Excepting Eugenia,
those leaves with short petioles or with petiole:blade ratios 1:10 or
larger, all had tapering blade bases with the leaves mostly arranged up-
ward in attitude. Cleyera albopunctata has short petioles but the leaves
have a noticeable curvature. Marcgravia sintenisii, again with a short
petiole, also has a slight curvature and a drip tip.
The frequency of taxa having leaves of strongly curved form suggests
a selective value can be attached to this growth form. The blades may
be noticeably curved longitudinally as well as laterally or in but one plane.
This curved form is particularly evident in taxa of Calycogonium, Cleyera,
Gesneria, Gonocalyx, Hornemannia, Ilex, Miconia pycnoneura, Symplocos,
Tabebuia, and Torralbasia, that is in 10 of the 40 taxa of dicotyledons or
25% of the flora. ;
A heavy upper cuticle was found in 22 taxa or 55% of the dicotyledon-
ous plants. Ardisia, Cleyera, Clusia, Calyptranthes, Calycogonium, Eu-
genia, Gonocaylx, Haenianthus, Hillia, Hornemannia, Ilex, Marcgravia,
238 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Miconia foveolata, Miconia pachyphylla, Miconia pycnoneura, Micropholis,
Ocotea, Psychotria guadalupensis, Symplocos, Trichilia, Torralbasia, Tabe-
buia, and Wallenia, that is, all woody taxa except Mecranium amygdali-
num, Grammadenia sintenisii, Hedyosmum arborescens and Psychotria
berteriana possess a heavy upper cuticular layer.
Junger found that leaves with drip tips were less frequently overgrown
with algae, fungi, lichens and bryophytes than those without. He be-
lieved that the presence of these epiphyllae interfered with assimilation
to such an extent as to be a serious handicap to the plant. Micropholis
garciniaefolia and Eugenia borinquensis, which stand out as the only taxa
of the 40 dicotyledons or 29 woody plants in which the leaves were rounded
or cordate at the base and rounded at the apex and appear to lack any
special adaptation for getting rid of surface water, seemed to support
the larger populations of epiphyllous plants. The abundance of epiphyl-
lous leafy Hepaticae on different species will be considered later in this
paper in relation to the metabolism of the forest.
In a superficial classification of the texture of leaves within the forest
components, the leaves would be considered as membranaceous in all of
the monocotyledons except Anthurium, which had leathery leaves. Among
the dicotyledonous plants 13 taxa would have the leaves classified as
membranaceous, 8 taxa would be described as fleshy or succulent, and 21
taxa as having the leaves leathery or coriaceous. The high percentage
of water in the tissues or the relatively small amount of material forming
dried weight will be considered later and is indicated in TABLE 1, column 8.
The heavy texture of the leaves, the thickness of the blade, and the
amount of succulence all support previous suggestions that the flora of
the mountain summit shows many xeromorphic characteristics. Bews re-
gards the rain forest type of leaf as xeromorphic and associates its charac-
ters with the low specific conductivity of the wood for water. Shreve re-
marks that the prolonged occurrence of rain, fog, and high humidity at
relatively low temperatures places the vegetation of a montane rain forest
under conditions which are so unfavorable as to be comparable with the
conditions of many extremely arid regions. Xeromorphy is usually in-
terpreted from such anatomical characteristics as cuticle, hypodermis, thin
palisade layers, pubescence or idioblasts.
Wylie (1954) noted that a xerophytic flora may have a high proportion
of representatives with leaves having a hypodermis. In his studies of
plants of North Island in New Zealand, Wylie, even though avoiding “ex-
treme xeromorphs and succulents,” concluded that the species studied re-
vealed a high average thickness of leaves, extensive spongy mesophyll and
palisade parenchyma areas, great cuticular depth, and “the proportion hav-
ing a hypodermis were greater than for any group previously studied.”
Wylie (1946) compared his studies of the New Zealand plants with
previous ones of his own, based on plants of Florida and of other temper-
ate areas.
Wylie reported that the leaves of the New Zealand species studied,
ranged in blade thickness from 731 » for Pseudopanax to 172 p for Olearia,
1969 | HOWARD, ELFIN FOREST, 8 239
and the 38 species averaged 406 ». This was much greater than the corres-
ponding thickness of 216 » for 121 Florida dicotyledons and 80 » for 80
species of northern dicotyledonous trees. Philpott reported a mean blade
thickness of 234 p for 24 species of Ficus growing in Florida and Cooper
found a mean laminar thickness of 336 » for 19 species of woody dicotyle-
donous plants in the climax chaparral in western California.
Within the Pico del Oeste forest the woody plants by comparison had
leaves ranging in thickness from 787 p» in Clusia grisebachiana to 146 p in
Psychotria berteriana and averaged 379.6 » in thickness. The herbaceous
flora had leaves ranging in thickness from 625 p» in Peperomia hernandii-
folia to 141 » in Sauvagesia erecta, and all herbs had leaves averaging
281.6 w in thickness (TABLE 2, column 1).
Wylie reported that a hypodermis was found in 24 or 63% of the 38
species examined in the New Zealand study area. Eighteen species had a
hypodermis on both the upper and lower surface, 5 species had only an
upper hypodermis, and one species is described as having only a hypoder-
mis on the lower side.
In the Pico del Oeste elfin woodland 19 of 40 taxa or 47% have a hypo-
dermis. Two taxa, Begonia decandra and Hillia parasitica had both an
upper and a lower hypodermis. No plant was observed with only a lower
hypodermis. Seventeen taxa had an upper hypodermis alone. Twenty-
one taxa did not possess a hypodermis (TABLE 2, columns 2, 3).
The presence of a hypodermis is often regarded as a xeromorphic charac-
ter, although a multiple hypodermis is also an anatomical characteristic
of taxonomic value. Carlquist noted that “continued periclinal division
of the epidermis is of taxonomic importance in certain families such as the
Piperaceae.” Within the plants of Pico del Oeste, multiple hypodermal
layers were found in taxa of Peperomia (Piperaceae), Hedyosmum,
(Chloranthaceae), Ocotea (Lauraceae), Clusia (Guttiferae), Calycogo-
nium (Melastomataceae), and Hornemannia (Ericaceae).
A ratio was determined between the thickness of the upper epidermis and
that of the upper hypodermis in this mossy forest. Ratios varied from 1:1
in most plants with a hypodermis, to 1:15 in Psychotria guadalupensis
and averaged 1:4.1. Although Wylie did not use such a figure calculation,
the figures in table 2 of his paper (1954) suggest a ratio range in the New
Zealand plants from 11:1 to 1:21.3 but an average of 1:3.8, or less than
that found in the Puerto Rican vegetation.
Stalfelt considers the mechanical strengthening of leaves through the de-
velopment of sclerenchyma as particularly common among xerophytes as
a means of reducing the injurious effect of wilting. Branched idioblasts
were found in but four taxa within the mossy elfin forest (TABLE 2, col-
umn 8 and Prats IIa).
Watson concluded that the formation of palisade tissues in leaves might
be a morphological response to light. He suggested that the cigar-shaped
palisade cells are formed in increasing number with increasing light in-
tensity during leaf development. We examined the leaves on Pico del
Oeste to see if palisade mesophyll development was reduced with the re-
240 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
duced light intensities we have reported there (Baynton 1968, 1969). Two
taxa, Lobelia portoricensis and Mikania pachyphylla, seemed to be without
a definite palisade layer in the leaves examined (PLATE Id). Further,
19 of the remaining 38 taxa possessed a palisade mesophyll of but a single
cell in thickness. The remaining 19 taxa had a palisade mesophyll in
part exceeding a single cell layer to 3 to 4 cells in thickness (TABLE 2, col-
umn 9). In taxa which could be measured, the palisade layer exceeded the
spongy layer in thickness in only 4 taxa, while the ratio of palisade to
spongy, in 34 taxa, ranged from 1:1 to 1:7.4 and averaged 1:2.4 (TABLE
2, column 10). Referring to Wylie’s study of New Zealand plants, of the
38 taxa he examined 7 had a thicker palisade layer than spongy layer
and the comparable ratios determined from the figures he gave show a
range from 1:1 to 1:3.3 with an average of 1:1.5.
On the basis of limited comparative data it appears that the leaves of
the plants growing in the elfin forest on the summit of Pico del Oeste are
thicker than usual and approach leaves of admittedly xeromorphic type.
The frequency of a hypodermal layer or multiple hypodermal layers is
high. The ratio of thickness of palisade and spongy mesophyll layers
suggests that the plants surviving on Pico del Oeste have adjusted to the
low light values through a reduction in the palisade mesophyll zone and
an increase in the amount of spongy mesophyll.
LEAF DEVELOPMENT
Richards has reviewed the earlier literature which claimed that a few
species in Buitenzorg were ever-growing and showed no foliar periodicity
whatever. On further study it was shown that one plant at least was
in continuous leaf production when young, but when older leaf produc-
tion was distinctly periodic. Richards (1964, p. 193) concluded that “it
is certainly true that most rain-forest trees produce new leaves, not con-
tinuously, but in periodic flushes, so that a single shoot bears several ‘gen-
erations’ of leaves at the same time.”
Our observations on the development of stems were in relation to the
production of leaves (Taste 3). Initially we observed that certain plants
did grow in obvious flushes where the young leaves were brightly colored
or soft in texture in comparison with the mature leaves. We recognized
20 taxa which grew in flushes, 8 of which had conspicuous terminal long
shoot-short shoot growth patterns. Twelve taxa were considered to be
in continuous production of leaves but this varied from branch to branch
on a given plant. A large specimen of Clusia grisebachiana, for example,
failed to produce a single new leaf during the period of this study.
marked plant of Trichilia pallida did not add a single leaf, or lose any,
for a period of three years after the plant was tagged for observation. //ex
sintenisii which appeared to have young green leaves all of the time proved
to have only some of the individual shoots on the plant in a stage of
growth or expansion at any given time. A shoot of J/ex tagged for observa-
tion was shown to produce a flush of leaves and then remain in a mature
1969 | HOWARD, ELFIN FOREST, 8 241
but quiescent stage before renewing its growth. Some of the shoots re-
newed growth with no change in the size of the leaves while others had
an initial renewal of growth in the production of leaves, or a single leaf
of smaller size or reduced to cataphyll proportions. When the internodes
along a stem were measured carefully there was evidence that growth
of the internodes had been reduced in some areas giving further evidence
to a periodicity of growth.
Clearly, it is difficult to determine that a given shoot has not added a
leaf, but the majority of plants observed in the elfin forest did exhibit
some degree of periodicity of growth and leaf production during the period
of study.
The suggestion has been made that leaves develop quickly in tropical
forests. Studies of leaf expansion within a temperate area at the Arnold
within a 10-day to three-week period in most native and cultivated species.
Although many species studied within the elfin forest did complete the
expansion of leaves within that period, there were notable exceptions.
Tagged shoots where fairly large leaves were counted and observed at
regular intervals showed the following times for development from a no-
ticeable leaf primordium to full expansion.
Symplocos micrantha 5 weeks
Clusia grisebachiana 7 weeks
Miconia pycnoneura 14 weeks
NUMBER AND PERSISTENCE OF LEAVES PER PLANT
Regular observations of the elfin forest components impressed upon
us the fact that some plants had many leaves and that others, equally
characteristically, had few leaves. Although the leaves may have been
produced in flushes of growth or seemingly continuously, there was a leaf
fall that in most plants seemed to equal leaf production. We selected 24
plants of comparable size and age of Miconia pachyphylla, Wallenia
yunquensis and Dilomilis montana, counted the leaves, and found less
than 5% variation in the number of leaves on a given plant of the species.
Branches of Miconia foveolata or Psychotria berteriana characteristically
had but 3 pairs of leaves at the end of a shoot. When new growt
curred the new shoot had a comparable number of leaves and the leaves
of the former growth generation abscissed. The largest number of leaves
on a mature plant was found on Micropholis garciniaefolia with 10,487,
while Brachionidium parvum characteristically had but 4 leaves per plant.
The following table indicates the plants with the greatest number of
leaves in comparison with the total photosynthetic area represented on the
plant, and the rank of the plant in frequency counts for transects reported
in the first paper of this series. Leaf numbers and total photosynthetic
area for all species is given in TABLE 1. The leaf count was obtained as a
242 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
by-product of gathering foliage material for a chemical survey of the
plants within the elfin forest.
TOTAL
TOTAL NUMBER OF LEAVES PHOTOSYNTHETIC AREA FREQUENCY *
Micropholis 10,487 Prestoea 289,460 cm.” Pilea krugi
Tlex 8,684 Micropholis 96,480 Wallenia
Calyptranthes 4,539 Tabebuia 60,040 Calycogonium
Tabebuia 2,680 Hedyosmum 44,908 Vriesea
Hedyosmum 2,339 Eugenia 23,000 Ocotea
Calycogonium 1,345 Lobelia 20,300 Calyptranthes
rdisia 1,857 Calyptranthes 18,609 Pilea obtusata
Gonocalyx 1,345 Psychotria 18,093 Dilomilis montana
berteriana
Haenianthus 1,294 Haenianthus 17,339 Miconia pachyphylla
Marcgravia (adult) 1,280 Ardisia 17,458 Tabebuia rigida
Cleyera 678 Ilex 15,631 Eugenia borinquensis
* Frequency = descending order of frequency in transects.
Holttum noted that in the uniform climate of Singapore, trees of a
number of deciduous species change leaves annually, many in February,
others in August, apparently because of leaf senescence.
Within the elfin forest of Pico del Oeste the greatest noticeable leaf fall
in the dominant plants of the forest occurred in February for Eugenia,
Ocotea, and Tabebuia and was conspicuous in March for Lobelia. In each
of these plants the leaf fall preceded the development of new year’s growth,
and the plants presented a barren appearance for a short period in con-
trast to their normal condition. Other species developed new growth before
the erratic abscission of the older leaves.
Richards notes the many widely different types of behavior among trop-
ical trees in regard to leaf fall and leaf persistence. According to Warming
and Graebner the average length of leaf life of tropical species is about
13-14 months. We made an attempt to mark branches and to record
the number of leaves, the nature of the new growth, and the length of
time individual leaves persisted (TaBLe 3). In general, the results were
unsatisfactory. Often the tagged branch failed to develop any new leaves
during the period of observation, while an adjacent branch of the same
plant for which data had not been recorded produced a flush of leaves, oF
flowered, or died. It is not possible to report with accuracy that the
growth flush in a long shoot-short shoot growth pattern represented an
annual increment of growth as may be done in temperate areas with
deciduous or bud-forming plants. We did observe that some leaves re-
mained on the plant during two full years of observation. Branches of
Ilex sintenisti which appeared to have two flushes of leaves per year re-
tained some leaves through 20 internodes, which represented 7 flushes as
determined by areas of short internodes and by cataphylls. Torralbasia
cuneifolia, which also grows in flushes with the production of many cata-
phylls, also retained leaves for 20 nodes representing 7 flushes in this
plant. Only one or two of the larger leaves persisted while cataphylls and
1969 | HOWARD, ELFIN FOREST, 8 243
smaller leaves abscissed. Gonocalyx produced 3 to 4 leaves per flush and
retained 20 leaves in 5 recognizable flushes with all leaves persisting. Tabe-
buia tended to retain only 1 pair of leaves of each flush of 2 pairs and the
oldest persisting leaves were 10 internodes below the apex, suggesting that
some leaves have persisted for five years.
In general, younger plants in the undergrowth tended to hold more
leaves per shoot for a longer period of time than did the plants with shoots
exposed in the canopy.
FACTORS OF PRODUCTIVITY OF THE LEAVES
Although it has been suggested that the persistent cloud cover, high
humidity saturated soil, and the growth form of individual plants all in-
fluence growth rate or development of the forest, we found additional
factors worthy of mention.
The very slow growth of some component trees within the elfin forest
has been recorded by Wadsworth and Bonnet in their comparative study
of the tabonuco (Dacryoides excelsa) rain forest and the colorado (Cyrilla)
forest in Puerto Rico. Although Cyrilla racemiflora was not encountered
in the elfin forest of Pico del Oeste, four other taxa of the colorado forest
were. No distinctive growth rings have been seen in the woody trunks
of the Pico del Oeste plants. Wadsworth and Bonnet grouped the trees
in diameter-size classes and estimated the age of the trees by summing the
period required for a plant to pass from one diameter class to another.
They concluded that a 4” trunk of Ocotea spathulata was 200 years old;
one of Micropholis garciniaefolia, 170 years old; and one of Calycogonium
squamulosum, 80 years old. The annual growth rate for saw timber and
polewood species in the Luquillo Mountains was 0.07 inches for Tabebuia
rigida, 0.05 inches for Micropholis garciniaefolia, and 0.04 inches for
Calycogonium squamulosum and Ocotea spathulata. They concluded that
the soil is the common factor most important in the forests they studied.
The saturated, poorly aerated organic soil inhibited root penetration and
the absorption of water and resulted in the very slow growth.
The cloud and fog cover, the high humidity and abundant rain docu-
mented by the studies of Baynton suggest that photosynthetic activity
in the elfln forest is low. We were unable to test the amount of photo-
synthesis carried on by the component species. Tests of evaporation with
potometers and of transpiration with cut branches within the forest were
complete failures. Gates, however, demonstrated by infra-red temperature
measurements that transpiration did occur during brief periods of sun-
shine and clear sky. As there were longer periods, even days of full sunshine
on the peak, the plants grew even though the growth rate was slow.
A survey was made of the stomatal types, size, and distribution to
determine any specializations that might occur within the elfin forest
components (TaBLEs 3, 4). Sinnott suggested that xerophytes tend to
ave a high stomatal frequency but cites no reference. Regrettably, we
have failed to find any comparative data for other forest zones.
244 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Although stomatal apparatus types are commonly associated at the
family level, there are variations and exceptions as reported throughout
the work of Metcalfe and Chalk. We found the anomocytic type of
stomatal apparatus (PLATE IIIa) to be most common as represented in
16 taxa of dicotyledons and 4 taxa of monocotyledons. The paracytic
type (PLATE IVa) was present in 13 taxa of dicotyledons and 3 monocoty-
ledons. Anisocytic type (PLATE IIIc) was present in 9 taxa of dicotyledons.
The gramineous type (PLATE IVd) was present in all 6 taxa of Cyperaceae
and Gramineae. A didymocytic stomatal apparatus (PLATE IIIb) was
represented in 1 taxon of monocotyledons and 1 of dicotyledons (TABLE
3, column 9; TABLE 4, column 3).
Stomatal openings of varying sizes were found in Justicia martin-
soniana where large numbers of the stomatal apparatus appeared to abort
before the final cell division which, we suspect, would have formed the
guard cells. The openings, therefore, were of varying sizes. Pilea krugit
(Pirate Va) also had stomatal apparatus of varying size with very small
guard cells approximately 0.002 mm. long appearing over the veins, while
mesophyll tissue was surmounted by guard cells averaging 0.023 mm. in
length. Marcgravia sintenisii with heteroblastic growth showed the same
number of stomata per square mm. for juvenile and adult foliage, but the
guard cells were 0.037 mm. long on the juvenile leaves and only 0.028 mm.
long on the adult leaves. The stomatal apparatus, however, appeared to
be broader in the adult leaves.
Stomata occurred in definite patterns in many of the monocotyledons,
as expected, but they were found in groups of 2 to 3 or 3 to 7 in Gram-
madenia (PLATE Vb) and in groups of 6 in Gesneria sintenisii and with
2 to 3 very closely associated, almost united, in Sauvagesia.
On leaves of Pilea yunquensis stomata were found only on the upper
surface of the leaf. Metcalfe and Chalk refer to work of Mohler, who found
stomata on the lower surface in the species of Pilea he examined except
for Pilea spruceana, where they were on the upper surface. Stomata
tended to be oriented around the long hairs on Cleyera.
Accessory or subsidiary cells to the guard cells were usually clearly
defined and commonly contrasted with those of the epidermis. Unusually
shaped subsidiary cells appeared in Alloplectus (Pitate IIIc) and in
Justicia martinsoniana and in Renealmia antillarum (PLATE I1Id).
he subsidiary cells had a characteristic homogeneous yellow-brown
pigmentation in Clusia in contrast to the adjacent epidermal cells. In
Tabebmuia rigida the subsidiary cells were generally clear in contrast to
the mottled appearance of the epidermal cells. The walls of the subsidiary
cells of Micropholis were straight in conspicuous contrast with the sin-
uous walls of the other epidermal cells.
No conspicuous elevation of stomatal apparatus was discerned in the
components of the elfin forest. Torralbasia was the only taxon with the
guard cells noticeably sunken and overlain by 6 epidermal cells (PLATE
er.
The length of the guard cells was measured and those found in 14 taxa
“Y
1969 | HOWARD, ELFIN FOREST, 8 245
of monocotyledons averaged 0.035 mm. in length while in 39 taxa of
dicotyledons the guard cells averaged 0.028 mm. in length. Within the
monocotyledons the largest guard cells were in Eleocharis, measuring 0.048
mm. long, while /sachne had the smallest, 0.023 mm. long. Within the
dicotyledons the longest guard cells were found in Hedyosmum arbores-
cens and Peperomia emarginella, each 0.048 mm. while the smallest
were those of Miconia pycnoneura, 0.010 mm. in len
In considering the length of the guard cells in ise to the habit of
the plant we found the following lengths:
8 taxa of herbaceous plants average 0.031
2 taxa of woody epiphytes average 0.030
24 taxa of trees or shrubs average 0.028
5 taxa of woody climbers average 0.027
The number of stomatal openings ranged from 18 per square mm. in
Guzmania berteroniana to 230 per square mm. in /sachne angustifolia.
Within the dicotyledons Peperomia emarginella had 22 stomata per square
mm. while Miconia pycnoneura had 2230. While Guzmania had only 18
stomata per square mm., there were 96 stellate hair clusters in the same
area.
Vriesea sintenisii, another bromeliad, was examined in several sections
of the leaf. The upper portion of a mature leaf showed 11.4 stomatal ap-
paratus per square mm. with 41.4 stellate hair-glands in the same area;
the middle portion of the leaf had 28 stomata per mm.” and 19 stellate
glands, while a basal portion above the water level showed 49.4 stomata
per mm.” and 19 glands in the same area.
It has been suggested that metabolic activity of individual plants or
leaves might be impaired by the presence of epiphyllous algae and leafy
hepatics. The young leaves of most species within the forest are a bright
green color when they first develop. In other species the young leaves
were colored when young or expanding and developed a green color when
near maturity. Young leaves of Rajania and Ipomoea, herbaceous vines,
were bronze in color when young. Young leaves of the woody climbers
Gonocalyx and Hornemannia were pink to red or orange-red in color. Cleye-
ra and Symplocos also produced young leaves bronze in color, while Caly-
cogonium had the young leaves reddish. The herbaceous Peperomia hernan-
diifolia had reddish young leaves. Miconia pachyphylla was unique in
losing the green pigments and having the leaves turn a bright red or
Orange immediately before falling.
Upon reaching mature size, the leaves of most species in the Pico del
Oeste elfin woodland acquired a covering of epiphyllous non-vascular
plants. Spores, gemmae, gemmalings and sporelings, or fragments of liver-
worts are wind-borne and settle on the new leaves of most species. They
appeared most quickly on species with depressed midrib or veins such as
Marcgravia, Ilex, Symplocos, Gonocalyx or Tabebuia, and were rarely seen
on the pubescent leaves of Miconia foveolata or the rugose leaves of M7-
conia pycnoneura. Dr. Margaret Fulford, in work to be reported later,
246 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
examined a collection of leaves from plants within the forest and in 94
collection numbers found approximately 680 specimens belong to 40
genera and more than 75 species. There appeared to be an average of
eight species per sample with a maximum of 18 species. Preliminary data
showed the following epiphyllous species distribution on representative
leaves:
Anthurium dominicense
Ardisia luquillensis
Calyptranthes krugii
Eugenia borinquensis
Gonocalyx portoricensis
Grammadenia sintenisii
Ilex sintenisit
Micropholis garciniaefolia
Miconia pachyphylla
Miconia pycnoneura
Ocotea spathulata
Peperomia emarginella
Symplocos micrantha
Trichilia pallida
Wallenia yunquensis
FOr RK DO UI
——
COOnmarThy Kw OW +
_
The number of epiphyllous taxa on any given leaf is not indicative of
the leaf size or the percentage of the surface covered. The speed with
which epiphyllae grew and covered the surface of the host was startling.
Leaves of Eugenia borinquensis were completely and densely covered with
liverworts in less than five months after leaf expansion. The amount of
light reaching the photosynthetic area of the leaf is certainly reduced
by the abundant epiphyllous growth. Epiphyllous growth occurred on
leaves exposed at the summit of the canopy although the number of seem-
ingly dead or desiccated plants was high. The leaves of the lower and
inner branches of the forest components were more densely covered.
Epiphyllae were less common on leaves of the truly herbaceous species.
LEAF DAMAGE
An additional factor in reducing the potential metabolic production of
the plants in the elfin forest is evident in the amount of damage to the
foliage of individual species. This has generally been attributed to wind.
Damage done by animals as found in the Pico del Oeste forest has not
been recorded.
The sheared effect and directional growth of woody plants along sea
coasts have been attributed to wind and to salt spray. Beard and Gleason
and Cook have suggested the same factors are important in the shaping
of the mountain-top forests in the West Indies. The effects of wind were
observed in the canopy of Pico del Oeste. Within a few feet of the roof
of our observation tower a slender stem of Eugenia borinquensis had worn
a circular opening in the canopy of surrounding species. Branches of
Tabebuia rigida were worn smooth through the cambium to the xylem
by friction against each other due to movement in the wind. The leaves
of Prestoea montana were broken and lacerated when they exceeded the
shelter of the lee forests. The soft leaves of Psychotria berteriana were
severely lacerated on a few plants growing in open areas. The soft flush
growth of Hornemannia, Gonocalyx and Marcgravia was broken and
leaves torn when the leading branches were whipped about in gusting
winds.
nice, _einenieenaeemmeneaninntin
1969 | HOWARD, ELFIN FOREST, 8 247
Microscope slides which were exposed to collect wind-blown particles
also revealed crystals of salt. We failed to find any quantities of salt
crystals on leaves or any indications of leaf damage due to salt spray
from ocean storms. Apparently the large amounts of rain water or pre-
cipitation from the clouds washed the leaves free of salt.
The succulent young leaves were severely damaged by the populations
of insects which existed on Pico del Oeste. In column 6 of TABLE 1 is
recorded the percentage of leaves of each species that was affected by
animal damage.
The program of collecting foliage material for drying and future chem-
ical tests permitted an assessment of damage to leaves on representative
plants. Among the monocotyledons, animal damage was relatively light
and only plants of Rajania cordata and Brachionidium parvum appeared
to be severely affected. Many dicotyledonous species, however, were eaten
with great regularity. In one plant of Clusia grisebachiana selected for
study 98% of all leaves had been partially eaten and the reduction in leaf
surface was 24%. These figures were computed by an actual count of
the leaves which had been eaten by insects, and the degree of surface
loss computed by reconstructing the outline where possible, and measur-
ing the area lost by planimeter. Eighty percent of all leaves on repre-
sentative plants of Haenianthus salicifolius, Wallenia yunquensis, Eu-
genia borinquensis and Miconia foveolata were similarly eaten. Miconia
showed a reduction in leaf surface of 35% and Eugenia borinquensis ex-
hibited loss of 25%. The following table shows the plants most susceptible
to insect damage.
PERCENT PERCENT REDUCTION pH PERCENT
DAMAGED IN SURFACE AREA WATE
Clusia grisebachiana 98 24 3.9-4.3 66
Haenianthus salicifolia 86 19 5.1-5.2 62
Wallenia yunquensis 84 16 3.5-4.2 7S
Eugenia borinquensis 81 25 4.8 30
Miconia foveolata 35 3.9-4.0 71
Miconia pachyphylla 79 21 3.7-4.3 60
Grammadenia sintenisii 77 _— 4.1-4.5 84
aan = racemosa 73 — 3.9-5.3 67
Rajania cordata 70 — 49 =
Peberomin hernandiifolia 70 oa 4.6-5.1 —_
The nature of the leaf damage varied. In most cases the insect began
on the margin and ate for varying distances towards the midrib. In other
cases the apex of the leaf was chosen as the point of initial attack. Mi-
conia pachyphylla and Miconia foveolata, with the characteristic reticu-
late network of veins of the family, were characterized by holes in the
leaves. No evidence was found of insect or animal attacks on the petioles
or pulvini. Insect attacks seemed to be present at all months of the year.
Damage to unexpanded primordia was rarely seen. Young leaves of a flush
might be consumed completely as quickly as they began to expand in
po Cn and Eugenia. Other leaves were attacked only as the
248 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
lamina developed. The majority of the damaged leaves persisted on the
plant following the insect attack. The leaves subsequently developed cal-
lous tissue or what appeared to be cork in many instances along the mar-
gin of laminar tissues that had been eaten.
Regrettably, we have been unable, up to this stage, to obtain scientific
names or determinations for the insects seen or collected during this
study. The following tabulation suggests that the food habits of the
insects were often specific:
Gray caterpillar which stings: Cleyera, Grammadenia, Miconia pycnoneura,
Gray caterpillar with tufts of orange hairs and longer white hairs: JJex, Clusia.
Slender green walking stick: Eugenia, Marcgravia.
Tan-colored stouter walking stick: Miconia pycnoneura.
Large spiny walking stick: Ardisia, Calycogonium, Micropholis, Wallenia.
A weevil: Calycogonium.
Spittle bugs: Eugenia.
Green grasshopper with white lines: Tabebuia.
Black grasshopper: Tabebuia.
Leaf hopper: Cyathea.
Leaf miners: Hornemannia.
A slug (Gaeotis nigrolineata Shuttleworth): Lobelia.
A snail (Luquillia luquillensis Shuttleworth): Lobelia.
Gall-producing insects: Ocotea.
No domatia were encountered in leaves of species within the elfin for-
est, although domatia were found in other species of plants in forests of
lower elevations.
The nature of the attraction in the leaves of the component species
to the insects cannot be determined. It was evident that the leaves varied
in their texture, the amount and the color of the liquid within the tissues,
their aromatic constituents and the pH of the cell contents. Clusia (Gut-
tiferae), Micropholis (Sapotaceae), and Lobelia (Campanulaceae) pos-
sessed a latex, as is characteristic for the families involved. Ninety-eight
percent of the leaves of Clusia, 33% of the leaves of Micropholis and 11%
of the leaves of Lobelia were damaged by insects or snails. [pomoea reé-
panda also has cells containing a yellow material, although this did not
flow when the leaf tissue was broken, and 61% of the leaves of this
plant were damaged by insects. Hedyosmum arborescens, Calycogonium
squamulosum, Miconia foveolata, and Symplocos micrantha could be clas-
sified as “bleeders,” for the leaves, petioles or stems exuded a clear liquid
when cut or broken. The percentage of leaves damaged by insects in these
taxa were: Hedyosmum, 61%; Calycogonium, 21%; Miconia foveolata,
81%; and Symplocos, 14%. Aromatic principles were present in the
leaves or bark of some species and can be described as follows: Calycogo-
nium — cider odor; Llex—odor of hay; Mecranium — sweet; Miconta
foveolata — rank; Miconia pachyphylla — sweet; Miconia pycnoneura —
sweet; Symplocos — rank and acidic; Tabebuia — medicinal; and Wal-
lenia — odor of spinach. The nature of insect damage in these taxa 1S
reported in TABLE 1, column 6.
4
1969 | HOWARD, ELFIN FOREST, 8 249
In the preparation of several pounds of dried material of leaves or
branches for shipping and subsequent chemical analysis, it was evident
that the species of the elfin forest contained different amounts of liquid
and solid materials. Large quantities of certain plants would produce
only a few pounds of dry weight material while other species clearly had
less liquid to evaporate. Standard weight samples of leaves were obtained
and dried in an oven to obtain the percentage of water in the leaves of
each species. For mature leaves the percentage of water ranged from 93%
in Psychotria guadalupensis and Begonia decandra to 44% for Calyp-
tranthes krugii. Only 1% of the leaves of Psychotria guadalupensis were
damaged by insects, 26% of the leaves of Begonia and 41% of the leaves
of Calyptranthes. In the list of 10 most severely damaged species pre-
viously given, the percentage of water in leaf tissue varied from 60% to
84% of the taxa for which we have data. Clearly some factor other
than the amount of liquid in the plant tissue was responsible for the in-
sect damage.
The abundance of liquid in some leaves led us to a simple measurement
of the pH of the plant liquid which could be extracted (TABLE 1, column
7). Further details on these tests will be given in a later paper. For each
species, leaves of a size normally eaten by insects were crushed between
clean microscope slides and several drops of liquid were tested immediately
with a Beckman pH meter. The acidity varied from 2.4 in Begonia de-
candra to 6.5 in Justicia martinsoniana. The values of the plant sap in
the 10 most commonly eaten species ranged from 3.5 in Wallenia yun-
quensis to 5.3 in Hornemannia racemosa but averaged 4.4.
LITERATURE CITED
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Table 1
CoLtumNn 1— Total number of leaves per plant. Cotumn 2— Total photosynthetic area in cm.” CoLtuMN 3—average blade are
cm.” Corumn 4— Ratio, blade length:width. Corumn 5— Ratio, blade length: Laden length. Corumn 6 — Percent of gre prover
by animals. Cotumn 7 —pH of leaf sap. Cotumn 8 — Percent of water in leaf tiss
COLUMN NUMBER 1 2 3 4 5 6 7s 8
GRAMINEAE
Arthrostylidium sarmentosum 168 523 12 ipa sessile ms — —
Ichnanthus pallens 12 97 8.1 re | sessile 0 — —
Tsachne angustifolium 18 144 8.0 10:1 sessile 0 = —
CYPERACE
Carex nae chya 27 2240 11.9 38:1 sessile 0 — —
Eleocharis yunquensis 54 609 — terete sessile 0 — —
Scleria secans 14 3101 62.0 70:1 sessile 0 — —
PALMAE
Prestoea montana 10 289,460 28,946.0 — -— 0 5.1-6.2 —
ARACEAE
Anthurium dominicense 10 918 91.8 6.8:1 Ast 0 5.0-5.8 _
BROMELIACEAE
Guzmania berteroniana 20 4,392 225.0 — sessile 0 —
Vriesea sintenisii 19 1,581 80.0 Svial sessile 0 oct 83
DI0SCOREACEAE
Rajania cordata 20 370 18.0 2 S01 Lt 70 4.9 —
ZINGIBERACEAE
Renealmia antillarum 8 736 92.0 S071 constr. 0 4.8-5.3 —
8 ‘LSAYOA NIATA ‘GUVMOH [6961
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COLUMN NUMBER 1 2 4) 4 5 6 | 8
SAPOTACEAE
Micropholis garciniaefolia 10,487 96,480 9.1 1,6:1 8.0:1 33 4.1-4.8 52
SYMPLOCACEAE
Symplocos micrantha 32 268 8.3 233i 20.0: 1 14 4.0-4.8 60
OLEACEAE
Haenianthus salicifolius 1,294 17,339 13.4 V foal a 8.0:1 86 5.1-5.2 62
CONVOLVULACEAE
Ipomoea repanda 18 205 113 25 5.031 61 4.5-6.2 —_
BIGNONIACEAE
Tabebuia rigida 2,680 60,040 22.4 19:1 8.3:1 60 5.0-5.5 ip
GESNERIACEAE
Alloplectus ambiguus 16 168 10.5 22°) 9.5:1 43 4.6-5.5 _
Gesneria sintenisii 256 9,022 35:2 2321 8.9:1 67 5.3-5.7 84
ACANTHACEAE
Justicia martinsoniana 9 47 52 2521 11.4:1 0 4.1-6.5 —
RUBIACEAE
Hillia parasitica 48 564 SB vy ie | 5.6:1 29 4.7-4.9 83
Psychotria berteriana 305 18,093 59.3 20:1 4.5:1 62 5.0-5.7 85
Psychotria guadalupensis 213 410 1.9 2.0:1 1.4:1 1 4.9-5.0 93
CAMPANULACEAE
Lobelia portoricensis 416 20,300 48.7 3.4:1 4.5:1 11 4.7-5.0 79
COMPOSITAE
Mikania pachyphylla 46 274 5.9 1.4:1 4.5:1 10 5.1-6.0 —-
SZ
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Table 2
Cotumn 1—Leaf thickness in ». CoLumMn ead a oe . x, absent 0). Corumn 3 — Lower hypodermis. oo 4—
Ratio, upper epidermis:upper hypodermis. sear 5 — Upp e. CoLuMN 6 —Lower cuticle. Cotumn 7—Crystals (ra
raphides, rh rhombic, d = druses, f = vas CoLu bee ee? asts or sclereids present. CoLtumNn 9 — Multiple edi jayer.
CoLUMN 10 — Ratio of palisade layer to spongy mesophyll. Isodiam. = isodiametric cells only
COLUMN NUMBER 1 2 3 4 5 6 7 8 9 10
PIPERACEAE
Peperomia emarginella 275 Ps 0 1:10 0 0 d,ra 0 0 4.
Peperomia hernandiifolia 625 x 0 1:3.0 0 0 d 0 0 isodiam.
CHLORANTHACEAE
Hedyosmum arborescens 311 2% 0 i332 0 0 0 0 0 ey.
MoRACEAE
Cecropia peltata 91 0 0 —_ 0 0 d 0 0 1:0.39
URTICACEAE
Pilea krugit 146 0 0) —_ 0 0 f re) 0 jal
Pilea yunquensis 146 0 0 _ 0 0 f 0 O 131
LAURACEAE
Ocotea spathulata 475 2x 0 Mia x x 0 0 x RE oe:
MELIACEAE
Trichilia pallida 209 0 0 _ x Ps rh 0 0 1322
AQUIFOLIACEAE
lex sintenisi 421 0 0 —_ x x d 0 x 1:1.8
CELASTRACEAE
Torralbasia cuneifolia 458 0 0 — X = d 0 x 1:4.5
OCHNACEAE
Sauvagesia erecta 141 X 0 io .; 0 0 d 0 0 is
[6961
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COLUMN NUMBER 1 2 3 4 5 6 7 8 9
SAPOTACEAE :
Micropholis garciniaefolia i A A DB 2-6 20-4 027 230 Anomocytic
SYMPLOCACEAE
Symplocos micrantha S A A — 2-6 - .030 230 Paracytic
OLEACEAE
Haenianthus salicifolius sg DS — Se - 025 327 Anomocytic
CONVOLVULACEAE
Ipomoea repanda wc A A,T DB 9- 032 110 Paracytic
BIGNONIACEAE
Tabebuia rigida T s D DB 1-2 pr 10-4 .023 230 Anomocytic
GESNERIACEAE
Alloplectus ambiguus H A M DB i 22- 032 56 Anisocytic
Gesneria sintenisii S A M DB c 20- 035 168 Anisocytic
ACANTHACEAE
Justicia martinsoniana H si D — — - 025 170 Paracytic
RUBIACEAE
Hillia parasitica S,E T D — a - 037 100 Paracytic
Psychotria berteriana S 9 D — 2 pr 3- 029 210 Paracytic
Psychotria guadalupensis S,E 4 YD — 8- .025 110 Paracytic
CAMPANULACEA
Lobelia portoricensis SH T A DB — 37- .040 190 Anomocytic
COMPOSITAE
Mikania pachyphylla & A,T M — —_ - 035 196 Anomocytic
~S a “=
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1969 | HOWARD, ELFIN FOREST, 8 261
Table 4
STOMATAL STOMATA TYPE
TAXON SIZE IN MM. NUMBER OF STOMATAL
MM * APPARATUS
GRAMINEAE
Arthrostylidium sarmentosum 0.027 205 Gramineous
Ichnanthus pallens 0.043 140 Gramineous
Isachne angustifolium 0.023 230 Gramineous
CYPERACEAE i
Carex polystachya 0.041 102 Gramineous
Eleocharis yunquensis 0.048 120 Gramineous
Scleria secans 0.030 110 Gramineous
PALMAE ;
Prestoea montana 0.025 140 Paracytic
ARACEAE ;
Anthurium dominicense 0.046 56 Paracytic
BROMELIACEAE J ;
Guzmania berteroniana 0.044 18 Didymocytic
Vriesea sintenisii 0.039 28 Didymocytic
DIOSCOREACEAE -
Rajania cordata 0.035 120 Anomocytic
ZINGIBERACEAE ‘
Renealmia antillarum 0.027 140 Paracytic
ORCHIDACEAE , ; .
Brachionidium parvum 0.039 47.5 Didymocyt “i
Dilomilis montana 0.032 120 Anomocytic
ARNOLD ARBORETUM
HARVARD UNIVERSITY
262 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
EXPLANATION OF PLATES
PLATE I
a. Terminal bud of an adult branch of Marcgravia sintenisii. b. Stem of Hillia
parasitica showing the flattened stipular sheath. c. Stem apex of /lex sintenisu
i a eria
and enclose and protect the terminal bud. g. Three views of the stem apex an
terminal leaf pair of Calyptranthes krugii, left figure shows the mature leaves ;
central figure shows the plicate stipule pair separated at the base; right figure
shows the plicate stipule pair separated at the apex.
PLATE II
PLATE III
Epidermal cells and stomatal apparatus of: a, Miconia pachyphylla; b,
Brachionidium parvum; c, Alloplectus ambiguus; d, Renealmia antillarum.
P PLATE IV
Epidermal cells and stomatal apparatus of: a, Anthurium dominicense ; be
Hillia asitica; c, Torralbasia cuneifolia; d, Scleria secans; e, Haenianthus
salicifolius var. obovatus; £, Gonocalyx portoricensis.
PLATE V
a. Lower epidermal surface of Pilea krugii showing the cluster of small sto-
mata over a vascular bundle, a multicellular gland, and two of the larger stomatal
apparatus. b. The lower epidermal surface of a leaf of Grammadenia sintenisu
showing the stomatal apparatus grouped in clusters of three.
Jour. ARNOLD ARB. VoL. 50 PiaTeE I
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268 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
LECTOTYPIFICATION OF CACALIA L.
(COMPOSITAE-SENECIONEAE) +
BERYL S. VUILLEUMIER AND C. E. Woop, Jr.
Tue INTERNATIONAL Cope of Botanical Nomenclature (1966) provides,
by means of nomenclatural types, stability in the application of names at,
or below, the rank of family. The use of a name is determined by the no-
menclatural type of that name, and changes may result when, or if, the
choice of a type is shown to be incorrect. To avoid disadvantageous
changes in the application of names provision is also made in the Code
to preserve current usage and to avoid the confusion which could result
from varying opinions concerning the choice of a lectotype. The sene-
cionid genus Cacalia, as circumscribed by Linnaeus (1753, 1754) was
quite heterogeneous, and the process of choosing a lectotype has been
both complicated and subject to individual interpretation. Three dif-
ferent species have been proposed as lectotype, and even now there are
conflicting opinions (Cuatrecasas, 1960, and Pippen, 1968, vs. Pojarkova,
1961, and Vuilleumier, 1969). It seems expedient to review the typifica-
tion of this genus once more. Cacalia as typified here (by C. hastata L., a
choice made by Kitamura, 1942) provides an example in which the choice
of a lectotype in accordance with the guides outlined in the Code also
complies with the recommendation (7B) that a lectotype be so chosen
as to preserve current usage.
In the first edition of Species Plantarum (2: 834-836. 1753), Lin-
naeus described and named ten species of Cacalia which he divided into
two groups: ‘‘Frutescentes,”’ consisting of four shrubby species, and
‘“Herbaceae,” with six herbaceous species. In 1754, Miller (Gard. Dict.
Abr. ed. 4. 2: ord. alph.) split Cacalia, placing the four shrubby species
in Kleinia Mill. Cacalia was thus restricted to the herbaceous species,
and it is from the six original species of this group that all choices of
lectotype have been made. To our knowledge, no one has suggested that
Cacalia be typified by one of the species removed to Kleinia.
There has been, however, considerable disagreement as to which of
the six herbaceous species should be designated as lectotype. Rydberg
(1924) concluded that Cacalia alpina should be the type species; Cuatre-
casas (1955, 1960) and Pippen (1968) have concurred in this choice.
e of an informal series of peripheral papers arising from research toward a
Generic Flora of the Southeastern United States being carried on through the gen-
erous help of the National Science Foundation (Grant GB-6459X, C. E. Wood, Jr.,
principal investigator).
We acknowledge with thanks the assistance of Dr. Bernice G. Schubert and Dr.
Elizabeth Shaw and we are indebted to Andrey I. Baranov for his translation of a
portion of A. Pojarkova’s treatment of Cacalia in Flora URSS.
1969 | VUILLEUMIER & WOOD, CACALIA 269
Hitchcock and Green (1927) chose another type, C. atriplicifolia, ene
Pojarkova (1960) also adopted. Kitamura (1942), however, ma
third choice, C. hastata, and Pojarkova (1961), changing her pact
agreed with him. Quite independently, Shinners (1950) came to the
conclusion that either C. suaveolens or C. hastata should be the lectotype
species. After a careful review of the arguments and of the nomen-
clatural and taxonomic history of Cacalia in conjunction with the appli-
cation of the Rules and Recommendations of the Code, we are convinced
that C. hastata should be the lectotype species.
Since the nomenclature of species of vascular plants begins in 1753 and
that of genera in 1754, in most instances the application of a name prior
to 1753 should be given little weight relative to a post-1753 application,
especially since Linnaeus not infrequently reapplied older names in a
completely different sense. However, one of the principal arguments ad-
vanced, first by Rydberg, and then by Cuatrecasas and Pippen for the
selection of Cacalia alpina as the type species of Cacalia is an historic one.
Rydberg wrote (loc. cit., p. 370), “Of the species of the second [herba-
ceous| group only the last two, Cacalia atriplicifolia and C. alpina, had
been known as Cacalia before Linnaeus’ time.2 The oie Cacalia, applied
to the last one, dates back to Vaillant and L’Obel. C. alpina L. or Ade-
nostylis alpinus is therefore the historical type of Cacalia.” Cuatrecasas
(1960, p. 182) reiterated, “There is no doubt that the name Cacalia was
first applied to C. alpina and that Linné had this species in mind when
he established the genus in his Genera Plantarum. Therefore, Cacalia
alpina is the type of Cacalia.’ Pippen added (1968, p. 377), “It seems
clear that C. alpina L. is the most logical lectotype of Cacalia. This
species, named C. alpina by Linnaeus (1753), embodied the Linnaean
and pre-Linnaean concept of Cacalia in that essentially all of the species
of Cacalia described by pre-Linnaean botanists (L’Obel, 1581; Clusius,
1601; Bauhin, 1623; —— 1699; Tournefort, 1700) actually repre-
sented the same species
The adoption of this ‘historical argument would restrict the name Ca-
calia to a genus consisting of four or five species of Europe which has
been known since 1816 as Adenostyles Cassini. Contrary to all three
authors, however, the use of arguments concerning the application of names
before the starting point for botanical nomenclature can result only in
further confusion, as is shown below
In the first edition of Genera Plantarum (1737, p. 252), Cacalia in the
sense of Tournefort (that is, C. alpina L.) is found in the synonymy of
Tussilago: “TUSSILAGO*. Tournef. 276. Vaill. A. G. 1720. f. 46. Cacalia
*In the next paragraph Rydberg added, “. . . only Linnaeus himself had used
Cacalia for C. suaveolens in his Hortus Upsaliensis.” Rydberg probably was misled
by Linngeus’s mace viet to Hortus Upsaliensis immediately following the diagnostic
phrase name cif ame) 0 itp suaveolens in Species omaha (1753, p.
835). In Hortus UgeaBions eaten p. - a byeevgy this a species of
Kleinia (“Kleinia caule Shee ceo,” etc. ); the diagnos the same in sage ies Plan-
tarum, except that Cacalia has been Sou wea for Kleinia, We have not found any
mention of Cacalia in Hortus Upsaliensis.
270 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Tournef. 258. Petasites Tournef. 258. Vaill. A. G. 1719.” Following the
description the observation is added: “‘Cacalia T. caule ramoso est, &
corollulis hermaphroditis quadrifidis, sine radio ligulato.” On the same
page is found Kleinia: “KLEINIA. Cacalianthemum Dill. elth. 54. 55.
An Tithymaloides ? Klein. monagr.”” The corolla is described as with the
limb “quinquefido, erecto,” and the stigmas as “duo, oblonga, revoluta.”
In the second edition (1742, p. 401), the synonymy of Tussilago is am-
plified by the addition of “Vaill. A. G. 1719.” to the reference to Cacalia,
and to that of Kleinia (p. 394) is added “Porophyllum Vaill. A. G. 1719.
t. 20. f. 39.” which Linnaeus had maintained as distinct in the Hortus
Cliffortianus (1738, p. 494). Neither description was changed in any
way in this edition. The synonymy and descriptions of Tussilago and
Kleinia in the editions of 1743 and 1752 are identical with those of the
second, but these editions were not prepared by Linnaeus.
In the fifth and crucial edition (1754, p. 362), Linnaeus treated Cacalia
of Vaillant and Tournefort as congeneric with Kleinia and combined the
two under the name Cacalia: “CACALIA.* Vaill. A. G. 1719. Tournef.
258. Kleinia edit. prior. Cacalianthemum Dill. elth. 54. 55. An Tithyma-
loides ? Klein. monagr. Porophyllum Vaill, A. G. 1719. #. 20. f. 39.”
Most interestingly, the generic description is identical with that of
Kleinia of the first four editions of Genera Plantarum! There is no
mention of the tetramerous corolla of Cacalia alpina which had appeared
in earlier editions under Tussilago.
Rydberg argued (/oc. cit.) that “Linnaeus’ description of the genus
[Cacalia] points to this species [C. alpina] especially the description of
the style tips: ‘Stigmata duo, oblonga, revoluta.’ This is characteristic
of Adenostylis alpinus which on account of its oblong style branches had
been placed in the tribe Eupatorrear, but which Dr. B. L. Robinson
rightly restored to the SENECcIONEAE. C. atriplicifolia as well as C.
suaveolens has a true Senecioid style, with truncate style-branches.”
Cuatrecasas further argued, “Among all the species of Cacalia in Linné’s
Species Plantarum, Cacalia alpina is the only one with elongate, curled
stigmas and 4-merous corollas.” Cuatrecasas is correct, but Linnaeus’s
description (1754) of the corollas of Cacalia as five-fid and the stigmas
as “‘duo, oblonga, revoluta” does not apply to C. alpina but to the species
which he had formerly placed in Kleinia, a name which he abandoned in
favor of Cacalia in 1753. Moreover, the description of the corolla and
stigmas is precisely the same as in Senecio (Gen. Pl. ed. 5. 373).
A note under Cacalia alpina in Species Plantarum (1753, p. 836) reads:
“Hanc speciem genere cum antecedentibus convenire docuit autopsia, hinc
genere conjugenda: Cacalia cum Kleiniis.” In the second edition (1763,
p. 1171) this note is clearer and is amplified: “Hanc speciem genere cum
antecedentibus convenire docuit autopsia, hinc genere conjugendae Ca-
caliae cum Kleiniis. Calyx hujus speciei flosculis 3 s. 4 tantum.” We
translate this to read: ““My observation has shown this species to agree
generically with the preceding ones; hence the Cacalias are to be joined
in a genus with the Kleinias. The involucre of this species with only 3
1969 | VUILLEUMIER & WOOD, CACALIA 271
or 4 florets.” From this comment and the placement of the species last
in the genus it appears that Linnaeus regarded C. alpina as somewhat aber-
rant in, but belonging to, the genus which he had formerly called Kleinia.
The use of the plural of Cacalia also suggests that he had in mind more
than one species.
Linnaeus did not change the description of Cacalia in the sixth edition
of the Genera, but the authors of the seventh and eighth editions noted
the departures of C. alpina from the others of the genus. Reichard (Gen.
Pl. ed. 7. 411. 1778) observed, “C. alpina foliolis calycis conglutinatis
corollulisque quadrifidis differt.”’ Schreber (Gen. Pl. ed. 8. 545. 1791) in-
cluded in the description of the corolla “limbo quadri-f. quinquefido,
erecto,’ and Haenke in his edition (Gen. Pl. ed. 8. 709. 1791) had pre-
cisely the same description and observation as Reichard. Cuatrecasas
(1960, p. 182) attributes a comment to Schreber (1791, p. 545) which
we have been unable to locate in the copy available to us: “ ‘Cacalia dif-
fert a Senecione flosculis quadrifariam scissis.’ ”’
To return to 1753, Cacalia as set forth by Linnaeus in the works which
are the starting points for botanical nomenclature has a protologue which
is that of Kleinia of the first four editions of the Genera Plantarum, with
the exception of the pre-Linnaean Vaillant and Tournefort references to
Cacalia, both of which apply to C. alpina. Cacalia alpina does not agree
with the generic description in either corolla or stigmas, but a number
of the other species do, among them species currently assigned to Cacalia
and those removed by Miller to Kleinia. It appears that this is but an-
other example of Linnaeus’s changing the name of a genus (Kleinia) to
one which he liked better (Cacalia), even though the species which had
borne that name historically was somewhat aberrant within an already
heterogeneous genus. In the interests of nomenclatural stability, it seems
to us most unwise and unwarranted to do anything but to begin the no-
menclatural and taxonomic history of Cacalia at 1753 and to proceed
from that year in the choice of a type species. The species chosen should
be in agreement with the protologue and must be one of the ten described
in Species Plantarum in 1753, taking into consideration those which have
been removed to other genera. In reaching the conclusion that Cacalia
hastata must be the lectotype species, our reasoning follows essentially
the same arguments as those succinctly presented by Shinners (1950).
All of the ten original species have been transferred to one or more
other genera at one time or another. The chronological sequence of the
more important of these transfers follows:
C. papillaris, C. anteuphorbium, C. kleinia, and C. ficoides: Segregated as
the genus Kleinia by P. Miller (Gard. Dict. Abr. ed. 4. 2: ord. alph. 1754),
although the combinations under that genus were not made until much later
by Haworth (1812) and De Candolle (1838). f
C. alpina: Transferred to Tussilago L. by Scopoli (Fl. Carniol. ed. 2. 2: 156.
1772) as T. Cacalia Scop. (not T. alpina L.). Placed in a new genus,
Adenostyles, by Cassini (Dict. Sci. Nat. Paris 1(Suppl.): 59. 1816).
C. Porophyllum: Removed by Cassini (Dict. Sci. Nat. Paris 43: 56. 1826)
to Porophyllum Guett. as P. ellipticum Cass.
272 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
C. suaveolens: Placed in Senecio L. by Elliott (Sketch Bot. S. Carol. & Ga.
2: 328. 1823); later placed in Synosma Raf. ex Britton & Brown (Illus. FI.
No. U.S. Canada 3: 474. 1898)
C. sonchifolia: Tentatively removed to Crassocephalum Moench (= Gynura
Cass., nom. cons.) by Lessing (Synop. Comp. 395. 1832). Later placed in
Emilia Cass. by De Candolle as E. sonchifolia (L.) DC. ex Wight (Contr.
Bot. India 24. 1834).
C. hastata L.: Tentatively referred to Ligularia L. by Lessing (Synop. Comp.
390. 1832), but the combination under Ligularia not made. Later trans-
ferred to Senecio as S. sagittatus by Schultz Bipontinus who united Cacalia
with that genus (Flora 28: 498. 1845).
C. atriplicifolia L.: Transferred to Mesadenia Raf. (nom. superfluum =
Arnoglossum Raf.) by Rafinesque (New Fl. N. Am. 4: 79. 1838). Treated
as a Senecio by Hooker, who combined Cacalia with Senecio (Fl. Bor.-Am.
1: 332. 1834).
By the beginning of 1838 only Cacalia hastata and C. atriplicifolia of
the original species had not been transferred formally to other genera.
Early in 1838 appeared Volume six of A. P. de Candolle’s Prodromus,
which included a revision crucial in the typification of Cacalia. De Can-
dolle divided Cacalia into four sections, retaining only three of Linnaeus’s
original species: C. hastata and C. suaveolens, which he placed in section
Eucacalia DC., and C. atriplicifolia, which was a member of section Cono-
phora DC. Thus, he effectively limited the choice of lectotype to either
C. hastata L. or C. suaveolens L. Since C. suaveolens had been trans-
ferred to Senecio by Elliott (1823), who left C. atriplicifolia in Cacalia,
C. hastata becomes the lectotype species. This species is quite in accord
with the original description of the genus (Linnaeus, Gen. Pl. ed. 5. 362.
1754), and we can see no reason for choosing another species.
The selection of Cacalia hastata as lectotype for Cacalia L., primarily
on the basis of the removal of the other species to different genera, is in
accordance with both the Guide for the Determination of Types set forth
in the International Code and the recommendation (7B) that the lecto-
type should be so selected as to preserve current usage. As so typified,
Cacalia is a genus of North America and Asia (including easternmost
Europe). The question of whether a number of genera should be segrega-
ted from Cacalia as now circumscribed taxonomically is not yet settled,
but no matter what the outcome of future investigations, Cacalia as typl-
fied by C. hastata, will be stable and a minimum of new combinations will
have to be made.
We regard the species of Cacalia in eastern North America as belonging
to two sections: CAcaLtA, represented in North America by C. suaveolens
L. and in Asia by C, hastata L. and a number of allied species; and CONO-
PHORA DC., restricted to eastern North America. (Cf. Vuilleumier, 1969.)
The pertinent synonymy of these sections is shown below:
Cacalia L. Sp. Pl. 2: 834 1753; Gen. Pl. ed. 5. 362. 1754.
Sect. Cacalia.
1969 | VUILLEUMIER & WOOD, CACALIA 273
Cacalia sect. Eucacalia DC. Prodr. 6: 327. 1838. LectoTYPE SPECIES: C.
hastata
Synosma Raf, ex Britton & Brown, Illus. Fl. No. U.S. Canada 3: 474. 1898.
Type sPEcIES: S. suaveolens (L.) Raf. ex Britton & Brown (C. suaveolens
Hasteola Raf. ex —— Not. Syst. Leningrad 20: 380. 1960, nom. super-
uum. TYPE SPECIES: H. suaveolens < Pojark. Se suaveolens 1.) [ Not
validly published by Rafinesque, New FI. N. Am. 4: 79. 1838; as validated
by Pojarkova includes the type species 7 Cacalia.]
Sect. Conophora DC. Prodr. 6: 329, 1838. LecrotyPE species: C.
atriplicifolia L.
a aplaag Raf. Fl. Ludovic. 64. 1817. Type species: A. plantagineum
Raf. C. plantaginea (Raf.) Shinners (C. tuberosa Nutt.).
Mesadena Raf. New Fl. N. Am. 4: 78. 1838, nom. superfluum. LEcTOTYPE
ECIES: M. atriplicifolia (L.) Raf. (C. atriplicifolia L.). [Includes the
“eto species of Arnoglossum Raf. |
LITERATURE CITED
CuatTrREcasas, i A new genus and other novelties. Brittonia 8: 151-163. 1955.
[ Cacalia, :
: = a on Andean Compositae —IV. bid. 12: 182-195. 1960.
Hircucocx, A. S., & M. L. Green. Standard-species of Linnaean genera of
Phanerogamae. Pp. 111-199 im International Botanical Congress, Cam-
bridge, England, 1930. Nomenclature. Proposals by British Botanists. 1929.
[Cacalia, 180.] (Reprinted in Brittonia 6: 114-118. 1947.)
Kiramura, S. Compositae Japonicae. Pars tertia. Mem. Coll. — Kyoto
Univ. B. 16: 155-292. pls. 1-7. 1942. [Lectotype of Cacalia, 170.]
Pippen, R. W. Mexican “Cacalioid” genera allied to Senecio (Compositae).
Contr. U.S. Natl. Herb. 34: 363-448. 1968.
Poyarkova, A. Notae criticae de genere Cacalia L. s. l. (In Russian.) Not.
Syst. Leningrad 20: 370-391. 1960.
. Cacalia L. Fl. URSS 26: 683-697. 1961.
RypBERG, 2 A. Some senecioid genera —I. Bull. Torrey Bot. Club 51: 369-
378.
ibaa Tk H. The Texas species of Cacalia. Field Lab. 18: 79-83. 1950.
VuILLEUMIER, B. The genera of Senecioneae in the southeastern United
States. Jour. Arnold Arb. 50: 104-123. 1969. [ Cacalia, 115-119.]
Gray HERBARIUM ARNOLD ARBORETUM
OF
IN
HARVARD UNIVERSITY HarvARD UNIVERSITY
274 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
A REVISION OF THE MALESIAN AND PACIFIC RAINFOREST
CONIFERS, I. PODOCARPACEAE, IN PART
Davip J. DE LAUBENFELS
THE RAINFOREST FLORA of a considerably submerged land area extend-
ing from southeast Asia through Indonesia and New Guinea to the Tonga
Islands includes an important conifer element for much of which systematic
examination is essentially lacking. It will be the purpose of this study
to present in three parts a critical account of the genera of Coniferales
that occur primarily in this area, that is to say, all of the tropical rain-
forest conifers beyond the mainland of Asia except two species of Pinus
whose ranges extend into Indonesia and the Philippines, plus such species
in southeast Asia as belong to the island rainforest groups.
Conifers are, in general, strongly divided into northern hemisphere
and southern hemisphere elements (Li, 1953). The characteristically
southern hemisphere families are Podocarpaceae and Araucariaceae, both
of which reach their greatest luxuriance in the area under consideration.
In addition, several genera of Cupressaceae occur in the southern hemi-
sphere as does one genus of Taxodiaceae, but of these only Libocedrus 1s
truly a part of the rainforest and included here. Taxaceae, formerly con-
sidered in the Coniferales, is not important, posing no taxonomic prob-
lems here, and will be omitted also, The rainforest conifers to be studied
involve twelve genera and well over one hundred species. New Caledonia
alone, centrally located with respect to the floristic region but recently re-
markably isolated, has preserved some forty species, all endemic, while
the extensive forests of New Guinea have yielded nearly as many again.
Indonesia in general has a conifer flora of equal richness to New Guinea,
sharing many species, while the rainforests of Queensland, Fiji, and lesser
areas have fewer elements many of which are endemic. :
More than a third of the species being described, both new and prev!-
ously recognized, were studied in their natural state during two extensive
field trips to the Pacific in 1957 and in 1964—65. In addition, the directors
and personnel of many herbaria contributed greatly to the completeness
of the study by their sympathetic help, and I should like to extend my
deep gratitude to them. The following key identifies the herbaria whose
specimens were consulted:
A Arnold Arboretum of Harvard University, Cambridge
BM ___ British Museum (Natural History), London
BRI Botanic Museum and Herbarium, Brisbane
FI Herbarium Universitatis Florentinae
GH Gray Herbarium of Harvard University, Cambridge
iLL ~——sdU niversity of Illinois Herbarium, Urbana
1969 | DE LAUBENFELS, PODOCARPACEAE 275
ad Royal Botanic eee Kew
L Rijksherbarium, Lei
LAE Department of sei Division of Botany, Lae
NA U.S. National Arboretum, Washington
Nsw National Herbarium of New South far Sydney
NY New York Botanical Garden, New Y
P Muséum National d’Histoire unin oe
RSA Rancho Santa Ana Botanic Garden, Claremont
spt Hortus Botanicus Bergianus, Stockholm
Us U.S. National age (Department of Botany), Smithsonian Institu-
tion, Washin De
Z Botanic Pearcy pie Institute of Systematic Botany of the University,
Ziirich
In addition, I should like to thank M. Corbasson, Director, Bureau
address, M. Schmid, Centre O. R. S. T. O. M., Nouméa, Lucien Lavoix
of Nouméa, and J. W. Parham, Department of Agriculture, Suva, Fiji,
for oes help both in the field and i in obtaining additional important speci-
men
The following comments apply to the citation of specimens:
1. Each specimen is accompanied by a symbol denoting its development and
sex. These include ‘“‘j” for juvenile, ‘‘s” adult but sterile, “2” female structures
present, and 3” a structures present When more Ren one stage is in-
cluded on the same sheet, more than one symbol will be used. Where appro-
priate and available, elevation figures will ae included (“m. ig for meters or “ft.”
for feet).
2. Where collections are numbered in series (sometimes without a collector’s
name) the standard abbreviation will be used. These include:
ANU Herbarium ee Seine agape
BRUN Brunei, Forest Departm
BSIP__ Bri itish Solomon inode Pade
BW Boswezan, Forestry Division, Netherland New Guinea
NGF N Guinea Forest Departm
NIFS_ Netherlands Indies Forest ice bb: series bossen buitengewesten —
islands outside Jav:
SAN North Borneo Forest Department, Sandakan
SFN Singapore Field Number
Podocarpaceae is a well differentiated family that is distinguished by
holz, 1941). The seeds in many genera are produced on structures so
modified from the cone morphology that one can not easily refer to them
as cones, although true seed cones and intermediate structures are found in
the family. Pollen cones are always truly cone-like and for all but two
276 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
genera are developed strictly on separate plants from the seed structures.
Nine of the twelve genera being recognized here occur in the tropics and
four of these are endemic to the tropics. All but a few species of the
family grow in very moist areas, some, particularly in Tasmania, being
found beyond the tree line as small or prostrate shrubs. Most, however,
are large forest trees, many with broad leaves quite unlike the usual
conception of conifers.
Within Podocarpaceae there has been great variation in the size and
complexity of the recognized genera. The genus Podocarpus as generally
treated involves up to eight sections and well over one hundred species,
the differences between some of these sections being every bit as great
as those which separate most other genera. It is being proposed here
to divide Podocarpus into five separate genera in order to produce a more
balanced treatment of the family. In addition, one new genus is separated
from Dacrydium because of the different form of the fertile shoots and
the strikingly different foliage morphology. The result is a total of twelve
genera in the family, of which nine are to be considered in this study, six
in part I and three in part IT.
KEY TO THE GENERA OF PODOCARPACEAE
1. Seed cone cdi seeds not subterminal.
2. Ovules in ;
a produce on ordinary foliage branches; adult leaves in the
form of scale
4. Seed ned covered by an epimatium; leaves opposite, decussate.
igi e teen en ns a icrocachrys, not tropical).
4. Seed completely enveloped in the fertile scale or epimatium;
MOOVES GOUEMIEY GUIANBED. <5 cso ce nnumi sen vneswendenenes KE
on ee BEA ea aes ee (some species of Dacrydium, not tropical).
3. ag shoot specialized; adult leaves linear, flat, constricted at the
OCI eee Treen) renee (Saxegothaea, not tropical).
a. pedis
a Fertile as lacking; adult leaves developed. ............---- 002°
ie hosts, Seta ie See ines PA eid al eit Re: Microstrobus, not tropical).
5. Fertile scale an epimatium; adult leaves suppressed in favor of phyl-
i: a SION OF Pore ane Ee SPU bee Se EAE ee Phyllocladus.
1. Seeds one or a few, subterminal or dispersed near the end of a fertile
branch.
6. Seed free, projecting above an epimatium (fertile scale).
7. Seed structures terminal on ordinary foliage branches; leaves crowded,
awl-like, linear, or scale-like. ......... Decrydium (most species).
7. Seed structures cia on specialized shoots; leaves bilaterally flat-
OE SE I a es is ins ae eee en aten's Falcatifolium.
6. Seed covered by or aed with the scale.
8. Fertile bract forming a terminal crest over seed complex; leaves awl-
PRO RIES HT aint at Rt onl ge Sea arene Dacrycarpus.
8. Fertile bract separate from the seed complex; leaves
9. Seed complex becoming erect; leaves bilaterally flattened. -
vig Pee Ae RtU ees eee es AE Acmopyle.
1969 | DE LAUBENFELS, PODOCARPACEAE 277
9. Seed complex remaining inverted, leaves bifacially flattened.
10. Fertile shoot terminal on ordinary foliage branches; leaves
scale-like; parasitic shrub. [Podocarpus sect. Microcar pus}. r
Specialized fertile shoot, usually A oi leaves broad and
flat, usually distichous; not parasitic.
11. Fertile shoot scaly; leaves never with both hypoderm
and accessory transfusion tis
12. Seed with a beak; paary with hypoderm, usually
amphistomatic and decussate, oval or lanceolate. .
BR ry ieee remanent ee Orne Decussocarpus,
12. Seed without a beak; leaves without hypoderm,
spirally placed and hypostomatic, linear. ..........
2 seat nee tok leashed Ee ee a rumno pitys.
. Fertile shoot divided into a naked peduncle and a spe-
cialized fleshy receptacle; leaves with both hypoderm and
accessory transfusion tissue. ............ Podocar pus.
_
°
~_
—
Phyllocladus L. C. & A. Rich. ex Mirbel, Mém. Mus. Hist. Nat. Paris
13: 48. 1825, nom. cons. Type species: Phyllocladus billardieri Rich.
ex Mirbel [Phyllocladus aspleniifolius (Labill.) Hooker].
Podocarpus Labill. Nov. Holl. Pl. Sp. 2: 71. t. 221. 1806. Type species: Podo-
carpus aspleniifolius Labill. [ Phyllocladus aspleniifolius (Labill.) Hooker].
Brownetera L. C. Rich. Ann. Mus. Paris 16: 299. 1810. Nomen nudum based
on Podocarpus aspleniifolius.
Thalamia Sprengel, Anleitung zur Kenntniss der Gewiichse. ed. 2. 2: 218.
1817, based on Podocarpus aspleniifolius.
Small to large trees; bark dark brown or blackish and smooth, reddish
and fibrous within, shed in large thin flakes; abundantly branched,
branches often in whorls; juvenile leaves linear or slightly broader near
the apex, acute or rounded but with a small spine-like point, 1 mm. or
more wide and about 1 cm. long, changing rapidly on small plants to flat-
tened leaf-branch complexes or phylloclads with scale-leaves on non-
foliage branches; leaves represented by small spurs on the margins of
the phylloclads, ‘strongly keeled on the dorsal side, triangular in cross
section and on older plants scarcely or not distinguishable; phylloclads
extremely variable in shape, broad, dorsiventrally slightly differentiated
in some cases, reaching several cm. in length or aggregated along branches
in complexes to more than 20 cm. long or transitional as a large deeply
lobed phylloclad; monoecious, but individual trees may be verge
pollen cones in clusters but the central axis of the cluster in most cas
continuing growth, nearly sessile or stalked; seed cone consisting of seat
or numerous scales some of which are sterile, single ovules erect in the axil
of a scale: seed cones terminal or marginal on fully grown or reduced
phylloclads or clustered as are the pollen cones, becoming swollen, fleshy
or leathery; erect seeds as many as 20 per cone but usually only - or 3,
with a filmy aril (symmetrical but rough edged epimatium) growing as
*To be taken up as a genus elsewhere.
278 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
a cup around the lower half, protruding beyond the scale when ripe, oval,
wider than thick, with the micropyle as a crooked tip, about 3 mm. long.
The genus consists of five closely related species in mild to cool and
very moist climates, three in New Zealand, one in Tasmania, and one in
mountain areas from the Philippines to New Guinea. PAyllocladus is
sharply distinguished from related taxa by the distinctive phylloclads
which give it the popular name of “celery-topped pine.” The bark con-
tains abundant tannin and the wood is of good quality but, because all of
the forest species grow as scattered individuals, its commercial value is
limited. One species in New Zealand and part of the population in Tas-
mania grow as bushy pioneer plants around mountain meadows. The one
tropical example of this genus is the only podocarp species growing in
the tropics whose seeds are produced in a recognizable cone. This cer-
tainly suggests that the family Podocarpaceae, so abundantly developed
within the tropics, had its origins in cooler climate areas.
1. Phyllocladus hypophyllus Hooker f. Icon. Pl. ¢. 889. 1852. Type:
n., Mt. Kinabalu.
ge arwred hypophyllus var. protracta Warb. Monsunia 1: 194. 1900.
Synt ane: ey S. Mindanao, mountain forest of Dagatpan and
18272, Batjan (not seen).
Phyllocladus pesthstoen ‘(Warb.) Pilger, Pflanzenreich IV. 5 (Heft 18): 99.
Phyllocladus major Pilger, ig a 54: 211. 1916. Type: Ledermann 9872,
Lordberg, NE. New Gui
Common small tree on ridges or becoming quite large in the forest, 30 m.
or more high; bark hard, rough with large lenticels, dark brown, breaking
off in large scales; inner bark straw color; hence: more or less whorled
around the main stem and densely ramified: foliar buds on young plants
with long thin and somewhat spreading bracts, these becoming tighter and
more globular on older plants; phylloclads sometimes glaucous, particularly
underneath, variable in shape, deeply lobed on young specimens but be-
coming ese lobed in maturity, margins nearly entire to wavy with indi-
vidual lobes ca. 5 mm. wide and 2 mm. long, oval to triangular, 3 or 4
cm. long and 2 cm. wide, single or aggregated alternately along lateral
branches of limited growth: pollen cones clustered around a shoot that
continues growth, peduncle 5-25 mm. long; mature pollen cones to 15 mm.
long, 3 mm. in diam.; seed cones clustered on stalks about 1 cm. long oF
terminal on a slightly modified phylloclad or any possible intermediate con-
dition, small, with 1-3 or more fertile scales, first red when mature, then
brown and leathery.
DistRIBUTION. Luzon and Borneo to New Guinea, scattered and often
common in moist forests and on ridges generally, from 1,500 to 3,200
meters, and occasionally from 900 to 4,000 meters. Map 1.
Sarawak. Mt. Poi, upper cave, Clemens 20026 j (Ny). Mt. Laiun, Richards
ee eee
1969 | DE LAUBENFELS, PODOCARPACEAE 279
out loc. Beccari 2391 s (kK), 3220 2 (kK). Brunei. Mt. Ulak, Ashton BRUN
1033 s 4,300 ft. (K, L). North Borneo. Jesselton, Kumu Rengis, Wyatt-Smith
[?] 71650 2 80 ft. [sic] (x, 1, Us). Penampang, Leano-Castro 5992 s 6,000
ft. (k, L), Clemente 6217 s 5,000 ft. (K). Ranau, Meijer SAN 21968 2 5-
6,000 ft. (kK), Mikil 56277 s 7,000 ft. (K), Burgess SAN 25167 s 4,500 ft. (xk).
Mt. Kinabalu, Low s.n. 2 8,000 ft. (K-holotype), s.2. 6 10,000 ft. (Kk), Gibbs
4088 j 7,000 ft. (BM, K), 4152 2, j 6,000 ft. (Bm, K), 4238 s (BM), 4273 @
9-12,000 ft. (Bm, kK), Clemens 10556 & (A), 10565 @ (A, GH, K), 10654
2 (A), 10957 s (BM), 27930 2 6,000-13,500 ft. (A, BM, K, L, NY), 29328
? 10,000 ft. (A, BM, ILL, K, L, NY), 29743 2 8-9,000 ft. (BM, K, NY),
30029 2 7,000 ft. (A), 30030 2 10,500 ft. (K, Ny), 31838 & 7,000 ft. (A, L),
31927 8 89,000 ft. (Ny), 32459 s (BM, L), 50626 s (BM), 50784 2 7-9,000 ft.
(A, BM, L), 50797 2 10,000 ft. (BM, L), 51220 s (Bm), Haviland 1092 2,
11,000 ft. (A, BM, K, L), Sinclair & Kadim 9053 s 6,950 ft. (L), Chew & Corner
RSNB 710 2 7,500 ft. (k, Ny), RSNB 4172 2 5,000 ft. (x), RSNB 4824 9
6,000 ft. (K), Smythies S10622 2 9,000 ft. (K, L), Wyatt-Smith 80370 s (x),
80371 2 (kK, L), Anderson S27089 8 11,800 ft. (K), 527090 2 11,300 ft. (kK, L),
Meijer SAN 22114 s 4,000 ft. (k), SAN 29271 & 9,700 ft. (K, L). Nicholson
SAN 17823 2, 6 8,800 ft. (K, L), Fuchs & Colenette 21430 2 3,375 m. (Kk),
Carr SFN 27617 s 11,500 ft. (sm). Trusmadi Kudat, Mikil SAN 31784 8 (1).
Sobong Peak, Lobb (1857) s 4,000 ft. (BM, K). Borneo. W. Region, Bengka-
jang, Banan, NIFS 669671 j 1,400 m. (Lt), bb24777 j 1,200 m. (a, L). B. Raja,
Winkler 1036 s 1,600 m. (x). Ulu Kelan, Molengraaf B3477 s (t). Top of
Semedum, Hallier 697 @ (A, K, L, Ny). Mt. Palimasan, W. Kutei (Belajan R.),
Kostermans 12894 5 700 m. (pM, K). Mt. Niapa on Kelai R., Kostermans
21482 $ 1,000 m. (x, L). Philippines. Luzon: Mt. Panai (Benguet), Gillis
27257 s (A, K, L, us), Merrill 4753 j (K, L, NY, US), Quisumbing & Sulit 82404
Ss 7,700 ft. (Ny). Mt. Sifigakalsa (Benguet), Sulit 7669 8 2,500 m. (A, L).
Benguet, Alvarez 18364 s (pm). Lepanto Dist., Curran 10957 & (kK, L, NY, US).
Mt. Data, Steiner 2150 j 2,200 m. (L). Mt. Pukis (Bontoc), Ramos & Edano
37757 8 (a, us). Mt. Tabuan-Buan (Cagayan), Ramos 77401 2 5,800 ft. (xk,
Ny). Center, Loher 4851 s (K), 5203 2 (aA, K, Ny, US). Mrnporo: Mt. Halcon,
Merrill 5788 s (xk, Nx, us). Mt. Dulangan, Whitehead (1896) s 5,000 ft. (BM).
Minpanao: Mt. Katanglad (Bukidnon), Sulit 10052 2 2,200 m. (a, L), 10124
$ 2,300 m. (A). Mt. Candoon (Bukidnon), Ramos & Edano 38738 2 (A, US).
Kaatoan Chinchona (Bukidnon), Britton 439 2 1,380 m. (L). Mt. Apo (Davao),
Elmer 11463 s (a, BM, FI, K, L, NY, US, Z), Clemens 15675 j (A, NY), Mearns
Hutchinson 4679 s (L). Mt. KcKinley, Kanehira 2676 j (NY). Celebes. Mt.
Tampai, Palu (Menado), NIFS bb15154 2 2,500 m. (L). Parigi Lombok
(Menado), NIFS 6b15026 s 1,100 m. (tL). Sawito (Enrekang), NJFS 6b20782
S 1,750 m. (L). Mt. Tahole, Labu (Malili), Burki bb24089 2 1,500 m. (L).
Porehu (Malili), NJFS 6619564 2 1,500 m. (A). Makale-Toloko (Manggala),
NIFS bb20270 s 1,200 m. (a, L). Moluccas. Batjan, de Haan bb23236 s 2,199 m.
(L). Obi, de Haan 6b23812 s 700 m. (x). Buru, N/JFS 6621509 s 800 m. (t),
Binnendyk s.n. j (K, L). Middle Ceram, G. Sofia, Stresemann 133 s 2,200 m.
(L). New Guinea. VocELKop: Mt. Nettoti, van Royen 3873 s 1,960 m. (L),
van Royen & Sleumer 7403 s 1,750 m. (K, L), Versteegh BW 10407 s 1,700 m.
280 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
(L). Neentjapaki Mts., Kebar Valley, Kalkman BW 6373 s 1,090 m. (L). Adjar,
Kebar Valley, Koster BW 6887 2 1,110 m. (L). Tobie Mts., Kebar Valley,
Schram BW 7972 s 720 m. (x). Anggi Lakes, Gibbs 5992 2 7~9,000 ft. (BM,
K), Versteegh BW 248 s 2,000 m. (A, K, L), BW 253 2 2,100 m. (A, K, L), BW
281 s (A, L), Kanehira & Hatusima 13704 s (A), 14096 s (A), Stefels BW 2008
j 1,875 m. (L), BW 2010 s 1,860 m. (t), BW 2031s 2,200 m. (L). Koebri Ridge,
Gibbs 5657 s 8,500-9,000 ft. (pm, K). Ransiki, Sioriep, Mangold BW 2262 s
1,200 m. (K, L). Mt. Mundi (Ransiki), Mangold BW 2254 s 1,900 m. (tL).
WeEsTERN Harr: Mt. Genofa (E. of Arguni Bay), Salverda bb22564 s 750 m.
(L), Versteegh BW 7596 2 1,000 m. (L). Wissel Lakes, Eyma 4954 ¢ 1,750 m.
(A, K, L), 5228 6 (a, K, L), 5371 2 (A, K, L), Versteegh BW 3009 @ 1,750 m.
(A, L), Johannes BW 3262 s 1,750 m. (L), Vink & Schram BW 8764 2 1,500
m. (L), BW 8945 s 1,850 m. (1). Nassau Mts., Docters v. Leeuwen 10906 s
2,600 m. (A, K, L). Mt. Doorman (Mamberamo R. Region), Lam 1628 &
3,250 m. (1), 1647 2 3,500 m. (L), 1742 2 3,250 m. (x), 1984 s 2,560 m. (z).
Lake Habbema, Brass 9058 & 3,225 m. (A, BM, K, L), 9090 2 (A, BM, K, L),
10528 % 2,800 m. (A, BM, K, L), Brass & Meyer-Drees 10432 @ 3,225 m. (a, 1),
Brass & Versteegh 10446 9 2,840 m. (A, BM, L), 10446A 8 3,200 m. (A, BM,
L). Barnhard Camp, Brass & Versteegh 11931 2 1,850 m. (A, BM, K, L), 12523
s 1,100 m. (A, BM, L), 12523A @ 1,150 m. (A, K, L), 13520 @ 900 m. (A, BM, L),
135204 2 (a, L), Brass 12191 2 2,100 m. (A, L). Cycloop Mts., van Royen
isotypes of Phyllocladus major), Wabag-Maramuni Road, Saunders 1025 s
10,000 ft. a Rjcoenge (Mt. eae pee & Pullen 5871 s 7,600 ft. (A,
BM, K, US). Wichmann, Pulle 982 s 2,500 m. (Kk, L), 1018 s (kK, L), 1042
é - 100 m. He i, 2). Upper Minj ae. Pullen 273A j 9,000 ft. (A, L). Al
River Mts. (Nondugl), Womersley NGF 5351 2 7,000 ft. (A, BM, XK, L). Mt.
ae Sirimbki, Walker ANU 859 2 9-9,500 ft. (A, K, L), 859A j 9,500 ft.
(A, K, L). Chimbu, Cavanaugh NGF 3333 2 (kK). Wa imambuno (Chimbu u),
Saunier 824 s 9,000 ft. (A, BM, K, L, us). Mt. Wilhelm, Page 5651 s 2,600
m. (K, L, z). Lake Inim, har haps "ANU 2177 5 8,300 ft. (kK, L). sissegeraigs
Clemens 4942 s 6-7,000 ft. (A, z), 5117A 2 6,000 ft. (A). Samanzing, Clemens
9384 3 7-8,000 ft. (a), 9549 3 8~9,000 ft. (a). Mt. Enggom, Sarawaket Range,
van Royen NGF 16182 & 11,000 ft. (k, L), Mannasat, Cromwell Mts., Hoo8-
land 9482 2 7,600 ft. (kK). Bolan, Lauterbach 303 s 2,400-3,000 m. (pM). Mt.
Kaindi (Bulolo), Brass 29692 2 2,150 m. (A, K, L, Ny, us), Millar & Womers-
ley NGF 12255 s 7,000 ft. (a, K), McVeagh NGF 7580 2 3,000 ft. (A, BM, KE,
L), de Laubenfels P4381 4 6,500 ft. (A, K, L, RSA, sBT), P431A j (a), Toropai
NGF 17153 @ 6,900 ft. (k, L), Havel & Kairo NGF 17341 @ 7,000 ft. (K).
Wau-Salamaua Road, Millar NGF 22785 92 6,400 ft. (xk). Mt. Amungwiwa,
Wau, Womersley NGF 17946 s 11,400 ft. (tL). Wagau, Buang Region,
Womersley NGF 17902 s 4,500 ft. (kK, x). Papua: Mt. Giluwe, Schodde 2014
? 8,800 ft. (k, L). Mt. Tafa (Cent. Div.), Brass 4035 : ay ny). Murray Pass,
Wharton Range, Brass 4578 s 2,840 m. (A, BM, NY), 4584 @ (A, K, L, NY, US).
Mt. Obree, Owen Stanley Range, Lane-Poole (1923) s “mn 000 ft. (A, K). Mt.
Dayman, Maneau Range, Brass 22453 2 2,230 m. (A). Mt. Maneao, —
519 s 7,500 ft. (K). Mt. Mon [Mau?], Crutwell 896 j 6,800 ft. (K).
Vinevo, Goodenough, Crutwell 1423 s 7,000 ft. (kK).
ILLUSTRATION. Hooker, f. Icon. Pl. t. 889. 1852.
1969 | DE LAUBENFELS, PODOCARPACEAE 281
°
—— ee ea eres te si —<—$$__ ~——+- 7 — e's
: a
| tte s
a
\ BALANSAE SS
atum (Roxburgh) Wallich (dots west o :
i tensi benfels, known only from the
3, D. pectinatum de Laubenfels (dots west of line), D. nidulum de
lansae Brongniart & Gris, known through-
Fiji Islands; ;
Laubenfels (dots east of line , D. ba
out New Caledonia, and D. cupressinum Solander ex Lambert, known from
d
New Zealand.
282 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
In addition to its completely disjunct distribution, Phyllocladus hypo-
phyllus can be differentiated from all other species of the genus by its
distinctly larger phylloclads. In other species the larger structures are
deeply lobed and transitional to branch systems, those without deep lobes
are less than 20 mm. wide or 25 mm. long. PAyllocladus hypophyllus is
unique also in frequently having the seed cones terminal rather than lateral
on the phylloclads, and in having peduncles on both pollen and seed cones
up to twice the length observed in other species. The two species with
intermediate sized phylloclads approaching the lower limit of those of P.
hypophyllus are P. glaucus and P. aspleniifolius, both of which share the
glaucous habit with P. hypophyllus. The former has seed cones with
numerous fertile scales and the latter has particularly small and nearly
sessile pollen cones. The species P. major and P. protractus have been
differentiated from P. Aypophyllus on the basis of the shape of the cladodes,
the position of the seed cone, and by their glaucous aspect. These dif-
ferences, however, are found within local populations related to age of
the tree or even on different parts of the same specimen.
Dacrydium Solander ex Lambert, Descr. Genus Pinus 1: Appendix 93.
1807. Type species: Dacrydium cupressinum Solander ex Lambert.
Lepidothamnus Phil. Linnaea 30: 730. 1860. Type species: Lepidothamnus
fonkit Phil. [Dacrydium fonkii (Phil.) Benth.].
Shrubs and trees varying considerably in stature; juvenile leaves awl-
shaped (falcate needles), longer than the adult, or in some species bi-
facially flattened and linear; adult leaves quite variable among the
species from scale leaves to leaves resembling the juvenile needles and
with either gradual or abrupt transitions uniting the different forms dur-
ing their ontogeny; dioecious (or rarely monoecious in some New Zealand
species); pollen cones cylindrical, terminal, or lateral and sessile, or both;
seed cones much reduced, with bracts hardly modified from foliage leaves,
often becoming fleshy when ripe, terminal, often on a short lateral branch;
ovules inverted on bracts in a nearly terminal position and partly covered
by an epimatium; seeds usually becoming erect, projecting well beyond
the apex of the modified cone, occasionally occurring in pairs or three
together, sometimes surrounded by the leaf-like extremities of the cone
bracts, oval with the micropyle forming a small tip, usually somewhat flat-
tened, on some species remaining inverted and covered by the fertile scale.
The genus Dacrydium occurs in a wide range of temperature and soil
conditions but rarely in anything less than a very moist climate. It is
readily divisible into two subgroups based on the internal morphology of
the wood, leaves, and pollen (Tengnér, 1965). In one of the groups, called
by Florin (1931) Group C, the adult leaves are more or less overlapping;
broad, bluntly keeled scales (in one species, a prostrate alpine shrub,
plants with juvenile type short flat leaves sometimes are fertile). The
other group, called Group B, lacks scale leaves in all but two species
where the scale is narrowly and sharply keeled and strongly appressed.
1969 | DE LAUBENFELS, PODOCARPACEAE 283
The seeds in this group always become more or less erect, while in Group
C some species have inverted mature seeds covered by the fertile scale.
Group C is entirely extra-tropical and will not be treated here. Group
B is primarily tropical, the two groups overlapping in New Zealand
where most of the Group C species are found. In Group B the juvenile
leaves are scarcely distinguishable between the various species, being
lanceolate, slender, and bifacially flattened on the seedling but soon be-
coming strongly keeled and awl-shaped. For the most part, the seeds are
also very similar throughout, so that the species are distinguished pri-
marily by the form of the seed and pollen cones, and by the adult leaf
form. Four common leaf types occur, one with fairly short needles (2-
5 mm.) changing gradually from the juvenile form (cupressinum, balansae,
nidulum, pectinatum), a longer type with more flexible needles (beccarii) ,
a type with narrow, flat, and lanceolate leaves (xanthandrum), and one
with scale leaves changing abruptly from the juvenile needles (elatum,
novo-guineense). In addition, there are a number of local species, usually
with distinctly bifacially flattened leaves and, in most cases, rather rare.
Most of the species are too small in growth form or are too rare to be
useful, but a few, as D. cupressinum, are valuable lumber trees.
KEY TO THE SPECIES OF DACRYDIUM
1. Trees or prostrate shrubs with adult leaves broad, imbricate, bluntly keeled
scales (Group
1. Trees or bushes with adult leaves narrow, appressed, sharply keeled scales or
longer spreading leaves (Group
2. Abrupt change between juvenile and adult leaves, which are minute (not
more than 1.5 mm. lon
3. Bracts in the fertile area similar to scale-like foliage leaves. ...
3. aay in the fertile area distinctly longer than the foliage aor or
1. en and pollen cones not small; foliage leaves scale-like.
Henin deduces Mo hiats ath auc niet ee eee are 3. D. novo- guineense.
4. Seeds and pollen cones small; foliage leaves spreading. .
. D. nausoriense.
2. Gradual change from juvenile to adult leaves, which are at least 2 mm.
ong.
_ Bracts in the fertile area not surpassing the epimatium and not
longer that the foliage leav
me
6. sroerreca aa og ate lanceolate; leaves pe (0.6 mm.)
eee ele ee ee D. seen
6. tin ab long triangular; leaves less than 0.4 mm. thic
curved upwards at the ti
Pp.
7. Slender, linear oo. with - og turned upwards.
Bet hades atc aes D. pectinatum var. ‘pectinatum.
k, ly taperin, ny spreading leaves
page Send : es : een 5b. D _ pectinatum v. var. - robustum.
Bracts in the fertile area distinctly longer than the epimatium and,
where the foliage leaves are not long, distinctly elongated by contrast.
-
JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
8. Bracts.in the fertile area distinctly longer than the foliage leaves
of the subtending branch; gin ye a triangular
9. Bracts, in the fertile area, and foliage — strongly keeled,
triangular or quadrangular in cross sect
10. Developing seed extending well oa the elongated
bracts of the fertile area; foliage leaves not more than
0. wide.
11, Leaves staging Me as thick as wide, tip more or
less Blunt and Hat incurved, ......2ciesccvcaends
RCs aE era oe Bib ay a nidulum var. nidulum.
_
—
D.
. Leaves noticeably wider than thick, tip distinctly
pai with a slight prickle, generally incurved and
rowded. ........ 6b. D. nidulum var. araucarioides.
Hire a completely surrounded by elongated
racts, the tip protruding slightly on maturity; foliage
leaves ae ee than 1.0 mm. wide.
12. Pollen cones 2.0 mm. in diameter; leaves curved
upwards but not phe wg markedly tapering, quad-
rangular in cross section. .......... 7. D. balansae.
. Pollen cones 2.5-3. aan mm. in diameter; leaves strong-
ly incurved, linear, axial surface concave towards
EAs 65< oF vv Acs aes ou8s 8. D. araucarioides.
9. Bracts in the fertile mar and foliage leaves distinctly flat, more
than twice as wide a:
13; it and alien: cone small: leaves srg 3-4.5 mm.
Re eat a td oat ers al oak Iycopodioides.
13. Sed and pollen cone not small; leaves Tina, 4-7 m
SE ae ee tee eee Pere spore
8. Bracts in ee fertile area no longer than the ee eis of the
subtending branch; microsporophylls elongated, lanceolate.
14, iy bract not surpassing the mature seed, leaves 5-10 mm.
—
=
_
i)
1S. » Pollen cone 20-25 mm. long by 5-7 mm. in diameter;
re less than 0.8 mm. wide
Seed cone terminal on ordinary foliage —
mature seed surrounded ne bracts. 11. D. magm
. Seed cone terminal on shoots with reduced ee
seed well exposed when mature.
17. Leaves quadrangular or triangular in cross sec-
tion, imbricate.
18. Leaves uniform, more than 5 mm. long, at
least ten times as long as wide.
19. Leaves spreading outward. ......----:
12a. D. beccarii var. beccarii.
19. Leaves ante and compact. ...---
Qn oe 12c. D. beccarii var. rudens.
Leaves variable, sometimes less than 5 mm.
long, less se eight times as long as wide.
sees Seta . beccarii var. subelatum.
17. Leaves twice as eae as thick, spreading at
nearly right angles to the stem. .....-----°°
13. D. xanthandrum.
—
ON
—_
2
1969} DE LAUBENFELS, PODOCARPACEAE 285
15. Pollen cone 20-25 mm. long by 5-7 ee in diameter;
leaves at least 1.0 mm. wide. .......... gay
Nene bract much longer than the seed, for 12- 20 m
~
- guillauminii.
comosu
2. Dacrydium elatum (Roxburgh) Wallich, London Jour. Bot. 2: 144.
1843.”
Juniperus i aia Fl. Indica 3: 838. 1832. Lectotype: Wallich 6045,
Malay
iendians sung Miquel, Pl. Junghuhn, 1: 4. 1851. Type: Junghuhn
s.n., Sumatra.
Decrydium ed Hickel, Bull. Soc. Dendr. France 76: 74. 1930. Lecto-
type: Pierre 1396, Cochin China, Phu Quoc Island.*
Tree to 40 m., much branched with masses of erect twigs forming a
dome-like crown; bark furrowed and flaky, reddish-brown, inner bark
pink; juvenile leaves acicular, to at least 12 mm. long, gradually becom-
ing shorter and more robust before changing abruptly on young trees,
about 6-8 mm. long, sharply keeled on four sides and nearly straight,
spreading, acute; mature foliage branches cord-like, 1-2 mm, in diam
covered with imbricate scales which are acute and sharply keeled, 1—1.5
mm. long by 0.4-0.6 mm. wide, occasionally passing through a semi-adult
or transitional stage of short spreading leaves about 1.5 mm. long; branches
with juvenile leaves occasionally fertile; pollen cones terminal, usually on
short lateral branches, thus sometimes almost lateral, cylindrical, 4-5 mm
long and 1.2 mm. wide: microsporophylls triangular, acute; seed cone
terminal, generally on short lateral branches, bracts of the cone becoming
slightly enlarged, red and fleshy when mature; the solitary naked seed
becomes almost erect, tapering to a blunt apex, reaching 4-4.5 mm. in
length
DistrisuTion. In humid mountain forests from north central Thailand
and Tonkin to central Sumatra and Sarawak, from 500 to 1,700 meters 1n
elevation or even down to sea level where suitable conditions exist. Map 2.
Thailand. NortuH CENTRAL: Loei, Phu Krading, Tham Nam, Royal Forest
Dept. 3631 j, 1,045 m. (us), Kerr 8727 j (K), 8727A @ (kK), 8727B 3 yee a
2263 j 1,300 m. (A). Without loc., Smitinand 19058 j 1,200 m. (XK). TRAL:
N. akhaun Nayok, Phengkhlai 691 8, j (K, L). Cambodia. Plateau Pega
Gulf of Siam, Showe (1927) s, j 3,000 ft. (sm). North of Kampot, Poilane
14707 2 (ny). Near Komplon (Phnom Penh), Bejoud 717 ° (1LL). Without
loc., Pierre 19074 s (k). Tonkin. Than Moi, Balansa 596 ¢, j (ILL, K). An-
nam. Summit Mt. Bani, near Da Nang, Clemens 4280 j (K, NY, US). Bana,
near Tourane, Poilane 1539 s, j 1,200 m. (N¥-syntype of D. pierrei), 7095 2 (A-
Specie cutively through the whole paper.
Thee I did ay siete a bbe: but list ed many specimens of which Balansa 576
is the first. The one specimen collected by Pierre is here chosen as the lectotype be-
Cause of the specific epithet.
286 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
syntype of D. pierrei). Kontum Prov., Poilane 33351 s 1,200 m. (ILL). Nha-
trang, Poilane 25 j (A-syntype of D. pierrei), 3455 2 (a-syntype of D. pierret),
3782 Q (A, K-syntypes of D. pierrei), 4411 2 (a-syntype of D. pierret).
Cochin China. Phu Quoc Island, Gulf of Siam, Pierre 1396 8, j (A, K, Ny-iso-
types of D. pierrei), Harmand (Godefroy) 901 é (a-syntype of D. pierrei).
Without loc., Poilane 32825 8 (ILL), Godefroy-Lebeuf sn. 8 (k). Malaya.
Thailand border (Botong), G. Ina, Kerr 7554 s, j (K). Penang, Wallich 6045
s (BM-lectotype of Juniperus elata; K-isotype), Sinclair 39094 s, j (K, L),
Walker 70 j (kK), Maingay 2262 s (kK), 2753 s (kK), Curtis 2880 s, j (K). Perak,
Ernst 1213 s, j (z). G. Butu, Wray 1028 j (x), 3899 s, j (kK). Pahang, G. Tahan,
Hanif & Nur sn. 8 (x), SFN 7959 s, j 5,500 ft. (a, K), Wray & Robinson
5354 s 3,300 ft. (kK), 5380 j (kK). Pahang, G. Lesong, Wakau 4155 j (kK). Ja-
hore, Mt. Ophir, Maingay 1503 2 (FI, GH, K, L), Moxon s.n, s (L). Sumatra.
Between Tapanuli and Silindong, Junghuhn s.n. j 2,000 ft. (L-holotype of D.
junghuhnii). Pajakumbuh, W. Taram, Meijer 6938 8, j 500-1,000 m. (Xk, £),
7040 2 (L). Poya Kombo, Teysmann 21647 8 (Kk), s.n.s (kK). Without loc.,
Praetorius sm. j (L). Sarawak. Merurong Plateau (Bintulu), Brunig S9991 s
750 m. (L). Mt. Dulit, Richards 1962 s 1,250 m. (BRI, K, L, US). Between Biak
R. and Sut, Pickles 2991 8 2,360 ft. (L, us). Lawas, Brunig 510673 4, j 900
m. (L). Borneo. Without loc., De Vriese s.n. j (L).
ILLustrations. Riptey, H. N. Fl. Malay Peninsula ¢. 227. 1925.
Corner, E. J. H. Gard. Bull. Straits Settlements 10: ¢. 5. 1939
Dacrydium elatum differs from D. novo-guineense in the form of the
female cone, in the form of the juvenile leaves, size of the pollen cone,
size of the mature tree, and in its occurrence generally at lower elevation.
Specimens of D. pectinatum have been much confused with D. elatum be-
cause the pectinatum foliage is similar to the juvenile foliage of D. elatum
but, the leaves of D. pectinatum are, in fact, shorter and distinctly curved.
The known range of these two species overlaps only in Sarawak. The
name elatum has further been applied to almost any uncertain Dacrydium
specimen from Borneo to the Fiji Islands. Hickel described D. pierrei,
contrasting it with D. beccarii, which he mistook for D. elatum.
3. Dacrydium novo-guineense Gibbs, Contrib. Phytogeography and
Flora of the Arfak Mountains 78. 1917. Lectotype: Gibbs 5648,
New Guinea, Arfak Mountains.
Tree to about 10 m. with branches rigidly ascending into a rounded
crown; juvenile leaves acicular, spreading and incurved, lanceolate, acute,
keeled on the back, to 7 mm. long by 0.7 mm. wide but variable in size,
changing abruptly to the adult form, occasionally passing through a semi-
adult or transitional stage of short spreading leaves about 1.0-1.5 mm.
long; mature foliage in imbricate scales, acute and sharply keeled, 1.0-
1.5 mm. long by 0.4-0.6 mm. wide; foliage branches 1.0-2.0 mm. in di-
ameter, penultimate branches becoming larger; pollen cones terminal,
usually on short erect lateral branches, cylindrical, 8 mm. long, micro-
sporophylls triangular; seed cones terminal on short curved lateral
branches, bracts long and spreading, reaching 3 mm. at the cone apex,
1969 | DE LAUBENFELS, PODOCARPACEAE 287
the whole cone becoming red and fleshy when mature, the single apical
seed becoming almost erect and extending well beyond the cone bracts,
5 mm. long, edges slightly keeled, tapering to a small blunt apex.
DistriBuTion. In open to mossy forests, often on ridge tops from 1,300
to 2,750 meters in elevation, occasionally lower. Locally common but
apparently localized; from Obi and the mountains of western New Guinea
at least as far as the Western Highlands of the Territory of New Guinea.
The collections from the Celebes are tentatively included here until it
can be determined whether these represent Dacrydium elatum or D. novo-
guineense. Map 2.
Celebes. Manado, Poso, Eyma 1623 s 1,700-1,800 m. (L), 3642 4 (tL). Ma-
samba, Kuniapu, N/JFS bb24964 s 1,500 m. (L). Masamba, Omboan, N/FS
6626288 j 1,800 m. (L). Enrekang, NJFS bb20786 j 1,900 m. (L). Moluccas.
W. Buru, Stresemann 395 s, j 1,800-2,000 m. (L). Buru, Martin s.n. j (1).
Obi, de Haan bb23813 s 700 m. (L), bb23814 s, j (L). New Guinea. VocELKoP:
Tamrau Mts., Van Royen & Sleumer 7219 s, j 2,000 m. (L). Kebar Valley,
Van Royen 3857 s, j 1,980 m. (L). W. of Mt. Nettoti, Van Royen & Sleumer
7948 2 2,100 m. (K, L, LAE), 7948B j (L). Arfak Mts., Gibbs 5648 2 9,000 ft.
(sa-lectotype; K-isotype), 5508 s, j 7,000 ft. (sm, K-syntypes), Kanehira &
Hatusima 13518 s 2,000 m. (a), Gjellerup 1032 s, j 1,800 m. (L). Anggi Lakes,
Versteegh 256 2 2,100 m. (1), 262 2 (1), 269 (1), Stefels BW 2015 j 1,860
m. (L), BW 2033 s, j 2,100 m. (L). WESTERN HALF: Wissel Lakes, Eyma 4422
S, J 1,750 m. (a, K, L), 4519 2 1,760 m. (A, K, L), Vink & Schram BW 8746
S 1,820 m. (L). Hellwig Mts., Pulle 663 2 1,300 m. (L), 966 s 2,600 m. (K, L).
Barnhard Camp, Brass & Versteegh 11967 $ 1,520 m. (A, BRI, kK, L), 12507
j 2,100 m. (A, BRI, L), TERRITORY oF NEw GUINEA: Western Highlands, Mt.
Hagen, Cavenaugh NGF 3337 s, j (A, BRI, L). Tagen R., Jimmi Valley, Womers-
ley & Millar NGF 7680 2 4,300 ft. (A, BRI). Minj-Jimmi Divide, Robbins 598
3, j 6,500 ft. (A, BRI, K, L, us). Nondugl, Womersley NGF 4420 &, j (A, BRI,
K, L). Papua: Sibium Range, Pullen 5930A j (L).
ILLUSTRATION. Gipps, L. S. Contrib. Phytogeography and Flora of the
Arfak Mountains, t. 3. 1917.
Being one of the scale-leaved species of Dacrydium, D. novo-guineense
cannot be distinguished in the sterile form from D. elatum from which it
differs in the elongated bracts of the seed cone and to a lesser extent in
the form of the juvenile leaves and the size of the pollen cone. Juvenile
specimens can often be separated from D. beccarii, with whose range it
overlaps, by their coarser and less dense growth. From D. nidulum the
juvenile leaves differ in their variable size including, for the most part,
greater length. The rapid change from juvenile to adult form is so strik-
ing and comes when the tree is yet quite small so that collectors generally
include mature leaf forms when dealing with D. novo-guineense. The
rigid ascending branches are another distinctive character.
4. Dacrydium nausoriensis de Laubenfels, sp. nov.
Arbor ad 25 m. alta, ramosissima. Folia plantarum iuvenilis acicularia,
ad 9 mm. longa, ad formam adultam abrupte convertentes; folia plantarum
288 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
—v
—_
—
—
—
—
ee
ee
Eccl
7
a, Dacrydium anion ig de Laubenfels, portion of the holotype,
de Laubenfels P302 (A), enlarged; b, ena de i _ nfels var. pectima-
tum, portion of the isotype, Nicholson SAN 292 (L), ged: c, D. pectina-
tum, var. mee a de Laubenfels, portion a the boi: ce "Meijer SAN 3790
(L), enlarged; b and c are at same magnificati
adultarum parva, patula, acuta, dorsaliter carinata, densa, 1 mm. longa,
0.4 mm. lata. Strobili masculi cylindracei, terminales vel laterales, saepe
utroque, parvi (?), ad 2.5 mm. longi. Strobili feminei ad apicem ramulorum
saepe brevi; folia ad basem seminis longiora, ad 2 mm. longa; semen pro-
trudendum, 3.5—4 mm. longum. Holotypus: de Laubenfels P302 (A) ), Fiji,
Nausori Highlands. Fic. la
DistRiBuTION. In slightly open forest on the leeward sides of the large
islands of Fiji and apparently of limited extent.
Fiji. gs Levu: Nausori “ee de Laubenfels P302 2 1,900 ft. (A
holotype RSA, SBT-lsotypes), P303 j (A, RSA, ait P304 & (A, RSA, SBT),
ae NH19 2 (x), NH23 & (x), Bas 29 _ Kuruvoli 13326 & (K).
A Levu: Lambassa, Sarava, Damanu L14 5 (e) E), pri FD832 2 400 it.
o
1969 | DE LAUBENFELS, PODOCARPACEAE 289
The species of Dacrydium with sharp scale-leaves, changing abruptly
from juvenile to adult form (D. elatum and D. novo-guineense) stand
apart from the other species, with D. nausoriensis representing a somewhat
transitional position. The abrupt change from fine juvenile needles to the
more robust and very short adult leaves is in accord with the scale-leaved
species, while the still spreading orientation is the common condition for
other species. Occasional specimens of D. elatum and of D. novo-guineense
have transitional leaves abruptly marked off from the juvenile leaves and
closely resembling the adult leaves of D. nausoriensis. The bark of this
new species is virtually the same as in all other species of the group, with
large thick flakes, fibrous and brown within but with a tough smooth
surface generally well supplied with lenticels and weathering gray. The
seeds are also of the usual type showing a slight marginal keel and be-
coming a rich brown color. The pollen cones seen may not be fully grown.
5. Dacrydium pectinatum de Laubenfels, sp. nov.
Arbor ad 40 m. alta, ramosissima; cortex canus vel rufulus; folia brevia,
oblique adscendentia, patentia pectinatum, apice paulo incurva, dorsus
carinata, 2-5 mm. longa, 0.4-0.8 mm. lata (iuvenilis ad 20 mm. longa).
Strobili masculi cylindracei, terminales, 9-12 mm. longi, 2 mm. lati. Stro-
bili feminei ad apicem ramulorum, saepe ramulorum brevium; folia ad
basis parviora; folia strobilorum sub semine maturo parva, crescentes
carnosa rubra; 1-2 folia ultima fertilia. Semen 4.5 mm. longum, non
tegens foliis strobilorum. Holotypus: Nicholson SAN 17292 (a), North
Borneo, Sandakan. Fics. 1b and 2.
The short bracts in the fertile area not even surpassing the epimatium
and not longer than the foliage leaves, distinguish this new species from
all but two others, one of which, Dacrydium elatum has distinctly smaller
pollen cones and scale-like foliage leaves abruptly marked off from the
juvenile leaves, while the second, D. cupressinum, has very elongated
microsporophylls and thick straight stubby foliage leaves. The short
spreading needles distinguish sterile specimens of D. pectinatum from
other species with which its range overlaps. Two varieties have been
recognized because of rather marked differences in leaf form.
Sa. Var. pectinatum.
Folia gracilia, linearia, acicularia, 2-5 mm. longa, ssidinedl saacianane
DistrisutioN. From Hainan through the Philippines to Billiton Is-
land, at low elevations up to 1,500 meters but mostly below 600 meters.
Several specimens are reported from sandy soils. Map 3.
Hainan. Yaichow, Liang 62041 s (NY), 62619 j (Ny), 62670 a (NY, US),
63214 s top of mt. (a, Ny, US). Hung Mo Mt., Tsang & Fung LU 18100 9 (A,
NY), LU18152 2 (a, Ny), McClure 18303 j 1,000-1,500 m. (Ny). Po-ting, How
72869 2 (a). Five Finger Mt., Chun 1367 j (A), 2089 s (a). Dai Land, Dung
Ka, Chun & Tso 4380 2,400 ft. (A, Ny). Chim Fung Mt., Lau 5283 $ (A).
290
JOURNAL OF THE ARNOLD ARBORETUM
®
ARNOLD ARBORETUM |
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Attend: 2s Hedin ee
HARWARD UNivesei* HER BA TS
E 2. Dacrydium pectinatum oS eens rar. pectinatum, photogré aph
of rt pera Nicholson SAN 172
Without loc., hae ce Chun 70144 2 2,000 ft. (A, ny, us), Liang 63693 3 (NY,
US), 65094 3 (A , Wang 33651 s (A, NY), 36532 & (a, NY), Tang 457 @ (A);
Hance 22162 j (pm). Billiton, NIFS bb32284 92 (a, L), Rossum 122 (L),
784 2 (L). Sarawak. Bako National Park ~ NE. of Kuching), Purseglove
P5066 j 400 ft. (K, L), P5553 & 350 ft. (x, ), Brunig §12073 3 120 m. (L),
S12074 s 13 , Sinclair & Kadim pita al s (A, K, L), Sing JC/59 s 300 ft.
(kK), Rashid S9546 5 | hap (L), Nicholson 1319 s 200-300 ft.
Brunig S1101 2 1,4 (K, 1).
K),
‘
* Pa’
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\Comosum { -
» °
e : - «@ x :
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se LYCOPODIOIDES
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( a TAXOIDES
Maps showing distribution of: 4, Dacrydium beccarii Parlatore (dots), D
guillauminii Buchholz, known only from New Caledonia; 5, D. xanthandrum
Xe
1969] DE LAUBENFELS, PODOCARPACEAE 303
Haviland 2070 s (x). Philippines. Leyte: Biliran, Sulit 21694 s 1,350 m. (L).
Gros: Dumaguete, Or, Herre 1150 s 4-6,000 ft. (A, ny). Mt. Canlaon, Edano
21936 j 1,860 m. (L). Mt. Marapara, Curran & Foxworthy 13612 s (L, NY, US).
Mt. Silay, Everett 4227 j (Ny, us). Without loc., Britton 343 s 1,700 m. (t).
Mrinpanao: Mt. Malingdang, Mearns & Hutchinson 4547 s (K, L, ce US),
4731 s (NY, US). Moluccas. Taliabu, Hulstijn 126 2 (L). New Guinea. Vogel-
kop, Upper Aifat Valley, Moll BW 12853 s 870 m. (L); Tamrau Ra., Van Royen
& Schram 7791 s 920 m. (K, L, LAE). ohn Mts., Gjellerup 572 s 600-1,500
m. (A, K, L). Hellwig Mts., Lorents 1698 s 2,100 m. (K, L). Wissel L., Maiare,
Eyma s.n.s (L). Norman “ I., Brass 25660 2 mt. crest (A, K, L, LAE, US). New
Britain. Mt. Tangis, Frodin N GF 26902 s 3,500-5,000 ft. (L). Solomon Is. Santa
Ysabel, Baea BSIP 2475 : well above 3,000 ft. (ridge top) (kK, L, LAE), Brass
3264 4, } 1,400 m. (A, Hex, 1). Guadalcanal, Mt. Popomansiu, Braithwaite
4810 2 (kK), Hill 9004 j *,000 ft. (K).
ILLUSTRATION. CorNER, E. J. H. Gard. Bull. Straits Settlements 10:
t. 6. 1939
The branches of this variety have a definite bushy aspect because of the
fine dense growth of needles. Several specimens with leaves more robust
than normal for the species have been included here, although their status
is a little uncertain. These include Van Steenis 8357, Brass 3264, and
Brass 25660.
12b. Var. subelatum Corner, Gard. Bull. Straits Settlements 10: 243.
1939. Type: Corner SFN 33224, Malaya, Pine Tree Hill.
Adult leaves noticeably less bushy than in the typical variety, variable in
length, 3-6 mm. long; up to three seeds in a fertile structure.
DistrIBUTION. Mixed with var. beccarii in the mossy forests and ex-
posed ridges of Malaya, from 1,200 to 2,300 meters.
Malaya. G. Bubu (Perak), Wray 3875 2 5,000 ft. (a, K). G. Tahan (Pahang),
Hanif. & Nur SFN 7994 & 5,500-7,000 ft. (K). G. Tapis (Pahang), Symington
& Kiah s.n. 2 4,600 ft. (K). Fraser Hill (Pahang), Cubitt 6519 s (x). Pine Tree
Hill (Pahang), Corner SFN 33224 s 4,200 ft. (K-isotype). G. Padang (Treng-
ganu), Moysey SFN 31072 s 4,000 ft. (K), SFN 31841 s 3,800 ft. (x).
ILLUSTRATIONS. CorNER, E. J. H. Gard. Bull. Straits Settlements 10:
t7&8
Only the shorter needles distinguish this variety from variety beccari,
and intermediates between them can be found
12c. Var. rudens de Laubenfels, var. nov.
Folia patula incurvata conferta in forma rudenti. Holotypus: Brass
27821 (A), Sudest Island. Fic. 4b.
Pilger (dots), D. comosum Corner, known only from the Malay agp D.
lycopodioides Brongniart & Gris, known only from New Caledonia; 6, Falca
tifolium falciforme (Parlatore) de Laubenfels (dots west of line), F po bieasiesme
de Laubenfels (dots east of line), F. taxoides (Brongniart & Gris) de Lauben-
fels, known only from New Caledonia
304 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
DIsTRIBUTION. New Guinea to Sudest I., from 300 to 3,000 meters in
elevation.
New Guinea. WESTERN Hatr: Mt. Goliath, de Kock 42 s 3,000 m. (L). With-
out loc., Brandenhorst 132 s (L), 133 s (L), van Romer 1233 s (1). Sudest
(Tagula) I., Brass 27821 2 500-600 m. (A-holotype; K, L, Us-isotypes), 28187
9 300 m. (A, K, L, US), 28188 j (A, L, US).
This variety with incurved leaves forming a compact and smooth rope-
like branch system contrasts strongly with the two varieties which have
spreading leaves and a ragged appearance. Otherwise var. rudens does not
differ significantly from the remaining varieties of this species.
13. Dacrydium xanthandrum Pilger, Bot. Jahrb. 69: 252. 1938. Lec-
totype: Clemens 4504, New Guinea, Morobe District.
Tree to 30 m. high, sometimes stunted on ridges; densely branched;
bark chocolate brown or reddish, peeling in thick flakes, bearing lenticels;
leaves spreading obliquely, slightly incurved, linear-lanceolate, generally
wider than thick, keeled on the back, acute, 6-10 mm. long, or longer on
vigorous branches and when juvenile, 0.6-0.8 mm. wide, not crowded;
pollen cones lateral or terminal and subtended by several reduced leaves,
oval to cylindrical, 5-13 mm. long; microsporophylls narrowly triangular
to lanceolate, acute, 2—2.5 mm. long; seed cones terminal, often on very
short branches, fertile bracts in the form of reduced leaves; seeds rich
tan, 2-angled, 5 mm. long, more or less protruding when mature, Fic. 5.
DistripuTion. The island of Borneo and the Philippines to the Solo-
mons, in the mountains from 1,000 to 2,400 meters, rarely down to 500
meters above sea level. Map 5.
Sumatra. Road from coast to Tapanuli (Toba L.), Bangham 1070 2 4,100-
4,500 ft. (A, K, NY). Sarawak. Mt. Luiga, Beccari 3948 6 (Fr). Baram, Ander-
son 4545 2 4,800-7,000 ft. (k, L). G. Mulu, Hotta 14597 8 1,200-1,600 m. (L).
orth Borneo. Kinabalu, Nicholson SAN 17827 2 8,800 ft. (BRI, K, L), Clemens
32502 s 6,000 ft. (A, K, L, NY), 34341 2 5-6,000 ft. (A, K, L, NY). Ranau, Nichol-
son SAN 39768 2 8,000 ft. (k), Meijer SAN 29153 s 7,000 ft. (kK, L). Tambu-
nan, Mikil SAN 32086 8 montane (x, L). Penampang, Clemente 5980 s 5,000
ft. (k, L), Leano-Castro 5985 s (kK, L). Mt. Alab, Keith 5965 j 6,000 ft. (K, L).
. Borneo. B. Raja, Winkler 1037 & 1,600 m. (L). Philippines. Mt. Umingan
(Nueva Ecija), Luzon, Ramos & Edafo 26510 2 (a, K, us). Mt. Halcon, Min-
doro, Rabor 20485 &, j 1,600 m. (t), Edafio 3265 s 780 m. (A), Merrill 5714 s
(us), 5789 j (NY, Us). Calapan, Mindoro, Vidal 3910 2 (a, K). New Guinea.
Cycloop Mts., Karstel BW 5440 s 510 m. (L, LAE). Sepik region, Ledermann
9395 s (L). Chimbu, Cavenaugh NGF 3334 j (a). Morobe District, Ogeramnang,
Clemens 4504 8 (a-lectotype; z-isotype), 5390 2 5,900 ft. (A-syntype), 6398
5,850 ft. (a-syntype), 6408 s 5,850 ft. (A), 6488 s 4,500 ft. (a). Bougainville Is.
Kajewski 1694 & 950 m. (A, BRI), 1709 2 1,000 m. (A, BRI, L). Solomon Is.
Guadalcanal, Walker BSIP 247 2 1,500 ft. (A, BRI, K, L), Kajewski 2652 s 1,200
m. (A, BRI, L).
The range of this species overlaps that of Dacrydium beccarii with
which it is often confused, both being found for example, at Ranau and
1969 | DE LAUBENFELS, PODOCARPACEAE 305
er
‘
if
id
“wniasoguy aAONUy |
PLANTS OF NORTHEASTERN NEW GUINEA
Me. Mga M.S. Comens
wine aatee panThbs disc
hee . & Mosebe Thstnet
Ficure 5. Dacrydium xanthandrum Pilger, photograph of the lectotype,
Clemens 4504 (a).
on Mt. Kinabalu, D. xanthandrum differs in the noticeably flattened
leaves which are widely spreading and distinctly less dense. It also grows
with D. gibbsiae and its leaves strongly resemble the transitional leaves
306 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
of that species, but, not only are the adult leaves of D. gibbsiae much more
robust, the pollen cone is much larger, and the fertile shoots have unre-
duced leaves. The specimen from Sumatra cited here has much more
robust leaves similar to nearly mature leaves of D. gibbsiae and may,
with more material, be found to represent a distinct taxonomic unit.
14. Dacrydium gibbsiae Stapf, Jour. Linn. Soc. Bot. 42: 192. ¢. 4.
14. Type: Gibbs 4162, North Borneo, Mt. Kinabalu.
Dacrydium beccarii var. kinabaluense Corner, Gard. Bull. Straits Settlements
10: 244. 1939. Type: Carr SFN 26437, North Borneo, Mt. Kinabalu (not
seen; photo included in description).
Much branched tree to at least 12 m. high; juvenile leaves acicular,
12-20 mm. long, spreading but slightly incurved; mature foliage leaves
becoming wider and thicker but distinctly flattened, incurved and im-
bricate (an angular keel on the dorsal side), acute, aggregated into rope-
like shoots about 8 mm. in diameter, individual leaves 5—7 mm. long, 1.0-
1.3 mm. wide, rigid; pollen cones terminal or lateral, cylindrical, 20-25
mm. long by 5—7 mm. in diameter; microsporophyll lanceolate, 5 mm. long;
seed cone terminal, often on a very short lateral branch, formed of largely
unmodified leaf- ile structures and with one or two fertile apical leaves,
becoming reddish when mature; seeds becoming almost erect, surrounded
by but spreading apart the subtending leaves, oval and tapering slightly
towards the apex, 4.5 mm. long.
DistrIBuTION. On the slopes of Mt. Kinabalu, in serpentine soils
where it is common from 1,500 to 3,300 meters.
North Borneo. Mt. Kinabalu, Gibbs 4162 2 over 6,000 ft. (BmM-holotype; K-
isotype), 4050 j (BM), Clemens 1 10685 @ (A, GH, K), 10879 j (a), 11091 6 (A),
28542 s 11,000 ft. (K), 30922 j 45,000 ft. (a, L, Ny), 33037 2, j 5,000 ft. (A,
, a i: 40151 2 6,500 ft. (A, NY), Griswold 67 j (A), Haviland 1183 s 6,600 ft.
(x), C hew & Corner 4303 j (K), 4361 j 7,000 ft. (kK), 8024 2 (xk), Nicholson
SAN 17826 2 9,000 ft. (art, L), Meijer SAN 21097 s 5,500 ft. (x), SAN 21098
j, 5,000 ft. (k), SAN 23500 s 6,000 ft. (kK), Colenette 543 s 8,000 ft. (K). Pinosok
Plateau, Colenette 542 2 5,100 ft. (x).
ILLustRaTION. CorNeR, E. J. H. Gard. Bull. Straits Settlements 10:
t. 9. 1939, as Dacrydium beccarii var. kinabaluense.
This is one of the many distinctive endemics of Mt. Kinabalu and, like
many, is characteristically robust in form. The pollen cone is unique.
With the discovery of fertile Dacrydium xanthandrum specimens well up
on Mt. Kinabalu, many of the “juvenile” specimens may actually be that
species.
15. Dacrydium guillauminii Buchholz, Bull. Mus. Hist. Nat. Paris il.
21: 282. 1949. Type: Buchholz 1728,° New Caledonia, Riviere
des Lacs.
"In the — of this species the collection number given is 1278, clearly a
typographical e
1969} DE LAUBENFELS, PODOCARPACEAE 307
Erect bush 1-2 m. high; bark with small dark rough flakes, fibrous
brown within, surface more or less smooth at first and covered with
numerous small lenticels, developing many small cracks with age; pro-
fusely branched; leaves becoming denser and less spreading with age but
not at all reduced in size, acute, needle-like or slightly compressed, bushy
imbricate, 13-17 mm. long, 1.0 mm. wide; pollen cones terminal and
lateral, the lateral ones at the base of a terminal cone and smaller, 8—14
mm. long, tapering from the base; microsporophylls with a long lanceolate
tip from 5 mm. at the base of the pollen cone to not more than 2 mm.
long near the apex; seed cones terminal, sometimes on very short lateral
branches; bracts of the seed cone unmodified or slightly reduced leaves;
seeds up to five in a cone, subterminal, oval, wider than thick, laterally
keeled, the tip rounded with the micropyle projecting, 4.5 mm. long.
DistRIBUTION. Probably the most restricted species of the genus, found
only for a few kilometers along the Madelaine River (Riviére des Lacs)
and on the margins of Lac en Huit, from which that river flows, and only
at the very edge of the water.
New Caledonia. Riviére des Lacs, Buchholz 1728 $ (1Lt-holotype; K, P-
isotypes), de Laubenfels P341 2 (A, RSA), P341A & (A, RSA), Bernier 323 j (P),
sn. 8 (p), Sarlin 242 s (p), Déniker 205 p.p. (z), Baumann-Bodenheim & Guil-
laumin 11798 s (p, z), Hiirlimann 3471 s 146 m. (z), Bernardi 9360 s (P, 2),
Blanchon 1162 s (p). Lac en Huit, de Laubenfels P116A é (spt), P116B 2 (x,
sBT), McKee 3385 4 (A, K, P, US).
ILLUSTRATION. SARLIN, P. Bois et Foréts de la Nouvelle-Calédonie,
t. 21, 1954.
This distinctive bush, a component of the serpentine maquis, bears
strong resemblances to Dacrydium beccarii and probably represents an
endemic pedomorphic variant of that species.
16. Dacrydium comosum Corner, Gard. Bull. Straits Settlements 10:
244. 1939. Type: Corner 33222, Malaya, Pine Tree Hill.
Tree 4-12 m. high; profusely branched with an umbrella-shaped crown;
bark in small flakes; foliage branches bushy, densely leafy; leaves spread-
ing at an angle and then incurved near the base, lanceolate-pungent, dis-
tinctly flattened, 12-20 m. long and 0.7-1.3 mm. wide; juvenile leaves
up to 33 mm. long; pollen cones unknown; seed cone on a short lateral
branch, often with two seeds; seeds 4-5 mm. long.
DistriBuTION. Mossy forest on exposed ridges, from 1,200 to 2,000
meters elevation in parts of Malaya, common locally but of restricted
range.
Malaya. Pahang. Pine Tree Hill, Corner SFN 33222 s 1,500 m. (k-isotype),
Burkill & Holttum 8536 s (a, K), Melville & Landon 4814 s (x). G. Tahan,
Hanif & Nur SFN 8307 s 1,500-2,000 m. (A, K).
308 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ILLUSTRATION. Corner, E. J. H. Gard. Bull. Straits Settlements 10:
t. 10. 1939.
Like Dacrydium guillauminii, D. comosum is apparently a_pedo-
morphic variant of some other species, perhaps D. xanthandrum. The
distinctly flattened and much longer leaves set it apart from D. beccarit
which grows in the same area. The relationships between the flattened
but falcate-leaved Dacrydium species (xanthandrum, comosum, gibbsiae,
spathoides, and lycopodioides) are unclear. They may be a group with
a common origin or each may have developed separately from other stock.
It is worth noting that, where known, their juvenile leaves at inter-
mediate stages have an unflattened form. Thus the flattening, for
some at least, does not represent a continuation of the seedling flattened-
leaf condition. This is in distinct contrast to the flat and not falcately
incurved leaves in other genera of the family.
Falcatifolium de Laubenfels, gen. nov. Type species: Falcatifolium
falciforme (Parlatore) de Laubenfels.
brevissimis. Strobili feminei in ramulis brevissimis, axillares; squamula
ultima sola ovulifera; ovulum unicam inversum, epimatio involutum; se-
tandem suberectum, epimatio cristato basi breviter involucrato,
crista lateraliter prominens; strobili maturi carnosi.
This new genus was previously included as a part of Dacrydium, identi-
fied as group A by Florin (1931, pp. 256-259) because of its differences
from other members of that genus. Tengnér (1965) also discussed the
distinctions between Florin’s group A and the rest of Dacrydium. Sev-
eral basic differences justify the separation of Falcatifolium as a new
genus. The fertile structures in Falcatifolium are produced on specialized
axillary shoots whereas in Dacrydium they grow terminally on ordinary
foliage branches. The epimatium of the new genus has a pronounced
hump which projects laterally from the mature cone, in contrast with
the smaller epimatium of Dacrydium which becomes a cup-like fringe
at the base of the mature seed, not projecting at all. Very striking In
Falcatifolium are the bilaterally flattened leaves which spread out dis-
tichously, contrasting not only with the fertile shoots and basal scales
of new growth, but also with the bifacially flattened juvenile leaves which
give way rapidly to the adult form at about the second year of growth.
In Dacrydium bilaterally flattened leaves do not occur. The name Falcatt-
folium reflects the basal falcate curvature of the leaves away from the
branch. Tengnér (1965) further reports a lack of vascular fibers and pollen
differences which separate this new genus from Dacrydium. Four spectes
can be differentiated, primarily on the basis of leaf form, distributed from
Malaya to New Caledonia in moist mountain forests, where they occur
as undershrubs or small understory trees.
we
1969 | DE LAUBENFELS, PODOCARPACEAE 309
KEY TO THE SPECIES OF FALCATIFOLIUM
1. Leaves broad and flat.
2. Leaves blunt to acute, normally more than 20 mm. long and 3 mm. wide.
3. Pollen cone 20-30 mm. long by 2-3 mm. in diam.; upper edge of the
leaf normally curved upwards, leaf variable in size er a more
Chvae: DS naee, De gods SS i os See F. falciforme.
. Pollen cone 15-25 mm. long by 1.5~2.0 mm, in Ps m.; upper edge
of leaf cle even slightly curved upwards, leaf rarely as much as
5 mm. 18. F. taxoides.
2. Leaves a ieais: 12-17 mm. long by 2-3.5 mm. wide. .. 19. F. papuanum,
ke Awe pra Tere i 5 Svc a ek onc os 20. F. angustum.
Ge
17. Falcatifolium falciforme (Parlatore) de Laubenfels, comb. nov.
Podocarpus falciformis Parlatore in DC. Prodr. 16(2): 685. 1868. Lectotype:
Beccari 2437, Sarawak, Mt. Poe.
Nageia falciformis (Parl.) Kuntze, Rev. Gen
Dacrydium falciforme (Parl.) Pilger, ce ; oe 18): 45. 1903.
Tree 3-10 (rarely to 25) m. tall; bark more or less smooth, rich purple-
brown, inner bark dark reddish; leaves variable in size, on mature fruit-
ing trees from 20 to 65 mm. long and 5-7 mm. wide, smoothly curved out-
ward from near the base to the widest part (about one third of the length
from the base), then tapering and curving more or less gradually towards
the acute tip, smaller leaves which may be almost straight and probably
not fully developed, sporadically occurring along with normal leaves, nar-
rowed at the base to a short, angled petiole and then decurrent; pollen
cone axial or terminal on a short, 2-3 mm., scaly stalk, cylindrical, 20-30
mm. long and 2—3 mm. in diam.; microsporophyll small, triangular-acute;
seed cone on a short scaly shoot up to 5 mm. long, the cone made up of
about a dozen larger, acuminate scales, the apical one fertile, the whole
cone becoming fleshy on maturity; seed with a humped epimatium at the
base, oval, flattened and narrowed to a blunt apical ridge, 6 mm. long, 5
mm. wide, and 4 mm. thick.
DistripuTion. Mostly an understory tree in open rainforests from
600 to 1,650 meters in elevation, from Malaya and Luzon to Obi in the
Moluccas. Map 6
Malaya. Mengkuang, Wyatt-Smith 93115 8 5,000 ft. (kK, L, us). Batu Gajah,
Perak, Ridley 5695 8 (x). G. Tahan, Hanif & Nur — 7851 & (xk), Ridley
16026 & (kK), 16178 s (k). Pine Tree Hill, Penang, Poore 6228 s 4,300 ft. (K).
Fraser Hill, Nur 10507 s 4,000 ft. (A). Lingga Archipelago. Teysmann 169 9
(L), Hullett 5695 8 (a, BM). Sarawak. Santubong top, Beccari 2126 3 (F),
Haviland (1890) $ 2, 800 ra (Kk). Mt. Dulit, ae 1834 $ 900 m. (A, BM, K,
L), 1836 j (pM, kK). Mt. Poe, Beccari 2437 2 (rI-lectotype; A, k-isotypes),
Clemens 20238 s 6,000 ft. (Ny), 20263 s 5,000 ft. (A, Ny). Mt. Mattang, Bec-
Cari 1331 s (Ft), 1697 s (FI), 2941 2 (FI), Koley 11669 s (k). Trusan, so
S8743 s (K, L). Meruong Plateau, Brunig 59994 s 800 m. (L). Without loc. An-
derson 8365 & 2,000 ft. (K, L), Gibbs 4400 s 3,000 ft. (BM, K). Brunei. Ash-
310 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ton BRUN 1031 s 4,300 ft. (K, L), 1066 s 4,750 ft. (K, L). North Borneo. Kina-
balu, Clemens 10962 s (A, K), 27851 j 7,000 ft. (BM, K, NY), 33078 & 5,000 ft.
(A, K, L, NY), 5.%. s 4-5,000 ft. (A, L, NY), Gibbs 4067 s (kK), Chew & Corner
1863 & 5,500 ft. (K), 4847 s 5,000 ft. (x). Lahad Datu Dist. (Mt. Silam), Wood
SAN A4179 s 2,500 ft. (A, BRI, L), Meijer & Anak SAN 37497 6 2,000 ft. (kK,
L), SAN 22705 s (k). Penampang, Clemente 5995 s (kK), Leano-Castro 5986 s
(x). Ranau, Meijer SAN 20953 s (K), Anon. SAN 20279 j 4,000-4,500 ft. (1).
Borneo. Bengkajang, NJFS bb9664 s 1,400 m. (L), bb24778 s 1,200 m. (A, L),
bb25157 s 1,100 m. (L). G. Damus, Hallier 506 s (L). Mt. Palimasan, Koster-
mans 12779 s 500 m. (L). Lianggagang, Hallier 2688 s (L). Philippines. Mrn-
DANAO: Davao, De La Cruz 27746 j (us). Minporo: Mt. Halcon, Merritt 4425
2 (F, us), Merrill 5744 s (kK, L, Ny, US), Rabor 20482 s 1,600 m. (L). Without
loc. Whitehead (1896) s, j (BM). Luzon: Mt. Umingan, Nueva Ecija, Ramos
& Edaiio 26394 & (a, ny, us). Mt. Camatis, Tayabas, Edaio 4508 2 (A).
Celebes. Manado, Eyma 3671 j (Lt), NIFS bb17544 s 1,400 m. (A, L), 0b21294
2 1,200 m. (L), bb24778 s (a). Obi. de Haan bb23815 j 700 m. (1).
ILLUSTRATIONS. PiicER, R. Pflanzenreich IV. 5 (Heft 18): fig. 4 D-G.
1903; Nat. Pflanzenfam. ed. 13: fig. 227 D-G. 1926; Gress, L. S. Jour.
Linn. Soc. Bot. 42: ¢. 8. 1914, all as Dacrydium falciforme.
Shape of pollen cone and mature leaf size and shape distinguish Falcati-
folium falciforme from other species in the genus. In contrast, F. taxoides
has a more slender pollen cone and mature foliage leaves with only
sporadically the slightest upward curve of the upper leaf margin, while
in F, falciforme such a curve is normally pronounced and only sporadical-
ly absent. The mature leaf size of F. papuanum is completely below the
great size range of F. falciforme, differing also in a straight profile and
apiculate tip. The larger, probably deep-shade-grown leaves of F. falct-
forme with the sweeping curve of their upper part are attractive and
quite unique, paralleled only in F. angustum whose leaves are quite narrow.
18. Falcatifolium taxoides (Brongn. & Gris) de Laubenfels, comb. nov.
Dacrydium taxoides Brongn. & Gris, Ann. Sci. Nat. Paris V. 6: 245. 1866.
Lectotype: Vieillard 1259 p.p. New Caledonia, Balade.
Podocarpus taxodioides Carriere, Traité Conif. 2: 657. 1867. Type: Vieil-
lard 1259 p.p. New Caledonia, Wagap.
Podocarpus taxodioides var. gracilis Carriére, ibid. 658. Type: Vieillard 1259
p.p. New Caledonia, Balade.
Nageia taxoides (Brongn. & Gris) Kuntze, Rev. Gen. Pl. 800. 1891; as N.
taxodes.
Bush or small tree from 2 to perhaps 15 m. high, bark thin, more OF
less smooth, scattered with lenticels, light reddish brown and fibrous
within, occasionally breaking off a flake; loosely branched; juvenile
leaves bifacially flattened, long ovate, almost linear, tapering to a sharp
tip, keeled on the lower surface, 15-20 mm. long and 1.5 mm. wide; ma-
ture foliage leaves somewhat variable, smoothly curved outward at the
base and expanding to the greatest width at about one third their length,
then tapering slightly toward the rounded or acute apex, sometimes al-
aw
1969 | DE LAUBENFELS, PODOCARPACEAE 311
most linear, the tip usually straight and pointing directly outward or
occasionally bent slightly towards the branch apex without a corres-
ponding bend in the upper leaf edge (or rarely a slight curve), more or
less narrowed at the base to a petiole and then decurrent; pollen cone
axillary or terminal, often with several on a short axillary branch with
minute scales, cylindrical, 15-25 mm. long and 1.5—-2.0 mm. in diam.;
microsporophyll with a minute acuminate tip; seed cone on a slender
scaly branch up to 6 mm. long, the cone with about a dozen larger
elongated scales up to 2 mm. long, the apical one fertile, the whole cone
becoming fleshy on maturity; seed with a humped epimatium at the base,
oval, strongly keeled on the sides with an elongated blunt tip, 7 mm, long,
4 mm. wide, and 3 mm. thick.
DIsTRIBUTION. In moist rainforests (but not mossy forests) as an
understory shrub or small tree throughout New Caledonia wherever these
conditions occur, which is most commonly in the 800 to 1,200 meter
range but occasionally reaching almost to sea level and to at least 1,400
meters.
New Caledonia. Balade, Vieillard 1259 p.p. s (p-lectotype of Dacrydium
taxoides and holotype of Podocarpus taxodioides var. gracilis). Ignambi, Comp-
ton 1571 9 3,500 ft. (Bm). Upper Diahot, Hiirlimann 1887 3 (p, z). Mt. Col-
nett, Hiirlimann 1965 & (p, z). Tao, Baumann-Bodenheim 15881 s (P, Z).
P), Thorne 28705 s (Pp), Baumann-Bodenheim 5654A s (P, Z), 15680 4 (Pp, 2),
Baumann-Bodenheim & Guillaumin 11259 s (P, Z), 11262 s (P, z), 11286 s (P, Z),
11287 s (Pp, z), 11292 s (P,z), 11296 s (P,z), Blanchon 340 s (P). Mt. Dzumac, Bar-
ets 7 s (Pp), Blanchon 1247 s 700-900 m. (Pp). Dumbea, Sunshine Mine, a
1587 s 650 m. (Pp, z), 1609 s (Pp, z). Mt. Koghis (Mone, Bebo), Pancher 379
(p), Blanchon 566 s 300 m. (P). Upper R. Bleue, Bernier 301 s (P). agrees
Bodenheim 15021 2 (p, z), de Laubenfels P400 3 800 m. (RSA, SBT), / on
ville & Heine 187 s, j (ve), Bernardi 9404 s (P, z). Upper Mois de Mai, a 2
1390 s (ILL, P). NE. of Lac Naoué, Hirlimann 3180 s 500 m. (Z). Bois : ud,
Baumann-Bodenheim 12492 s (P, Z), 14996 @ (P, z). Upper Kuébini, Hur a
3542 2 265 m. (z), 3543 s (z). Without loc., Balansa 184 & (Pp), Deplanche
169 s (Pp), Mueller s.n. s (p), Sarlin 229 s (e), Baudouin 387 s (P).
312 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ILLUSTRATIONS. BroNcGNIART & Gris, Nouv. Arch. Mus. Hist. Nat.
Paris 4: t. 3. 1868, as Dacrydium taxoides; Pircer, R. Pflanzenreich
IV. 5 (Heft 18): fig. 4 H-L. 1903, as Dacrydium falciforme; Nat. Pflan-
zenfam, ed. 2. 13: fig. 227 H-L. 1926, as Dacrydium taxoides; SARLIN,
P. Bois et Foréts de la Nouvelle-Calédonie, t. 19. 1954, as Dacrydium
taxotdes.
From Falcatifolium falciforme this species differs in its smaller leaves
and pollen cones and in the straight rather than upwardly curved leaf
tips. From F. papuanum it differs in lacking a pungent leaf tip and hav-
ing distinctly larger leaves. These two species and F. taxoides are clearly
quite closely related, being geographic segregates. F. taxoides is sometimes
the host to another conifer as a root parasite (de Laubenfels 1959).
19. Falcatifolium papuanum de Laubenfels, sp. nov.
Arbusculus vel arbor ad 22 m. altus; folia patentia, ad apex apiculata,
linearia vel ovato-linearia, 12-17 mm. longa, 2—3.5 mm. lata. Strobili
masculi ignoti; strobili feminei cum ramulis brevissimis, squamis lanceo-
latis, 1.0-1.5 mm. longis, bracteis strobilorum ca. 2 mm. longis; semen
lateraliter et terminaliter carinatum, 6 mm. longum, 4.5 mm. latum, 3
mm. crassum. Holotypus: de Laubenfels P483 (a), New Guinea, Morobe
District. Fic. 6a, b.
DistRIBuTION. In moist rainforests as an understory plant in the east-
ern part of the island of New Guinea (possibly in the Vogelkop), from
2,000 to 2,400 meters in elevation. Map 6.
New Guinea. VocELKop: Mt. Nettoti, Van Royen & Sleumer 8203a j 1,920
m. (L), Terr. New Guinea: Al R. Mts., Womersley NGF 5354 s 7,000 ft. (4,
BRI, K, L). Mt. Hagan Sta., Hoogland & Pullen 5891 2, j 7,600 ft. (A, BRI, i.
t. Kum, Womersley NGF 9419 s 7,000 ft. (BRI, K, L). Nondugl, Womersley
NGF 4483 s 7,000 ft. (a, K, L). Morobe Dist., Edie Creek (Mt. Kaindi), de
Laubenfels P483 2 6,500 ft. (A-holotype; K, RSA, SBT-isotypes), Brass 29127
s, 7,200 ft. (L), Womersley NGF 11038 s 6,700 ft. (BRI, K, L), NGF 13922 g
7,200 ft. (Kk, L). Papua: Mt. Tafa, Cent. Div., Brass 5107 s 8,000 ft. (BRI, NY).
Ridge betw. Adai and Turui Rivers, Lane-Poole 397 s (A, K).
The apiculate and somewhat small leaves, whose mature size is com-
pletely below the considerable range of both Falcatifolium falciforme and
F. taxoides, distinguish this new species. The leaf profile is straight as
in F, taxoides, but without the rounded tip of that species. The juvenile
leaves reach 22 mm. in length and 4 mm. in width. The bark, gray to
dark brown, and flaky with large lenticels, and a red-brown inner bark,
is not unusual. A remarkable specimen from the Vogelkop, an entire
small plant of Falcatifolium, has distinctly smaller leaves, 6-10 by 2 mm.
(Fic. 6b). Inasmuch as juvenile leaves are usually distinctly larger than
those of the adult, it may be that this isolated specimen represents a
distinct entity.
20. Falcatifolium angustum de Laubenfels, sp. nov.
1969 | DE LAUBENFELS, PODOCARPACEAE 313
py Wy
HW
Seeman
=
~—
—
eae
:
-_ =
a
oe
_
a, Pee let papuanum de ecieen are portion of the holotype,
de Lawbenfel P483 slightly enlarged; b [in he same, fragment of Van
Royen & Sleumer $2030 from the Vogelkop, ew Guinea (L), enlarged.
Arbor ad 20 m. alta; folia plantarum iuvenilis acicularia, crassiora
quam lata, lanceolata, falcata, patentia, e basi curvata extrinsecus, ad
apici Curvata sursum, ca. 7 cm. longa, basem versum 1.2 mm. crassa; folia
plantarum adultarum minus curvata vel quasi recta, pungentia, carinata
314 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
¥¥
ep
me
FicurE 7. a, Falcatifolium angustum de Laubenfels, portion of the holotype,
Brunig S8866 (1); b, Dacrycarpus expansus de Laubenfels, portion of the holo-
type, Hoogland & Schodde 7463 (1): a and b. approximately natural size.
a latere, 18-35 mm. longa, 1—2.5 mm. crassa: strobili masculi terminales
vel laterales, immaturi ovati, 8 mm. longi, 2 mm. diametro; strobili
feminei ignoti. Holotypus: Brunig S8866 (1), Sarawak, Bintulu. Fic. 7a.
DIstRIBUTION. At low elevation along the coast of Sarawak.
Sarawak, Bintulu, Brunig $8860 & 300 ft. (L), 58866 8 400 ft. (t-holotype),
S963 j 500 ft. (kK, L). Kuching, Anderson 12448 s 800 ft. (K).
This distinct new species with its narrow but nevertheless bilaterally
flattened leaves is intermediate between the other species of F alcatifolium
and Dacrydium, and seems to represent an early stage of the development
of the genus. In the transition between seedling leaves and norma
foliage leaves of F. taxoides are found leaves of identical morphology to
the adult leaves here. The bark is purplish-brown, irregularly flaky to
scaly, weathering gray.
[To be continued |
VotumE 50 NuMBER 3
JOURNAL
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HARVARD UNIVERSITY
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D. A. POWELL
CIRCULATION
| fi 23 908
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52. CONTENTS OF NUMBER 3
A Revision oF THE MALESIAN AND Paciric RAINFOREST CONIFERS,
-ACEAE, IN PART (Concluded). David J. de Lau-
Sia Mastsvane AND Paiesenioke oF PrI-
iM. H. Zimmermann and P. B. Tomlinson ....
a ee AND FLOWERS IN
NA (Pazatat. Natale W. 1) Cees
315
370
411
JOURNAL
OF THE
ARNOLD ARBORETUM ._—,,:°
VoL. 50 Jury 1969 NUMBER 3
A REVISION OF THE MALESIAN AND PACIFIC RAINFOREST
CONIFERS, I. PODOCARPACEAE, IN PART *
Davin J. DE LAUBENFELS
Dacrycarpus (Endlicher) de Laubenfels, stat. nov.
Podocarpus sect. Dacrycarpus Endlicher, Syn. Conif. 221. 1847. Type species:
Podocarpus imbricatus Blume [Dacrycarpus imbricatus (Blume) de Lau-
benfels].
Podocarpus sect. Dacrydioideae Bennett ex Horsfield, PI. jav. rar. 41. 18 8.
ype species: Podocarpus dacrydioides Rich. [Dacrycarpus dacrydioides
(Rich.) de Laubenfels].
Folia parva vel squamata, Strobili feminei terminales; receptaculum
verruculosum, tandem carnosum; unus vel duo bracteae terminalae fer-
tiles, cum ovulo connatum in forma crista superans; ovulum inversum
epimatium contingens.
The distinguishing character of Dacrycarpus is the union of the bract
with the seed and seed scale on one side, forming a projecting crest
particularly noticeable on immature fruit. As in most of the family, the
seed is inverse and Dacrycarpus resembles Podocarpus, of which it has
long been treated as a section, because of the union of the fertile scale
with the seed and its distinct receptacle. In addition to the fusion of the
fertile bract with the corresponding scale and seed, however, is the fact
that the cone is produced terminally on leafy branches and not on
specialized shoots or peduncles. The leaves of Dacrycarpus also differ
markedly from Podocarpus and resemble those of Dacrydium, being
sometimes difficult to distinguish in the sterile form. There are, in addi-
tion, New Zealand species of Dacrydium in which the seed is covered by
-
* Continued from volume 50, p. 314
316 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
the leaves are distinctly bilaterally flattened, often spread out distichously,
yet this character is usually lost in the mature form. Elsewhere among
the conifers only in Falcatifolium and Acmopyle are bilaterally flattened
leaves found but without change in the adult form. Where the foliage
leaves are bilaterally flattened or acicular, sterile specimens of Dacrycar-
pus can be identified by the sharply dimorphic leaves because the penul-
timate branches always have bifacially flattened leaves or scales.
Dacrycarpus is composed of nine species ranging from Burma to New
Zealand. Some of the species are geographically isolated and the one
species in New Zealand (D. dacrydioides) is not tropical in habitat. The
various species are distinguished primarily by the shape of the involucral
leaves subtending the receptacle and the shape of the foliage leaves. Most
discussion of specific differences in the morphology of the seed-complex
simply involves degrees of maturity. Among the various species there
are several wide ranging groups. A more or less scale-leaved type in-
volves D. dacrydioides and D. imbricatus and is the most important for
lumber and afforestation. Longer acicular leaves and a relation to moist
habitats characterize D. steupii and D. vieillardii. Long involucral leaves
and a mid-mountain distribution are characteristic of D. cumingii and
D. cinctus. The remainder of the species are mostly localized and are
found at high elevations.
KEY TO THE SPECIES OF DACRYCARPUS
1. Involucral leaves spreading and not oo the seed and receptacle at all;
mature leaves not distinctly flatten
2. Leaves short to scale-like (less nee 2m
3. Pollen cones terminal, linear; ni leaves very short,
longer than the foliage leaves. ................ D. Hoong are
3. Pollen cones lateral, ovoid; involucral leaves about as long as the re-
ceptacle and longer than the foliage leaves
4. Leaves slender (0.4-0.6 mm. wide).
5. Leaves imbricate......... 21a. D. imbricatus var. imbricatus.
5. Leaves spreading. .......... 21b. D. imbricatus var. patulus.
4. Leaves robust (0. 61 .O mm. as,
eaves spreading. .......... 21c. D. imbricatus var. robustus.
6. Leaves Sib acate eRe are ee 21d. D. imbricatus var. curvulus.
2. Leaves elongate (at least 2 mm.).
7. Involucral leaves less than 2 mm. (shorter than the foliage leaves);
foliage leaves strongly variable and more or less imbricate. ....----
a eh > on Cae ens oh SEs ely 72. D. vieillardii.
7. Involucral leaves more than 3 mm. (longer than the foliage leaves) ;
foliage leaves more constant and strongly spreading. ......------*:
a Cue Np an eee ea te gt Ss gel ER Cie Pee, CoM AAA Glen EN A 23. D. steupii.
1. Involucral leaves clasping the seed and receptacle; mature foliage leaves
ttened,
8. Leaves bilaterally flattened
9. Involucral leaves long ‘(7-10 _ surpassing the mature seed; fo-
liage leaves slender. 24. D. cumingit
1969] DE LAUBENFELS, PODOCARPACEAE 317
9. Involucral leaves short (5-7 mm.), not covering mature seed; fo-
Hage leaves POuuUSt. . <5 isnds cwieoems sccee die, 25. D. kinabaluensis.
8. Leaves bifacially flattened.
10. Involucral leaves long (5-6 mm.); foliage leaves long and narrow
(2-$ mm, by GAO6 mii Ve oboe cs ac no oon bce, 26. D. cinctus.
d
wide (2-4 mm. by 0.6—1.0 mm.).
11. Pollen cone lateral; seed not large (5-6 mm. long); foliage
SOAVOR SURCACH. Ss o55 a de sk cnn wae cubes : nSUS.
- Pollen cone terminal; seed large (7-8 mm. long); foliage leaves
She eo adda osha eee eee 28. D. compactus.
—
ji
—
21. Dacrycarpus imbricatus (Blume) de Laubenfels, comb. nov.
Podocarpus imbricata Blume, Enum. Pl. Javae 1: 89. 1827. Lectotype: Blume
5.n.,. W. Java.
Podocarpus cupressina R. Br. ex Mirb. Mém. Mus. Hist, Nat. Paris 13: 75.
1825 (nomen); R. Br. ex Horsfield, Pl. Jav. Rar. 1: 35. ¢. 10. 1838. Type:
Horsfield 5.n., Java.
Podocarpus horsfieldii Wallich, Cat. No. 6049. 1832. Nomen nudum,
Nageia cupressina (R. Br.) Muell. Phyt. New Hebr. 20. 1874.
Tree up to at least 30 m. tall; bark dark brown or blackish on the
surface, weathering gray, inside a rich red-brown and granular (slightly
fibrous), breaking off in small thick scales with a rough surface; juvenile
leaves bilaterally flattened and distichous, nearly linear, curving outward
from the base and upward at the tip, narrowing rapidly to a fine mucro,
10-17 mm. long and 1.2-2.2 mm. wide, shorter toward the branch tip
and base, the first leaves at the branch base short and acicular, the whole
foliage branch of limited growth; leaves on seedlings and on penultimate
branches quite distinct, bifacially flattened, lanceolate, mucronate, im-
bricate, decurrent, 2-4 mm. long and 0.7—1.0 mm. wide; terminal shoots
On young plants sometimes very long, whip-like, up to 20 cm.; on older
plants more compact, the foliage leaves becoming progressively smaller,
fertile specimens sometimes having distichous and bilaterally flattened
leaves 3-5 mm. long and 0.6—-0.8 mm. wide; foliage leaves in older trees
eventually becoming short and needle-like or more or less scale-like,
about 1-1.8 mm, long, strongly keeled and acute but neither flattened
nor distichous; pollen cones lateral or rarely terminal, subtended by a
few scale leaves on a branchlet 1-3 mm. long, oval but elongating with
the shedding of pollen, to 6-12 mm. long and 2-2.5 mm. in diam. (about
5 mm. long before elongating); microsporophyll triangular, acute to
apiculate; seed cone terminal, often on a short lateral branch bearing
scales which become elongated just below the receptacle, forming an in-
volucre, the involucral leaves spreading and generally less than 4 mm.
long, acicular and sharply pointed; seed cone a short, warty, glaucous
receptacle 3-4 mm. long, formed of enlarged bract bases, the tips of one
Or two bracts (resembling the involucral leaves) projecting from the
receptacle, one or two terminal bracts fertile, the whole receptacle be-
318 JOURNAL OF THE ARNOLD ARBORETUM [vor. 50
coming red upon maturity; mature seed globose, slightly ribbed on the
back with a blunt crest, 4-6 mm. in diam., 5—6 mm, lon
The short acicular or scale leaves and the lateral more or less oval
pollen cone distinguish this widespread species from other members of
the genus. Longer leaves occur on fertile specimens but, if present, are
less than 5 mm. long and very robust (var. robustus) or are distichous.
Dacrycarpus imbricatus can be subdivided into four varieties on the basis
of mature foliage leaf form (Fic. 8). The reproductive structures and
immature leaves of the varieties are indistinguishable. When identifying
these varieties, care must be taken to compare the leaves of only the
ultimate foliage branchlets and not the distinct penultimate scale-cov-
ered branches (the penultimate branches of all varieties resemble the
foliage branches of var. imbricatus).
21a. Var. imbricatus.
Mature foliage leaves strongly appressed, slender, about 1.5 mm. long
and 0.6 mm. wide, the whole foliage branch 0.75—-1.25 mm, in diam.; in-
volucral leaves 2-4 mm. long. Fic. 8a.
DisTRIBUTION. Scattered and common in rainforests from low elevation
up to about 3,000 meters, but particularly from 700 to 2,400 meters
(agriculture has commonly destroyed the forests at low elevation); in
Java and the Lesser Sunda Islands, and occasionally in Celebes and
Borneo. Map 7.
s (NY). G. Pangranggo, Schiffner 1475 & 1,900 m. (L), Palmer & Bryant 988
j 2,900 m. (us), Winkler 1866 j 2,400 m. (x). Gegerbintang (Preanger), Den
Berger 549 2 1,100 m. (x). Mt. Tankuban Prau, Anderson 67 s (K).
Lembang, Junghuhn sn. 2 (1). Bandang, Junghuhn sm. s (L). Takokok
(Preanger), 1,150 m. Koorders 15535 @ (A, L), 27704 s (L). G. Besser, Winckel
sm. (L). G. Ungaran (Semarang), 1,000-1,350 m. Koorders 1283 s (L), 1284
j (L), 1285 j (1), 27705 @ (x, L). G. Kukusan (Lawu), Elbert 52 j 1,500-1,700
m. (L). Ngebel (Madiun), 1,450 m. Koorders 1278 s (1), 1279 j (L), 1280 ¢ (L),
1281 s, j (L), 1282 j (t), 29188 j (L), 29189 2 (a), 38626 j (L), 38652 j ().
G. Ardjuno (Pasuruan), Koorders 38187 2 2,100-2,400 m. (L). Ngadasarl,
Koorders 37922 s (t), 37923 j (K, L). Ngadiwono, La Rinere s.n. s 1,600 m-
(t). Idjen Plateau (Besuki), 1,700 m. Koorders 1290 j (L), 1296 2 (2); 1297
j (t). G. Kendeng, Koorders 28507 9 (a). G. Tapandajan, Coert 637 j 1,750
m. (L). G. Tenga (Parverua), Dugeh 1382 j 1,600 m. (L). Parverua, Oillerings
175 s (ut). G. Guntar, Anderson 429 j 4-6,000 ft. (x). E. Java, Coert 1437 $
VIEILLARDII
a Mars showing distribution of: 7, Dacrycarpus imbricatus (Blume) de Lau-
nfels, var. imbricatus and var. patulus de Laubenfels; 8, D. nape ws
tae de Laubenfels (dots north of line), and var. "curvulus (Miq
ubenfels (dots south of line); 9, D. steupii (Wasscher) de pr a
D. vieillardii known only from New Caledonia.
320 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
(L), Went s.n. j (L). Without loc. Blume 492 @ (tL), s.n. 2 (L-lectotype of
Podocarpus imbricata), Junghuhn s.n. j (L), Horsfield sn. 2 (K-holotype of
Podocarpus cupressina; GH-isotype), 108 s (K), 1166 2 (Kk), Korthals s.n, }
(L), van Hasselt s.n. s, j (L), Simmoro s.n. s 3-4,000 ft. (L), Coert 1209 s (L),
Zollinger 2262 2, j (A, 2), De Vriese sn. 2 (kK), Miquel sn. s (K). Lesser
Sunda Is. Batt: Mt. Batukan, Kostermans, Kuswata, Sugeng & Supadmo KK
& SS 138 2 1,300 m. (A, K, L), Sarip 371 j 1,930 m. (1). Buleleng, N/FS
bb17269 8 1,300 m. (A, L). Lompox: Mt. Rindjani, Elbert 2266 j 700-1,700 m.
(A, L). Lenek (mid.), NJFS bb15504 2 700 m. (K, L). Plambi (SW.), Elbert
2428 j 200-400 m. (1). SumBawa: Batu-Lanteh Mts. (N.), Elbert 4191 j 1,500-
1,700 m. (A, L, us). Frores: Rensch 1307 j (x). G. Kasterso, Posthumus 3235
j 1,800 m. (1). Sumpa: Lairondja (E.), [but 547 j (L), NIFS bb9003 j 1,000 m.
(k, L). Trmor: Nenas (mid.), NIFS 6b11803 2 1,600 m. (1). Mt. Perdido
(cent. Port.), Van Steenis 18267 8 1,600-1,750 m. (L). Without loc., Forbes
3855 8 (A, L). Sarawak. Kuching, Clemens s.n. j (Ny). Mt. Dulit, Richards
1768 j 1,300 m. (K, L). Kapit, Upper Rejang R., Clemens 21066 j (NY). Brunel.
B. Ulak, Ashton BRUN 1032 j 4,300 ft. (kK, x). B. Pagon, Ashton BRUN 1065
8 4,750 ft. (x, L). North Borneo. Ranau, Sadau 42890 2 4,920 ft. (K, L),
Sario SAN 32246 & (x), Lajangah SAN 33085 j (x). Tambunan, Mikil SAN
32070 j (K). Borneo. Sakumbang, Korthals s.n. s, j (1). B. Raja, Winkler
1035 j 1,700 m. (L). Celebes. G. Bantaeng, Biinnemeyer 11903 s 2,300 m. (K,
L), 12019 2 2,060 m. (A, L), NIFS bb5460 2 2,000 m. (L), Everett 42 j 7-
10,000 ft. (x). Roto (Masamba), NJFS bb24957 j (L). G. Kambuno (Ma-
samba), Eyma 1369 j (L). Enrekang (Rantelmo), NJFS bb29195 j 1,600 m.
, L). Upper Binuang, NJFS 6b20202 j (A, K, L, NY). Mt. Mambuliling, De-
Froidville 173 j (x). Betw. Angin-Angin and Pintealon (Enrekang), Eyma 570
j 1,550-2,600 m. (A, x, L).
ILLusTRATIONS. BENNETT, J. J., Pl. Jav. Rar., ¢. 10. 1838, as Podo-
carpus cupressina; BLuMer, K. L., Rumphia 3: t. 172 & t. 172B. 1849, as
Podocarpus cupressina; Pricer, R., Pflanzenreich IV. 5 (Heft 18): fig.
7E. 1903; Nat. Pflanzenfam. ed. 2. 13: ¢. 124E. 1926, as Podocarpus im-
bricatus; Koorprers, S. H., & Tu. VALeton, Atlas der Baumarten von
Java 3: t. 585 & 586. 1915, as Podocarpus imbricata; WASSCHER, J.,
Blumea 4: ¢. 4, fig. 2, 1941, as Podocarpus imbricata.
The variety imbricatus is well known in Java and the Lesser Sunda
Islands and is widely cultivated. Because there have been only scat-
tered collections elsewhere, the possibility of artificial introduction must
be considered. Juvenile plants can not be identified to variety in this
species so they have been assigned to whatever mature form is known
in the vicinity. Some rather large juvenile leaves appear in the col-
lections from the Lesser Sunda Islands.
21b. Var. patulus de Laubenfels, var. nov.
Podocarpus kawaii Hayata, Bull. Econ. Indochine 20: 439. 1917. Type:
Hayata in 1917, Tonkin.
Folia patula, acicularia, falcata, basi carinata, acuta, 0.8-1.5 mm. longa,
0.4-0.6 mm. lata; ramuli foliis inclusis 1-2 mm. diametro; folia involu-
1969 | DE LAUBENFELS, PODOCARPACEAE 321
tae
fii)
i i
obidudatedes
dlitiliidwulia Hitliuai
wily
ijubudlinda
9
stout
nln vata
uth
it
IGURE 8. a, iia iis eg ml (Blume) de Laubenfels var. chigtonn
fragmenta showing mature foliage form; b var. oe “yon de Lauben a
of holotype de Laubenfels P- ma hg ete < 0.9 _ var. robustus de ” jc
fels, fragments showing mature foliage d, var. Curt — ee ) de
Laubenfels. fragments, ane ace nea form, c and x 0.8
=
a.
cralia 1-3 mm. longa. Holotypus: de Laubenfels P328 (A), Fiji Nandari-
vatu. Fic. 8b
DistriButTion. Scattered and common in rainforests from low eleva-
tion up to 2,500 meters, particularly from 700 to 1,700 meters and lower
where moist forests occur: from Upper Burma to Fiji, particularly from
South China to Sumatra, otherwise apparently in a more or less discon-
322 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
tinuous distribution overlapping with other varieties of the species and
in isolated populations east of New Guinea. Map
Burma. Hukong Valley, Hole 21 s (kK). Serpentine Mines (S. of Hukong
Valley), em 5007 j 1,600-2,600 ft. (cH, K). Tampyu (Kachin), Thompson
(1896) ¢ (xk). Northern Triangle, Arahku, Kingdon-Ward 20626 j (A, BM),
21295 j (A, BM), 21393 s 4-5,000 ft. (A, BM), 21626 j (A, BM). Thailand. Nakhan
Rachasima, Phengkhlai 568 j (K). Pulom Lo, Dan Sai, Kerr 5788 2 1,000 m.
(BM, K). Kao Kuap, Kerr 17715 s 500 m. (B BM, K). Kao Soi Dao, Trang, Kerr
19435 j 500 m. (kK). Botong, Pattani, Kerr 7648 6 600 m. (K). Laos. Betw.
Dasia and Cateng, Fania Prov., Poilane 16092 @ (a). Tram-la, Tranninh
Prov., Poilane 2147 j (A, K, L). Boloven near Attopeu, Poilane 15922 2 (NY).
Without loc., ace 1932) s (L). Cambodia. Kuang Repoe, Opong Prov.,
Pierre 5528 s (in mts.) (A, K). Sckral Mts., Samrongtong Prov., Pierre
5528 PP. j a x, NY). Phnom Penh Forest, Bejaud 718 3 (ILL). Elephant
Podocarpus kawaii). Annam QUANG Trr PROV.: Dent de Tigre, Poilane 10293
Q@ (A, K, L). Bach Ma (N. of Da Nang) Poilane 29960 2 (ILL). Dong-tri Mas-
sif, Poilane 10995 j (a, L). Dong-co-pah Massif, Poilane 11110 2 (A, L, NY).
Without loc. Poilane 13644 2 (a, K). SOUTH: ‘Near Dakto, Pozlane 35595 j
(1), Dalat (Lang Bian Massif), Evrard 1779 @ (a, NY), 238 j (A, NY), Che-
valier 30027 j 1,400 m. (A), Poilane 4038 j (A). Nonh-hoa (near Nhatrang),
Poilane 6509 2 (a, NY). Nhatrang, Poilane 3387 j (A, K), 9103 j 500-1,500 m.
(ILL). Chapu, Petelot sm. 2 1,500 m. (NY). Without loc. Delacour & Low
(1927) 2 (Bm), Kloss s.n. s 5,200 ft. (pM), Vim sn. @ 1,500 m. (us). China.
Kwangtung-Tonkin border, Tsang 27332 j (A, K). Chen Pien Dist, Kwangsi,
Ko 55900 j (a). Tsin Hung Shan, N. Him Yen, Ching 7034 j 4,000 ft. (A, NY,
us). Kwangsi, Wang 39608 2 (a). Hainan. Fan Ya (5 Finger Mt.), Chun
& Tso 44250 s 4,000 ft. (a, 1, Ny), McClure 8705 2 700-1,000 m. (A, BM).
Seven Finger Mts., Liang 61783 j (a, Ny). Dung Ka, Chun & Tso 43955 &,
j 2,400 ft. (A, Ny). Kan-en Dist., Lau 3556 j (A). Without lo oc. McClure 18304
j 1,000-1,500 m. (ny), 18279 9 (xy), How 72870 2 (Bm), Chun 1390 2 (A);
Liang 65187 j (A, NY), 65257 j near summit (A, NY, us), Tang 438 j (a), Wang
35591 2 (NY, US). Malaya. Kedah Peak, Low 28 j (K), Kochumen 70988 j
3,200 ft. (Kk, L). Penang, Curtis s.n. j (us). Gov't Hill, Penang, Maingay 2239
s (Kk). G. Batu Pateh, Perak, Wray 1198 2 (K). G. Benom, Pahang, Whitmore
3268 2 (K); ahs Telom, Strugnell 23931 j 2,800 ft. (A); Kluang Terbang,
Barnes 10907 2 (x). Selanger, Pahang Track, Ridley 8636 s 1,500 ft. (A).
Fraser Hill, Deris 22563 2 (xk). Batang Padang, Selangor, Murdoch 11964 j
K). B. Etam, Selangor, Kekall 19814 s (K). Malacca, Mt. Tapah, Werner
13509 @ 1,000-1,600 m. (x). Karoland, Sigurunggurung, NJFS 6b5443 g
1,500 m. (L). Karoland, Tongkoh, NJFS bb28147 s (A, K, L). Karoland, NIFS
bb27 68 9 1,400 m. (L), bb7708 @ (x). Simelungun, Yates 2148 j (t, Ny), Esche
bb35321 s 1,200 m. (1), Lérzing 11508 j 400 m. (L); Marehat Huta, N/FS
664866 j 700 m. (t). re Yates 1987 j (t). Mt. Singalan, Upper Padang,
Beccari 49 j 2,000 m. (Ft, K, L), Schiffner 1473 j 1,700 m. (L), 1474 s 2,500 m.
(L), Ernst 851 j (z). Solok, NIFS 6b4130 j 1,000 m, (x). Kerintji Indrapura,
1969 | DE LAUBENFELS, PODOCARPACEAE 323
NIFS bb18752 j 1,200 m. (A, L). Siolok Daras (G. Kerintji), Robinson & Kloss
S.m. $ 3,000 ft. (kK). G. Tudjuh (G. Kerintji), Meijer 6584 é 1,500-1,700 m.
(L), 7267 j 1,500-2,000 m. (L), Jacobs 4483 j 2,000-2,200 m. (k, L). Taram, R.
Tjampo, Meijer 6780 j 500-1,000 m. (L). Bengkulen, Redjang, Paja Magelang,
Renwarin bb2436 $ (1). Kriu, Waimengaku, N/FS 6b8737 j 950 m. (L). Beng-
kulen, G. Pesagi, Rappard P19 j 1,700 m. (a, 1). G. Pesagi, Liwa, De Voogd
119 s 1,800 m. (L), 134 j 1,700 m. (L). Lae Pondom, Surbeck 532 j 1,600-1,800
m. (L). Leaukavear, Balten Pooll s.n. s 1,630 m. (L). Palembang, Seminung,
Rappard S28 s 1,800 m. (a). Philippines. MrnpANAo: Mt. Katanglad, Bukidnon
Prov., Sulit 9896 2 1,800 m. (A, L). Lanao Prov., Alvarez 25176 j (A, us). Mt.
Malindang, Mizamis Prov., Mearns & Hutchinson 4666 & (K, L, NY, US). Mt.
Batangan, Warburg 14721 j (NY). Celebes. Pamula dama, B. Koroué (Masam-
ba), NIFS 6b24951 2 2,000 m. (a, L). Ululu (Masamba), N/FS 6b24956 s
1,700 m. (A, L). Palu, Wuka Tampai Mt. (Masamba), NJFS 6b15155 s 2,500
m. (L). Porehu (Malili), NJFS bb19559 8 1,200 m. (A, L). Moluccas, Buru,
Fakal, Toxopeus 485 j 1,100 m. (L). Morotai, Kostermans 1215 j 1,000 m. (a).
Middle Ceram, Stresemann 158 j 1,000 m. (tL), 354 8 1,450 m. (t), 363 j 1,100
m. (L). Guinea, Cycloop Mts., Karstel BW 5441 $ 510m. (L). Terr. New
Guinea, E. Highlands, Osaka, Womersley NGF 24928 2 4,000 ft. (LAE). New
Britain. Mt. Tangis, Frodin NGF 26889 j 3,000-4,500 ft. (L). New Hebrides.
Erromanga, Corbasson 18123 j 200 m. (Pp). Aneityum, Kajewski 849 Q 500 ft.
(A, NY, US, z). Fiji. Nandarivatu, Gibbs 775A & Bs (BM), Smith 4901 j 800-
900 m. (L, us), 6245 2 850-970 m. (a, L, US), Degener 14315 2 (ny, us). Gil-
lespie 4263 } 900 m. (ny), Lam 6876 8 850 m. (t), Vaughn 3258 s (BM), de
Laubenfels P328 2 2,000 ft. (A-holotype of Dacrycarpus imbricatus var. patu-
lus; K, RSA, SBT-isotypes)/ P331 j (A, RSA). Nausori Highlands, de Laubenfels
P306 j 1,900 ft. (A, RSA). Namboutini, de Laubenfels P310 j 1,000 ft. (a, RSA).
From its distribution, one might guess that Dacrycarpus imbricatus
variety patulus is the primitive representative of the species which has
been largely displaced over much of its range by other varieties, but sur-
vives alone both on the western and the eastern parts of the range.
Specimens from Fiji, when compared with specimens from Sumatra and
southeast Asia, can not be distinguished. In the Philippines, Celebes, and
New Guinea where overlap with other varieties occurs, specimens are
difficult to identify because juvenile and transitional stages are indis-
tinguishable and all too often are all that is collected. In the Philippines
forms transitional to var. robustus apparently occur, while in Borneo and
Celebes the very few mature specimens seem transitional between var.
imbricatus and var. patulus. Several specimens from lower elevations in
New Guinea do not have a robust form and have been referred to the var.
patulus,
21c. Var. robustus de Laubenfels, var. nov.
Podocarpus papuanus Ridley, Trans. Linn. Soc. London. II. 9: 158. 1916.
Syntypes: Kloss in 1913, New Guinea, Mt. Carstensz and Giulianetti &
English in 1897,8 Wharton Range.
Podocar pus leptophylla Wasscher, Blumea 4: 414. 1941. Type: De Kock 39,
New Guinea, Mt. Goliath (not seen).
* Ridley refers only to Giulianetti.
324 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Folia brevia, patula, acuta, ad apicem incurvata, fortiter carinata, ro-
busta, 1.2-1.8 mm. longa, 0.6-0.8 mm. lata, ramuli foliis inclusis 1.5—2.5
mm. diametro; folia involucralia 2-3 mm. longa. Holotypus: Brass 30568
(A), New Guinea, Mt. Wilhelm. Fic. 8c.
DIsTRIBUTION. Scattered and common in moist rainforests from near
sea level to 3,300 meters, but mostly 1,000 to 2,700 meters from North
Borneo and the Philippines to the eastern end of New Guinea. Map 8.
Sarawak. Mt. Poe, Beccari 2431 j 3,000 m. (FI), Clemens 20134 j summit (A,
Ny). Mt. Mah, Beccari 2812 2 (FI, K). North Borneo. Penampang, Clemente
5981 j 5,000 ft. (A, K, L), 6216 s (K), Leano-Castro 5988 j (K, L), 5991 j 3,500
ft. (kK, L). Tenompok Pass (Kinabalu), Smythies S10601 2 4,500 ft. (K, L),
Clemens 28631 8 5,000 ft. (A, ILL, K, L, NY), 29779 j (A, K, L, NY), Melegrito
A471 j 4,700 ft. (K, L). Pentaran Basin (Kinabalu), Clemens 33618 9 8,000
ft. (A, K, L, NY). Masilan R. (Kinabalu), Clemens 51635 2 8,000 ft. (A, K, L).
Mt. Gedeh (Kinabalu), Clemens 30371 j 6—9,000 ft. (Ny). Kinabalu, Colenette
579 s 8,000 ft. (kK), Clemens 28954 j 8,000 ft. (Bm, K), Chew & Corner RSNB
4084 j (x). Tiong Pass, Keith 5930 6 5,500 ft. (K, L), 5967 j 5,300 ft. (K, L).
Philippines. Luzon: Mt. Santo Thomas (Benguet), Elmer 6550 2 (K, NY, us),
6551 Q (kK, Ny, us), Williams 1298 2 (cH, K, NY, US), 1299 2 (Ny). Panai
(Benguet), Mearns 4405 2 7,000 ft. (x, us), Santos 31817 2 (a, us), Gillis
27255 @ (a, us), Sulit 7586 & (srt). Mt. Osdung (Benguet), Quisumbing &
Sulit 82481 j (Ny). Benguet Dist., Leafio 20673 s (us), 20674 j (us). Lepanto
Dist., Curran 10960 s (us), Darling 14498 j (L), Vidal 1818 s (K). Mt. Data
(Lepanto), Alcasid 1847 $ (1), 1897 j (L), Merrill 4503 j (K, NY, US), 4546 J
(K, L), Stern 2242 j 7,050 ft. (tL), Stern & Rojo 2289 j 7-8,000 ft. (ILL), 2292
j (ILL). S. of Bontoc, Walker 7526 j 6,000 ft. (us). Mt. Banahao, Barthe (1857)
s (A). Mrnporo: Merrit 8529 j (k, Ny, us). Mrnpanao: Mt. McKinley
(Davao), Kanehira 2652 j (Ny), 2726 s (xy). Tupi, Mt. Matutum (Cotabato),
Sumajit (1966) 2 293 ft. (L). New Guinea. VoceLKop: Nettoti Ra., Versteegh
W 10411 4 1,700 m. (L), Van Royen & Sleumer 7948A j 2,100 m. (L). Kebar
Valley, Van Royen 3895 j 1,750 m. (L). Anggi Lakes, Gibbs 5540 7-9,000 ft.
(kK), Versteegh BW 250 2 2,000 m. (a, L), Kostermanns 2197 s 2,000 m. (t),
Stefels BW 2014 j 1,860 m. (1), BW 2006 j 1,875 m. (L). Arfak Mts., Hatam,
2,100 m. (kK, L). Cycloop Mts., Versteegh & Koster BW 14 s 750 m. (A, K, L)-
Terr. New GumneA: 12 miles N. of Wabag, Womersley NGF 11260 @ 7,000
ft. (K, L), 11067 j 7-8,000 ft. (L, Nsw). Wabag, Saunders 1048 j 7,100 ft. (Z,
LAE). Tambul (Mt. Hagen), Womersley NGF 14253 j 8,000 it. (L). Mt. Kum
(Mt. Hagen), Womersley NGF 9430 s 7,000 ft. (a, L, Nsw). Wankl (Mt. Ha-
1969] DE LAUBENFELS, PODOCARPACEAE 325
gen), Hoogland & Pullen 5868 s, ] 2,300 m., (A, K, L, Us). Mt. Hagen, Cava-
naugh NGF 3322 j (a, kK). L. Inim, Flenley ANU 2176 s 8,300 ft. (K, L). Al
R. Mts. (Nondugl), Womersley NGF 5338 j 7,000 ft. (A, K, L, Nsw), NGF 5353
j (A, K, L). Waimambuno (Chimbu), Saunders, 823 j 9,000 ft. (A, L). Mt. Wil-
helm, Brass 30568 2 2,650 m. (A-holotype of Dacrycarpus imbricatus var.
robustus; K, L, NY, USs-isotypes), 30570 j (x, L, NY, US). Chimbu, Cavanaugh
NGF 3332 j (A, K, L), Stauffer 5652 j 2,600 m. (K, L, z). Fatima R., Marafunga-
Chimbu Div. (Goroka), Womersley NGF 24563 s 7,700 ft. (K, L). Marafunga,
Upper Asaro Valley (Goroka), Womersley & Sleumer NGF 14013 2 8,200 ft.
(K, L), Anden JARA 7 s 8,300 ft. (K). Danlo (Goroka), Saunders 861 j 8,500
ft. (L), 865 (L). Above Goroka, Womersley & Floyd NGF 6138 & 8,300 ft.
(A, K, L). Purosa (Okapa), Brass 31660 j 1,950 m. (a, L, NY, US), 31852 & (a,
K, L, NY, US). Wagau, Sayers NGF 21613 j 4,500 ft. (L). Samanzing, Clemens
3323 2 4,600 ft. (a, z), 5473 j (a), 8848 j (A). Sarawaket, Clemens 5586 2
7,000 ft. (A). Mt. Rawlinson, Hoogland & Craven 9553 2 6,000 ft. (kK), 9354
} (K), 9355 j (kK). Wau, Womersley & Millar NGF 8324 s 5,500 ft. (A, L, NSW),
Mt. Kaindi (Edie Creek), McVeagh NGF 7581 & 5,850 ft. (A, K, L, NSW),
Womersley & de Laubenfels NGF 19460 (P485) 2 7,500 ft. (A, K, L, RSA, SBT),
de Laubenfels P482 j 6,500 ft. (A, K, RSA, SBT), Brass 29577 s 2,060 m. (L, US),
29598 s 2,250 m. (A, L, NY, US). 29599 s (A, L, NY, US), Havel & Nauari NGF
17134 s 7,300 ft. (kK, L). Morobe Dist., Anon. NGF 3128 j (L). Papua: Anga
Valley near Ebenda (S. Highlands), Schodde 1561 s 6,500 ft. (K, tL). Alola,
Carr 14194 2 6,000 ft. (A, L, Ny). Boridi, Carr 13264 j 4,700 ft. (a, Ny). Mt.
Mau, Crutwell 897 j (kK). Murray Pass, Brass 4768 j 2,840 m. (NY). Mt.
Scratchley, Giulianetti (1896) s 12,200 ft. (K). Wharton Ra., Giulianetti &
English (1897) @ 11,000 ft. (K-syntype of Podocarpus papuanus). Mt. Tafa,
rass 4962 s 2,400 m. (A, NY), 5115 j (Ny). Owen Stanley Ra., Lane-Poole 264
j 5,000 ft. (A). Sibium Ra., Pullen 5914 j 2,650 ft. (a, L), 5930 8 3,520 ft. (a,
K, L). Mt. Dayman (Milne Bay), Brass 22582 Q 2,000 m. (A, K, L), 23393 3
1,700 m. (A, K, L, us).
ItLustrations. Grpss, L. S., Contrib. Phytogeography and Flora of
the Arfak Mountains, ¢. 4. 1917, as Podocarpus papuanus; WASSCHER, J.,
Blumea 4: t. 4, fig. 3. 1941, as Podocarpus papuana.
There has been a great deal of difficulty in separating this variety,
when treated as a species (Podocarpus papuanus), from the type
(Podocarpus imbricatus) (Gibbs, 1917; Wasscher, 1941). When speci-
mens of fully mature forms are placed side by side they are definitely dis-
tinct, but the various juvenile and transitional forms so often met with
can not be distinguished and have been greatly confused in the her-
baria. Inasmuch as the reproductive structures are essentially identical,
it seems best to maintain it in varietal status. Certainly where var. ro-
bustus occurs, other varieties are usually rare or absent. A few mature
specimens in the Philippines are more or less intermediate between
varieties robustus and patulus, including Leano 20673 and (see var.
patulus) Mearns & Hutchinson 4666. Perhaps these two varieties tend
to merge in the Philippines. Certainly typical var. robustus specimens
have come from Borneo. Variety robustus differs most from the typical
variety, imbricatus. From var. curvulus it differs in the same way that
var. patulus differs from var. imbricatus. From var. patulus it differs in
326 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
the same way (except for habit) that var. curvulus differs from var. imbri-
catus. The status of Podocarpus leptophylla is uncertain as I have not
seen the type. From its description it appears to belong to this species
but perhaps not to this variety.
21d. Var. curvulus (Miquel) de Laubenfels, comb. nov.
Podocarpus cupressina var. curvula Miquel, Pl. Junghuhn. 1: 4. 1851. Lecto-
type: Junghuhn s.n.,
Podocarpus imbricata var. curvula (Miquel) Wasscher, Blumea 4: 398. 1941.
Mature foliage srs strongly adpressed, robust, ca. 1.2—-2.0 mm. long
and 0.8-1.0 mm. wide, the whole foliage branch 1-1.25 mm. in diam.
and drooping; eine leaves 2.5-4.5 mm. long, more or less clasping
the receptacle. Fic. 8d.
DistripuTION. Generally on mountain ridges, often in solid stands and
sometimes dwarfed or procumbent, from 1,350 to 3,300 meters in eleva-
tion but mostly above 2,000 meters, from Sumatra and Java. Map 8.
Sumatra. Atjeh, Gajoland, Van Steenis 8423 2 2,100-2,250 m. (a, K, L) and
8 (K, L, Nsw). Java. Mt. Gedeh (Pengalengan), Junghuhn s.n. 8 4-~7,000 ft.
(L-syntype). Dieng Mts., Mt. Prahu, Junghuhn s.n. 8 5-7,000 ft. Ae
Kedec, Wonosobo, Zwart O5te. 3 (t). Without loc., Junghukn sn. @ (L), %
(Ny), 4 j (L), Blume s.n. Q (1).
ILLUSTRATION, WasscHER, J., Blumea 4: ¢. 4, fig. 28. 1941, as Podo-
carpus imbricata var. curvula.
The most striking character of var. curvulus is its weeping habit, but
herbarium specimens can be readily distinguished by their robust branches
with adpressed scale leaves.
22. Dacrycarpus vieillardii (Parlatore) de Laubenfels, comb. nov.
Podocarpus taxodioides var. tenuifolia Carriére, Traité Conif. 2: 658. 1867.
Type: Vieillard 1260, New Caledonia, Paita (juvenile form).
Nevada elatum Wallich var. compactum Carriére, ibid. 693. Type: Vieil-
lard 1262, New Caledonia, Paita
Dacrydium elatum Wallich var. tenuifolium Carriére, ibid. Type: uncertain.”
ger otiey vieillardii Parlatore in DC. Prodr. 16(2): 521. 1868. Type:
Vieillar
Podocarpas tenuifolia (Carriére) Parlatore, (based on Dacrydium ela-
m var. tenuifolium) T:
N pore vieillardi (Parlatore) Kuntze, Rev. a Pl. 800. 1891.
Nageia tenuifolia (Carriére) Kuntze, ibid.
Tree to ca. 25 m., often much less; bark hard, slightly rough with scat-
tered low lenticels, breaking off in small thick flakes or short strips, dark
but weathering gray, brown and slightly fibrous or granular within; juve-
arently Carriére meant to replace this by his Podocarpus taxodioides var.
Pa “Cae the same type specimen) but failed to delete it from the manus cript.
1969 | DE LAUBENFELS, PODOCARPACEAE 327
nile leaves bilaterally flattened and distichous, up to 10 mm. long and
1.0 mm. broad, spreading and acute with a minute spine turned upward
more or less parallel to the branch, smaller towards the base and apex
of a branch, gradually reduced in size, thickened and losing the dis-
tichous habit: adult leaves acicular, sometimes not bilaterally flattened,
Straight, spreading at an angle of about 30°, acute, with a minute spine
turned upward, not distichous, from 2 to at least 4 mm. long in the mid-
dle, but beginning as scales at the base of a branch growth unit, ca. 0.4—
0.6 mm. wide, 0.4-0.8 mm, thick, sometimes continuing growth into addi-
tional growth units; non-foliage leaves of main shoots scale-like, appressed,
bifacially flattened, at least 2 mm. long; pollen cones lateral and sub-
tended by a short stalk with a few small scales or rarely terminal on a
short branch, linear, 7-12 mm. long and 1 mm. in diam.; microsporophyll
triangular and acute; seed cone on a lateral or terminal scaly shoot 6-8
mm. long, the scales 0.6—0.8 mm. long and appressed, the cone subtended
by 6-10 spreading involucral leaves 1-2 mm. long, robust, keeled, acute,
the cone itself formed of a small warty receptacle 2-3 mm. long with one
projecting sterile bract and one or occasionally two apical ovules; seed
oval or globular, generally with a blunt double crest and somewhat
elongated at the base, 4 mm. in diam., 5.5-6 mm. long.
DistRIBUTION, Throughout New Caledonia, particularly in areas of
serpentine rock, along river banks and in moist draws generally where
flooding is common, from sea level to 800 meters.
w Caledonia. Mt. Paéoua, McKee 17029 s 600-900 m. (Pp). Mt. Boulinda,
McKee 17199 j 750-850 m. (P), 17176 j (vp), Stauffer, Blanchon & Boulet 5778
2 (P, z), Veillon 142 j 750 m. (P). Baraua R., McMillan 5173 4 & K, P),
g (s K, NY, z), 2419 & (A, K, NY, P, Z). errs y pean 181 2 (K, P),
Pancher s.n. 2 (Bm, Pp), White 2112 o (a), 2285 s, j (A, K, P), Franc 35 j (K,
7, gig 69 s (Pp), 205 j (P), 253 j (P), 257 3 (®), ' Buchhols 1140 s (ILL, K, P),
oe K, % Sere 1040 s (P, z), McKee 2353 8 (P), 2567 6 ),
mann 1533 s (Pp, z), de hairs P389 Q 165 m. Pr k, re P389a j . RSA),
P4448 160 m. (a, ee Baumann-Bodenheim 15040 s (P, Z), 15041 s (P, 2)
Aubréville & Heine S R. Blanche (Upper Yaté), Bernier 206 s (P),
non, 241 s (P), Deckiok 1349 2 (ILL, K, - oes Q (ILL, K, P), 1464 ‘ ut,
K), 1465 s (ILL, K, Pp), 1553 @ (ILL, P), 1 Q (ILL, K, P), de Lau =
P11 s (spr), Baumann-Bodenheim & pee 10843 9 (Pp, z). Mare
328 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Hiirlimann 3109 j (z), 3158 6 (z), 3159 j (z). Canyon of Yaté R., Bernier 254 j
(Pp), 255 s, j (2), 256 j (P), 258 2, j (P). Plaine des Lacs, McKee 1142 2 (A).
Prony, Le Rat 222 j (p), 1719 s (p). Southwest, Moore 4 é (xk). Without loc.
Balansa s.n. 2 (BM, K), Pancher 4s (p), Mueller 68 s (Pp), Sarlin 237 s (P), 341
s (p), Baudouin 335 j (P).
ILLUSTRATIONS. PincER, R., Pflanzenreich IV. 5 (Heft 18): fig. 7F.
1903; Nat. Pflanzenfam. ed. 2. 13: fig. 124F. 1926; Sar in, P., Bois et
Foréts de la Nouvelle-Calédonie, ¢. 25. 1954, all as Podocarpus vieillardii.
The elongated pollen cones distinguish Dacrycarpus vieillardit from
other species except D. dacrydioides (the pollen cones of D. steupii are
not known). The leaves of D. dacrydioides are far shorter than those of
D. vieillardii and the pollen cone is normally terminal rather than lateral.
The leaves of D. steupii are shorter and more spreading while the involu-
cral leaves are longer than the foliage leaves, opposite to the condition in
D. vieillardii. The species with leaves resembling those of D. vieillardii
have much longer involucral leaves and sharply spreading and not im-
bricate foliage leaves. The low elevation river-bank habitat is also a
unique character.
23. Dacrycarpus steupii (Wasscher) de Laubenfels, comb. nov.
Podocarpus steupii Wasscher, Blumea 4: 405. 1941. Type: NIFS 6622837,
Celebes, Rantelmo (not seen).
Tree to 36 m. but usually much less; bark brown or gray, inner bark
pink, peeling in thin strips; juvenile leaves bilaterally flattened and
distichous, up to 8 mm. long and 0.9 mm. thick, becoming shorter and
not distichous, transitional leaves (sometimes fertile) variable in length,
the longest in the middle of a shoot, 3-4 mm. long, acicular, tip pungent
but turned upward parallel to the branch, becoming more constant in
size at 2 or 2.5 mm. in length as a mature form, strongly keeled on the
sides and back and spreading at an angle of 60° or more from the stem,
leaves on non-foliage branches lanceolate, bifacially flattened, almost ap-
pressed, 2-3 mm. long; pollen cones unknown; seed cone on a short
leafy lateral shoot 3-5 mm. or more long, the involucral leaves at the
base of the cone elongated and becoming widely spreading as the seed
develops, 3-5 mm, long, the cone made up of a small warty receptacle
2-3 mm. long with a sterile bract protruding on one side, one or two
terminal bracts fertile; seed globular with a small crest, 5—6 mm. long
and 4.5-5 mm. in diameter.
DistriBuTIoN. Locally common but widely dispersed on high wet
peaks or in bogs from 1,000 to 3,420 meters in elevation, mostly 1,600 to
3,000 meters, from Borneo to eastern New Guinea. Map 9.
Borneo. Peak of Balikpapan, Kostermans 7350 2 1,000 m. (A, K, L). Philip-
pines. Luzon, Benguet, Curran 10829 s (L). Celebes. ENREKANG: near Pin-
tealon, spur of Pokapindjang, Eyma 572 2 2,350 m. (A, BRI, K, L). Tinabang,
. side of Rante Mario, Eyma 675 2 3,000 m. (A, BRI, K, L, LAE), 778 j (® tL),
~~
1969] DE LAUBENFELS, PODOCARPACEAE 329
Manado: Palu, E. of Linden Sea, Blumbergen 3976 s 2,250 m. (A, L), 3977 j (L).
New Guinea. VocELKop: Aifat Valley, Moll BW 12820 s 860 m. (L), 12840 Q
920 m. (L), 12876 s 1,050 m. (L). WesTERN HALF: Wissel L., Eyma 5101 s
1,750 m. (A, K, L). Kadaitadie, E. of Motito, Wissel L., Vink & Schram BW
8667 s 1,900 m. (L, LAE). Baliem R., Brass & Versteegh 11187 2 1,600 m. (a,
K, L, LAE). Terr. NEw GUINEA: Wabag near L. Inim, Flenley ANU 2175 s
(K, L), 2769 2 8,300 ft. (kK, L). Aiyura, Womersley NGF 4428 2 6,000 ft. (a,
BRI, K, L, LAE). Sattleberg, Sambanga, Clemens 7258 s 5,000 ft. (A), 7562A s (a),
7902B 2 6,000 ft. (A). Mt. Amungwiwa, S. of Wau, Womersley NGF 17939 s
11,400 ft. (L). Papua: Ialibu, L. Buneh (S. Highlands), Pullen 2716 2 6,950 ft.
(BM, L, LAE), 2716A j (L). Uriko, road from Woitape to Kosipi (Cent. Div.),
Van Royen NGF 20289 ° 6,500 ft. (x; 4).
ILLUSTRATION. WasscHER, J., Blumea 4: ¢. 4, fig. 4. 1941, as Podo-
carpus steupii.
The preference for wet conditions which appears to characterize this
species probably explains why it is only occasionally found over its broad
range. Many specimens have been filed with other species. Sterile speci-
mens are distinguished by the short spreading acicular leaves becoming
nearly uniform in size on mature specimens. The leaves are generally
shorter and less (or not at all) bilaterally flattened than for comparable
stages of Dacrycarpus cumingii. On the other hand the leaves of D. com-
pacta are short and uniform but differ in being distinctly bifacially flat-
tened, fairly broad, and nearly appressed. The seed cones in each case
Sive positive identification. The one specimen from the Philippines is
somewhat uncertain because it is sterile and more or less juvenile.
24. Dacrycarpus cumingii (Parlatore) de Laubenfels, comb. nov.
Podocarpus cumingii Parlatore in DC. Prodr. 16(2): 521. 1868. Lectotype:
Cuming 803, Luzon, Mt. Banahao.
Nageia cumingii (Parlatore) Kuntze, Rev. Gen. Pl. 800. 1891. :
Podocarpus imbricatus Blume var. cumingii (Parlatore) Pilger, Pflanzenreich
IV. 5 (Heft 18): 56. 1903.
Tree to at least 20 m.; juvenile leaves bilaterally flattened and dis-
tichous, up to 12 mm. long and 1.2 mm. thick, the tip curved and parallel
with the branch, soon losing the distichous habit and becoming coarser;
Mature foliage leaves bilaterally flattened, spreading, somewhat falcate,
acute with a fine spine curved upward, strongly variable in length, the
longest in the middle of a branch unit, 6 mm. long and 0.6 mm. thick;
leaves on non-foliage branches bifacially flattened, lanceolate, nearly ap-
Pressed, 2-4 mm. long and 0.6 mm. wide; pollen cones lateral on short
shoots 2-5 mm. long, oval, 8-10 mm. long and 2-3 mm. in diam., micro-
sporophylls lanceolate; seed cone on a short, usually lateral shoot 6-10
mm. long or more, leaves elongated greatly at the base of the cone so
that the curving involucral leaves surround even the mature seed, the
longest at least 10 mm. long and 0.5 mm. thick, the cone formed ofa small
warty receptacle 2-3 mm. long with one or rarely two apical fertile
330 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
bracts; the mature seed with a distinct asymmetrical crest, 4.5—-5 mm. in
diam. and 5—6 mm. long.
DistRIBUTION. In mountain forests up to 3,000 meters in the Philip-
pines, Borneo, and (according to Wasscher, 1941) in Sumatra. Map 10.
Sarawak. Mt. Penrissen, S. of Kuching, Jacobs 5017 2 1,400 m. (Kk, L, US).
Philippines. Luzon: Mt. Polis (Mountain Prov.), Steiner 2207 s 2,040 ft. (L).
Mt. Pulog (Benguet), Curran, Merritt & Zschokke 18049 2 (1), Ramos &
Edano 45005 s (a), Steiner 2032 j 2,400 m. (L). Mt. Banahao (Tayabas), Cum-
ing 803 @ (a-lectotype; F, K, L-isotypes), Foxworthy 2387 & (L). Loher 7137
Q (kK, us), 7138 8 2,250 m. (kK), Curran & Merritt 7886 2 (Ny, us), Ramos
19557 s (us), Klemme 66 6, j (a), 874 2 (ny, us), Whitford 951 2 (x, NY,
us), Holman 4 @ (a), Vidal 623 9 (a, K, L), Barthe (1857) @ (a-syntype),
Ocampo 27926 s (A), Robinson 5656 2 (srt), Sulit 30051 s (Bri), Lucban
(Tayabas), Elmer 7465 2 (A, K, L, z). Mt. Mahaihai (Luconia), Wilkes s.n. s
(cx), Central Luzon, Loher 4852 2 (A, K, us). Without loc. Loher 2138 s, J
(us). Mrnxporo: Mt. Halcon, Merrill 5563 s (Ny). Panay: Mt. Midiaas (An-
tique), Yoder (1905) 2 (L). Necros: Canlaon Volcano along lake (E. Negros),
Edano 21935 j 1,860 m. (L), 21944 j (L). Mrnpanao: Mt. Apo (Davao),
Elmer 11684 @ (A, K, L, NY, US, z). Mt. McKinley (Davao), Edano 993 s (A).
ILLUSTRATION. WaASSCHER, J., Blumea 4: t. 4, fig. 5. 1941, as Podo-
carpus cumingii.
The long involucral leaves are the most distinguishing character of
this species, being approached only by Dacrycarpus cinctus which has
very different leaves. The bilaterally flattened mature foliage leaves are
the same as D. kinabaluensis but not as robust. Juvenile leaves of D.
steupit resemble mature leaves of D. cumingii.
25. Dacrycarpus kinabaluensis (Wasscher) de Laubenfels, stat. nov.
Podocarpus imbricatus Blume var. kinabaluensis Wasscher, Blumea 4: 400.
41. Type: Clemens 27854, North Borneo, Mt. Kinabalu.
Small tree or shrub down to 2 m. high; juvenile leaves bilaterally
flattened and distichous at first, at least 10 mm. long and 1.2 mm. thick,
falcate with a long upturned acuminate apex; mature foliage leaves bi-
laterally flattened, robust (stiff), falcate, spreading at about a 30
angle, strongly curved upwards at the apex and pungent, the spine not
projecting, markedly variable in length, becoming reduced on older
plants so that the longer spreading leaves may be as short as 2 mm., 0.5
mm. thick, and nearly quadrangular in cross section; leaves on non-
foliage branches bifacially flattened, lanceolate and pungent, curved up-
wards, 1-2 mm. long and 0.5-1.0 mm. wide, the size varying with the
robustness of the branch but not within a given branch; pollen cones
lateral on a short branchlet about 3 mm. long, globular, 8 mm. long an
3 mm. in diam.; seed cone on short lateral or terminal shoots 5-15 mm.
long and bearing, as is usual for the genus, the non-foliage type leaves;
at the base of the cone the leaves elongated to the size of foliage leaves
“ ~~ 6 ae
8 G ats Te 3
& c
‘ ra POA
iS e:
na : 4
pr ry) \ é ee
/ Ie Pi Z ~ .
. a a Fhe yj e
re ss set € bd ay, &
ee, oe a BS See °
AL SS" . ' : rs n
~~ COMPTONII! ig KO Be Tey & i 5
Maps § a er distribution of: 10, Dacrycarpus cumingii (Parlatore) de ar (dots west of line), and D. cinctus (Pilger) de
oe (east of line); 11, D. compactus (Wasscher) de Laubenfe ts), D.
b , known
12, De see sede vitiensis (Seemann) de epithet (dots), D. comptonii (Buchholz) de Laubenfels, known
ad from New Caledonia; 13 ichi i (Hickel) 4 ee (dots north of line),
, D. nagi (Thunberg) de Laubenf
S32 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
and nearly surrounding the young ovule but not reaching beyond the
middle of the ripe seed, up to 7 mm. long and 1.0 mm. thick, the cone
made up of a small warty receptacle 2-4 mm. long with one or two
sterile protruding bracts and an apical fertile bract; the mature seed
with a distinct asymmetrical crest, 5—5.5 mm. in diameter and 7 mm.
long.
DIsTRIBUTION. In mountain dwarf forests sometimes forming almost
pure stands on Mt. Kinabalu from 2,750 to 4,000 meters at the timberline.
North Borneo. Mt. Kinabalu, Paka Cave area, Clemens 10636 2 (A, GH, K),
10662 2 (A), 10686 s (A), 27092 s 13,000 ft. (A, K, L, NY), 27854 @ (A, K, L-
isotypes), 28910 2 11,000 ft. (A, K, L, NY), Wood & Wyatt-Smith SAN A4493
s 10,500 ft. (a, BRI), Meijer SAN 21988 2 9,500-11,000 ft. (k), SAN 29265 6
10,000 ft. (x). Mt. Kinabalu, Marai Parai, Clemens 32316 2 10-11,000 ft. (4,
K, L, NY), 32317 @ 11,000 ft. (a, L, NY), 32318 j 11,000 ft. (a, L, Ny). Mt.
Kinabalu, Gurulau Spur, Clemens 51201 2 (L). Mt. Kinabalu, side of Granite
Dome, Clemens 29914 2 12,500 ft. (K, L); S. slope, Jacobs 5755 2 3,600 m.
(K, L, us). Mt. Kinabalu, Clemens s.n. 2 11,000 ft. (pm), Nicholson SAN
17825 2 10,000 ft. (BRI, k, L), SAN 39766 2 9,000 ft. (K), Sinclair & Kadim
9146 2 10,700 ft. (kK, L), Haviland 1094 j 11,000 ft. (K), 1095 2 11,000 ft.
(a, K), Chew & Corner RSNB 868 2 10,500 ft. (kK), RSNB 5887 @ 9-11,000 ft.
(K), Gibbs 4216 4 12,000 ft. (x), Anderson $27079 @ 11,300 ft. (K).
ILLUSTRATION. WASSCHER, J., Blumea 4: ¢. 4, fig. 2y. 1941, as Podo-
carpus imbricata var. kinabaluensis.
The robust form of this high-elevation species is characteristic of conifers
in such places, and in general habit Dacrycarpus kinabaluensis resembles
D. compactus from high mountains in New Guinea, although in detail their
leaf form is, of course, quite different. D. kinabaluensis is most closely
related to D. cumingii, differing in the markedly robust foliage leaves and
distinctly shorter involucral leaves. The pollen cones also differ somewhat
in shape. Perhaps it could be treated as a variety of D. cumingii but not,
certainly, of D. imbricatus.
26. Dacrycarpus cinctus (Pilger) de Laubenfels, comb. nov.
Podocarpus cinctus Pilger, Bot. Jahrb. 69: 253. 1938. Type: Clemens 5261,
New Guinea, Busu River. 5
Podocarpus dacrydiifolia Wasscher, Blumea 4: 410. 1941. Type: NIF
bb13633, Celebes, Pawreang Mts.
Shrub of less than 4 m. to a tree up to 30 m. high; bark brown to black,
hard and uneven, inner bark reddish, breaking off in rough scales or plates;
juvenile leaves slightly bilaterally flattened and distichous at first, the
longest 12 mm. long and 0.8 mm. thick, falcate and bending upward to the
pungent tip, gradually changing to resemble the mature leaves; mature
foliage leaves of uniform size on a branch, slightly bifacially flattened,
lanceolate, falcate, eventually reduced to 2-3 mm. in length and spreading
1969 | DE LAUBENFELS, PODOCARPACEAE 333
at an angle to give the branch system a diameter of 3—4 mm., about 0.4-0.6
mm. wide, mature specimens including leaf sizes ranging up to 5 mm. in
the center of a branch unit, often glaucous; leaves of non-foliage branches
the same or more distinctly bifacially flattened, pollen cones terminal or
lateral on a very short branch 2—3 mm. long, globular or oval, 4-10 mm.
long and 2-3 mm. in diam., microsporophylls acuminate; seed cone lateral
on a short branch or terminal, 5—15 mm. long, involucral leaves much longer
than foliage leaves and clasping the young seed but generally not reaching
past the middle of the mature seed, the longest 6-7 mm. long, the cone
formed by a small warty receptacle 3-4 mm. long, with one or two project-
ing sterile bracts, seed and receptacle becoming red when ripe; mature
seed with a small asymmetrical crest, 6-7 mm. in diam. and 6—7 mm. long.
Fic. 9a.
DistripuTiIon. Mountain forests to high mountain shrubbery from 900
to 3,600 meters but mostly 2,200 to 3,200 meters, from the Celebes to the
high mountains of New Guinea. Map 10.
Celebes. Masamba, N/FS bb24958 s 900 m. (L). Pawreang Mts., Ulu Salu
(Upper Binuang), NJFS 6b13633 2 1,800 m. (L-holotype of Podocarpus dacry-
diifolia). Pinapuang (Manado), Eyma 3873 s (£4). Ceram. G. Sofia, Central
Mts., Stresemann 125 j 1,300 m. (Lt). G. Pinaia, Middle Ceram, Eyma 2276 s
3,030 m. (L), Stresemann 251 s 3,010 m. (L), 276a j 2,530-2,750 m. (L). New
Guinea. WesTERN HALF: Mamberamo R. (Mt. Doorman), Lam 1773 ? 3,260
m. (L). Hellwig Mts., Pulle 964 2 2,600 m. (kK, L), van Romer 736 s (L). Lake
Habbema, Brass 10513 2 2,800 m. (A, K, L), 10514 j (A, L), 10675 2 3,000 m.
(A, K, L), Brass & Versteegh 10447 2 2,840 m, (A, L). L. Quarles, Versteegh
BW 2537 s 3,600 m. (x, L). Terr. New Guinea: Wapu R. (Wabag), Hoogland
& Schodde 7166 s 9,500 ft. (A, L). Tomba, Mt. Hagen—-Wabag Road, Flenley
ANU 2819 2 8,900 ft. (K, L), Robbins 238 2 8,000 ft. (A, K, L, LAE, US). Lal
Valley (Wabag), Robbins 3112 @ 7,500 ft. (A). Minj-Nona Divide, Pullen
5052 2 10,600 ft. (L), 5267 s 9,500 ft. (K, L). Keglsugl (Chimbu), Saunders
804 s 8,000 ft. (L, LAE). Toromambuno Mission (Upper Chimbu), Pullen 313
2 9,000 ft. (A, BRI, K, L, LAE, US), 313A j (K, L, LAE). Mt. Wilhelm Track,
Chimbu Valley, Robbins 673 2 9,000 ft. (A, BM, L, LAE). Mt. Wilhelm, Brass
30412 s 2,770 m. (A, K, L, NY, US), 30707 2 3,180 m. (A, K, L, LAE, NY, us),
Stauffer 5670 8 3,250 m. (z). Kerigomna Camp (Goroka), Hoogland & Pullen
tw. Mt. Kerewa and Mt. Ne, Vink 17188 j 2,890 m. (L).
2 2,880 m. (x). Mt. Giluwe, above Klareg, Schodde 2021 ¢ 8,800 ft. (K, L, oe
2104 j 9,100 ft. (x, LAE). Woitapi-Kosipi Road (Cent. Div.), Van Royen
334 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
20309 2 6,300 ft. (K, L). Murray Pass, Wharton Range, Brass 4688 2 2,840
m. (A, K, L, NY, US).
ILLUSTRATIONS. WasscHER, J., Blumea 4: ¢. 4, fig. 6, as Podocarpus
cincta, and fig. 7 as Podocarpus dacryditfolia. 1941.
Confusion has existed between this species and Dacrycarpus compactus
with which it overlaps in range, although they are not at all the same. The
involucral leaves of D. cinctus are long and narrow, clasping the smaller
seed, while those of D. compactus are short and triangular, barely reach-
ing the base of the distinctly larger seed. Foliage leaves contrast in the
same way. The terminal position of the pollen cone which is usual in D.
cinctus is a character shared only with D. compactus and D. dacrydioides.
The essential non-dimorphic quality of the mature foliage is shared only
with D. compactus, D. expansus, and D. imbricatus (in part). There are
several specimens of D. cinctus which differ from the typical form in the
direction of D. compactus and could, perhaps, be recognized as forming a
variety (Fic. 9b). These are: Brass 4688, 10513, 10514, 10675, Brass
& Versteegh 10447, Versteegh 2537, Pulle 964, Hoogland & Pullen 5574,
Pullen 5052, 5267, Lam 1773, Schodde 2021, 2104, van Romer 736, Vink
NGF 12430, and Womersley NGF 14018. They differ in that the foliage
leaves are somewhat broader (up to 0.8 mm.), as are the involucral
leaves (up to 1.0 mm wide). The possibility exists that these are hybrids,
being found at and not far below the lower elevation limit of D. compactus.
27. Dacrycarpus expansus de Laubenfels, sp. nov.
Arbor ad 25 m. alta; cortex squamosus. Folia plantarum iuvenilium
dimorpha, ad ramulos breves compressa bilateraliter, patentia, falcata,
pungentia, ad 12 mm. longa, 1.5 mm. lata, biseratim expansa; ad ramulos
magis elongatos compressa bilateraliter, imbricata, lanceolata, pungentla,
ad 4 mm. longa, in basi 0.8 mm. lata; folia plantarum adultarum com-
pressa bifacialiter, expansa, falcata, acuta, dorso carinata, 2-4 mm. longa,
0.6-1.0 mm. lata. Strobili masculi laterales ad ramusculis 1-2 mm. longis,
ovoidei, 6 mm. longi, 3 mm. crassi. Strobili feminei ad apicem ramulorum
saepe brevi 4-5 mm. longi, foliis parvis; folia involucra longiora, 3-4 mm.
longa, 0.6 mm. lata; receptaculum parvulum, verruculosum, 2-3 mm.
longum; semen globosum, cristatum, 4 mm. diametro, 5—6 mm, longum.
Holotypus: Hoogland & Schodde 7463 (L), New Guinea, Yobobos Grass-
land. Fic. 7b.
Distripution. Locally common in disturbed forests in the highlands es
New Guinea at 2,600-2,670 meters.
New Guinea. Terr. New Gurnea: Yobobos Grassland, Laiagam Subdistrict
(Wabag), Hoogland & Schodde 7463 2 8,500 ft. (L-holotype; BRI, Lak-isotypes),
7440 s (L, LAE), 7682 j (L, LAE), Robbins 3214 8 8,700 ft. (BRI, L, LAE). PAPUA:
E. foot of Mt. Ambua, Tari Subdist., Vink 17502 2, é 2,670 m. (L), 17499 J
(L), 17500 j (L), 17501 j (x).
we
Cro
nn
1969 | DE LAUBENFELS, PODOCARPACEAE
=
~
2
s
~
FIGURE 9, a. Dacrycarpus cinctus (Pilger) de Laubenfels, portion of Hartley
13263 (A), typical form of the species; b, the same, fragments showing variation
toward D. compactus: c, D. compactus (Wasscher) de Laubenfels, fragments.
—
The short distinctly bifacially flattened involucral leaves clasping the
receptacle, but not the somewhat small seed, distinguish this new species
from all others in the genus. The only other species which have distinctly
bifacially flattened involucral leaves are Dacrycarpus cinctus and
compactus. Those of D. cinctus are twice as long, clasping the seed,
336 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
while those of D. compactus are up to twice as broad below a much larger
seed. As in both of these species, D. expansus lacks dimorphic foliage
when mature, contrasting in appearance because of the strongly spreading
rather than imbricate leaves which are also distinctly broader than those
of D. cinctus. The normally lateral pollen cones also distinguish D. ex-
pansus from these two species. Except for the great contrast in shape of
the involucral and foliage leaves D. expansus resembles D. steupii in
gross morphology, differing in habitat preference.
28. Dacrycarpus compactus (Wasscher) de Laubenfels, comb. nov.
Podocarpus compacta Wasscher, Blumea 4: 411. 1941. Type: Brass 4284,
New Guinea, Mt. Albert Edward.
Small tree 2-15 m. high; bark hard, rough, warty, dark gray, breaking
off in scales, inner bark reddish straw color; juvenile leaves bilaterally
flattened, lanceolate, falcate and curved upward at the tip, acute, strongly
keeled laterally, not distichous, 2.5 mm. long and 0.6 mm. thick; mature
foliage leaves not dimorphic, bifacially flattened, spreading slightly, fal-
cate, lanceolate, pungent, keeled on the back, 2—3 mm. long, 0.6-1.0 mm.
wide (the wider probably on older plants); pollen cones lateral on a short
branch about 3 mm. long or more usually terminal, 7-8 mm. long and
3 mm. in diam., microsporophylls lanceolate, acute; seed cone terminal,
generally on a short branch 6-17 mm. long, involucral leaves robust,
4-5 mm. long and 0.8-1.2 mm. wide, clasping the receptacle, the cone
itself made up of a small warty receptacle 3-4 mm. long with a sterile
bract protruding; seed globular, with a blunt crest, 7-8 mm. long and 7
mm. in diam. Fic. 9c.
DistripuTion. In high mountain forests and shrubberies often as an
emergent, and sometimes the dominant tree at the tree line in New
Guinea, from 3,200 to 3,900 meters. Map 11.
New Guinea. WESTERN Hair: L. Habbema, Brass 9291 2 3,225 m. (A, K, L),
21104 2 3,225 m. (A). Terr. New Gurnea: Mt. Kinkain, Cent. Kubor Range
(Minj), Saunders 708 2 11,800 ft. (a, L), Pullen 5111 8 11,770 ft. (a, K, 2);
5138 @ 12,000 ft. (K, x). Mt. Wilhelm, Robbins 718 2 12,000 ft. (1), Brass
29861 2 3,650 m. (A, K, L, NY, US), 29935 2 13,100 ft. (A, K, L, NY, US), 5.” §
3,320 m. (us), Millar NGF 14671 2 12,000 ft. (x, L), Womersley NGF 8852
Q 11,870 ft. (A, K, L), 8861 8 (a, K, L), Pullen 338 12,500 ft. (a, L), Hoogland
& Pullen 5650 & 11,700 ft. (A, BRI, K, L, US), 5703 2 12,500 ft. (A, BRI, K, L, us);
Havel NGF 17421 2 11,500 ft. (x), Stauffer 5670 8 3,250 m. (x, L), Balgooy
287 2 3,650 m. (L). Mt. Otto (E. Highland), Brass & Collins 31021 ¢ 3,460 m.
(A, K, L, Ny). Mt. Piora, Kaimantu Subdiv. (E. Highland), Henty & Carlquist
NGF 16566 2 10,500 ft. (k, x). Papua: Mt. Dickson, Goilala Subdist., Hartley
TGH 12958 2 11,500 ft. (L). Mt. Albert Edward, Brass 4284 @ 3,630 m. (A,
ny-isotypes), 4284A j (A, Ny), 4347 j (Ny), 4348 s 3,680 m. (A, NY).
ILLUSTRATION. WaAsSCHER, J., Blumea 4: ¢. 4, fig. 8. 1941, as Podo-
carpus compacta,
1969] DE LAUBENFELS, PODOCARPACEAE 337
The particularly large seed completely free of the involucral leaves
and the small bifacially flattened not widely spreading leaves distinguish
Dacrycarpus compactus from other species. D. expansus, with rather
similar, though spreading leaves, has much smaller seeds and more lan-
ceolate involucral leaves. The wild tree of D. compactus is a very strik-
ing plant, often rising above the other shrubs near the tree line and stand-
ing out with a dark green color. The short juvenile leaves are the most
primitive in the genus and apparently only in this species are the juvenile
leaves never distichous.
ADDITIONAL SPECIES:
Dacrycarpus dacrydioides (Rich.) de Laubenfels, comb. nov.
Podocarpus dacrydioides Rich. Essai d’une Flore de la Nouvelle Zéland, 358.
t. 39, 1832. Type: D’Urville in 1827 (not seen).
Podocarpus thujoides R. Br. ex Mirb. Mém. Mus. Hist. Nat. Paris 13: 75.
1825 (nomen).
Dacrydium excelsum Cunn. Ann. Nat. Hist. 1: 213. 1838 (nomen illeg., based
on Podocarpus dacrydioides).
Nageia excelsa (Cunn.) Kuntze, Rev. Gen. Pl. 800. 1891.
Acmopyle Pilger, Pflanzenreich IV. 5 (Heft 18): 117. 1903. Type
species: Acmopyle pancheri (Brongn. & Gris) Pilger.
Small trees; foliage leaves linear, bilaterally flattened, distichous, with
scale-like, triangular and bifacially flattened; pollen cones terminal and
lateral together; seed cones on short branches which are lateral or terminal
Or grouped together, becoming enlarged and warty as a receptacle with
a single subterminal seed; the ovule at first inverted and partially covered
by the epimatium, eventually becoming nearly erect, fused with the
epimatium, and fleshy.
This genus is characterized by the unique combination of the seed fused
with the epimatium (fertile scale), together with an inverted ovule which
becomes gradually nearly erect as it matures. The seeds of other genera
of Podocarpaceae which are fused with the fertile scale do not become
erect. Acmopyle shares bilaterally flattened and distichous leaves with
Falcatifolium and juvenile forms of Dacrycarpus. With the latter it also
Shares a warty receptacle. Two species are known, differing in the charac-
ter of seed and receptacle as well as in size of the pollen cone and details
of leaf-form. Both are island endemics.
29. Acmopyle pancheri (Brongn. & Gris) Pilger, Pflanzenreich IV. 5
(Heft 18): 117. 1903.
Dacrydium pancheri Brongn. & Gris, Bull. Soc. Bot. France 16: 330. 1869.
Type: Pancher in 1869, New Caledonia, Mount Mou.
338 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Nageia pancheri (Brongn. & Gris) Kuntze, Rev. Gen. Pl. 800. 1891.
Podocarpus pectinatus Masters, Gard. Chron. III. 9: 113. 1892. Type: Hort.
Sander s.n., of New Caledonian origin.
Acmopyle alba Buchholz, Bull. Mus. Hist. Nat. Paris II. 21: 281. 1949. Type:
Buchholz 1704, New Caledonia, Bois de Mois de Mai.
Tree from 5 to 25 m. high; bark hard and smooth, weathering to a gray
color, brown to tan and fibrous within, with scales breaking off on older
trees; foliage leaves bilaterally flattened and decurrent, distichous, linear
and spreading 60° to 75° from the branch, tapering somewhat towards
the apex which is turned slightly in the direction of the shoot apex, or
slightly falcate, at first with two glaucous bands on each surface asso-
ciated with the stomatiferous areas but with further development this
condition suppressed on the upper surface but the white bands remaining
prominent below, the midrib marked by a faint line on the upper surface
and more pronounced below, leaves shorter at the beginning and end of
a sequence of growth with the leafy shoots never producing a second
cycle of leaves but commonly continued into fertile shoots; shade leaves
spread out into a flat and almost solid plane, except for the smaller
leaves at either end of the branch, 16-21 mm. long and 2.8-3.0 mm. wide,
slightly revolute on the margins; leaves exposed to the sun less regularly
placed and more noticeably keeled, often overlapping and weakly spread
into a plane, 10-15 mm. long and 1.8—2.2 mm. wide, with intermediate
forms sometimes found; non-foliage leaves scale-like, triangular, bifacially
flattened, keeled on the back, less than 2 mm. long, on main branches
bearing foliage shoots broadly decurrent and dispersed, on fertile branches
(and occasionally at the base of a leafy shoot) more or less crowded; pol-
len cones terminal or often a pair (one of which is lateral) produced at
the apex of a leafy shoot or on a scaly shoot which may itself be terminal or
lateral either at the apex of a leafy shoot or a main branch bearing leafy
shoots (on vigorous trees all of these together), subtended by a few small
scales, 10-20 mm. long by 2-3 mm. in diam. (fide Hooker, 1902, to 35 mm.
by 4 mm.), the microsporophylls small and triangular; seed cones terminal
or lateral at the apex of a leafy shoot, or on a main shoot, or terminal or
lateral on a scaly shoot which may be either terminal on a leafy shoot, or
lateral on a main shoot (on vigorous trees a combination of these) ; the seed
cone subtended by a peduncle 9-22 mm. long, densely covered by small
overlapping scales and slightly enlarged toward the cone to a diameter of
about 2 mm.; the cone formed by a fleshy warty receptacle 8-18 mm. long
involving about 4 to 8 bracts whose free tips each surmount a bulge; one
ovule inverted and protruding from the enveloping epimatium in the axil
of a sub-apical bract, becoming almost erect and fleshy, the epimatium
completely fused to the mature ovule and apparently attached for about
half its length (marked by a roughened area on the seed and a charac-
teristic ridge on the dried fruit); seed globular, 10-11 mm. in diameter,
thick and hard.
DIsTRIBUTION. Scattered in moist rainforest over serpentine rocks 1n
1969 | DE LAUBENFELS, PODOCARPACEAE 339
most of New Caledonia from near sea level to at least 1,200 meters.
Growing as a canopy tree in drier areas and sometimes found in the
understory within the mossy forest where it is fully fertile.
New Caledonia. Upper Diahot, Tendé Forest, McKee 17540 j 500 m. (P). Mt.
Colnett, Hiirlimann 1964 2 1,200 m. (Pp, z). Mt. Paéoua, McKee 17057 s 900-
1,100 m. (Pp), Bernardi 10151 2 900-950 m. (P, z). Mt. Boulinda, Veillon 136
$ 1,200 m. (P), Schmid 137 s (orstom). Crest W. of Col de Rousettes, de Lau-
benfels P429 s 700 m. (A, RSA). Me Arembo, Bernier 1007 s (K, Pp). Mt. Koun-
gouhaou N., McKee 17954 2 1,000-1,100 m. (P). Mt. Mou, Pancher (1869) @
1,200 m. (p-holotype of Dacrydium pancheri), Balansa 2862 2 (BM, K, NY, P),
Compton 485 2 (BM), Franc 170 @ (A, BM, K, NY, P, Z), Virot 10s (a, Pp), Le Rat
697 s (P), 980 Q (kK, P), 2594 s (A, P), Bernier 278 2 (Pp), 1309 2 (P), Buchholz
1451 2 (ILL, K, P), 1587 9 (ILL, K, P), 1587S j (ILL, P), 1593S j (IL, P), 1790
é (ILL, K, P), McMillan 5013 2 (P), 5014 s (P), de Laubenfels P130 2, 8 1,140
m. (sBT), P355 @ (A, K, RSA), P356 3 (a, RSA), Brousmiche s.n. s (P), Thorne
28704 s (p), Baumann-Bodenheim & Guillaumin 11260 s (P, z), Baumann-Boden-
heim 15632 s (Pp, 2), 15633 9 (Pp, z), McKee 3517 @ 1,100 m. (A, K, P). Mt.
Ouin, McKee 9795 s (K, p). Col de Mt. Dzumac, McKee 9773 s (K, P), 9774 j
(K, P), 12922 8 (p), de Laubenfels P447 2 900 m. (A, RSA), Baumann-Bodenheim
& Guillaumin 12714 s (ve, z), Blanchon 930 s (P). Mt. Koghis, Pancher sn. &
800 m. (P), Alleizette 142 2 (P), Brousmiche 9 2 (with Prumnopitys ferrugi-
noides), Hiirlimann 1657 s 1,050 m. (Pp, z). Mt. des Sources, Hiirlimann 911 s
(P, z). Mois de Mai, Bernier 276 s (P), 277 s (P), 279 s (P), 280 8 (P), 281s (P),
321 j (P), Buchholz 1354 s (ILL, P), 1388 s 200-250 m. (ILL, P), 1388A j (ILL),
1388M 8 (Pp), 1698 s (tL, P), 1698L (shade) s (Pp), 1704 2 (11L-holotype of
Acmopyle alba; x, p-isotypes), McKee 3454 s 200 m. (A, K, P), Baumann-Boden-
heim 13964 s (p, z), 14258 & (P, Zz), 14263 2 (P, 2), 14988 s (P, Z), 14992 s (®,
Z), 15096 & (p, z), 15097 9 (P, z), 15098 s (P), 15130 s (P, Z), 15208 s (P, Z),
15213 s (z). Slope N. of R. Bleue, de Laubenfels P136 s 700 m. (sBt), P382
250 m. (A, RSA), P383 6 (A, K, RSA, SBT), P383A (shade) s (A, RSA), P4465 @
770 m. (A, RSA), Baumann-Bodenheim & Guillaumin 10929 s (P, 2), Baumann-
Bodenheim 15043 j (z), 15055 s (Pp, z), McKee 12653 s 200 m. (Pp). Bois Elec-
trique, Foster 206 9 (p), de Laubenfels P377 2 240 m. (A, RSA), P378 s (A, RSA),
Hiirlimann 3411 s 220 m. (z). Without loc., Mueller 44 s (111, P). Cult., Hort.
Sander s.n, s (K-holotype of Podocarpus pectinatus).
ItLustraTIons. Hooker, J. D., Bot. Mag. t. 7854. 1902, as rine
pus pectinata; PiiceR, R., Pflanzenreich IV. 5 (Heft 18): fig. 24. yes :
Gray, Jour. Arnold Arb. 28: i IB. 1947; SarLIN, P., Bois et Foréts de
la Nouvelle-Calédonie, t. 22 & 23, as Acmopyle alba. 1954.
The difference between shade and sun growth forms suggests that two
entities are included under Acmopyle pancheri but this difference occurs
regularly on single plants. A. alba differs from A. pancheri ean!
larger pollen cones (18-20 mm. long by 3 mm. in diam. in A. @ : -
10-13 mm. long by 2 mm. in diam. in A. pancheri). There are absolutely
no other differences between these two taxa, while even larger pollen
cones are described by Hooker (1902). Because only a few examples of
pollen cones are available to show variations in size and because speci-
340 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
mens can not be distinguished in the absence of pollen cones, it is felt
that A. alba should not be separated from A. pancheri at this time. In the
future it might seem advisable to separate them at the varietal level.
30. Acmopyle sahniana Buchholz & Gray, Jour. Arnold Arb. 28: 142.
1947. Type: Gillespie 3273, Fiji, Mt. Vakarogasiu.
Small and gnarled tree 3-5 m. high; leaves bilaterally flattened, dis-
tichous, spreading at an angle of 60° to 80°, linear, slightly tapering and
curved in the direction of the branch apex near the blunt tip or falcate,
decurrent, 10-19 mm. long and 2.0-3.2 mm. wide (wider according to
Buchholz and Gray but not according to their illustrations or to herbarium
upper surface, midrib faintly marked on both surfaces, margin slightly
revolute, leaves smaller at the begining and end of a sequence of growth
with one sequence often continuing into a subsequent growth unit; non-
foliage leaves on main branches scale-like, bifacially flattened, long tri-
angular, 1.5—-2.5 mm. long, keeled on the back and broadly decurrent,
more crowded at the base of a foliage branch and on the peduncle of the
seed cone; pollen cones terminal, 5 mm. long and 2 mm. in diam.; micro-
sporophylls triangular, acute; seed cones lateral or terminal on a foliage
branch, with a short (5 mm. ) scaly peduncle, the cone formed by a fleshy
warty receptacle involving two bracts, the uppermost being fertile with
a single inverted ovule partly covered by a broad epimatium; seed becom-
ing nearly erect, rounded and elongated into a conical point, with the
epimatium fused along one side, its margin forming a fringe about half-
way to the micropyle, mature need not known.
DistrIBuTION. Known only from two isolated mountains on either
side of Viti Levu in dense low forest 800 to 1,050 meters in elevation,
where it is locally common.
Fiji. Vitr Levu: Mt. Vakarogasiu (Namosi), Gillespie 3273 s 900 m. (a-holo-
type; K-isotype), Koroiveibau 14598 2 2,600 ft. (kK). Mt. Koroyanitu (Mt.
Evans Range), Smith 4122 $ 950-1,050 m. (A, BRI, ILL, K). Without loc., Horne
Sn. S (K).
Intustration. Bucuuotz, J. T., & N. E. Gray, Jour. Arnold Arb.
28: ¢. IA. 1947.
This rare species is of interest because its only relative, Acmopyle
pancheri, occurs in New Caledonia, not a common combination.
Decussocarpus de Laubenfels, gen. nov. Type species: Decussocarpus
vitiensis (Seemann) de Laubenfels.
Nageia Gaertner, De Fruct. et Sem. 191. 1788. Type species: Nageia japonica,
nomen illeg. (description confused).
Folia opposita, decussatim vel spiraliter inserta, lanceolata vel rotun-
1969 | DE LAUBENFELS, PODOCARPACEAE 341
data, ad basim contracta, uni- vel multinervata. Strobili masculi solitarii
vel fasciculati. Strobili feminei pedunculati; pedunculi cum squamis (vel
foliis); semina saepius singula, globosa, inversa, squama fertilia cum
ovulo conjuncta.
The new genus Decussocarpus is composed of three sections formerly
treated as a part of Podocarpus. A group of characters unite these three
sections while distinguishing them from Podocarpus. Ovules are pro-
duced subterminally on a scaly shoot not divided into a naked peduncle
micropylar end of the inverted seed extends distinctly downward (towards
the base of the fertile complex) so that the mature seed appears to be
attached at an angle on the end of the fertile shoot. As a result the seed
displays a projecting curved beak in contrast with all related taxa. As-
sociated with the elongated attachment is the tendency for the fruit to
fall with the fertile shoot still attached. In contrast with Podocarpus, a
cluster of five or more pollen cones may occur in some species on a single
shoot.
The leaves of Decussocarpus have a number of distinguishing character-
istics. Opposite decussate leaves are found throughout the genus with the
exception that in the section AFRocARPUS some branches have spirally
placed leaves (herbarium specimens therefore may lack this character
which, nevertheless, can be readily found on any mature living specimen
of § Arrocarpus). A unique leaf orientation further occurs in all sections
of the genus, although it may be absent in a few species of the section
Dammaroiners. The distichous leaves being amphistomatic, instead of
making unequal twists on opposing sides of the branch to bring the axial
surface of the leaf upwards at all times, always turn in the same manner
with respect to the axis so that on the left side the abaxial surface is
uppermost. This can be seen in section AFROCARPUS even on branches
without decussate leaves. Unlike Podocarpus, the leaves of Decusso-
carpus have no accessory transfusion tissue and unlike Prumnopitys they
do have a hypoderm. Also unlike Prumnopitys, the leaves are not linear,
but oval or lanceolate.
Most species of Decussocarpus are large trees, some of which are valuable
timber trees and others are in demand for decorative planting in the
warmer parts of the world. The genus is divided into three sections based
on the relative width and venation of the leaves. Section DECUSSOCARPUS
has single-veined but relatively wide leaves compared to section AFRO-
CARPUS whose leaves are more than ten times as long as they are wide.
Section Dammaroues has broad multiveined leaves.
Section DEcUSSOCARPUS.
Podocarpus section Polypodiopsis Bertrand, Ann. Sci. Nat. V. 20: 65.
Type species: Podocarpus vitiensis Seemann [ Decussocarpus vitiensis (See-
mann) de Laubenfels].
342 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Trees with opposite decussate leaves which are ovate or lanceolate, ses-
sile, sharply narrowed to a decurrent base, single veined, amphistomatic,
and not more than about five times as long as wide; pollen cones sessile,
solitary or grouped on a special scaly shoot; seed cones in the form of a
scaly or leafy shoot with one or rarely two fertile subterminal bracts; ovule
inverted and covered by the seed scale which makes an apical crest over
the inverted base of the ovule; seed large, globular, blunt at one end but
elongated into a curved beak at the micropylar end (at the base of the
fruit) and covered by the fleshy seed scale.
An account of this section is given in Wasscher (1941), who shows
some uncertainty about how to treat it. Once its characters were com-
pletely known, there was agreement that it is most closely related to
section Dammaroies (Nageia of most authors). The difference in leaf
size and venation, however, made it advisable to separate these two taxa
into different sections which have until now been treated as a part of the
extensive genus Podocarpus. The new genus, Decussocarpus, is here be-
ing proposed to accommodate the two groups and the more recently named
section ArrocarPus in order to recognize the considerable morphological
differences that previously have been merged in the one genus Podocarpus.
Section Decussocarpus extends from eastern Indonesia to South America,
and fossil specimens of it from Chile were once the basis of reports of
Sequoia in the Southern Hemisphere (Florin, 1940). There are four
species.
KEY TO THE SPECIES OF SECTION DECUSSOCARPUS
1; Sa heal branches bearing scales.
Scales on non-foliage branches appressed, thin; eps leaves with a sharp
narrow midrib; mature pollen cones elongated. .............-------: 7
GE Lk RS ea a 31. D. vitiensis.
2. Scales on non-foliage branches spreading, thick; mature pollen cone globu-
lar iad elongate
e foliage leaves with a raised central band narrower than the
ad leaf margins; forest tree. ................ 32. D. comptonii.
3. Mature foliage leaves with a broad raised central band broader than
adjacent leaf margins; small tree at water’s edge. .......------ 007°
1. Both primary and secondary branches bearing leaves. .... (D. 17 ospigliosii).
31. Decussocarpus vitiensis (Seemann) de Laubenfels, comb. nov.
Podocarpus vitiensis Seemann, Jour. Bot. 1: 33. ¢. JJ. 1863. Type: Seemann
, Fiji.
Podocarpus filicifolius N. E. Gray a Patt), Jour. Arnold Arb. 43: 74. 1962.
Type: Kostermans in 1949, Morot
Tree to 43 m. high; bark brown to red brown, weathering to blackish
or gray, fibrous, fissured and peeling in short vertical strips; foliage
branches opposite or alternate on non-foliage branches and subtended by
a short 1-2 cm. scaly base, sometimes with both lateral and terminal
1969 | DE LAUBENFELS, PODOCARPACEAE 343
foliage branches together on the same base, the foliage branch not nor-
mally branching again; foliage leaves distichous and equally twisted at
the base, lanceolate with a small blunt tip, a narrow but distinct rib
marking the vascular bundle on both surfaces, juvenile leaves up to 40
mm. long by 8 mm. wide, adult leaves 15-25 mm. long and 3—5 mm. wide;
non-foliage branches with appressed and thin scale leaves which are
broadly decurrent and dispersed, 1-2 mm. long; pollen cones single and
terminal or grouped with terminal and lateral cones together, either one
to three at the apex of a foliage branch or one to three at the apex plus
opposite pairs of groups of one to three along a scaly branch, cylindrical,
10-24 mm. long and 1.8—2.2 mm. in diam., microsporophylls triangular,
about 1 mm. long; one or two ovules subterminal on a scaly shoot, 6-10
mm. long (which may be terminal or axillary on leafy or scaly branches
and solitary or grouped); ovule inverted with the micropyle lying close
to the attachment of the seed complex with the micropyle at the end of
an elongated beak that may extend more than 2 mm. below the attach-
ment, the fertile scale completely enveloping the ovule and forming over
the young seed an apical crest which sometimes persists on the mature
fruit; mature seed globular, pear shaped, 13-16 mm. long including the
curved beak, 8-10 mm. in diam., covered by the deep red fleshy scale
and usually accompanied when it falls by the fertile shoot on which some
of the scales may still persist.
DIstTRIBUTION. Scattered and locally common in a discontinuous se-
ries of regions from Morotai to the Fiji Islands in rainforests, from near
sea level to 1,800 meters. Map 12
Moluccas. Morotai, Kostermans (1949) j (t8-holotype of Podocarpus filici-
folius; (a, K- isotypes). New Guinea. WESTERN HatF: Wissel Lakes, Mt. Barara,
Eyma 5155 j (Lt). Wissel Lakes, Motito, Vink & Schram BW 8730 j 1,800 m.
(L). Barnhard Camp (Idenburg R.), Brass & Versteegh 12534 s 1,200 m. (A, B),
Brass 12787 2 1,200 m. (A, BM, K, L), 12787a j (A, L), 12912 2 (L). Cycloop
Mts., Versteegh BW 913 2 1,100 m. (K, L, LAE), Van Royen & Sleumer 6073
s 1,220 m. (kK, z). Papua: Koroba Station, Pullen 2840 2 5,300 ft. (LAE). Alola,
Carr 14160 8 6,000 ft. (a, BM, L, NY). Lala R., Carr 15666 6 5,000 ft. (A, BM,
L). New Britain. Mt. Tangis, ‘Talasea Dist., Frodin NGF 26292 j 3,500 ft. .
LAE), NGF 26917 s 2,400 ft. (L). Benim, Kandarian Dist., Henty & Frodin NGF
27359 8 1,000 ft. (L, Lar). Fullerborn Harbor (Kandarian), Hammermaster &
Sayers NGF 21842 & 100 ft. (L). Santa Cruz Is. Vanikoro, Walker BSIP 1580
j (L). Fiji. Vitt Levu: Nandarivatu, Degener 14483 j 750-900 m. (A, K, L, NY,
US), 14496 9 (a, Ny), Gillespie 3863 2 (K, NY, US), Gibbs 674 2, 6 (BM, K),
Vaughn 3254 j (am, K), Mead 1964 j (x), 1974 é (K), 1982 s (kK). Nausori,
Damanu NH15 s (x). Namboutini (Serua), de Laubenfels P309 j 1,000 ft. (a,
RSA), Damanu R10 s (k), R15 s (K), R32 s (K), Qoro & Kuruvoli s.n. (k).
Serua, Bola 10 s (k), Damanu NLS s (x), NL10 s (x), NL12 s (K), G7 s@,
“The Leiden specimen is accompanied by an unattached seed of D. wallichianus,
a Napa which was also collected in the area by Kostermans. Podocarpus ogee
d on the presumed pairing of the alien seed with the accompanying leaves
(de Linibetifeds, 1967).
344 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Naikorokoro, Damanu KU22 s (x). Galva Forest, Damanu 152 2 (kK). VANUA
Levu: Mt. Kasi (Thakaundrove), Smith 1796 s 300-430 m. (a, K, US). Without
loc.: Seemann 576 j (K-holotype of Podocarpus vitiensis; A, BM-isotypes), Horne
531 s (kK), Tothill 844 s (K), 845 s (kK), Graff 33 s (K).
ILLUSTRATIONS, SEEMANN, B., Jour. Bot. 1: ¢. JJ. 1863; Fl. Vitiensis,
t. 78. 1868, as Podocarpus vitiensis.
While in most ways typical of the genus of which Decussocarpus vi-
tiensis is the type species, the dimorphic foliage, shared by two other spe-
cies in section Decussocarpus is rather unusual, found elsewhere in the
family only in Dacrycarpus and Acmopyle. The compound clustering of
specimens of D. vitiensis from other closely related species. It is of inter-
est to note that D. vitiensis broadly overlaps D. wallichianus in its dis-
tribution; the latter extends throughout New Guinea as well as further
west.
32. Decussocarpus comptonii (Buchholz) de Laubenfels, comb. nov.
Podocarpus comptonii Buchholz, Bull. Mus. Hist. Nat. Paris. II. 21: 284. 1949.
Type: Buchholz 1684, New Caledonia, Mt. Mou.
Tree to at least 30 m. high; bark tan to gray-brown, weathering to
gray or dark gray, fibrous, becoming very rough and fissured on older trees,
breaking off in short vertical strips or rough fragments; foliage branches
opposite or alternate on non-foliage branches or one to several at the
apex of an older foliage branch, subtended by one or two pairs of spread-
ing scales; foliage leaves on young plants distichous and equally twisted
at the base, lanceolate with a blunt tip, the midrib marked below by 4
sharp narrow ridge and above by a slight groove, up to 30 mm. long by
6 mm. wide; adult leaves becoming not distichous but still equally turned,
coriaceous, the midrib marked by a raised strip narrower than the leaf
margins, the edges of the strip when drying appearing as two parallel
ridges on both leaf surfaces, ovate-lanceolate, 6-15 mm. long by 2.5-4
mm. wide; non-foliage branches with dispersed spreading scales which
are coriaceous, rounded, 1-2 mm. long on young plants and up to 4 mm.
long as reduced leaves on fertile specimens; pollen cones single in the
axils of foliage leaves, or from one to five or more at the apex of a
foliage branch, or in terminal or lateral groups on non-foliage branches
(not in compound groups), ovate, 4-6 mm. long (rarely to 12 mm. ) an
2.5—3 mm. in diam., microsporophylls short triangular with large spreading
edges to the open spore sacs; seed complex terminal on foliage branches
or on lateral scaly branches and involving 2—3 decussate pairs of spread-
ing scales or bracts followed by two unequal bracts one of which is fertile,
or rarely both are fertile and equal; micropyle of the inverted ovule at the
1969] DE LAUBENFELS, PODOCARPACEAE 345
end of an elongated beak extending about 2 mm. below the spreading
fertile bract, the fertile scale completely enveloping the ovule and form-
ing an apical crest which sometimes persists on the mature fruit; mature
eaten by some bird), the surface of the seed with low scallops and ridges.
Distr1BuTION. In rainforests throughout New Caledonia mostly from
750 to 1,450 meters, but also lower where lower rainforests occur. Prob-
ably the most common conifer in New Caledonia but always scattered in
the forest.
ew Caledonia. Ignambi, Compton 1524 s (BM), 1587 & (BM), Foster 160 j
(P, z). Mt. Panié, McKee 15594 2 1,000-1,400 m. (p), 15639 j 800 m. (Pp). Mt.
Tchingou, Hirlimann 1220 j 1,250 m. (Pp, z). Mt. Paéoua, McKee 17032 j 900—-
1,100 m. (P), 17056 2 (P), Bernardi 10131 s (Pp, z), 10149 s 900 m. (P, z). Mt.
Boulinda, McKee 17354 j 1,150-1,300 m. (P), 17357 ¢ (P), Veillon 120 j 1,100
m. (P). Mt. Me Maoya, McKee 13037 s 1,350 m. (P), 13492 j 1,400-1,450 m.
(Pp). Ridge W. of Col des Rousettes (Me Maoya), McKee 9886 2 800-900 m.
(K, P). Bourail, below Téné, Balansa 1381 2 (xk, Pp). Mt. Nekandi (Thio),
McKee 17908 j 1,200 m. (Pp). Dent de St. Vincent, oy 11 j (»). Mt. Hum-
boldt, Schlechter 15331 & 1,400 m. (BM, K, P, Z), 15332 2 (P), Buchholz 1578
$ 1,300 m. (ILL), Baumann-Bodenheim 15393 s ee m. eo z), 15411 s (z). Mt.
2), 11200 5 (p, z), 11301 s (P, z), Bernardi 9879 s (P, z), Blanchon 341 s (r).
Couvelée, Brousmiche 697 s (p). Mt. Dzumac, de Laubenfels P153 2 (sBT),
P415 2 760 m. (A, K, RSA), Baumann-Bodenheim & Guillaumin 12725 j (P, Z),
ries 2 800-900 m. (p). Upper Ouinné Valley, Bernier 267 j 750 m. (P), 268
Ae! Baumann-Bodenheim & Guillaumin 12815 s 700 m. ie 12843 s 700 m.
1573 j 530 m. (Pp, z), Bernardi 9445 9 600 m. (Pp, z). Bois de Mois de Mai
(Walker’s Place), Bernier 203 2, j (P), 269 j ), ie s (P), Buchholz 1350 s
ILL, es hy 1350A j (ILL, aul 1359 j (ILL, z=, &), 7 a ae P), 1367 s —
P
Bodenheim 15028 s (p, z). Inland from Bay of Pirogues, White 2120 j (a).
Without loc. Sarlin 228 j = a. 552 j (P).
ILLusTRATION. SARLIN, P., Bois et Foréts de la Nouvelle-Calédonie,
t. 26. 1954, as : Restor comptonii.
346 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The juvenile form of Decussocarpus comptonii has a great deal in com-
mon with the adult form of D. vitiensis and the ecology of the two is
identical. The adult form of D. comptonii is in a number of ways differ-
ent from its juvenile form. The fact that D. comptonii is strictly endemic
to New Caledonia while D. vitiensis extends for several thousand miles
and both to the east and to the west of New Caledonia is a clear illustra-
tion of the curiously isolated flora of New Caledonia. There are many
closely related species of conifers between the two areas mentioned, but
none are common to the two.
33. Decussocarpus minor (Carriére) de Laubenfels, comb. nov.
Nageia minor Carriére, Traité Conif. ed. 2. 641. 1867. Type: Vieillard 1275,
New Caledonia, Lake Arnaud.
Podocarpus minor (Carriére) Parlatore in DC. Prodr. 16(2): 509. 1868.
Podocarpus palustris Buchholz, Bull. Mus. Hist. Nat. Paris. II. 21: 284. 1949.
Type: Buchholz 1421, New Caledonia, Plaine des Lacs.
Small tree or shrub 2-3 m. high; bark tan to dark brown (often
stained with iron oxide from flood waters), very rough, fissured, fibrous,
slightly scaly, breaking off in short thick vertical strips or ragged frag-
ments; foliage branches opposite or alternate on non-foliage branches or
single to grouped at the apex of an older foliage branch, subtended by one
or two pairs of spreading scales; juvenile leaves distichous, equally twisted
at the base, not crowded, lanceolate, the midrib marked by a broad
raised area that may appear as three ridges when dry, up to 39 mm. long
by 3-4.5 mm. wide, on young plants smaller, more crowded, not distichous
but still equally twisted; mature foliage leaves almost imbricate and
crowded but still (in part) with a slight equal turning, coriaceous, the
midrib marked by a broad raised area, wider than the not raised margins
(the raised area upon drying either irregularly wrinkled or appearing as
three ridges), ovate, blunt, 7-20 mm. long by 2.5—-5 mm. wide; non-
foliage branches with dispersed spreading scales which are thick, rounded,
1.5-2.5 mm. long, smaller on juvenile specimens; pollen cones solitary
or clustered up to five or more, terminal and lateral in the axils of spread-
ing scales on a deciduous shoot at the apex of foliage branches, ovate,
4-8 mm. long by 2-2.5 mm. in diam., microsporophylls triangular with
an elongated point; seed complex terminal on foliage branches and in-
volving 2-3 decussate pairs of crowded spreading scales or bracts fol-
followed by two unequal bracts one of which is fertile (or rarely both
fertile and equal); micropyle of the inverted ovule at the end of a blunt
beak which extends up to 2 mm. below the fertile bract, the fertile scale
completely enveloping the ovule and forming an asymmetrical apical
crest which persists on the mature fruit; mature seed globular pear-
shaped, about 20 mm. long including the curved beak and 11-12.5 mm.
in diam., covered by the glaucous fleshy scale which sometimes becomes
deep red when ripe and after drying tends to crack and flake off the seed
(which may persist on the tree for some time or break off anywhere from
1969] DE LAUBENFELS, PODOCARPACEAE 347
the base of the fertile bract to the base of the fertile shoot), surface of
the seed rough and porous (making it very buoyant)
DistrisuTion. Along lake and river banks in shallow water over soils
derived from serpentine, in the headwaters of the Yaté River and along
small streams closer to the coast in the southernmost part of New Cale-
sa at low elevations (up to 200 meters).
ew Caledonia. Pirogues R., White 2261 @ (A, K, P, Us). R. Blanche (Mois
de MD, McMillan 5120 s 600 ft. (A, K, P), Baumann- Bodenheim 13923 s (P, Z).
Upper Yaté R. (22 km. station), Bernier 204 S,j (), 265 9, 5 @), 257 4, 3
(Pp), Buchholz 1347 s (Pp), 1348 9 (ILL, K, P), 1421 9 (111-holotype of Podo-
carpus palustris; K, P-isotypes), 1705 j (ILL, K, P), Sarlin 73 2 (P), de Lauben-
fels P112 2, 8, j (sz), P160 2 (ssr), Foster 200 8 (p). Marais Kiki (Yaté
R.), McKee 1118 s (A), 1119 2 (a, Pp), Baumann-Bodenheim 6370 j (z), Hiirli-
mann 3157 } (z). Creek Pernod, Guillaumin 8339 j (z), 8345 2 (z), Blanchon
1160 8 (Pp). R. des Lacs, bridge, Thorne 28565 2 (GH, Pp), Baumann-Boden-
heim & — 6511 5 (z), 6580, j (P, Z), 6766 & (P, z), Hiirlimann 3113
s (z). R. acs (near Madeleine Falls), Bernier 125 j (Pp), 246 s (P), 249 s
(P), 250 3 es Le Rat 2587 2 (pe), Buchholz 1474 2 (ILL), 1719 2 (ILL), 1729 2
(ILL, kK, p), Ddniker 228 2 (p, z), 228a 2 (z), de Laubenfels P340 Q (A, RSA),
P340A 8 (A, RSA), Baumann-Bodenheim & Guillaumin 11749 s (P, Zz), 11811 s (P
z), Stauffer 5807 2 (Pp, z), Blanchon 736 s (P). Plaine des Lacs in general, Le Rat
607 @ (BM, P), 751 2 (P), 1040 % (K, P), 2621 2 (P), McMillan 5139 & 600 ft.
ee K, P), Baumann- Bodenheim & Guillaumin 6582 J (Pp, z), 6594 s (Pp, z), Aubré-
(ear), McKee 3382 2 (a, Pp, us), Rohrdorf 178 2 (z), Bernardi 9369 4 (P, Z).
bi nd ao ‘ee 658 s (A, ?). Plaine des Lacs, lake bank, Franc 207 s (a, “n
K, P, z). L. Arnaud, Vieillard 1275 4 (p- holotype of Nageia minor). La
Chate, Pa 208 s (P). Prony Bay (B. du Sud), Vieillard 1275 (apparently
not the same as the previous collection of the same number) ¢ (A, BM, GH, K,
See Sartin, P., Bois et Foréts de la Nouvelle-Calédonie,
t. 27. 1954, as ps ened palustris.
The peculiar habit of this species which grows in shallow water with
a swollen base to the trunk (somewhat like a bald cypress), immediately
sets it apart. In general morphology it strongly resembles Decussocarpus
comptonii, but close inspection reveals a different leaf morphology and
slight differences in the pollen cone and the seed. This general similarity
Prevented the recognition of D. comptonii as a species for many years.
ADDITIONAL SPECIES:
Decussocarpus rospigliosii (Pilger) de Laubenfels, comb. nov.
Podocar pus rospigliosii Pilger, Notizbl. Bot. Gard. Berlin 8: 273. 1923. Type:
Esposto 556, Peru, Oxapampa (not seen, A photo.
348 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Section Dammaroides (Bennett) de Laubenfels, comb. nov.
Podocarpus section Dammaroideae Bennett ex Horsfield, Pl. Jav. Rar
1838. Type species: Podocarpus latifolia Wallich [Decussocarpus sie
ianus (Presl) de Laubenfels
Podocarpus section Nageia Endlicher, Syn. Conif. 207. 1847. Type species:
Podocarpus nageia R. Br. [Decussocarpus nagi (Thunb.) de Laubenfels].
Leaves opposite decussate or subopposite, multiveined with the veins
converging to the apex, obovate to elliptic, acute, distichous, rather large;
terminal buds small, bud scales acute; pollen cones linear, solitary or
grouped in the axils of leaves; seed cone on a scaly shoot with one or two
subterminal bracts fertile; ovule inverted and enveloped by the fleshy
fertile scale; seed globular with a slight protrusion at the micropylar
end close to the spreading fertile bract but on the opposite side from the
fruit attachment.
The multiveined leaves immediately distinguish section DAMMAROIDES
from the remainder of the genus but, without fruit, trees of this group
are very similar to the genus Agathis of the Araucariaceae. These are
distinguished by their globular terminal buds quite different from the
acute scales of buds in the section DammaromeEs. It has been cus-
tomary for some time to call this section Nageia, ignoring Bennett’s name
apparently because he used an improper ending. Bennett described his
section quite adequately and both Pilger (1903) and Wasscher (1941)
refer to his name without comment. Gordon (1858) and others treated
this section as a genus, Nageia Gaertner, a name which if accepted would
have priority not only for the genus being proposed here, but also in the
genus Podocarpus (if the section were to be retained in that genus). In-
deed, Kuntze (1891) did substitute Nageia for Podocarpus but Pilger
(1903) pointed out that the original description of Nageia confused its
characters with those of Myrica (“stam. quattuor et styl. duo.’”) and,
therefore, the name must be rejected.
There are five species differing in the presence or absence of a fleshy
receptacle, the number and position of the pollen cones, and the orientation
and size of the leaves.
KEy TO THE SPECIES OF SECTION DAMMAROIDES
1. Seed with a fleshy receptacle.
2. Pollen cones grouped on a peduncle; leaves at least 6 ag ie gos Saas
4 Dd. 0 llichianus.
1 EE Ct BES FE at te Sa U7 isa mn eg ea 35. D. motleyi.
Seed lacking a fleshy receptacle.
3. Leaves amphistomatic and equally turned, 20-31 cm. long. .....-----°°
36. D. maximus.
3. Leaves hypostomatic and unequally turned, less than 20 cm. long.
4. Pollen cone cluster sessile; leaves at least CM TON eos Pe
a9. fleuryi.
1969] DE LAUBENFELS, PODOCARPACEAE 349
4. Pollen cone cluster on a peduncle; leaves usually less than 6 cm. long
er Re ne Ga cis ee ee 38. D. nagi.
34. Decussocarpus wallichianus (Presl) de Laubenfels, comb nov.
Podocarpus latifolia Blume, Enum. PI. Javae 1: 89. 1827, nomen illegit., non
unb. 3 e: Blume s.n., Java, Mt. Salak.
Podocarpus latifolia Wallich, Pl. As. Rar. 26. 1830, nomen illegit. Type:
Wallich 6050, India, Mt. Sillet.
Podocarpus wallichianus Presl, Bot. Bemerk. 110. 1844, based on Podocarpus
latifolia Wall., which is a later homonym.
Podocarpus blumei Endlicher, Syn. Conif. 208. 1847, based on Podocarpus
latifolia Blume.
Podocarpus agathifolia Blume, Rumphia 3: 217. 1849, based on Podocarpus
latifolia Blume,
Nageia blumei (Endl.) Gordon, Pinetum 135. 1858.
Nageia latifolia (Wall.) Gordon, ibid. 138.
Nageia wallichiana (Presl) Kuntze, Rev. Gen. Pl. 800. 1891.
Podocarpus latifolia Blume forma ternatis De Boer, Conif. Archip. Ind. 14.
1866. Type: Teysmann s.n., Moluccas, Ternate.
Tree up to 48 m. high; bark smooth, peeling in large, thin, very irregular
plates, tan to brown within, weathering to dark brown or gray and develop-
ing scattered large lenticels and irregular longitudinal markings; leaves
decussate, distichous, amphistomatic, equally turned so that the lower
surface is exposed on the left and the upper surface is exposed on the
right side of the branch, many parallel vascular veins, elliptic, acute to
acuminate, sometimes abruptly narrowed to the short (5-10 mm.) petiole,
mostly 9-14 cm. long by 3—5 cm. wide but sometimes smaller to 6 cm.
long and 2 cm. wide or (particularly for juvenile and shade leaves) up
to 23 cm. long and 6.8 cm. wide, the extreme sizes (both smaller and
larger) mixed with more usual sizes on the same tree (both extreme
width and extreme length not usually together, the ratio of length to
width varying from 2 to 6, so that the narrowest leaves are not usually
the shortest while the longest are not usually the widest); terminal buds
often 2-3 mm. beyond the last pair of leaf bases (lateral buds sessile),
abruptly but slightly wider than the stem and then tapering, bud scales
acute-acuminate and erect; pollen cones 1—7 on an axillary scaly peduncle
2-10 mm. long with one cone terminal and the remainder in decussate
pairs, cylindrical, 8-18 mm. long by 3-4 mm. in diam., microsporophylls
lanceolate, 2-3 mm, long; seed cone on an axillary scaly peduncle, 8-20
mm. or more long, the lanceolate scales deciduous as on the pollen cone
peduncle; receptacle enlarged and becoming blackish and very fleshy upon
maturing, 7-18 mm. long, composed of 4 to 7 sterile bracts, the curled ends
of which protrude from the receptacle, and 1 or 2 subterminal fertile
bracts with inverted ovules; seed smooth, globular with a small beak
next to the point of attachment, completely covered by the thin fertile
scale which accompanies the ripe seed, 15-18 mm. in diam.
DistriputTion. In rainforests from eastern India to Normanby Island
350 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
east of New Guinea, often as a common forest element at low elevation
and extending as high as 1,575 meters (one collection at 2,100 meters).
Map 13.
India. Mt. Sillet, Wallich 6050 4 (A-isotype of Podocarpus latifolia Wallich).
E. o Griffith 3005 6 (GH, P). Khasia, Hooker & Thompson s.n. s 3,000
ft. (a, Apan 191 2, & (GH). Assam, King sn. & (a, L, Us). Thailand.
Temscein Falconer s.n.s (Lt), Kao Lu as sa Stritamurat, Kerr 15445 s 600
K). Tamtieng, Ranaung, Kerr 11770 2 200 m. (kK). Kapor, Ranaung, Kerr
pe 2 50 m. (K). Kumpuam, Ranaung, — {O86 2 50 m. (K). Kuabun,
Ranaung, Kerr 16351 s 50 m. (K). Lem Dan, Kaw Chang, Rabil 19 s (K).
Klaung Non Si, Kaw Chang, Kerr 9324 s 10 m. (K). Kao Kuap, Kuat, Kerr
17714 s 500 m. (K). Kao Ri Yai, Kanburi, Kerr 10400 s 1,400-1,500 m. (kK).
Baw Rai Kinat, Kerr 9509 s 300 m. (x). Adang, Sulut, Kerr 14132 s 500 m (K).
Khas Yai, 105 km. E. of Saraburi, King 555A 2 780 m. (L). Laos. Pak Mu-
nung, Wengchau, Kerr 21215 & 1,400 m. (x, Pp). Cambodia. Phnom San Kas,
Miiller 499 s (p). Elephant Mts., Poilane 23216 s 200 m. (Pp). Kre-dek, Poilane
oe s 600 m. (Pp). Cochin China. Phu Quoc I., Pierre 5529 s (A, NY, P), 5530
9 (A, P). Annam. Vinh, Linhcam, Chevalier 38234 s (Pp); Ke Bon, ‘Chevalier
38127 s (P). Quang-nam (S. of a Nang), Poilane 31558 s 500 m. (p). Massif
Ngok Guga near Dakto, Poilane 35675 s 1,000 m. (p). Massif du Brian near
Djiring, Poilane 24234 s (Pp), ye s 1,500 m. (Pp). Phan Rang (Can-Wa),
Poilane 5963 2 900 m. (a, P). W. of Ca Na, Evrard 2422 s 1,200 m. (A, NY, P):
Malaya. Jerai Reserve, Kedah, (Mal.) 17848 s (K). Kledang Saiang Reserve,
Perak, Mead 12828 2 (x), Noakes 20133 9 (k), 22147 s (x). Kinta, Perak,
Low 64 @ (k). = Scortechini sn. j (A), Kebal Ayam, Kurantan, Pahang,
Loh 15065 s (x). Soga, Johore, Ridley 11223 s (Kk). Labis, Sinclair 38991
s (L). Johore, G. ra Sinclair 10578 j (K, L, NY). Sumatra. Mt. Sibajak, E.
Kisaran, Peete 238 j (A, ¥, US). E. Coast, Asahan Yates 2554 2 (A, NY).
ogg? + 4A, 2) pao 8108 s (us). Tapanuli, Sibolga, NJFS 6b1357 j
m. oh pea Angkola, NIFS bb31536 s 600 m. (L). Benkulen, Redjang,
Sots bbE1084 s 800 m. (1), Renwarin bb2450 2 (L), NIFS bb8842 s 900 m.
(L). Palembang, Paaioann, NIFS bbE1106 s 15 m. (1). Palembang, Pasemah
land, NIFS TB200 s 1,200 m. (1). Mig srg: G. Pakiwang (Ranau L.), Van
Steenis 3754 s 1575 m. (L), Java. G. Lajung, Koorders 1261 s 150-250 m. (L).
G, Salak (Batavia), Anon. (1845) s (L), resis 24181 s 1,000 m. (x). Pre-
anger, Parakan Salak, Koorders 39402 s (K, L), 39403 s 1,000 m. (A), 39404 s
(L), 39406 s 1,100 m. (A), 39407 s 1,350 m. (a), 39409 s 1,000 m. (£1), 39413
s 1,000 m. (L), 39415 s (a). G. Sys gears Junghuhn s.n.s (L). G. Panga-
ee Junghuhn sn, 2 3,000 ft. (L). G. Gedeh, Anon. 204 s (LZ), hate 346
$ (1). Takokak, Koorders 1262 s (t), 1264 s (L), 11909 s (1), 39592 s 1,200
m. (A), 39596 s (a, L). Preanger, Tjipatudja, Backer 8866 s 450 m. (L). W With-
out loc., Blume sn. 2 (t-lectotype, Podocarpus latifolia Blume), 5.7. § (u),
Hasskarl sm. j (L), Junghuhn sn. 5 (L), Miquel sn. 8 (1). Karimata. Sung
Tajan, Teysmann 11598 g (1). Billiton. Riedel (1876) @ (#1). Sarawak. Lundu,
G. Gading, Anderson 15391 2 2,600 ft. (k, L). B. Mersing, in from Tatau,
Luang $2217 6 2 900 m. (x). Bintulu, Merurong Plateau, Brunig S9999 s 820
m. (L), Lawas, Brunig S$12083 s 1,000 m. (x). Mt. Majan, Clemens 21822 }
faa North Borneo. Penampang, Leaio- Castro 5989 s, 5,000 ft. (K, L).
1969 | DE LAUBENFELS, PODOCARPACEAE sot
Kinabalu, Clemens s.n. j 7,000 ft. (Ny), Chew & Corner RSNB 4878 2 5,000 ft.
(kK). Elopura, Sandakan, Keith A7 s 10 ft. (K). Gompa, Kudat, Balajadia 4055
s sea level (K). Tawau, Ampon A1652 2 (K, us), Martyn SAN 18453 2 50 ft.
(K, L, Ny), Meijer SAN 19547 s 30 ft. (L). Without loc. Wood 1244 i (A, ©),
sm. 8 (US), Burbridge s.n. j (K). Borneo. Tidung’s Land (SE. Borneo), NJFS
bb18217 s 4m. (a, L). G. Beratus (Balikpapan), Kostermans 7464 s 900 m. (A,
L), 7486 2 900-950 m. (A, K, L). Samarinda, Kostermans (1956) 8 low (tL).
Mahakam, Amdjah 51 j (L). Mt. Palimasen (Cent. Kutei), Kostermans (1954)
3 900 m. (K, L). Philippines. Cagayan, Curran 16738 2 (us), 17200 2 (Ny, US).
Mt. Sulu (Apayao Subprov.), Fenix 28348 2 (A, Ny, us). Mt. Moises (Isabella),
Ramos & Edano 46333 2 (A, NY). Baguio, Williams 1035 s (K, Ny, Us). Lamao
R., Mt. Mariveles (Bataan), Williams 399 s 2,200 ft. (Ny, US), 624 2 3,000 ft.
(NY), 752 2 2,000 ft. (wy), 753 2 1,800 ft. (A, K, NY, US), Barnes 147 @ (kx,
NY, US), 194 2 (xk, Ny, US), Copeland 244 2 (K, Ny, us), Whitford 1353 2 (K, NY,
us), Curran 17616 s (L). Mt. Giting-Giting, Sibuyan I., Elmer 12360 @ (A, K, L, NY,
us). Celebes. Usu (Malili), NIFS Cel/III-80 s (A, K, L), Cel/III-143 s 10 m. (A, L),
Cel/III-146 s 25 m. (kK). Tebetano (Malili), NJFS bb24489 j 450 m. (a, L).
Tawingana (Malili), N/FS bb25541 s 120 m. (Lt). Tamborano (Malili), NJFS
bb9696 s 600 m. (L). Gorontalo Buladu (Manado), N/FS bb15602 s 400 m. (Lt).
Poso, Majoa (Manado), NJFS 6b31500 s 700 m. (tL). Manado, Kolonodale,
Bakomtefe, N/FS bb31506 s 100 m. (a, L). Lapo Lapo, SE. of Kandari, Bec-
cart (1874) s (FI). Moluccas. Obi, Ahasrip 118 s (K, L, NY). Ternate, Teys-
mann s.n, } (L-holotype of forma ternatis). Morotai, Kostermans 1660 s 5
m. (A, L). W. Ceram, NJFS bb19647 s (L). New Guinea. VoGELKoP: Warsam-
son Valley (E. of Sorong), Moll BW 11652 s 30 m. (L). Kebar Valley, Tobie,
Schram BW 7951 s 740 m. (L), Smit BW 2314 s 650 m. (L), Sijde BW 5579 s
750 m. (L). Nertoi, Kebar Valley, Mangold BW 2350 2 750 m. (t). Kebar
Valley, Van Royen 5058 2 550-700 m. (kK, L). Sidai (W. of Manokwari),
Schram BW 1785 s (L), BW 6174s 5 m. (L), Koster BW 6705 s (L), BW 6760
2 10 m. (K, 1), BW 6922 s 10 m. (Lt), BW 6977 s 5-10 m. (L). Manokwari,
Menusefer BW 8180 2 140 m. (z). Baru (Teminabuan), Hallewas BW 944
S 10m. (L). Beriat (S. of Teminabuan), Schram BW 6021 2 10m. (L). SERUI
Biak, NIFS 6b30717 s 50 m. (A, L), 6b30779 s 80 m. (A, L), 6b30813 @ 50 m.
(A, L), bb30887 s 50 m. (L), Moll BW 9574 s 35 m. (L), BW 9589 s 35 m. (L).
Mios Num. I., NJFS 6630939 s 200 m. (A, L), 6630947 s 200 m. (A, L), 6630961
S 200 m. (A, L). Japen, Sumberaba, Koster BW 11121 s 8 m. (L). Japen,
Aisao, Schram BW 10596 s 200 m. (L). Japen, NIFS 6630260 s 300 m. (L).
West: Armina (Babo), Moll BW 12968 s (Lt). Agondo (Babo), Lundguist
bb32983 (264) s 20 m. (K, L). Esania, Borowai (Fak Fak), Stefels BW 3147 s
20 m. (L). Tiwara, Arguni Bay, Telussa BW 5158 s (L). Tairi, Kaimana, Lou-
500 ft. (k, x). Gabensis (Lae), NGF W41 j (a). Papua: Fly R., D’Albertis
(1877) 5 (Ft). L. Daviumbu (Middle Fly), Brass 7909 s (A, L). Sibium Range,
Pullen 5932 s 2,300 ft. (t). Oriomo R., Hart 5005 @ (A, K, L), Brass 5878 s
(A, K, L, ny, us), 5880 j (A, Ny, US), 5906 s 5-10 m. (a, Ny), White & Gray
352 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
(Cent. Div.), Brass 3962 j 500 m. (A, Ny). Milne Bay Dist., Womersley NGF
19298 2 1,200 ft. (K, L). Normanby I., Brass 25824 j 600 m. (a, L).
ILLUSTRATIONS. BLUME, C. L., Rumphia 3: ¢. 173. 1849 (as Podocarpus
agathifolia): Prrcer, R., Pflanzenreich IV. 5 (Heft 18): fig. 9A (as Podo-
carpus wallichianus) & fig. 9B (as Podocarpus blumei). 1903; Nat.
Pflanzenfam. ed. 2. 13: fig. 1344 (as Podocarpus wallichianus) & fig.
134B (as Podocarpus blumei). 1926; Koorpvers, S. H., & TH. VALETON,
Atlas der Baumarten von Java 3: t. 588. 1915 (as Podocarpus blumet).
Podocarpus wallichianus and P. blumei have generally been treated as
distinct species but are here being united under the name Decussocarpus
wallichianus, Wasscher (1941), who treated Podocarpus blumei, analyzed
the two taxa and concluded they were probably the same. The differences
reported, as thickness of leaf and acuminate leaf tip, are variations that
may occur even on a single plant depending on age and exposure. Further,
considerable variation in leaf dimensions are frequently found on single
herbarium sheets. By comparing the predominant sizes within any re-
gion, it appears that this species is, in fact, rather consistent throughout
its considerable range.
35. Decussocarpus motleyi (Parlatore) de Laubenfels, comb. nov.
Dammara motleyi Parlatore, Enum. Sem. Hort. Bot. Mus. Florent. 26. 1862.
Type: Motley 1300, Borneo, Bandjarmasin.
Podocarpus beccarii Parlatore in DC. Prodr. 16(2): 508. 1868. Type: Bec-
cari 2649, Sarawak, near Kuching.
Nageia beccarii (Parlatore) Gordon, Pinetum 2: 186. 1875.
Agathis motleyi (Parlatore) Warburg, Monsunia 1: 185. 1900.
Podocarpus motleyi (Parlatore) Diimmer, Jour. Bot. 52: 240. 1914.
Tree up to 40 m. high; leaves opposite or subopposite, amphistomatic,
distichous, coriaceous, elliptical, acute or with a small acuminate tip, nat-
rowed at the base to a short thick petiole 2-3 mm. long, somewhat
variable in shape, usually 3-5 cm. long and 15-22 mm. wide but reaching
7.5 cm. long and 28 mm. wide; terminal buds compact, tapering at first,
the scales rounded to lanceolate, acute to acuminate, 3-5 mm. long; pol-
len cones axillary, solitary, sessile, cylindrical, 15-20 mm. long when
mature and 5—6 mm. in diameter; microsporophylls lanceolate, keeled, 2
mm. long; seed cone on an axillary scaly peduncle 2-5 mm. long with
3-4 pairs of decussate deciduous scales, receptacle becoming fleshy (on
some specimens with nearly fully developed seed there is no enlargement) ,
8-12 mm. long, composed of 5 to 9 spreading sterile bracts, the single
subterminal fertile bract with an inverted ovule covered by the fertile
scale; seed smooth, globular, with a small beak at the micropylar end
near the point of attachment, 13-16 mm. in diam.
1969] DE LAUBENFELS, PODOCARPACEAE 353
DistrIBuTION. Mostly in low poorly drained areas but extending to
500 meters elevation in enna from Sumatra and Malaya to the
southern part of Borneo. Map.
Malaya. G. Tebu (Trengganu), Sinclair & Salleh SFN 40798 s (z). Lumut
Dindings (Perak), Hadden 16554 2 (xk), Symington 27841 s (K). Legari Melin-
tang, Dindings, Strugnell 16568 s (K). Johore, S. Kayu, Mawai-Temulang Road,
Corner 21341 s (kK). Sumatra. Barus (Sibolga), Tapanuli, NIFS 6bb29532 s 25
m. (A, L), bb31596 s 1 m. (A, L). Near Banjunglintjir, Palembang, N/JFS
m. (L).
Arch., Karimon, NJFS bb17229 s 150 m. (A, L). Belimbing, NJFS 5628495
S 6m. (A, L). Sarawak. Near Kuching, Beccari 2649 2 (r1-holotype of Podo-
carpus beccarii; K-isotype). G. Perigi (Lundu), Anderson 13304 & 1,000 ft.
(A, K). Simunjan, Drahman $0316 s sea level (L). Without loc., Foxworthy
353 2 (kK). Borneo (SE. part). Tidung’s Land, S. Lebakis, NIFS 6b18328
SS i, (A, 2). ale noe Motley 1300 s (k-isotype of Dammara motleyi).
Puruk Tjahu, Tahudjan, NJFS 6621151 2 500 m. (A, L).
ILLUSTRATION. WaSSCHER, J., Blumea 4: ¢. 4, fig. 11. 1941, as Podo-
carpus motleyi.
The extremes of leaf sizes in Decussocarpus motleyi and D. wallichianus
approach each other and the latter occurs throughout the range of the
former so that the possibility of confusion exists. The pollen cones are
diagnostic and the terminal buds lying directly above the last leaf attach-
ment help to differentiate the two. By observing the most common leaf
Sizes of a tree (or even a specimen), however, the species can be readily
Separated. The lack of enlargement of the receptacle on some specimens
(see NIJFS 12T1P185) merits further observation.
36. Decussocarpus maximus de Laubenfels, sp. nov.
Arbor parva, 4.5 m. alta; folia magna, decussata, coriacea, elliptica,
acuminata, basi magis rotundata, in petiolum perbrevem angustata, 20—
34 cm. longa, 6-9 cm. crassa; gemmae parvae, acuminatae; strobili fem-
inei ramunculum axillarem 12 mm. longum formantes; squamae ra-
musculi decussatae, 3-4 mm. longae, deciduae; receptaculum nullum;
semen globosum, diametro ca. 16-18 mm. Holotypus: Anderson 3361/7
(L), Sarawak, Sibu. Fic. 10.
DistriBUTION. In low elevation swamp forest of Sarawak and possibly
Sumatra
Sarawak. Sg. Assan, Naman F. R. (Sibu), Anderson 3361/7 @ 12 ft. (1-
holotype ; K-isotype)./) Sumatra. Silo Maradja (Ashan), Bartlett 6805 s (K, L
NY, Us). Between Ey Tombak and Taratak, Tanah Pe (ancien
L).
Decussocarpus maximus is unique in the genus in combining a lack of
a fleshy receptacle with amphistomatic leaves, although, as mentioned
above, some specimens of D. motleyi with very small leaves may also
JOURNAL OF THE ARNOLD ARBORETUM [ VOL.
wees iceml eat] Malis Sg
eid
HORA AW,
teh yew hes Hee oF
Depart, Sanaa at
may? Dts
' “etoearrae
+ Sandie, pan se ond
; Lovalite Ste —
Not Qegl tow, 26% ie peat
Det Godelier Coll wt ier
Deertuted be
A. R,. Anderson 3361/7 (1)
50
egies: 10. Decussocarpus maximus de Laubenfels, photograph of the holo-
type
1969 | DE LAUBENFELS, PODOCARPACEAE 355
combine these characters. The leaves of D. maximus are without ques-
tion the largest of any conifer, being approached at their lower limits
by juvenile leaves of D. wallichianus and of various species of Agathis.
The largest leaves described above belong to a fertile specimen. Aside
from leaf size, the leaf form and terminal buds correspond to D. wallich-
ianus but the seed is produced on a shoot that is not fleshy, The range
of D. maximus is included within the range of closely related species
and the Sumatra specimens, being sterile, are distinguished by their leaf
size only (Bartlett 6805: 15-20 by 6.5-9 cm. and blunt, possibly from
damage; Bartlett 8226 and Teysmann (Bangka): 20-22 by 7-8 cm.). In
having neither hypostomatic leaves nor a fleshy receptacle, D. maximus
(and perhaps a part of D. motleyi) exemplifies best, among the species
in section Dammaroides, the special characteristics of the genus.
37. Decussocarpus fleuryi (Hickel) de Laubenfels, comb. nov.
Podocarpus fleuryi Hickel, Bull. Soc. Dendrol. France 75: 75. 1930. Lecto-
type: Fleury 38017, Tonkin, Phu Tho.®
Tree to at least 10 m. high; leaves opposite, decussate, coriaceous,
hypostomatic with unequal turning so that the upper surface of the leaf
is always uppermost, elliptic, acuminate, narrowed at the base more or
less to a petiole, 8-18 cm. long and 3.5—5 cm. wide; terminal bud large,
somewhat beyond the nearest leaves, tapering sharply, but scales lan-
ceolate, acute, erect; pollen cones grouped in an axillary sessile cluster
of three and subtended by several pairs of overlapping, broad, keeled
scales, long cylindrical, about 3.5 cm. long and 4 mm. in diam.; micro-
sporophylls small, triangular, acute; seed cone on an axillary scaly not en-
larged peduncle, 15—20 mm. long; ovule inverted in the axil of a subter-
minal bract; seed globular, 15-18 mm. in diam. Fic. 11.
DistripuTION. In mountain forests from northern Annam to Kwang-
tung. Map 14,
Kwangtung. Naam Kwan Shan (Tseng Shing Dist.), Tsang 20123 ¢ (A, NA,
NY, US), 25273 @ (a), Tonkin. Phu Tho, Chevalier 38017 2 (v-lectotype).
Hoa Binh, Ste. forestier 8408 s (p-syntype). Phu Huo, Chevalier 37512 s (P).
Annam. Vinh, Nghe An, Fleury 30180 s (p-syntype). Near Hue, Poilane 29808
2 1,300-1,400 m. (ILL, Pp). Mt. Bana (25 km. from Tourane), Clemens 4190 s
(A, K, NY, P),
The much larger leaves distinguish sterile specimens of Decussocar pus
fleuryi from D. nagi, although both have hypostomatic leaves. Otherwise,
the female peduncle of D. fleuryi is longer, the pollen cones are longer,
and, particularly, the pollen cone cluster is sessile. The general form
of the leaf corresponds with that of D. wallichianus and even Hickel, in
the original description, included a specimen of the latter (Poilane 5963)
among the specimens he listed. However, the stomatic condition and
® Chevalier is given as the collector for this specimen on the sheet in Paris.
356 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
FLORA OF RWANGT UNG
Barta rts
Petccobu ‘Waeicbioon> Jtae}
ft. Bligh: fre
needy Sommon; thioxet. . Ble
Ne
n Kwan Shan Hi 1 it
f Viliag 4 H
Pies 11. Decussocarpus fleuryi (Hickel) de Laubenfels, photograph of
sang 252 (A).
1969] DE LAUBENFELS, PODOCARPACEAE 557
orientation of the leaves at once distinguish even sterile specimens. The
sessile pollen cluster and the lack of a fleshy receptacle also separate these
two species.
38. Decussocarpus nagi (Thunberg) de Laubenfels, comb. nov.
Myrica nagi Thunb. Fl. Japon. 76. 1784. Type: ex herb Thunb., microfiche
no. 23381 (AGH).
Nageia japonica Gaertner, De Fruct. et Sem. 1: 191. 1788 (in part, nomen
illeg., description confused).
Podocarpus nag ela R. Br. ex Mirbel, Mem. Mus. Paris 13: 75. 1825 (based
japonica).
nec. ees Endl. Syn. Conif. 207. 1847. Hort.
Nageia cuspidata (Endl.) Gordon, Pinetum 136. 1858.
Nageia ovata Gordon, Pinetum, Suppl. 42. 1862. Type: Fortune in 1861,
Japan, Yeddo (not seen).
Podocarpus nageia R. Br. var. rotundifolia Maxim. Gartenflora 13: 37. 1864
(based on Nageia ovata Gordon
Podocarpus nageia R. Br. var. angustifolia Maxim. ibid. Hor
Podocarpus ovata (Gordon) Henk. & es et Nadelh. ae 1865.
Dammara veitchii Henk. & Hoch. ibi H
es japonica (Gaertner) Nelson, ge ee 1866, nomen illeg., non
Sie
Podo saree caesius Maxim. Meél. Biol. 7: 561. 1870. Hort.
Nageia nagi (Thunb.) Kuntze, Rev. Gen. Pl. 798. 1891.
Podocarpus nagi (Thunb.) Makino, Bot. Mag. Tokyo 17: 113 .
P oo nagi (Thunb.) Makino var. rotundifolia (Maxim.) ee ibid.
P ais nagi (Thunb.) Makino var. angustifolia (Maxim.) Makino, ibid.
Podocarpus formosensis Diimmer, Gard. Chron, III. 52: 295. 1912. Type:
Schmiiser 1357, Formosa, S. Cape (not seen, photo of type accompanies
descript.
Podocarpus “nankoensis aig, Ic. Pl. Formos. 7: 39. 1918. Type: Hayata
in 1916, Formosa,
Podocarpus nagi (Thunb.) oo var. koshunensis Kanehira, Trans. Nat.
Hist. Soc. Formosa 21: 145. 1931. Syntypes: Mori 25075, Sasaki 25076,
25077, Kanehira 26078, 22239, Matsuda 2594 (not seen).
?
Podocarpus koshunensis (Kanehira) Kanehira, Formosan Trees. Rev. ed. 36.
1936.
Tree to 25 m. high; bark smooth, peeling in thin flakes, dark brown
weathering gray; leaves decussate, distichous, hypostomatic, multiveined,
elliptic, acuminate, to rounded at the tip, the apex often showing evidence
of aborted growth, sometimes abruptly narrowed to a short wide petiole,
often glaucous especially on the underside, 4.5—5 cm. long or sometimes
longer, 10-20 mm. wide, somewhat variable in size and shape even on
individual specimens; terminal bud often 1-2 mm. beyond the last pair
of leaves, abruptly wider than the stem and then tapering to an acuminate
apex, bud scales long lanceolate; pollen cones 1-5 on an axillary scaly
peduncle 3-10 mm. long, subtended by a lanceolate scale up to 6 mm.
long, cylindrical, 10-20 mm. long, the longer ones terminal in the cluster,
358 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
microsporophylls small, acuminate, widely spreading and not crowded,
1 mm. long; seed cone on an axillary peduncle with deciduous lan-
ceolate scales, peduncle 5—10 mm. long, not enlarged; one or two seeds
developing from inverted ovules (rarely there are 3 ovules) in the axils
of subterminal bracts, completely covered by the seed scale, globular and
elongated into a hooked beak at the micropylar end, smooth, glaucous,
the seed itself 12-13 mm. wide and 15-16 mm. long, the fleshy bluish-
black covering at least 2 mm. thick but drying on the seed and wrinkling,
the seed often falling with the peduncle attached.
DIsTRIBUTION, Scattered from southeastern China and Hainan to
southern Japan in forests at low elevation, up to 800 meters in more
southerly parts. Because of the high degree of disturbance of forests al-
most throughout its range and its popularity in cultivation, it is most
difficult to distinguish between plants naturalized from cultivated sources
and truly native individuals. Probably some of the specimens cited are,
in fact, cultivated even where not indicated as such. Map 15.
China. Hatnan: Pak Shik Ling (Cheng Mai Dist.), Lei 745 2 (A, L, NY, US).
Pak Shek Shan (Lam Ko & Cheng Mai Dists.), Tsang 681 (L.U. 17430) s (A,
L, NY, Us). Nodoa, Sha Po Ling, McClure 8131 2 800 m. (A). Manning, How
73876 s 700 ft. (a, P). Kwancost: Ta Tse Tsuen, Yung Hsien, Steward &
Cheo 728 2 380 m. (A, NY, P). Sup-man-ta Shan, Liang 69401 s (A). Wah
Kong (Hing On Dist.), Chung (Tsoong) 83667 s (A), Kiancst: Tung Lei
(Kiennan Dist.), Lau 3964 2 (a, us). FuKiEN: Hinghwa, Chung 924 @ (A).
Yenping, Chung 2979 9 (A), 3570 s (A), Dunn 3523 Q (A). CHEKIANG: Ping-
yang Hsien, Ho 1554 @ (a). Tsingtien, Keng 99 @ (a), 20-40 miles W. of
Wenchow, Ching 1832 2 250-450 m. (a, Pp, us). Yentang Shan, Chiao 14685
s (A, NY, Us, z), Hu 241 2 (a). Without loc., Chen 4091 2 (a). Formosa.
SOUTHERN: Koshun (=Hengchun), Kanehira 28 s (a), 29 s (A). E. Coast:
Hualien, Kuntz 084 4 (us). Nanwo (Karenko Prov.), Wilson 11109 s (A, US).
=
sj
ze
5
aN
=
=
=
a
@
by
S
i~
wn
iy
5S
i
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Nm
mn
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mn
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S
=
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=>
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=
&
SS
8
Qa
Q
R
@
aS
=
a
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=
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& Kasugidani), Moran 5351 s 350 m. (us).
ILLUSTRATIONS. PitcER, R., Pflanzenreich IV. 5 (Heft 18): fig. 9C-E.
1903; Nat. Pflanzenfam. ed. 2. 13: fig. 134C-E. 1926, as Podocarpus
nagi; Dimmer, R., Gard. Chron. III. 52: ¢. 132. 1912, as Podocarpus
formosensis; KANEHIRA, K., Formosan Trees, rev. ed. t. 4. 1936, as Podo-
carpus koshunensis.
1969 | DE LAUBENFELS, PODOCARPACEAE 359
The large number of specific names which have been applied to De-
cussocarpus nagi, including several horticultural names, result from long
aquaintance with it in cultivation, and from its wide distribution. Most
of the differences noted for the various units proposed are within the
normal variation of a population or even of an individual. The variety
rotundifolia (equals Nageia ovata) of Podocarpus nagi possibly is defen-
sible as a distinct taxon, having leaves ovate, compared to the usual
elongated outline.
Section Afrocarpus (Buchholz & Gray) de Laubenfels, comb. nov.
Podocarpus section Afrocarpus Buchholz & Gray, Jour. Arnold Arb. 2
Type species: Podocarpus falcatus (Thunb.) R. Br. stearate
falcatus (Thunb.) de Laubenfels ].
Decussocarpus falcatus (Thunb.) de Laubenfels, comb. nov.
Taxus falcata Thunb. Prodr. Pl. paige 117. 1800. Type: Thunberg in
1773-1774, Cape of Good Hope (not s
ae Jateatu (Thunb.) R. Br. ex Mitb, Mém. Mus. Hist. Nat. Paris
aaa elo Stapf, Fl. Trop. sue “= Prain] 6(2): 343. 1917.
Type: Nelson 423, Transvaal, Houtschber
Decussocarpus gracilior (Pilger) de Laubenfels, comb. nov.
Podocarpus gracilior Pilger, sama IV. 5 (Heft 18): 71. 1903. Type:
Schimper 1160, Ethiopia, Che
Decussocarpus mannii (Hook.) de Laubenfels, comb. nov.
Podocarpus mannii Hook. Jour. Linn. Soc. 7: 218. 1864. Type: Mann 1065,
St. Thomas Island.
Nageia mannii (Hook.) Kuntze, Rev. Gen. Pl. 800. 189
Podocarpus usambarensis Pilger, ae la 5 "Glett 18): 70. 1903.
Lectotype: Holst 2467, Tanganyika, Usa
Podocarpus dawei Stapf, Fl. Trop. Afr. red. one 6(2): 342. 1917. Type:
Dawe 961, Uganda.
LITERATURE CITED
BucHuotz, J. T. Embryogeny of the Podocarpaceae. Bot. Gaz. 103: 1-37.
1941,
Dr orig otis D. J. Parasitic conifer found in New Caledonia. Science 130:
- 1959.
————.. The primitiveness of polycotyledony considered with special reference
to the cotyledonary condition in Podocarpaceae. Phytomorphology 1
296-300. 1962.
. Podocarpus vitiensis in the Moluccas (Taxaceae). Blumea 15: 440.
Florin, R. Untersuchungen sur Stammesgeschichte der Coniferales und Cor-
__ dhital les. Sv. Vet-akad. Handl. III. 10: 1-588. 58 pls.
————.. The Tertiary fossil conifers of south Chile and their phytogeographical
significance, Ibid. 19: 1-107. 6 pls. 1940.
360
JOURNAL OF THE ARNOLD ARBORETUM
[voL. 50
Grsss, L. S. A oboe i to the phytogeography and flora of the Arfak Moun-
tains. London.
Gorpon, G. The satay London.
1858.
Hooker, J. D. Podocarpus pectinata. bb5671,
bb6737, bb7077 (12a): bb7708,
bb8737 (21b); bb8842 (34): bb9003
(21a); bb9664 (17); bb9671 (1);
bb9696 (34); bb10748 (12a)
bb11803 (21a); bb13633 (26):
bb14390, bb14519 (6a):
bb15154. (1): bb15155 (21b)
bb15504 (2la);
bb17229 (35); bb17269 (21a);
[voL. 50
bb17544 (17); bb18217 (34);
bb18328 (35); bb18752, bb19559
(21b); 6619564 (1); bb19647 (34);
bb19709 (6a); 6b19869, bb19870
(4a); bb20202 (21a); bb20270 (1);
bb20535. (6a); bb20782 (1);
bb21151 = (35);
bb21294 (17); bb21509 (1) ; bb21511
(3); 0b24489 (34); bb24777
(1); 6624778 (17); 6b24779 (12a);
bb24934 (6a); bb24951, bb24956
(21b); bb24957 (21a); bb24958
(26); bb24964 (3); bb25157 (17);
bb25541 (34); bb26288 (3);
bb26589, bb27736 (5a); bb28147
(21b); 6b28495 (35); 6b28751,
bb28752, bb28753, bb28754 (5a);
bb29195 (21a); 0bb29532 (359%
bb30260 (34); 6b30321, bb30475
(6a); 6b30717, bb30779, bb30813,
bb30887, bb30939, —-bb30947,
bb30961, bb31500, bb 31506,
bb31536 (34); bb31596 (35);
6632284, bb32434, bb33046 (5a)
New Guinea Forestry Department
(NGF), the following by anony-
mous collectors: NGF-W41 (34);
NGF 3128 (21c); NGF 4503 (34)
Nicholson SAN 17292 (5a); SAN
17823 (1); SAN 17826 (14); SAN
17827 (13); SAN 39766 (25);
Nicolson 1319
Noakes 20133, 22147 (34)
Nur 10507 (17)
Ocampo 27926 (24)
Oillerings 175 (21a)
Omar SFN 376 (Sb)
Palmer & Bryant 988 (21a)
Pancher 4 (22); 379 (18); 380 (8)
Pascua 15692 -
Petit 138 (33); 177 (8)
Phengkhlai 568 nie 691 (2)
Pickles 2991 (2)
Pierre 1396 (2); 5528 (21b); 5529
5530 (34); se (2)
Pitard 2090
Poilane 25 (2); 320 (21b); 1539 (2);
2147, 3387 (21b); 3455, 3782 (2);
4038 (21b); 4411 (2); 5963 (34);
1969]
6509 (21b): 7095 (2); 9103, 10293,
16092,
23118 (21b); 23216, et 24314
(34); 29808 (37); 29960 (21d);
31558 (34); 32825, 33351 (2);
35595 (21b); 35675
Poore 6228 (17)
Posthumus 2175 mand i (21a)
Pringo Atmodjo 82 (21
Pulle 663 (3): 801 oa 964 (26);
966 (3); 982, 1018, 1042 (1)
Pullen 273A (1); 313, 313A (26);
338 (28); 2674, 2680 (6b); 2716,
2716A (23); 2840 (31); 5052 (26):
5111, 5138 (28); 5267 (26); 5914,
5930 (21c); 5930A (3); 5932 (34);
611
Purseglove P5006, P5553 (5a)
Quisumbing & Sulit 82404 (1); 82481
(21c)
Rabil 1
Rabor 20482 (17); 20485 (13)
pate 19557 (24); 77401 (1)
os & Edafio 26394 (17); 26501
Raap on 768 (21a)
9 (34
(12); 37757, 38738 (1); 45005
(24); 46333 3 (34)
Ramos & Pascasio 34497 (5a)
Rappard BW 697 (6b); BW 698
)
Rashid $9546 (5a)
Rensch 1307 (21a)
Renwarin 5b2436 (21b); 662450 (34)
Richards 1058 (1); 1059 (12a); 1628
(1); 1768 (21a); 1808 (12a); 1834,
1836 (17); 1962 (2): ao (12a);
1997 (10); 2421, 2476
Ridley 5695 (17); "8636 «ib, 11223
34); 16026, 16178
Robbins 238 (26); 598 (3); 673 (26);
718 (8). 3112 (26); 3214 (27);
3266
Beene 7
Rossum 122, 784 (5a)
Sadau 42890 (21a)
DE LAUBENFELS,
PODOCARPACEAE 367
Salverda 6622564
bb22576 (6b)
Santos 31817 (21c)
(1); 6622571,
)
Sarlin 73 (33); 228 (32); 229 (18);
237 (22): 242 (15); 244, 341 (22)
Saunders 708 (28); 804 (26); 823
(21c); 824 (1); 861 (21c); 1025
(1); 1048 (21c); 1088 (34)
Sayers NGF 21613 (21c
pen 1473, 1474 (21b); 1475
ations 15175, 13176 (8); 19331,
Schmid 137 (29)
Schodde 1561 (21c); 2014 (1); 2021,
26
Schram BW 1785, BW 6021, BW 6174,
BW 6705, BW 6760, BW 7951 (34);
BW 7972, BW 9271 (6a); BW
10596 (34)
Seemann 573 (6a); 576 (31)
Shockton 2699 (1
Sijde BW 5579 (34); BW 5596 (6a)
Sinclair 10578, 38991 (34); 39094 (2)
Sinclair & Kadim 9053 (1); 9146 (25);
10318 (5a)
Sinclair & Salleh SFN 40798 (35)
Sing JC/59 (Sa)
Singh SAN 24336 (Sa)
Skottsberg 202 (32)
Smit BW 2314 (34)
Smith, A. C. 1773 (6a); 1796 (31);
4122 (30); 4901 (21b); 5734, 6244
(6a); 6245 (21b); 7076 (31)
Smith, L. S. NGF 1352 (6a)
Smith, R. 66 (38)
Smitinand 19058
(
S10601 S10607
2)
(21c);
Sonohara, Tawada, & Amano 6290
(38
Spurway 376
Stauffer 5651 (1); 5652 (21c); 5670
(28); 5729 (18); 5807 (33)
Stauffer & Blanchon 5812
Stauffer, Blanchon & Boulet 5778 (22)
Stauffer & Kuruvoli 5841 (6a)
Stefels BW 2006 (21c); BW 2008,
BW 2010 (1); BW 2014 (21c);
368
BW 2015 (3); BW 2031 (1); BW
2033 (3); BW 2038 (21c); BW
147 (34
Steiner 2032 (24); 2150 (1); 2207
24
Stern 2242 (21c)
Stern & Rojo 2289, 2292 (21c)
Steup bb23045 (6a )
Steward & Cheo 728 (38)
Storck 906 (6a)
Stresemann 125 (26); 133 (1); 158
(21b); 251, 276A (26); 354, 363
21b); 395 (3)
Strugnell 16568 (35); 23931 (21b)
Sulaiman 2 (5a
Sulit 7586 (21c);
(21b); 10052, es (1);
(12a); 30051 (24)
Surbeck 107 (12a); 532 (21b)
Symington 27841 (35)
7669 (1): 9896
21694
Tagei 1795 (5b)
Tang 438 (21b); 457 (5a)
Telussa BW 5158 (34
Teysmann 169 (1 617 (Sa);
11598 (34); 11599 (5a): 21647 (2)
Thailand Royal Forest Department
631 (2
Thorenaar 12713 (35)
Thorne 28565 (33); 28568 (8);
28644 (18): 28704 (29): 28705
(18); 28734 (9)
Toropai NGF 17153 (1)
Tothill 553 (6a); 844, 845 (31); 854
6a
Toxopeus 427 (3); 485 (21b)
Tsang 681, L.U. 17430 (8); 20123,
25273 (37); 27332 (21b
Tsang & Fung L.U. 18100 =
Tuckwell W1553 (26)
Van Romer 736 (26); 1233 (12c)
Van Royen 3721 (1); 3857 (3): 3873
(1); 3895 (21c); 5058 (34); NGF
16182 (1); NGF 20289 (23); NGF
20309 (26)
Van Royen & Sleumer 6073 (31
6246 (6a); 7219 (3); 7403 (1).
7948 2); 79484 (21c); 7948B (3):
8203A (19)
JOURNAL OF THE ARNOLD ARBORETUM
[voL. 50
Van Royen, Sleumer, & Schram 7791
(12
a
Van Steenis 3754 (34); 8357 (12a);
23 (21d); 17544, 18267 (21a)
Vaughn 3254 (31); 3258 (21b)
Veillon 120 (32); 136 (29); 142 (22);
145, 511
Versteegh BW 248 (1); BW 250
(21c); BW 253 (1); BW 269 (3);
W
7596, BW 10407 (1); BW 10411
BW 12610 (21b);
15249 (6a)
Versteegh & Kalkman BW 5594 (6a)
Versteegh & Koster BW 14 (21b)
Vidal 623 (24); 3910 (13)
Vieillard 1259 (18); 1260, 1261, 1262
(22); 1275 (33); 1277 (8); 1278,
3262 (7); 3264 (32); 3265 (9)
Vink NGF 12430 (26); BW 15271
(6a); 17188, 17242 (26); 17499,
17500, 17501, 17502 (27)
Vink & Schram BW 8620 (6b); BW
8667 (23); BW 8730 (31); si
8731 (21c); BW 8746 (3);
8764 (1); BW 8796, BW i
(6b); BW 8945 (1)
ae oh 9 (9); 10 (29); 37 (7);
8 (32); 39 (18); 40 (9); 152
fy. 187 (7); 206 (22); 400 (7);
469 (18); 658 (33)
Wakau 4155 (2)
Walker, F. S. BSIP 212 (31); BSIP
247 (13
Walker 70 (2); ANU 859, ANU
859A (1); 5649 (38); 7526 (21c)
Wallich 6045 (2); 6050 (34)
ang 33651 (Sa); 35591 (21b);
36532 (5a); 39608 (21b)
Warburg 11119 (21a); 14721 (21b)
White 2001 (7); 2033 (32); 2112
(22): 2120 (32); 2122 (8); 2238
(7); 2261 (33); 2285 (22)
White & Gray NGF 10407 (6a); GF
10415 (34)
Whitford 951 (24); 1353 (34)
Whitmore 2368 (21b)
1969]
Williams 399, 624, 752, 753, 1035
(34); 1298, 1299 (21c
Wilson 6262, 8064, 10242,
11109
Winkler 512 (1); 1035 (21a);
(1); 1037 (13); 1866 (21a)
Womersley NGF 3704 (34); NGF
4420 (3); NGF 4428 (23); NGF
4483 (19); NGF 5338 (21c); NGF
10279,
1036
11067, NGF 11260 (21c); NGF
13922 (19); NGF 14018 (26);
NGF 14253 (21c); NGF 17621
(34); NGF 17902 (1); NGF 17939
(23): NGF 19298 (34); NGF
24563 (21c); NGF 24569 (26);
NGF 24928 (21b)
Womersley & deLaubenfels
19460 (21c)
Womersley & Floyd NGF 6138 (21c)
NGF
DE LAUBENFELS,
PODOCARPACEAE 369
Womersley & a NGF 7680 (3);
NGF 8324
Womersley & Sones NGF 14013
21¢
Wood 1244 (34); SAN 4172 (5b);
SAN A4179
Wood & Wyatt- ea SAN A4493 (25)
Wray 1028 (2); 1198 (21b); 3875
(12b); 3899
Wray & Robinson 5354, 5380 (2)
Wyatt-Smith 71650, 80370, 80371 (1);
93115
Yapp 493 (12a)
Yates 1987, 2148 (21b); 2554 (34)
Zollinger 2262 (21a); 3025 (34)
Zwart 6517 (21d)
DEPARTMENT OF GEOGRAPHY
SYRACUSE UNIVERSITY
Syracuse, New York 13210
370 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
THE VASCULAR SYSTEM IN THE AXIS OF DRACAENA
FRAGRANS (AGAVACEAE),
1. DISTRIBUTION AND DEVELOPMENT OF PRIMARY STRANDS.
M. H. ZIMMERMANN AND P. B. ToMLINSON 1
WE HAVE OUTLINED THE STATE of existing knowledge of the vascular
anatomy of monocotyledons with secondary growth in a previous article
which serves as an introduction to our present studies (Tomlinson &
Zimmermann, 1969). These have been concerned with a number of genera
in the Agavaceae. There are quantitative and often diagnostic anatomical
differences between the plants we have studied, but we believe that funda-
mental principles of vascular distribution is the same i all of them. We
have therefore restricted our description to one species, Dracaena fragrans
(L.) Ker-Gawl., of which we had abundant living material available for
a detailed study. The same species had also been used by earlier investi-
gators of this problem (e.g., Cordemoy, 1894; Meneghini, 1836; von Mohl,
1824) although often under the older, now incorrect name of Aletris fra-
grans L. Where necessary, however, we shall refer to other plant species.
For convenience, our results are presented as two separate articles, devoted
to primary and secondary tissues respectively, although such a separation
is somewhat arbitrary. Indeed, we will have cause to show that the two
types of vascular tissue are interdependent and often continuous.
MATERIALS AND METHODS
Sectioning. Specimens investigated were collected from a large clump
cultivated at the Fairchild Tropical Garden. Material was either sec-
tioned freshly or after fixation in FAA and subsequent washing. For the
study of the course of vascular bundles sequential series of tranverse
sections were cut on a “Reichert” sliding microtome from two lengths of
distal mature shoots, each representing a sympodium. Selection of ma-
terial needed some care because in some shoots the central tissue is easily
torn during sectioning. For the investigation of the vascular anatomy of
leaf-trace departure a complete series of sections 40» thick was prepared.
For the analysis of longitudinal continuity of vascular traces sections 33p
thick at intervals of 100. were used. These sections were stained in
safranin and Delafield’s haematoxylin and mounted permanently in
“Piccolyte.”’
For the study of vascular development in the crown, serial transverse
and longitudinal sections 10% thick of shoot apices were cut on a rotary
* Contribution to a study of the vascular system of monocotyledons by one of @
(P.B.T.), supported by N. S. F. grant GB-5762-X.
1969 | ZIMMERMANN & TOMLINSON, DRACAENA, 1 371
microtome from material embedded in “Paraplast.” Routine methods of
embedding, staining and mounting were employed. Because of the wide
diameter of these developing crowns, pieces of ribbon containing only
four sections were mounted on each slide.
Serial analysis. Cinematographic analysis of the three-dimensional
vascular structure was carried out with the series of sections from the
mature stem. These methods have been described in ample detail in
previous papers (Zimmermann & Tomlinson, 1965, 1966). The method
of plotting provascular strands in the series of sections from the meriste-
matic crown has also been described in previous papers (Zimmermann &
Tomlinson, 1967, 1968). The optical shuttle was employed for plotting,
according to the procedure described in the paper on the vascular develop-
ment of Prionium (Zimmermann & Tomlinson, 1968). A slight methodical
variation was necessary because each slide in this series contained four
sections. Most strands can easily be followed by matching the corre-
sponding section on two successive slides with an interval of 3 sections
between. In areas where provascular strands make sharp turns (immedi-
ately below the apical meristem) and require the use of each section,
optical alignment was achieved in the following way: 5d—6a, 5d—6b, Sd—
6c, 6c—7a, 6d-7a, etc., whereby the number indicates the slide, the letter
the section on the slide, and the italics mark the sequence of photography.
The reader can easily appreciate that alignment had to be achieved over
an interval of two sections (e.g., 5d-6c above) once every four sections.
With the Dracaena crown this was just possible without losing continuity
of the strand which was plotted. There is no question that the procedure
is easier to follow when each slide contains only a single section.
GENERAL MORPHOLOGY AND ANATOMY
Growth habit. Dracaena fragrans, a native of tropical Africa, is a com-
mon ornamental in South Florida. It has apparently been known in cul-
tivation in Europe at least since 1768 (Sims, 1808). In cultivation it
forms a diffuse shrub or rarely a low tree. Basal shoots are straight and
erect, but distal branches are often bent over by the weight of the
terminal cluster of leaves. Leaves are lanceolate, up to 75 cm. long, and
lack a petiole. Their insertion is open but broad and even overlapping.
On vigorous shoots, leaves persist for a long time, so that leafy shoots up
to 2 m. high may be present. Otherwise on suppressed and less vigorous
shoots leaves form the distinct terminal cluster which is so common in
many other woody monocotyledons. The longevity of leaves is of con-
siderable anatomical significance, a point which will be discussed later.
Each leaf subtends a minute axillary bud enclosed by its prophyll.
Inflorescences are always terminal (Fic. 1) and branching is closely
associated with flowering. Flowering and resultant branching begins in
plants 2 to 3 years old when they are about 1 m. high. The transition from
vegetative to reproductive state of the axis is marked by a gradual reduc-
372 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Fic. 1. Dracaena fragrans, Distal part of a flowering shoot, X 1/5.
tion in leaf size associated with an abrupt elongation of internodes (Fics.
2-5). Bracts which subtend the first-order flowering branches are white
and caducous although it is obvious from the transition upwards along
the shoot that they are homologous with foliage leaves. The detailed
structure of the flower-bearing branches has been described by Troll
(1962).
Branching is normally sympodial from a bud in the axil of one of the
transitional leaves immediately below the inflorescence. At the time of
anthesis this renewal bud is indistinguishable from other dormant buds
(Fic. 4). After flowering it grows out rapidly (Fic. 7), pushing the
terminal inflorescence into a pseudolateral position. Each axis branches in
this way and is, therefore, a sympodium consisting of many successive
growth units. The sympodium appears articulate both from the scars of
the dried inflorescence stalks, and a swelling which marks each joint
(Fic. 3). This articulation is most noticeable in distal, horizontal axes.
After flowering there is an evident competition among a number of
potential renewal buds, because the inhibition of more than one is always
released (Fic. 7). One of them usually becomes dominant and re-imposes
inhibition upon the others. However, two (rarely more) buds may grow
out simultaneously. This leads to a branched axis with an inflorescence
scar in the crotch of the fork. Plants in South Florida flower several times
in one year and although it is obvious that different axes flower at dif-
ferent times of the year it seems likely that a vigorous shoot can flower
1969]
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3, rie, . 2-5. Dracaena fragrans. Habit details. 2, Terminal inflorescence, < 1/2.
b ow ering shoot with all leaves detached, showing 3 units of sympodium and
ae uation of vegetative axis, X 1/2. Letters
Ciied i. & m distal unit shown in Fic. 3, their levels of insertion indi.
€d in that figu - . besinning with prophyll (a) and ending with most distal
transitional leaf ( (f), x
374 JOURNAL OF THE ARNOLD ARBORETUM [voL, 50
Fics. 6 and 7. Dracaena fragrans. Branching. 6, Erect axis with 2 branches,
renewal shoots at A and B from axillary buds whose inhibition is released by
destruction of apex of parent shoot at X, 2/3. 7, Normal renewal growth
below old inflorescence axis, X 2. Buds developing in axils of two most distal
coed ; prophylls conspicuous. This corresponds to Fic. 4 after lapse of 2
months.
two or three times each year. This is indicated by a close succession of
inflorescence scars. For a further morphological description of flowering
in arborescent Liliiflorae the reader is referred to the detailed work of
Schoute (1903, 1918).
The release of apical dominance is normally the result of flowering but
it may be induced in other ways. Decapitation releases from inhibition
the dormant buds immediately below the injury (Fic. 6). Apical domi-
nance is also released on the upper side of leaning stems where numerous
dormant buds may grow out, much as in woody dicotyledons (see Fig. 15
in Tomlinson & Zimmermann, 1969). Erect, rapidly growing suckers
commonly develop from the base of old plants, presumably for the same
reason. The influence of these various methods of growth on the distribu-
tion of vascular tissue is largely described in the second article of this
series.
Primary tissues (Fic. 8). Epidermis slightly thick-walled, covered
by a thin but conspicuous cuticle. Periderm in hypodermal or subhypo-
dermal layers developing early by etagen-like divisions of cortical cells,
the outermost derivatives suberized. Cortex, 1-3 mm. wide, of uniform
and fairly compact parenchyma; no independent cortical vascular system
. \ on
1969] ZIMMERMANN & TOMLINSON, DRACAENA, 1 375
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agrans, “ 36. All bundles in cortical area are leaf traces. Arrows point out
vertical bundles immediately above point of branching from leaf trace. They
appear small because they lack the fibrous sheath.
G. 9 (BELow). Transverse section of mature stem of Dracaena fragrans,
taken from the same ; w centimeters higher, below the sympodial
branch. A small amount of secondary tissue has been formed by the cambium,
xX 36.
developed. Central cylinder delimited by compact, often lignified ground
parenchyma and peripheral, congested vascular bundles. Central bundles
More diffusely distributed among thin-walled ground tissue resembling
376 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
parenchyma of cortex. Vascular bundles each with a sheath of narrow
compact angular cells, the sheathing cells thick-walled around phloem but
sheath becoming more uniformly sclerotic around peripheral bundles.
Vascular tissues collateral, including a wide strand of angular metaxylem
elements, V-shaped in transverse section, usually with narrow protoxylem
elements at the apex of the V and a single phloem strand in the angle of
the V. Peripheral vascular bundles with little or no protoxylem, the
metaxylem scarcely V-shaped. Leaf traces conspicuous in outer part of
central cylinder, the xylem represented largely by abundant protoxylem.
Xylem including fairly wide angular tracheids with indistinct end walls
and scalariform pitting. Protoxylem elements rounded, with annular or
spiral wall thickening. Metaxylem tracheids of the order of 5—10 mm.
long, and overlapping extensively. Phloem including long sieve-tube ele-
ments usually with transverse end walls and simple sieve plates, but sieve
plates commonly compound on oblique or very oblique end walls. Raphide
clusters common in otherwise unmodified parenchyma cells. Tannin cells
infrequent.
Secondary tissues (Fic. 9). This arises from an etagen cambium of
the type which has already been described in the earlier review (Tomlin-
son & Zimmermann, 1969). Ground tissue of compact tabular and radially-
arranged cells about 120, long, with slightly thickened and lignified cells,
the walls with abundant simple pits. Tannin deposits and raphide clusters
frequent. Secondary vascular bundles always amphivasal. Central phloem
strand including short sieve-tube elements with simple, more or less trans-
verse sieve plates. Phloem separated from xylem by short, thin-walled
parenchyma cells. Secondary tracheids conspicuously different from those
of primary vascular bundles; of the order of 3.6 mm. long and with
indefinite end walls; walls thick; bordered pits with crossed slit-like
apertures, more or less parallel to the axis of the cell. Short xylem
parenchyma cells infrequent.
The difference in length between secondary ground tissue cells and
secondary tracheids suggests that the latter undergo about a 30-fold
extension during development since both arise from similar initials.
COURSE OF PRIMARY VASCULAR BUNDLES
The distribution of primary vascular tissue in Dracaena fragrans is
similar to that of the palm Rhapis excelsa as described by us (Zimmer-
mann & Tomlinson, 1965) with slight quantitative differences. Each leaf
is supplied with a number of leaf traces which diverge from the stem at
varying depths. Major bundles diverge from the center, minor bundles
from near the periphery, and intermediate bundles from an intermediate
area of the stem. Outgoing leaf traces produce a number of derivative
bundles by branching. Most of these branches are short bridges which
link in an upward direction with nearby vertical bundles. Axial continuity
from each leaf trace is maintained by a continuing vertical bundle which
1969 | ZIMMERMANN & TOMLINSON, DRACAENA, 1 377
CENTRAL CYLINDER CENTRAL CYLINDER
<—__. <—
|
DRACAENA RHAPIS
Eat | |
usually diverges from the leaf trace at the very periphery of the central
cylinder. The newly released vertical bundle is normally very narrow if
the stem does not contain secondary tissue (Fic. 8). In this very peripheral
position it readily splits or anastomoses with similar neighboring bundles
on its way up the stem. This contrasts with the situation in Rhapis where
the vertical bundle is released near the stem center and accompanies the
parent leaf trace, on its outwardly diverging path, almost all the way to
the periphery of the central cylinder (Fic. 10). The upwardly continuing
vertical bundle, as in Rhapis, then gradually approaches, over a distance
of many internodes, the center of the stem whereupon the process of
bundle branching is repeated again in association with another, more
distal leaf. Major bundles have the longest, minor bundles the shortest
istances between two such successive leaf contacts. Thus, the overall
course of vascular bundles is similar to that illustrated for RAapis (Zim-
mermann & Tomlinson, 1965; Fig. 3, right). In Dracaena, in contrast
with Rhapis, the central bundles have no helical path. Major dorsal
bundles merely describe a turn of about 120° in the center, as described
378 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
in the next section of this paper. This turn, as with the helical twisting in
Rhapis, is in the direction of the phyllotactic spiral. This turn may be
governed by the same developmental principle which causes phyllotaxis.
The distribution of protoxylem changes throughout each bundle in the
same manner as in Rhapis. This change is less conspicuous in Dracaena
than in Rhapis because protoxylem and metaxylem elements are more
nearly of the same diameter. As one follows a vascular bundle of Dracaena
in a distal direction one finds protoxylem first not very far above its
divergence from the leaf trace as a vertical bundle. Continuing upwards,
the number of elements further increases to reach a maximum where the
bundle passes out into another leaf. Metaxylem is continuous into bridges
as well as the continuing vertical bundle but the leaf trace contains only
protoxylem. In this respect Dracaena is identical with Rhapis although
the “loss” of metaxylem from the outgoing leaf trace is less obvious be-
cause the two tissues are not so clearly distinguished.
Irregularities in the course of bundles throughout the stem are somewhat
more common than in Rhapis. The anastomosing tendency of the lower
part of vertical bundles, at the periphery of the central cylinder, has been
mentioned. If the stem consists of primary vascular tissue only, the
vertical bundles are quite small in their lowermost portion, at the periphery,
where they come off the leaf trace and without a well-developed fibrous
sheath (Fic. 8). In places where the primary vascular cylinder is covered
by a mantle of secondary tissue, the same vertical bundles are larger and
more conspicuous, because the fibrous sheath is better developed (Fic. 9).
Another irregularity which has been observed is the occasional forking
leaf traces. When such a bundle is followed upwards in the stem center,
the two branches diverge along two different radii. From these obser-
vations it appears that developmental processes are somewhat less rigid
in Dracaena than in Rhapis.
The important topic of the relation between primary and secondary
vascular bundles is reserved for the second article in this series.
DEVELOPMENTAL PATTERN OF THE PRIMARY VASCULAR SYSTEM
Observations. General aspects of the anatomy of the meristematic
crown are shown in the photomicrographs, FicurEs 11 and 12. Leaves and
leaf primordia are arranged in a phyllotactic spiral with a divergence
between 1/3 and 2/5, as can be seen from Ficure 11. The approximately
median longitudinal section through the crown shows the usual mono-
cotyledonous organization (Fic. 12). It is obvious from this longitudinal
section that primary thickening growth involves re-orientation of tissue
through about 90° as we have described for Rhapis and Prionium (Zim-
mermann & Tomlinson, 1967, 1968).
The developing vascular system of the meristematic crown is far too
complex to be demonstrated in individual microtome sections. Provascular
strands were, therefore, followed throughout a series of transverse sections
and their radial distance from the stem center plotted on graph paper 45
1969 | ZIMMERMANN & TOMLINSON, DRACAENA, 1 379
Fic,
eee crown of oo fragrans at the level of the apical meristem, X 2
Showing the phyllotactic arrangements of the leaves. Note the symmetrical ar-
rangement of the major leaf traces on the dorsal side.
1G. 12 (Bexow). Approximate median longitudinal section through the
meristematic crown of Dracaena fragrans, X 26. Because of their c ge three-
dimensional path, none of the indi vidual inh nr strands can be er
more than a very short distance. A thorough knowledge of their path, ~ ine
S$ quite easily. Note the sha arp turns of the major strands below the apica
meristem. Note also the minor leaf trace on the far right of the photograph.
380 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
had been done for Rkapis and Prionium. The results are shown in FIGURE
13. The three-dimensional arrangement of provascular strands in the
crown is difficult to represent on paper, our representation is therefore
simplified as follows. All radii are shown in a single plane. All leaves are
rotated into the same plane. This eliminates the 120° turn of the major
bundles. In order to reconstruct the three-dimensional pattern of the
crown from Ficure 13, the reader has to go through a mental exercise
which first involves rotating leaves back to their proper position, and
second involves re-establishing the 120° turn of the major bundles. This
process of simplification is very similar to the one used in our description
of the Rhapis crown. It has the advantage that one can more easily
appreciate the re-orientation of a major dorsal bundle during successive
developmental stages.
FicurE 13 shows the major dorsal leaf trace in leaf primordia P 1 to
P 17, P 1 being the youngest visible primordium. The pattern of vascular
development appears to be the same as the one found in Rhapis and
Prionium. Vertical bundles originate from major leaf traces of P 17.
Approximately below the base of P 14 they fuse into the meristematic
cap into which all blind-ending vertical bundles converge (cf. Zimmer-
mann & Tomlinson, 1967, 1968). From the diagram one can extrapolate
that the leaf-contact distance for a major bundle is about 20 to 25 inter-
nodes, although the series of sections was too short to show this directly. If
the section series had been longer and had included the insertions of older
(lower) leaves, the major leaf trace of P 1 would have been seen origin-
ating as a vertical bundle from a leaf trace diverging into a leaf at about
the level of P 20-25.
A rather unusual type of vertical-bundle branch was found in two
major leaf traces to P 11. Both vertical bundles ended distally immediately
below the apical meristem in what might be a leaf primordium younger
than P 1 and represented by an indistinct ridge. If this interpretation is
correct there would be a leaf-contact distance of 11 internodes between
P 0 and P 11. Only two such centrally located vertical bundle branches
were found and one of them is shown in Ficure 13. The developmental
meaning of this rare type of vertical bundle is unknown.
Ficure 13 shows some further irregularities which are of no funda-
mental significance, such as the apparent crossing over of the lower
portions of the major leaf traces of P 1 and P 2, P 5 and P 6, P 7 and P 8.
They could have resulted by comparing bundles on different radii of the
stem (the crown is not perfectly circular in transverse section), from slight
irregularities of development, or indeed, from the process of plotting.
The meristematic cap is similar in position and extent to that described
in the apices of Rhapis and Prionium. It is recognized as the umbrella-
shaped meristematic area, below the shoot apex proper, into the periphery
of which the blind-ending vertical bundles fuse. It is pierced by lea
traces already connected to vertical bundles. f
The primary vascular connection between an axillary bud and the
vascular system of the central cylinder has also been traced in this series
1969] ZIMMERMANN & TOMLINSON, DRACAENA, 1 381
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MILLIMETERS FROM STEM AXIS
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Fic. 13. The path of major leaf traces from leaf primordia 1 through 17,
obtained from plotting each individual bundle through a series of transverse
bu : :
bundle. The dashed line marks the approximate level of leaf insertion. INSET
BELOW. The 120° turn of the major leaf traces in the stem center, as seen in
successive transverse sections.
of sections. The provascular connection between axillary bud and stem
was established only in P 17 and older leaves. In P 15 the axillary bud
meristem was apparent but still entirely without discernible procambial
Strands. This suggests that vascular continuity between axillary buds and
main axis is established late, in the manner of minor leaf traces. A more
detailed discussion of the development of the vascular system of axillary
buds will follow in the second paper of this series.
Developmental inferences. The sequence of vascular development
is thought to be as follows. Leaf traces link up with a potential vertical
bundle in the cap, then differentiate out below the cap. Leaf traces which
develop early, i.e., those arising in a position near the center of the cap,
€come major bundles; those developing further out, near the cap
Periphery, become minor bundles. For comparison both a major and a
minor trace from P 7 are shown in Ficure 13.
382 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
These developmental processes have been discussed in detail in our
articles on the developmental pattern of the vascular system of Rhapis
and Prionium. We may merely point out here that there are no cortical
bundles in Dracaena, a fact which is of utmost significance as we shall see
in the next paper of this series.
DISCUSSION
In a previous review (Tomlinson & Zimmermann, 1969) we have noted
that von Mohl (1824) equated the primary vascular system of “Aletris
fragrans” and other species which he studied with that of a palm, in so
far as he understood the course of vascular bundles in the palm stem. Von
Mohl’s contemporaries and all subsequent investigators who studied ar-
borescent Liliiflorae at first hand claim to have confirmed his observations
(e.g. Meneghini, 1836; Millardet, 1865; de Cordemoy, 1894; and others).
However, our own more recent investigation of the palm stem (Zimmer-
mann & Tomlinson, 1965) has shown that von Mohl’s understanding was
incomplete, because he overlooked the axial continuity of vascular bundles
which is so important in long-distance transport. We have already given
our historical interpretation of the topic (Tomlinson & Zimmermann,
1966) and need not discuss it any further. The present study of Dracaena
fragrans has confirmed that von Mohl was right in principle. The primary
vascular anatomy of the axis of this plant does indeed correspond in all
essentials with that of a palm, but we now have a much more complete
understanding of the anatomy of the palm and its development. Dracaena
conforms in the pattern of primary vascular differentiation, a pattern
which we believe is fundamental for monocotyledons as a whole (Zim-
mermann & Tomlinson, 1965, 1967, 1968).
Our cinematographic analyses have included other species and genera
of arborescent Liliiflorae. Analysis of the mature axis of Cordyline ter-
minalis, Dracaena marginata and Pleomele (Dracaena) reflexa confirms
the course of vascular bundles described for Dracaena fragrans. Single
sections which we have prepared from the stems of several other genera
and species can also be interpreted according to our three-dimensional
analysis. This additional evidence puts our interpretation on firm ground.
SUMMARY
The primary vascular system of arborescent Liliiflorae was thought by
von Mohl and subsequent investigators to be equivalent in principle to
that of palms. An analysis of the system in the vegetative axis of Dracaen@
fragrans with the aid of cinematographic methods confirms this. In addi-
tion, however, it also shows that axial continuity of the palm type, OV&T
looked by these early anatomists, which has only recently been demon-
strated, also occurs in Dracaena and related plants. The origin of the
primary vascular system has been traced by plotting the course of
provascular strands in the developing crown. We regard, on the basis of
pee pene —_—
1969 | ZIMMERMANN & TOMLINSON, DRACAENA, 1 383
similar studies of other plants, this pattern as fundamental for the mono-
cotyledons. The basis has thus been laid for a future investigation of
secondary vascular tissues in these plants.
LITERATURE CITED
Corpemoy, H. J. pe. 1894. Recherches sur les Monocotylédones a accroissement
secondaire. Thesis. Paris. pp. 108. 3 pls.
MENEGHINI, G. 1836. Ricerche sulla struttura del caule nelle piante Monoco-
tiledoni. pp. 110. 10 pls. Minerva, Padua.
MItarbet, A. 1865. Sur l’anatomie et le rigour du corps ligneux dans
les genres Yucca et Dracaena. Mém. Soc. Sci. Nat. Cherbourg 11: 1-24.
Mont, H. von. 1824. De palmarum structura. In: K. F. P. von Martius, His-
toria Naturalis Palmarum 1: pp. I-LII. 16 is.
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ei Uber die Verastelung bei monokotylen Baumen. III. Die Veras-
telung einiger baumartigen Liliaceen. Rec. Trav. Bot. Néerl. 15: 263-335.
Sims, ae oo Dracaena fragrans, Sweet-scented Dracaena. In: Bot. Mag. 28:
081.
TROLL, . 1962. Uber die “Prolificitat” von Chlorophytum comosum. Neue
Hefte Morphologie 4: 9-68.
Tomtinson, P. B., _H. ZIMMERMANN, 1966. Vascular bundles in ee stems
Saithedy bibliographic evolution. Proc. Am. Philos. Soc. 110: 1.
1969. Vascular anatomy of monocotyledons Nan peal
rowth — an ‘introduction, Jour. Arnold Arb. 50: 159-179.
,&
————__,
excelsa, I. Mature vegetative axis. Jour. Arnold Arb. 46:
& 1966. Analysis of aia vascular ie in ren Optical
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——— . Anatomy of os palm Rhapis excelsa. IV. Vascular de-
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[M.H.Z.] BT.
HARVARD UNIver RSITY FAIRCHILD TROPICAL GARDEN
Casot FounDATION 10901 OLtp CuTLER Roap
PETE M Miami, FLorma 33156
MASSACHUSETTS 01366
384 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 50
COMPARATIVE MORPHOLOGICAL STUDIES IN DILLENIACEAE,
Iv. ANATOMY OF THE NODE AND VASCULARIZATION
OF THE LEAF
WILLIAM C. DICKISON
IN A CONTINUING EFFORT to provide comprehensive anatomical infor-
mation which might prove useful in elucidating taxonomic and phylo-
genetic relationships of the Dilleniaceae, an extensive investigation of nodal
and leaf vasculature was undertaken.
Aside from remarks pertaining to ovular structure by Cordemoy (1859),
and an occasional reference to internal structure by various other workers,
the earliest comprehensive anatomical investigations on Dilleniaceae are
the contributions of Baillon (1866-67, 1871) and Hitzemann (1886, cited
by Ozenda, 1949).
The first comparative morphological studies on the family to appear
were those of Parmentier (1896), who found the leaf to contain charac-
ters of diagnostic value, and Steppuhn (1895) who made an extensive in-
vestigation of stem, leaf, and root of some one hundred fifty dilleniaceous
species.
Solereder (1908) and Metcalfe and Chalk (1950) published additional
anatomical information, but contributed little to help clarify the phylo-
genetic position of the group. The most recent study on comparative
vegetative anatomy of the family was by Ozenda (1949) whose observa-
tions on seedling, nodal, and leaf anatomy were scattered among seven
genera.
All researches referring to the Dilleniaceae, therefore, are either in-
complete, or else were produced in the last century and thus warrant re-
investigation. This paper describes heretofore unreported anatomical data
of both taxonomic and phylogenetic significance.
MATERIALS AND METHODS
Material of over one hundred dilleniaceous species was examined. Speci
mens studied were received from, or are housed in: the Arnold Arboretum,
Harvard University (a); State Herbarium of South Australia, Adelaide
(ap); Arizona State University, Tempe (Asu); Botanic Museum and Her-
barium, Brisbane (pri); Commonwealth Scientific and Industrial Research
Organization, Canberra (caANB); Royal Botanic Garden, Edinburgh (£);
Gray Herbarium, Harvard University (crt); Royal Botanic Gardens, Kew
(k); Botanical Survey of India, Southern Circle, Coimbatore (aH); Mis
souri Botanical Garden, St. Louis (mo); Animal Industry Branch, North-
ern Territory Administration, Alice Springs (Nr); Western Australian
1969 | DICKISON, DILLENIACEAE, IV 385
Herbarium, Perth (pertH); Rancho Santa Ana Botanic Garden, Clare-
mont (RSA); Sarawak Museum, Kuching (sar); Botanic Gardens, Singa-
pore (sinc); University of California, Berkeley (uc); and the United
States National Museum, Washington (us). The assistance of the cura-
tors of these collections is gratefully acknowledged. I also wish to thank
Doctors R. D. Hoogland, H. Keng, and C. R. Metcalfe for providing seed
used in this study.
The study of lamina vascularization was accomplished entirely through
the use of cleared leaves. Clearing was carried out using the standard
NaOH method followed by safranin stain. Dried materials were initially
re-expanded in 5 percent NaOH prior to fixation and sectioning. Nodes
were serially sectioned and stained with a combination of safranin-fast
green. Petiole vasculature was followed by obtaining sections throughout
the length of the petiole as well as midway through the midrib.
NODAL ANATOMY
No detailed, comprehensive study of nodal anatomy in the Dilleniaceae
has previously been undertaken. Sinnott (1914) attempted to utilize the
nodes of several genera within the family (sensu Gilg, 1893) as evidence
to support his idea that the trilacunar node was primitive; furthermore,
that the unilacunar and multilacunar node was derived by reduction or
amplification. This author listed six genera of the Dilleniaceae (sensu
stricto) as having tri- or pentalacunar nodes.
Ozenda (1949) after an examination of Hibbertia, Dillenia, Schuma-
cheria, Tetracera, C uratella, and Davilla also concluded that the mature
nodes of the family were tri- or multilacunar; however, he was of the
opinion that the multilacunar condition was the primitive pattern. The
primitive nature of the multilacunar node in the Dilleniaceae has also been
advocated by Meeuse (1966, p. 49).
As a result of comparative morphological data from both fossil and
extant plants, in addition to ontogenetic considerations, the primitive na-
ture of the trilacunar node was questioned by Marsden and Bailey (1955)
and Canright (1955). These authors suggested that the unilacunar two-
trace system represented the primitive condition. The unilacunar two-
trace node is characteristically described as having two vascular traces
which arise from independent primary bundles and, therefore, do not
Tepresent the dichotomy of a single median trace.
Pant and Mehra (1964) re-evaluated nodal anatomy in many Pterop-
sida, and concluded that the statements of Marsden and Bailey (loc. cit.)
concerning nodal patterns in fossil ferns and gymnosperms were not always
substantiated, They then advised caution in accepting the unilacunar two-
trace node as primitive for all Pteropsida. Results from a study of devel-
opmental patterns in stem primary xylem indicated to Benzing (1967a, b)
that the odd-numbered trace, unilacunar one-trace or trilacunar, was more
likely to be primitive in angiosperms. A recent paper by Namboodiri and
386 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Beck (1968b) supports this view and regards the unilacunar one-trace
node as the primitive condition in the Coniferales.
My observations reveal that mature nodes of Dillenia (Fic. 8), Dides-
mandra, and Schumacheria (Fic. 7) are exclusively multilacunar.t | The
numerous leaf traces are associated with a corresponding number of pa-
renchymatous gaps in the cauline stele. The number of traces in Schuma-
cheria and Didesmandra was found to be stable at nine and seventeen
respectively. The number in Dillenia, however, varies from as few as seven
(D. puichella) to as high as twenty-seven (D. suffruticosa). Variability
is also evident within a single species, the number of traces apparently
reflecting the age and size of the node. The manner in which bundles de-
part the stele was found generally to be correlated with the presence or
absence of sheathing leaf bases. If the petiole does not sheath the stem,
all traces tend to depart simultaneously. If the leaves are amplexicaul,
the median trace passes out initially, with laterals departing in succession
at higher levels.
The leaves examined of the semi-herbaceous, rhizomatous genus Acro-
trema were supplied by three traces; i.e., the node was trilacunar (Fic. 4).
The median trace departs first with the resulting gap remaining open
above the level at which the two lateral gaps close. Large-leaved species
(e.g., A. arnottianum) should be studied when available to determine to
what extent leaf size affects the nodal pattern in this extremely variable
genus.
In contrast to the information presented by Ozenda (1949), all hib-
bertias are not uniformally trilacunar. Numerous species with reduced,
needle-like leaves possess unilacunar nodes (Fic. 2). In these cases, the
primary stele is composed of a continuous cylinder of vascular tissue with
no discrete bundles discernible. At the unilacunar node, a single trace
passes directly into the leaf. All broad-leaved hibbertias from New Cale-
donia and Fiji are trilacunar. Leaf size is not always indicative of nodal
patterns, however. Trilacunar Hibbertia huegelli (Fic. 45) and H. mono-
gyna (Fic. 46), for example, possess smaller leaves than H. nitida (Fic.
47) which is unilacunar.
The most reduced leaves in the family are encountered in the genus
Pachynema. The small, scale-like, lateral appendages were found to be
vascularized by a single prominent trace with a well defined gap in the
stele. From the flattened stem of P. dilatatum, leaves may be secondarily
supplied by weak cauline traces (Fic. 34).
The New World genera Curatella, Davilla (Fic. 6), and Doliocarpus
are mostly pentalacunar; but seven-trace nodes occur in Davilla aspera,
and Doliocarpus major is trilacunar. Trilacunar, three-trace nodes are
also uniform throughout the genus Tetracera where special effort was made
to examine representative species from the Old and New World tropics.
In Tetracera, three bundles are associated with three widely separated
“The report by Benzing (1967a) of unilacunar one-trace nodes in the mature
stems of Dillenia indica is in error. I have personally examined the sections used ™
this study and conclude that they were not taken from any member of the Dilleniaceae-
—— ae
1969 | DICKISON, DILLENIACEAE, IV 387
gaps (Fic. 3). The lateral bundles arc up and through the cortex where
they enter the petiole. This contrasts with the condition in the trilacunar
hibbertias with sheathing leaf bases, where the laterals enter the leaf di-
rectly from the stele (Fic. 5).
The seedling anatomy of Dillenia indica, Tetracera indica, Hibbertia
dentata, and H. scandens was examined. The cotyledonary nodes in the
first two species are of the 2:1 type, viz., two traces departing from a
single gap (Fics. 1, 40). My observation of the 2:1 cotyledonary nodal
pattern in Dillenia indica once again contradicts the information presented
by Ozenda (1949) who illustrated a single cotyledonary trace. Particular
attention was paid to the double traces at subnodal levels and in all in-
stances they originated from independent parts of the stele. In Dillenia
indica, a species with multilacunar nodes in the mature stem, the first
formed seedling leaves possess a trilacunar node (Fic. 41). Numerous
examples can be found in other dicotyledonous families of a similar pro-
gression, as in Magnoliaceae, Degeneriaceae, etc.
The cotyledonary node of Hibbertia dentata and H. scandens differs by
being of the unilacunar one-trace type (Fic. 12). No evidence of double-
ness could be observed in the single, broad strand of vascular tissue which
passes into the cotyledon. The occurrence of a 1:1 cotyledonary node in
Hibbertia is of particular interest, since it is a genus with trilacunar mature
nodes, and is generally considered to be more primitive in its characters
than either Dillenia or Tetracera. The question again arises whether an
even or odd number of nodal traces represents the primitive condition. A
thorough study of the cotyledonary node in other Dilleniaceae would be
worthy of careful attention.
It is perhaps significant, that more than one case was observed among
the seedling and mature nodes of Dillenia where an even number of traces
prevailed. This condition resulted from suppression of one of the lateral
bundles with the result that an even number of traces departed the stele.
Although this might be dismissed as abnormal, the fact that it was observed
more than once indicates that it may be of some significance.
Although the majority of plant families exhibit a combination of uni-
lacunar and trilacunar, or trilacunar and multilacunar nodes, it is relative-
ly uncommon for a single family to possess all three types (Bailey & Nast,
1944). The Dilleniaceae are, therefore, unusual in possessing four pat-
terns: unilacunar two-trace, unilacunar one-trace, trilacunar, and multi-
lacunar. Bailey (1956) points out that transitions from trilacunar to uni-
lacunar nodes may occur as the result of anatomical specialization in re-
sponse to the environment. The reduction of leaf size in Hibbertia is such
an adaptation. Thus, the mature foliage nodes in Dilleniaceae (sensu
stricto) demonstrate two distinct trends of specialization: (1) secondary
reduction and elimination of the lateral strands of the trilacunar nodes;
and (2) amplification of the trilacunar node by the addition of laterals.
Bailey and Howard (1941) note that trends of specialization in nodal
anatomy are not infrequently correlated with specializations elsewhere in
the plant (e.g., wood). No such direct correlations were noted in the dil-
388 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
lenias. In fact, those genera having the least advanced wood generally pos-
sess the more highly evolved nodes.
It is not possible to construct relationships within the family solely on
the basis of nodal structure due to the presence of a similar anatomy in
both Old and New World genera. Likewise, nodal anatomy is of limited
value in determining relationships beyond the family. When the nodes of
putatively related families are compared, it is evident that they are all
essentially tri- or multilacunar. There is, accordingly, no basis for ac-
cepting or rejecting alliances from this information alone.
A significant exception to the above generalizations is found in the
Theaceae where the node is uniformly characterized by a broad trace which
departs from a single gap. Keng (1962) agreed with Canright (1955)
in considering this pattern to be the result of phylogenetic fusion of several
separate traces. If Canright’s (Joc. cit.) suggested trends of nodal spe-
cialization are accepted, it leads one to the conclusion that the nodal
anatomy in the two families represents the culmination of distinct lines
of evolution. Therefore, although the wood and pollen of these groups
is similar (Dickison, 1967a, b), nodal anatomy suggests they may in fact
be only distantly related.
A similar conclusion might be reached regarding the predominantly
unilacunar, one-trace nodes of Ericaceae; however, the report by Philip-
son and Philipson (1968) of trilacunar nodes in Rhododendron gives cause
for re-evaluation.
PETIOLE VASCULARIZATION
An attempt to define the range of variability in petiolar anatomy of Dil-
leniaceae disclosed the following major patterns:
Species with Unilacunar Nodes (1:1).
(1) A single, slender, unbranched trace enters the lamina: numerous hibbertias
(Fic. 27).
Species with Trilacunar Nodes (3:3).
(1) Traces fuse and form a flattened arc: Hibbertia quadricolor.
(2) Traces fuse and form “V” shaped arc: Hibbertia coriacea (FIc. 17):
(3) Traces fuse and form cylindrical, flattened, or concave vascular ring, either
confluent or slightly dissected: Doliocarpus major; D. olivaceus; * Hib-
(4) Traces form a closed cylindrical ring with one or more medullary bundles
produced by invagination: Hibbertia lucida (Fic. 13). ;
(5) Traces form an abaxial arc of fused or dissected collateral bundles with
a separate adaxial trace derived from the inrolling and/or division of the
lateral bundles. The adaxial trace may be lost in the lamina: Acrotrema
* Nodes were not examined.
1969 | DICKISON, DILLENIACEAE, IV 389
sp.; A. bullatum; A. gardneri; A. lanceolatum; A. uniflorum; A, walkeri;
Hibbertia banksii; H. pancheri; H. wagapii: Tetracera boiviniana; T.
masuiana
Traces form an abaxial arc of fused or dissected collateral bundles with
a separate adaxial trace derived from division of the median bundle, The
adaxial trace may be lost in the lamina: Hibbertia scandens; Tetracera
indica.
=
Nn
—
Species with Multilacunar Nodes (five to many traces from an equal
number of gaps).
(1) Traces remain free forming a ring of widely dissected collateral bundles
(bundles are often of unequal sizes): Acrotrema costatum;* Didesmandra
(Fic. 15); Dillenia excelsa; D. luzoniensis; Schumacheria angustifolia.
(2) Traces fuse to form confluent or only slightly dissected ring, often “U”
or “V” shaped in outline: Davilla (Fic. 22); Dillenia bolsteri (Fic. 19);
D. eximia; D. indica; D. ovata; D. pentagyna; D. salomonensis; D. suf-
fruticosa; D. turbinata.,
Traces fuse to form confluent or only slightly dissected ring with an arc
(rarely superimposed) of fused or dissected medullary bundles: Curatella
americana (Fics. 10A,B,C); Dillenia alata; D. beccariana (Fic. 20); D.
castaneifolia (Fic. 18); D. megalantha; D. papuana; D. philippinensis;
D. reifferscheidia.
Traces form an abaxial arc of fused or dissected collateral bundles with
an adaxial enclosed siphonostele. The adaxial ring may subsequently open
laterally or invaginate to produce additional free bundles: Doliocarpus
coriaceus; D. dentatus; D. guianensis (Fic. 21); D. rolandri.
~~,
Ww
wa
“-o-~
ae
le
It is evident that petiole vascularization in Dilleniaceae is quite diverse
both between and within genera. In the present study, subtle deviations in
vascularization pattern, the general outline of the vascular cylinder, and
small, adaxial, subsidiary wing traces were ascribed little importance.
Despite acknowledged incompleteness, I feel the descriptions outlined
above will prove useful in future comparative studies relating to the family.
It should be emphasized, that the descriptions presented are not based
entirely upon observations from a single “characteristic” region. Wher-
ever possible, sections were examined throughout the petiole and midrib
aS suggested by Howard (1962). The importance of determining the
390 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
liocarpus, D. major and D. olivaceus are distinguished by their pubescent
ovaries and fruits. An absence of medullary bundles in the petiole was
also found to separate these taxa readily from all other species examined in
the genus. Hunter (1966) considers Doliocarpus rolandri Gmel. to be a
synonym of D. major Gmel. I have studied a collection from Brazil cited
as D. rolandri (Pires & Cavalcante 52254, us) and found the petiole to
possess medullary bundles, a character which is not encountered in D.
major. A re-examination of this genus taxonomically might yield addi-
tional basis for separation of the species. Other specific variation is found
in Acrotrema (where A. costatum, from Thailand and Malaya, is quite dis-
tinct from the Ceylonese species), Hibbertia, and Dillenia, though much
more material must be studied before the true value of these data can be
realized.
Petiole structure cannot be used to separate the Dilleniaceae into sub-
families or tribes. Moreover, there is little or no correlation between
petiole vascularization and nodal anatomy. When considered as a whole,
the vascular pattern in species with trilacunar nodes cannot be considered
more primitive than that in species with multilacunar nodes.
In plants with multilacunar nodes, petioles with widely dissected cyl-
inders tend to be correlated with slender venation lacking massive bundle
tive condition in multilacunar dillenias. Subsequent evolutionary pro-
gression has produced fusion of traces and the formation of more complex
medullary bundle patterns. These specializations have apparently oc-
curred more than once, since the same apparent trends are also evident in
species with trilacunar nodes.
These ideas of nodal and petiolar evolution in Dilleniaceae do not agree
with the conclusions of Decker (1967) who worked on the Luxemburgieae
(Ochnaceae). Within the Luxemburgieae, Decker considers the multila-
cunar node more primitive than the trilacunar, and petioles with numer-
ous, unfused bundles (some of which may be medullary), more primitive
than petioles with fused traces devoid of medullary bundles. In view of
the frequent derivation of the Ochnaceae from the Dilleniaceae, such
contrasting opinions are of special interest.
A foliar character of debatable morphological derivation is the presence
of petiolar wings in some Old World dilleniaceous species. Hooglan
(1952) attaches taxonomic importance to the presence of completely am-
plexicaul petiolar wings in certain species of Dillenia. Morphologically,
these wings are frequently considered to be stipules. Hoogland (/oc. cit.
does not accept this interpretation for the following reasons: (1) there
is often no sharp distinction between the petiolar wings and lamina, (2)
stipules of the usual morphological type do not occur in the dillenias, (3)
in caducous wings, separation from the petiole begins from the base of the
petiole and not from the apex as one would expect, and (4) Ozenda (1949)
describes the wings as being weakly vascularized in contrast to the situation
1969 | DICKISON, DILLENIACEAE, IV 391
in the Magnoliaceae where the stipules receive a separate trace from the
cauline stele.
I have found vascularization of the wings in Dillenia to vary from weak
(e.g., D. albiflos) to rather strong (e.g., D. philippinensis: D. suffruticosa).
In the latter case the venation is highly reticulate. In either situation the
wings are never supplied by independent traces from the cauline stele.
Although I do not have any original interpretation for these structures,
they do not appear comparable to true stipules.
VASCULARIZATION OF THE LAMINA
Major Venation. Although the prevailing type of major foliar vena-
tion in the Dilleniaceae is pinnate, with the secondary veins proceeding to
the margin of the blade, wide variation in leaf size, shape, and vasculariza-
tion is encountered in the genus Hibbertia. A study of leaf vasculature in
this genus showed that three basic venation patterns can be recognized:
(1) pinnate leaves in which the numerous, strong, parallel, lateral veins
extend diagonally outward from the midvein toward the margin of the
lamina where they are interconnected by curved peripheral venation (Fic.
43); (2) pinnate leaves in which the principal lateral veins are fewer in
number, irregular in their occurrence, more tenuous, and tend to sweep
upward upon departure from the midvein (Fics. 42, 44); and (3), a pat-
tern where two or more strong, terminal, lateral veins reflex back after de-
parture from the midrib to terminate, often very massively, at the leaf
ase. A varying number of prominent lateral veins may connect the mid-
rib with the reflexed lateral (Fic. 49). This specialized venation pattern
is exclusively associated with those hibbertias with reduced, needle-like
leaves. The physiological significance of this type of vasculature is not
clear.
Concomitant in Hibbertia with a general trend toward reduction in leaf
size as a response to xerophytic conditions, is a trend in reduction of leaf
vascularization. Theoretically, this specialization commences with the pro-
mination of the midrib was also observed in the mature leaves of the family.
392 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The first-formed seedling leaves of Dillenia indica, with their strong, paral-
lel, pinnate veins and serrate margins are sharply distinguished from the
cotyledons (Fic. 41).
Minor Venation. In addition to noteworthy features of major vena-
tion, the pattern and diameter of the minor veins, in association with
bundle sheathing, is often of diagnostic and perhaps of taxonomic signifi-
cance in the Dilleniaceae.
The occurrence of bundle sheaths around the veins is almost a universal
feature of dilleniaceous leaves. Sheathing is noticeably absent only in some
hibbertias and Acrotrema. When present, the sheath cells are either
unlignified and parenchymatous in nature, or lignified, pitted, sclerenchy-
matous elements.
Parenchymatous sheaths typically surround both the major veins and
terminal veinlets. These sheaths usually consist of cells elongated parallel
to the vascular bundles; however, occasionally they become considerably
lobed and oriented at right angles to the veins (Fic. 33). Sclerotized
bundle sheathing is recognized by the presence of lignified, extensively
pitted cells. When sclerenchyma occurs, it may form massive sheaths
over the veins and veinlets as in Hibbertia (Fic. 30), Tetracera (Fic. 32),
and some species of Doliocarpus. The formation of sclerified bundle
sheaths enclosing the terminal tracheids is an uncommon feature in di-
cotyledonous leaves (Esau, 1965). Of more frequent occurrence in the
family is sclerenchyma around the major veins, but with veinlets devoid
of sheathing or possessing only an incomplete sheath. The most striking
pattern is seen in Hibbertia banksii where the mature leaves exhibit an
interrupted sclerenchymatous sheath (Fic. 29).
arenchymatous sheath cells were observed in Curatella (Fic. 31) and
all species of Dillenia (Fic. 23), except D. philippinensis and D. reiffer-
scheidia where pitted elements are found. Also, the presence of lobed
parenchymatous sheathing around the terminal veinlets in Doliocarpus
dentatus (Fic. 33) and D. rolandri readily distinguishes them from all
other species of the genus. The variation present in Doliocarpus in the
node, petiole, and minor venation warrants further study.
Distinctions can also be made between genera and species on the basis
of the diameter of veins and veinlets. Very slender venation is present in
Acrotrema, Didesmandra (Fic. 26), Schumacheria, and some hibbertias
(H. scandens, H. dentata, H. tetrandra, etc.). Associated with slender
vascularization is weak bundle sheathing or its complete absence. Only in
Hibbertia is massive venation sometimes devoid of sheathing (Fic. 28).
There appears, nevertheless, to be in the family a rather distinct trend
toward increased vein size accompanied by intensification of the amount
of vein sheathing.
A restricted trend was observed in Dillenia toward the formation of
vein islets devoid of free vein endings. It is possible to trace this progres
sion from species with slender veins and numerous free vein endings (e.g.
1969] DICKISON, DILLENIACEAE, IV 393
D. salomonensis and Fic. 23) through species with an intermediate pat-
tern (e.g. D. quercifolia, D. ovalifolia, D. nalagi) to a pattern illustrated
by D. papuana (Fic. 24) where free vein endings are scarce. The terminal
condition in this sequence is seen in the massive, closed venation of D.
schlechteri (Fic. 25).
A taxonomic correlation of minor venation patterns in Dillenia is illus-
trated by similar closed venation types occurring in D. papuana, D. cyclo-
pensis, and D. schlechteri, all of which are considered closely related by
Hoogland (1959) on the basis of floral structure. The only other species
which were observed to possess comparable vasculature were D. beccariana,
and D. turbinata, On the basis of leaf venation, I was not able to segregate
Wormia as a distinct genus from Dillenia.
The leaves of the Dilleniaceae appear to display a rather distinct phylo-
genetic trend of specialization toward more massive vascularization, ac-
companied by an increase in bundle sheathing. The same fundamental
trends have also been described for the Winteraceae (Bailey & Nast, 1944).
When the venation pattern, size, type, and degree of bundle sheathing,
as well as petiole vasculature, are considered together, they offer excellent
diagnostic leaf characters at the family, genus, and in some instances,
species level. Additional material in all stages of maturity will have to
be examined to understand fully the taxonomic significance of this infor-
mation.
TERMINAL IDIOBLASTS
The occurrence of specialized terminal-veinlet elements in several wide-
ly diverse dicotyledonous families has been well established. In a recent
review of the literature, Tucker (1964) describes their presence in the
Magnoliaceae. The occurrence of terminal idioblasts is now reported for
the first time in the Dilleniaceae. ;
Specialized terminal cells were observed only in relatively few species
of Hibbertia. The diversity in vein endings is thus in accordance with
variation in leaf shape and venation. The terminal cells are all of the
basic tracheoid type (see Foster, 1956). Employing the classification
of Tucker (loc. cit.), one can recognize tracheoidal elements, viz. scalari-
form or scalariform-reticulate pitted cells, and dilated tracheids. Leaves
of H. scandens (cult. K, s.n.) and H. dentata (cult. K, s.m.) were found
to contain terminal elements which closely resemble tracheary cells in
general morphology. The elements in H. dentata (Fic. 36) tend to occur
singly and have exclusively scalariform pitting whereas the idioblasts
of H. scandens (Fic. 35) often occur in clusters where they assume more
irregular shapes and have reticulate pitting.
The terminal elements of Hibbertia pachyrhiza (C. L. Wilson 861 j Fic.
38), although of the basic tracheoid type, differ considerably in their
morphology. The latter cells are thick-walled, pitted to a much less degree,
and are characteristically spherical in outline. In comparison with sur-
rounding parenchyma these elements are significantly larger (75-140 » in
394 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
diameter). They occur singly or in clusters of three to four on each vein
ending.
Terminal elements in Hibbertia huegelli (C. L. Wilson 777), H. mono-
gyna (Maiden s.n.), H. elata (Ingram 19852), H. billardieri (Clemens
42584a), and H. linearis (White 8580) tend to be intermediate between
the elongated tracheoid element and the spherical one (Fic. 37).
Dilated tracheids were found in Hibbertia scandens (Fic. 39), H. den-
tata, H. nymphaea (Morrison s.n., a), and H. amplexicaulis (Pritzel 531).
In H. scandens and H. dentata they were often in the same leaf with
tracheoidal elements, and several veins were noted where the two types
were present at the same vein endings. Generally, however, dilated tracheids
seem to occur rather sparsely throughout the leaf and are not present at
every vein terminus.
The taxonomic usefulness of terminal idioblasts in the Dilleniaceae
appears limited in view of their rather infrequent occurrence. Phylo-
genetically it is of interest that the most diverse vein endings are found
in Hibbertia, which on the basis of other criteria, is considered rather
primitive. A similar situation has been reported in the Magnoliaceae
(Tucker, loc. cit.). The full phylogenetic value of terminal idioblasts
still remains to be developed; however, the trend toward the formation of
specialized terminal cells appears to be a distinct one, subsequently lead-
ing toward the reduction in size of the elements and in the amount of
pitting on the wall surface.
SUMMARY
A comprehensive study of nodal and leaf vascularization in Dillenia-
ceae has led to the following fundamental conclusions:
cotyledonary node is unilacunar two-trace or unilacunar one-trace.
Evidence from nodal anatomy appears to discredit a close relation-
ship between the Dilleniaceae and Theaceae.
(2) The petiolar anatomy of the family shows considerable diversity. De-
scriptions of major venation patterns reveal that, in general, vascular
cylinders composed of widely dissected bundles are more primitive
than petioles with fused bundles and more complex medullary traces.
(3) The vascularization of the lamina displays fundamental phylogenetic
trends of specialization in both major and minor venation. Bundle
sheath cells are either parenchymatous or sclerenchymatous and may
enclose the terminal tracheids. Slender venation patterns lacking
bundle sheathing are less specialized than coarser-veined leaves with
massive bundle sheathing.
(4) When considered together, nodal anatomy and foliar vasculature are
of excellent diagnostic value and frequently of taxonomic and phylo-
genetic significance in the Dilleniaceae.
1969 | DICKISON, DILLENIACEAE, IV 395
(5) The presence of specialized terminal idioblasts in the leaves of Hib-
bertia is a character of which the importance is yet to be determined.
MATERIAL EXAMINED
Acrotrema sp. Ceylon: Thwaites CP3899 (us). A. bullatum Thw. Ceylon:
Thwaites CP239 (us), A. costatum Jack. Thailand: Smitinand 2999 (us). A.
gardneri Thw. Ceylon: Thwaites CP253 (us). A. lanceolatum Hook. Ceylon:
Thwaites CP2660 (us). A. uniflorum Hook. Ceylon: Thwaites CP1014 (us). A.
walkeri Wight. Ceylon: Thwaites CP694 (us).
Curatella americana L. Brazil: Irwin 5470 (Ny); Nicaragua: Van der Sluijs s.n.
(preserved material).
Davilla aspera (Aubl.) Benoist. Trinidad: Howard 10502 (cu); Brazil: N.T.
Silva 16. D. multiflora (DC). St. Hil. Panama: Dodge & Allen 17360 (mo). D.
rugosa Poir. Brazil: A. de Mattos Filho s.n. (preserved material). Davilla sp.
Brazil: Irwin 5570 (Ny).
Didesmandra aspera Stapf. Sarawak: Burtt & Woods B.2540 (gE); S.18297 (sar);
Native collector (sar) s.2. (preserved material).
Dillenia alata (R.Br. ex DC.) Mart. New Guinea: P. van Royen 4677 (A, US).
D. albiflos (Ridl.) Hoogl. Malaya: Corner SING F 29369 (a). D. beccariana
Martelli. Sarawak: SAR 16272 (a). D. biflora (A. Gray) Martelli ex Dur. & Jacks.
Fiji: Gillespie 2182 (GH); A. C. Smith 8762 (us). D. bolsteri Merr. Philippines:
Wenzel 3112 (cH). D. castaneifolia (Miq.) Martelli ex Dur. & Jacks. New
Guinea: Womersley NGF 3768 (a). D. cyclopensis Hoogl. New Guinea: van
Royen & Sleumer 5812 (a). D. excelsa (Jack) Gilg. North Borneo: Ramos 1379
(A). D. eximia Miq. Borneo: NIFS bb 16830 (A). D. indica L. Australia: Cult.
BRI 5.2. (preserved material); India: Sastri s.n. (preserved material); Cult. E
C4388. D. luzoniensis (Vidal) Martelli ex Dur. & Jacks. Philippines: J. V.
Pancho s.n. (preserved material). D. megalantha Merr. Philippines: Quezon.
M. Q. Lagrimas s.n. (preserved material). D. monantha Merr. Philippines: Herre
1010 (a). D, montana Diels. New Guinea: Hoogland & Pullen 6265 (a). D.
nalagi Hoogl. Papua: Hoogland & Taylor 3438 (A). D. ochreata (Miq.) Teysm.
& Binn. ex Martelli. Celebes: NJFS bb 18085 (a). D. ovalifolia Hoogl. New
16655 (a). D. salomonensis (White) Hoogl. Solomon Islands: Walker & White
145 (a). D. schlechteri Diels. New Guinea: Womersley & Millar NGF 7000 (A).
D. suffruticosa (Griff.) Martelli. Singapore: Cult. sinc s.2. (preserved material) ;
Canright 978 (asv). D. turbinata Fin. & Gagnep. Hainan: How 72058 (A).
Doliocarpus coriaceus (Mart. & Zucc.) Gilg. British Honduras: Gentle 2892 (us);
Colombia: Cuatrecasas 16556 (us). D. dentatus (Aubl.) Standl. Bolivia: Krukof
396 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
10088 (uc). D. guianensis (Aubl.) Gilg. Surinam: uc 947180. D. lasiogyne
Benoist. Brazil: Klein 1.281 (us). D. major Gmel. Panama: von Wedel 2860
(mo); J. M. Johnston 1694 (mo). D. olivaceus Sprague & Williams ex Stand.
Panama: Stern et al. 11 (us). D. rolandri Gmel. Brazil: Pirés & Cavalcante
52254 (us).
a acicularis (Labill.) F. Muell. Australia: Queensland. Clemens aa
(A); C. T. White 9466 (a). H. altigena Schlechter. New Caledonia: H. S. M
3709 (aA). H. amplexicaulis Steud. Australia: Pritzel 531 (a). H. aspera De.
aseetiga New South Wales. Constable 42837 (a). H. aurea Steud. Aus-
C. L. Wilson 843 (us). H. aun oe Australia: Aston 359
©. “HL. banksti Benth. Papua: L. J. Brass 1 (a). H. baudouinii Brongn. &
Gris. New Caledonia: tee aril al ie? (a). H. bracteata (R.Br.)
Benth. Australia: New South Wales, C. T. White 5012 (a). H. billardieri F.
Muell. Australia: Queensland. Clemens 42584a (us). H. brongniartii Gilg. New Cale-
donia: Thorne 28580 (rsa). H. cistiflora Wakefield. Australia: New South Wales.
Helms 1290 (a). H. cistifolia R.Br. Australia: Specht 843 (us). H. coriacea
(Pers.) Baill. Madagascar: Humbert 5866 (us). H. crenata Andr. Australia:
C. L. Wilson 851 (us). H. cuneiformis (Labill.) Gilg. Australia: Cult. K, s.1. (pre-
served material); E. H. Wilson 297 (us). H. dealbata Benth. Australia: Specht
Australia: Royce 5760 (us). H. ebracteata Bur. ex Guillaum. New Caledonia:
H. S. McKee 3697 (a). H. elata Maiden & Betche. Australia: New South Wales.
Ingram 19852 (us). H. exutiacies Wakefield. Australia: Eichler 17965 (av). H.
fasciculata R.Br. ex DC. Australia: Aston 387 (a). H. furfuracea Benth. Aus-
ia: C. T. White 5382 Hi: mane Diels. Australia: C. L. Wilson 856 (us).
H. pi eee F. Muell. Australia: Perry 5379 (us). H. gracilipes Benth. Aus-
tralia: Royce 5792 (us). H. huegelli F. Muell. Australia: C. L. Wilson 777 (us).
H. hypericoides (DC.) Benth. Australia: E. H. Wilson 454 (a). H. pos i
Benth. Australia: C. L. Wilson = pee H. linearis R.Br. ex DC. Australia
Australia: C. L. Wilson 740 (us). H. microphylla Steud. Australia: C. T. White
5317 (A). H. miniata Gard. Australia: C. L. Wilson 782 (us). H. monogyna
R.Br. ex DC. Australia: New South Wales. J. H. Maiden s.n. (GH). H. montana
C. L. Wilson 861 oom i. pancheri (Porch. & sie Briq. New Caledonia:
Thorne 28585 (rsa). H. patula Guillaum. New Caledonia: H. S. McKee 3543
(A). H. procumbens DC. Australia: Long 209 (a). H. paces geo New
1969} DICKISON, DILLENIACEAE, IV 397
H. scandens (Willd.) Dryand. Australia: Cult. prt, s.2. (preserved material);
K, sm. (preserved material). H. sericea (R.Br.) Benth. Australia: Muir
855 (A). H. serrata Hotchkiss. Australia: C. L. Wilson 855 (us). H. stirlingii
C. T. White. Australia: C. L. Wilson 757 (us). H. stricta (DC.) R.Br. ex F.
Muell. Australia: Hoogland 8420 (cans). H. subvaginata (Steud.) Ostenf. Aus-
tralia: C. L. Wilson 764 (us). H. tetrandra (Lindl.) Gilg. Australia: C. L. Wil-
son 848 (us); Cult. K, s.2. (preserved material); Cult. £, C3544. H. tomentosa
R.Br. Australia: Specht 638 (A). H. tontoutensia Guillaum. New Caledonia:
McMillan 5060 (a). H. trachyphylla Schlechter. New Caledonia: Hiirlimann 846
(A). H. uncinata (Benth.) F. Muell. Australia: E. H. Wilson 155 (a). H. vaginata
(Benth.) F. Muell. Australia: C. L. Wilson 859 (us). H. vestita A, Cunn. Aus-
ia: New South Wales. NSW 55998 (a). H. wagapii Gilg. New Caledonia:
Thorne 28266 (GH).
Pachynema dilatatum Benth, Australia: Northern Territory. NT 6129. P. jun-
ceum Benth. Australia: Northern Territory. NT 6750.
Schumacheria castaneifolia Vahl. Ceylon: Abeywickrama s.n. (preserved ma-
terial). S. angustifolia Hook. f. & Thoms. Ceylon: us 597415.
Tetracera akara (Burm. f.) Merr. Borneo: Elmer 21314 (a). T. arborescens
Jack. Sumatra: Toroes 5293 (a). T. asiatica (Lour.) Hoogl. Hainan: Lau 3875
(a). T. asiatica (Lour.) Hoogl. ssp. asiatica Hoogl. China: Liang 69507 (a). T.
boiviniana Baill. Tanganyika: Tanner 2548 (uc). T. daemeliana F. Muell. Aus-
material). T. volubilis L. Mexico: Purpus 7647 (MO).
LITERATURE CITED
Bartey, I. W. 1956. Nodal anatomy in retrospect. Jour. Arnold Arb. 38: 269-
287
——, & R. A. Howarp. 1941. The comparative morphology of the Icacin-
aceae. I. Anatomy of the node and internode. Jour. Arnold Arb. 22: 125-132.
—, & C. G. Nast. 1943. The comparative morphology of the Winteraceae.
II. Ca : ; rb. 24: 472-481. :
—— & cali ve er fanharscntn morphology of the Winteraceae. IV.
Anatomy of the node and vascularization of the leaf. / bid. 25: 216-221. —
Barton, H. 1866-67. Observations sur l’anatomie des Dilléniacées. Adansonia
: 88-93,
———.. 1871. The Natural History of Plants. Vol. 1. (Transl. by M. M. Harteg).
L. Reeve & Co., London. :
Benzinc, D. H. 1967a, Developmental patterns in stem primary xylem of —
Ranales. I. Species with unilacunar nodes. Am. Jour. Bot. 54: 805-815.
398 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
. 1967b. Developmental patterns in stem primary xylem of woody Ra-
nales. II. Species with trilacunar and multilacunar nodes. /bid. 813-820.
Canricut, J. E. 1955. The comparative morphology and relationships of the
Magnoliaceae. IV. Wood and nodal anatomy. Jour. Arnold Arb. 36: 119-140.
Corpemoy, C. J. pe. 1859. Note sur les ovules de deux genres de Dilléniacées.
Bull. Soc. Bot. France 6: 409-411, 449-450.
DEcKER, J. M. 1967. Petiole Ve of Luxemburgieae (Ochnaceae).
Am. Jour. Bot. 54: 1175-1
Dicxison, W. C. 1967a. eee Pon aa studies in Dilleniaceae. I.
Wood anatomy. Jour. Arnold Arb. 48: 1-29.
. 1967b. Comparative neon ae ra studies in Dilleniaceae, II. The
pollen. /bid. 231-240.
Esau, K. 1965. Plant Anatomy, 2nd ed. John Wiley & Sons, Inc., New York.
Foster, A. S. 1956. Plant idioblasts: remarkable examples of cell specialization.
Protoplasma 46: 184-193.
Gite, E. 1893. Dilleniaceae. Nat. Pflanzenfam. III. 6: 100-128.
HiITzeMANN, C. 1886. Beitrage zur vergleichenden Anatomie der Ternstroemia-
ceen, Dilleniaceen, Dipterocarpaceen und Chlaenaceen. (Inaug. Diss.) Univ.
Kiel.
cageager R. D. 1952. A revision of the genus Dillenia. Blumea 7: 1-145.
959, Additional notes on Dilleniaceae 1-9. Ibid. 9: 577-589.
How en A. 1962. The vascular structure of the petiole as a taxonomic
aoe. Jn: Garnaud, Advances in horticultural is and their appli-
cations. Vol. III. pp. 7-13. Pergamon Press, New York.
Keno, H. 1962. Comparative morphological studies in Theaceae. Univ. Calif.
Publ. Bot. 33: 269-384.
Marspen, M. P. F., & I. W. Battey. 1955. A fourth type of nodal anatomy in
dicotyledons, illustrated by Clerodendron trichotomum Thunb. Jour. Arnold
Arb. 36: 1-51.
MeeusE, A. D. J. 1966. Fundamentals of Phytomorphology. The Ronald Press
Co., New York.
Mercatre, C. R., & C. CHALK. 1950, Anatomy of the Dicotyledons. 2 Vols.
The Clarendon Press, Oxford.
Nampoonrrr, K. K., & C. B. Beck. 1968b. A comparative study of the primary
vascular system of conifers. II. ea with opposite and whorled_ phy!l-
lotaxis. Am. Jour. Bot. 55: 458-4
OzenpA, P. 1949. Recherches sur i aiid: apocarpiques. Publ. Lab.
YEcole Normal Supérieure, Ser. Biol. Fasc. II. Paris.
Pant, D. D., & B. MenRa. 1964. Nodal anatomy in retrospect. Phy ‘tomorphology
14: 384-387.
PaRMENTIER, M. P. 1896. Contribution a l’étude de la famille des Dilléniacées.
Compt. Rend. Assoc. francaise. Avanc. Sci. [Sess. 24] 1895. pt. 2: 626-630.
Puivipson, W. R., & M. N. Puriipson. 1968. Diverse nodal types in Rhodo-
dendron. Jour. Armold Arb. 49: 193-224.
Stnnott, E, W. 1914. Investigations on the phylogeny of the angiosperms. *
The anatomy of the node as an aid in the classification of the angiosperms.
Am. Jour. Bot. 1: 303-322.
SOLEREDER, H. 1908. Systematic Anatomy of the Dicotyledons. (Engl. transl. by
oodle & Fritsch.) Vol. I. Oxford Univ. Press, London
mete H. 1895. Beitrage zur vergleichenden anatomie der Dilleniaceen.
Beih. Bot. Centralbl. 62: 337-342, 369-378, 401-413.
1969] DICKISON, DILLENIACEAE, IV 399
Tucker, S. C. 1964. The terminal idioblasts in magnoliaceous leaves. Am. Jour.
Bot. 51: 1051-1062.
DEPARTMENT OF BIOLOGY
VIRGINIA POLYTECHNIC INSTITUTE
BLACKSBURG, VIRGINIA 24061
EXPLANATION OF PLATES
PLATE .
Fics. 1-8. Dilleniaceae, nodal anatomy. Tetracera indica (seed received
from H. Keng, Sin gapore), transverse Ae of cotyledonary node showing
unilacunar range condition (c.t., cotyledonary trace), X 32. 2, Hibbertia
pungens (Royce 7640), transverse section of unilacunar node, X 32. 3, Tetra-
cera boiviniana (Tanner 2548), transverse section of trilacunar node illustra ating
widely separated gaps, X 13. 4, Acrotrema sp. (Thwaites CP3899), transverse
through
section through rhiz f leaf trace (It) an entitious root de-
parture, e scandens (cult. BRI, s.m.), transverse section of node
illustrating trilacunar condition with widely separated gaps t pet-
, ' vill gosa (de Mattos Filho s.m.), tranverse section of
pentalacunar node, . 7, Schumacheria castaneifolia (Abeywi a S$.M.),
ransverse section of weorree node. Note the rap trace departs paint
aii (It, leaf trace), 13, illenia ovata (cult. SING, 5.7.), transvers:
PLATE II
Figs. 9, 10. Dilleniaceae, petiolar anatomy. 9A, B, C, D, Acrotrema (Thwaites
CP3899), transverse sections of the petiole and midrib illustrating omaha n o
abaxial arc and adaxial bundle, &* 32. 10A, B uratella amer. chia oe rw
5470), eens sections of petiole illustrating formation of ull vee 7 beanies,
xX 1 transverse section of petiole at e amina showing
same
medullary indies Craps d by arrows), complete fusion of vascular cylinder,
and extraxylary fibers, X 30.
PLATE I
Fics. 11-14. Dilleniaceae, pears nd Bee anatomy. All figures < 30. 11A
verse section of petiole at base of lamina showing
bundles by invagination. 14, H. patula (McKee 3543), “transverse section
petiole at base of lamina depicting confluent — cylinder. Note ae
ber
PLATE IV
Fics. 15-22, Dilleniaceae, petiolar anatomy. 15, Didesmandra aspera (Sara-
wak, s.m.), transverse section of
transverse section of petiole at base 0 s
ected ose @) 32 ‘ ey coriacea (Humbert pon trans-
verse section of petiole at base 0 g vascular tissue, <
of lamina
18, Dillenia "castoneifotia: (B (Womersley N GF 3768), transverse section of. petiole
at base of lamina showing arc of medullary bundles, X 15. 19, D. bolste
400 * JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ge 3112), transverse section of Sag aes at base of lamina rapes confluent
— S pte “sa vascular cylinder, x 3 beccariana (SAR 16272), trans-
verse section of petiole at base Ste ‘showing superimposed medullary
bu a Gndicated by arrows), X 17. 21, Doliocarpus guianensis (uc 947180),
transverse section at base of petiole showing abaxial arc of dissected bundles
and adaxial siphonostele, X 17. 22, Davilla aspera (Howard 10502), transverse
section of petiole at base of lamina 7 ai nearly complete vascular cylinder.
Note abundant sclereids in cortex, & 3
PLATE V
. 23-26. Dilleniaceae, minor venation. All figures X 25. 23, Dillenia
. Q. Lagrimas not
ley & Millar NGF 7000), note complete absence of free vein endings accom-
panied by massive venation and bundle sheathing. Sheath cells extend into vein
islets. 26, Didesmandra aspera (SAR S.18297), note weak, slender venation
and incomplete bundle sheathing.
PLATE VI
Fics. 27-30. Dilleniaceae, minor venation. 27, Hibbertia pepe (Eichler
17965), a2 of leaf showing single leaf oe and termination of reflexed lateral
veins, 5. 28, H. subvaginata (C. L. Wilson 764), note massive cya lack-
ing bundle sheathing, X 35. 29, H. b anki ew 8431), venation showing char-
acteristic interrupted oo sheat 25. 30, H. wagapii (Thorne
28266), note terminal veinlets are ee enclosed by i es oer
bundle sheathing, « 2
PLATE VII
Fics. 31-34. Dilleniaceae, minor venation. 31, Curatella americana ({rwin
5470), minor venation showing abundant parenchymatous bundle sheathing, X
25. 32, Tetracera macrophylla (Canright 1127), note ctr See bundle
sheathin ng completely surrounds terminal tracheids, X 25. 33, Doliocarpus den-
tatus se f 10088), terminal veinlet with abundant parenchymatous ape
ing. sheath cells often orientated at right angles to vein, x 54.
Pachynena aia (NT 6129), scale-like leaf eae by weak jae
traces, X 5
PLATE VIII
isc 35-39. Terminal veinlet — in mtg a. 35, B. scandens Co
s.n.), X 130. 36, H. dentata (cult. K, s.n.), X 1 7. fi. “nuegelli cL
son ihy, x 100. 38, H. pachyrhiza (C. L. Wilson a S130. 39, H. scandens
(cult. K, s.m.), K 1
PLATE IX
Fics. 40-44. Dilleniaceae, major venation. 40, Dillenia indica (seed received
from H. Keng, rin ata pegs donary node and vascularization n of cotyle edon.
41, the — vasculariza of first foliage ge nd leaf. 42, route age
ta (cult. K, $.m.), natur. “ cae 43, ‘ tontoutensia (McMillan 5060), x1 4,
H. cisnesformis (Wilson 297), X 2
PLATE X f
Fics. 45-50. Leaf vascularization in Hibbertia. Due to i apeceteem ps —
size, magnifications 4.
H. huegelli (C. L. Wilson 777), X 3.5. . 46, H. mo yna (Maiden ns. ;
41, H. nitida (Fl. Novae Holl, 141), % §. 48, H. vestita (NSW 55098), x 13. 49,
H. exutiacies (Eichler 17965), x > oe H. see tas tase 387), X
Jour. ArNotp Ars. VoL. 50 PLaTE I
Dick1son, DILLENIACEAE, IV
Pirate II
Jour. ARNOLD Ars. VOL. 50
DickIson, DILLENIACEAE, IV
*
Jour. ARNOLD Ars. VoL. 50
12
Dick1son, DILLENIACEAE, IV
Prate Iii
Jour. ARNOLD ArB. VOL. 50 PraTE IV
Dickison, DILLENIACEAE, IV
Jour. ARNOLD Ars. VoL. 50 PLATE V
DickIson, DILLENIACEAE, IV
aot
Jour. ARNOLD Ars, VoL. 50 PraTe VI
Dickison, DILLENIACEAE, IV
Jour. ARNOLD Ars. VoL. 50 Piate VII
DickIsON, DILLENIACEAE, IV
|
|
t
Jour. ARNOLD Arp, VoL. 50 PiaTeE VIII
Dickson, DILLENIACEAE, IV
fis tae nna
Jour. ARNOLD Ars. VoL. 50
Dick1son, DILLENIACEAE, IV
PLaTE IX
—
1969 | UHL, NANNORRHOPS RITCHIANA 411
ANATOMY AND ONTOGENY OF THE CINCINNI AND FLOWERS
IN NANNORRHOPS RITCHIANA (PALMAE) !
NATALIE W. UHL
THE LARGE, TERMINAL, compound inflorescence of Nannorrhops ritchiana
(Palmae-Coryphoideae) is composed of unspecialized branch systems
(Tomlinson & Moore, 1968) which may serve as a model for the deriva-
tion of more specialized types of palm inflorescence. Observations on the
inflorescence of Nannorrhops ritchiana are continued here with a descrip-
tion of the anatomy and some aspects of the ontogeny of the rachillae, of
the ultimate flowering units, and of the flowers. Nannorrhops is especially
important because completely sheathing and vasculated bracteoles are
present throughout the ultimate flowering unit. Detailed studies confirm
Tomlinson and Moore’s tentative designation of this unit as a cincinnus
and reveal basic constructional principles that apply to many, if not all,
of the varied flowering units found in palms; e.g. the triad of a pistillate
and two staminate flowers, where interpretation has been difficult because
bracts are absent or lack vasculature (Uhl, 1966). The form and anatomy
of the carpel may also illustrate some primitive features for palms.
MATERIAL AND METHODS
Inflorescence branches from plants at the Fairchild Tropical Garden,
Miami, Florida, were available in various stages of development from the
following collections: Moore 6009, Read 735, and Tomlinson 14.X1.63 and
14.X1.66. These were fixed in formalin-acetic acid-ethanol, desilicified for
1 to 2 weeks with approximately 1/3 commercial strength hydrofluoric acid,
and embedded in Paraplast. Serial sections of flowers and rachillae were
made at 5, 7, 10, and 15 microns and were stained with safranin and fast
Sreen or safranin and aniline blue. Cincinni and flowers were also cleared
as described previously (Uhl, 1966), the number of cleared flowers ex-
amined exceeding 50. Two films were prepared for cinematographic anal-
ysis (Tomlinson & Zimmermann, 1965) of rachillae and mature flowers.
Some observations and photographs (Fics. 8-19) were made in polarized
light. Since growth is continuous, but not uniform, dimensions of the ma-
terial examined are included below.
RACHILLAE
Morphology. Structural patterns are simple despite the large size of
the inflorescence in Nannorrhops (Tomlinson & Moore, 1968, Fig. 42). Up
i *From work supported by National Science Foundation Grant GB-7758; principal
investigator, Harold E. Moore, Jr
412 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
to five orders of branches are formed monopodially. The visible flower-
bearing axes or rachillae are mostly branches of the fourth order, but
whatever the order, they are similar in size and in the number of flower-
clusters or cincinni produced. Rachillae taper slightly in diameter (from
1.5 mm. to 0.75 mm.) and are indeterminate in length and in potential
number of flower clusters. Fully expanded rachillae in the material ex-
amined range from 5 to 12 cm. in length and bear from about 20 to 45
cincinni. A few distal cincinni are usually abortive.
Development. Maturation of flowering axes within the inflorescence
is complex. Four different patterns can be recognized: one with reference
to the inflorescence as a whole, a second in the sequence of development
of lateral branches, a third on individual rachillae, and a fourth within
each flower cluster.
4
Fics. 1-7. Fics. . 1-3, Three successive developmental stages of a third yee
branch, am mee indicate sequence, further explanation in text. Fic. 1 immature
upper with one petal and one stamen removed, X 5; Fic. 6, part t of a cleared
rachilla to show bundles in —. pabcaslies stalks of first flowers of cincinni,
1969 | UHL, NANNORRHOPS RITCHIANA 413
Maturation is basipetal in the inflorescence as a whole. Upper first and
second order branches produce the first flowers (Tomlinson & Moore,
1968, Fig. 40).
The further expansion of specific lateral branches is not uniform but
can be related to the order of the branch. Third order branches mature
acropetally, but development of fourth order branches is irregular. Ma-
ture flowers are produced on some fourth order axes when others are
still in early stages of development as illustrated by Fics. 1-3. In early
Stages of growth, some fourth order branches are equal in size to the main
branch (third order axis) on which they are borne (Fic. 1). The result
is a digitate configuration, which may be useful in interpreting digitate
branching in mature inflorescences elsewhere in palms. In a later stage
(Fic. 2) some of the fourth order branches have matured while others are
still undeveloped.
Flowers may mature irregularly on a specific rachilla. Those in cincinni
at the middle of the rachilla often develop before those in cincinni nearer
the base or the apex (Fic. 3), but in general the order of development is
acropetal. Within each cincinnus there is still another acropetal series in
the maturation of individual flowers (Fic. 4).
Anatomy. The vascular system in rachillae is composed of a central or
subcentral group of about 10 (8-12) large vascular bundles with a num-
ber (ca. 13) of intermediate and smaller bundles peripheral to them (Fics.
8, 9, 12). The peripheral bundles represent strands which supply cin-
cinni, and they vary in number and position depending on the proximity
of the level examined to a cincinnus. Each large bundle has 1 to 4 large
vessels (Fic. 12) and a complete fibrous sheath which is approximately
5 to 7 cells wide over the xylem. Most commonly there are two large ves-
sels per bundle, but bundles about to branch have three large vessels and
small branch bundles only one. The narrow cortex is of small unspecialized
parenchyma 6 to 8 cells in width, and the epidermis is of smaller iso-
diametric cells.
BRACTS
Morphology. Two types of bracts may be present on rachillae. On
first, second, and third order branches there is usually an irregularly bi-
carinate prophyll which is inserted basally in an adaxial position. This
bract is commonly empty on first and second order branches but on third
order branches it subtends the first lateral branch.
Flower clusters are subtended not by prophylls but by irregularly fun-
nel-shaped bracts with attenuate dorsal tips. Similar but larger bracts
borne on one axis and subtending branches of the next order occur through-
out the inflorescence and may be arranged in a reduction series from a
foliage leaf (Tomlinson & Moore, 1968). Bracts subtending cincinni are
the smallest of the series and are all equal in size and shape at maturity.
On fully expanded rachillae, each bract is about 3 mm. long, the sheathing
Part extending for ca. 2 mm. of this.
414 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
—13. Successive transections of a cincinnus, taken in peter i
oS
at leve origin of pet ae be ey on second flower, X 36; Fic. *':
transection at level of o r trace to prophyll on sec cond flower, X 3
Fic. 12, transection of "Pchatle. An stalks of first, second, third, and fourth
1969 | UHL, NANNORRHOPS RITCHIANA 415
Anatomy. Each bract subtending a cincinnus is supplied by five vas-
cular bundles and by a large number of fibrous strands (Fics. 6, 8,9). The
vascular bundles, which may be designated as a midvein and two pairs of
lateral bundles, originate as small branches of peripheral stelar strands.
The continuing vertical bundles (Zimmermann & Tomlinson, 1965) from
which the midvein and first pair of lateral bundles originate usually enter
the stalk of the first flower. Vertical bundles providing the second pair
of laterals, however, continue in the rachilla. Lateral vascular bundles
branch and anastomose distally in the bract (Fic. 6). The numerous
fibrous strands (Fics. 6, 8) are wide tangentially and also branch and anas-
tomose distally. They are tapered somewhat proximally but are not con-
nected to the vascular cylinder of the rachilla.
FLOWERING UNITS
Morphology. With the initiation of the first flower, the growth pattern
of the inflorescence shifts from monopodial to a sympodial elaboration of
clusters, each consisting of five or six successively younger flowers (Fic.
4). The bract on the rachilla subtends the first flower. The stalk of this
flower in turn bears an adaxially situated bracteole which is completely
sheathing and has two subequal adaxial tips, thus differing from the bract
subtending the first flower and definable as a prophyll. The prophyll sub-
tends the second flower of the cluster. The stalk of the second flower
bears a similar bracteole which subtends the third flower. This pattern
is repeated up to five or six times in Nannorrhops (Fics. 8-13). Each
floral primordium is initiated on the opposite side of the appropriate
floral stalk and at an angle of approximately 75°. Although five or six
buds are present in the cluster, only three flowers usually mature. Because
flowers are successively younger and pedicels elongate successively during
maturation, the two-rowed condition of a cincinnus, though structurally
Present, is not readily evident macroscopically. Left-handed and right-
handed cincinni occur, depending on whether the second flower is initiated
on the left or right side of the first floral axis. A specific rachilla usually
bears predominantly left- or right-handed clusters — e.g. on a right-handed
rachilla only one or two basal and one or two median cincinni are left-
handed,
Anatomy. Bundles which supply the first flower of a cincinnus origi-
nate as branches of major axial bundles in the rachilla. The first such
branch originates at about the level of insertion of the second cincinnus
flowers, x 18; Fic. 13, transection through all flowers of a cincinnus, X 36
F .
Detatts: br, bract subtending first flower of a cincinnus; fl 1 to fl 6, successive
owers of a cincinnus; fs, fibrous bundles of bract; mv br, midvein of bract; lv,
On axis of the first flower; pr 2, prophyll of second flower, arrow points to lower
trace; pr 3, prophyll of third flower; pr 4 and pr 5, prophylls of fourth and
fifth flowers respectively; ra, rachilla.
416 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
below. About three more branches are derived from axial bundles at high-
er levels, and further branching of these provides the complete supply to
the first flower. At the level of origin of the midvein of the subtending
bract, this supply consists of a group of about 16 bundles. The exact
number of bundles is somewhat subjective unless the level is carefully in-
dicated, since bundles are frequently small, especially near their origin,
and fibrous bundle sheaths are often confluent for some distance.
Anatomically as well as morphologically prophylls are different from
other bracts in the inflorescence. The main vascular complement of each
prophyll is two vascular bundles, one supplying each tip (Fics. 9, 11, 13).
These traces are derived as small branches of marginal stelar bundles of
the floral stalk. The bracteole is obliquely inserted and irregularly bi-
carinate, one tip being slightly longer than the other. The trace to the
longer tip originates at a slightly lower level than that to the shorter (Fic.
11), and is a somewhat larger bundle which often branches distally. Un-
connected fibrous strands are also present in the prophyll (Fic. 11) and
occasionally a third vascular bundle (Fic. 11) is seen.
Above the origin of the traces to the first bracteole, the stelar bundles
of the first flower provide the vascular supply to the second floral stalk
(Fics. 8, 9, 20). Two of the ensuing bundles produce small branches,
each supplying one tip of the second prophyll (Fics. 10, 11), and the pat-
tern is repeated until up to five or six floral primordia are formed (Fics.
8-13, 20). Thus anatomically each flower, its axis, and bracteole are
identical to the others making up the cincinnus. Transections of the floral
axes of the first, second, third, and fourth flowers may be compared in
Ficures 10, 12, and 13 and their similarity noted. The pattern of origin
of the vascular supply to each floral stalk is also similar as can be seen
in Ficure 20 which is a camera lucida drawing of the major bundles in a
cleared cincinnus.
THE FLOWER AT ANTHESIS
Morphology. Among the palms, approximately 165 genera are monoe-
cious, about 39 are dioecious, and some 34 genera bear perfect flowers.
Nannorrhops belongs among the last, having perfect flowers with three
sepals, three petals, six stamens, and a tricarpellate gynoecium. Open
flowers (Fic. 5) are approximately 6 mm. long. The sepals are 3 mm.
long and are connate for two-thirds this length forming a sheath, above
which the membranaceous tips are free. Petals are ca. 5 mm. long, ovate,
somewhat fleshy, shortly imbricate near the base and then valvate. Stamen-
filaments are wide and fleshy basally (Fics. 5, 18), but taper to the at-
tachment of the versatile anthers which are subequal, basally divergent,
and laterally dehiscent. The three carpels are free in young stages, but 1m
mature flowers are connate by ventral faces through the ovarian and stylar
regions. Thus at anthesis the gynoecium is syncarpous with definite eX-
ternal grooves showing the limits of each carpel. Each carpel has a dis-
tinct stalk, an ovoid fertile part, and a long attenuate style through which
1969 | UHL, NANNORRHOPS RITCHIANA
of mature flowers, a in polarized light. Fic. 14,
Fic Sections
tangential Jongisction with two carpels, x 20; . 15, transection through sepal
a ty)
axis at level of origin of petal eae X 36; Fic. 16, transection
through base of flower above Fic. 15 3
; : Fic. 17, transection at higher level
where Carpel stipes are distinct, < 36; Fic. 18, transe ction of ovarian part of
ae he carpels connate, « 36; Fic. 19, transection of anthers and style,
ETAILS: ca s, carpel stipe; mi, micropyle; pe, petal; se, sepal tube; st,
stamen trace; sty, style.
-
418 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
a locular canal extends to open distally. There is no connection (com-
pitum, Carr & Carr, 1960) between locular canals of adjacent carpels. No
definite stigmas are present. Papillose stigmatoid tissue is apparently pres-
ent at anthesis around the stylar opening but is not developed until the
flower opens. An anatropous ovule is attached ventrally and basally in
each locule and is turned so that the micropyle is lateral rather than dorsal
in respect to the funiculus.
At anthesis about one-third the length of the flower consists of a tapered
solid basal part (Fic. 5) sheathed by the sepal tube and representing the
region of insertion of petals, stamens, and carpels. A very short petal-
stamen tube surrounds the free carpel stalks (Fics. 14, 17) but since all
organs are free just below the ovary in the mature flower, the short petal-
stamen tube does not seem to justify the term “perigynous.”
Anatomy. Floral anatomy in Nannorrhops ritchiana has been described
by Morrow (1965) and Gupta (1960). The present study confirms most
of the observations of these authors and provides further details of carpel
anatomy, organogeny, and histogenesis. The general outlines of the floral
vascular system can be seen in Fic. 7 which is a cleared preparation of
the central part of a flower. Just below sepal insertion, bundles present
in the floral stalk enlarge, extend peripherally, and branch forming a group
of bundles which provide traces to the floral organs. Further details of
this pattern are presented in a radial plot of one of the large axial bundles
(Fic. 23). The pattern is irregular in that traces to floral organs are
branches originating near the insertion of the organ or at a lower level.
Gupta (1960) reports two rings of bundles in the floral pedicels: an
outer of 11 or 12 and an inner of three larger ones. Morrow (1965) states
that 9 (8 to 10) strands enter the base of the flower. Three central strands
do mature first in floral stalks and are often larger (Fics. 12, 13). In
mature pedicels both large and small bundles are present with a gradual
transition in size. The number of bundles is somewhat subjective because
of the difficulty of getting exactly comparable levels. In the material I
studied, 10 to 15 bundles were present, five or six showing birefringent
xylem (Fics. 10, 12, 13).
Just below sepal insertion, larger bundles of the stalk extend toward
the periphery, become larger, and branch (Fic. 7). Smaller bundles may
fuse with larger ones or also branch. The floral stele, at the level of sepal
insertion, consists, therefore, of about 20 to 25 medium to small bundles,
arranged in a thick ring, the larger bundles toward the center. Fifteen (14
to 18) small sepal traces originate as branches of peripheral bundles of
this stele. Sepal traces near their origin consist of a few sieve elements
and two to four xylem elements and are very easily overlooked; but
slightly higher in the sepal-tube, fibrous caps are present on these bundles
and unconnected fibrous bundles are present between vascular strands.
Thus a ring of approximately 28 bundles is present in a transection of the
sepal-tube (Fic. 15). Five vascular bundles with four to six interspersed
==
1969 | UHL, NANNORRHOPS RITCHIANA 419
TR 20
Fic. 20. Wash drawing of a cleared cincinnus, done with Wild MS stereo-
microscope and drawing attachment, how major vascular rg to flowers.
ike
a two he fl 1 and fl 2, abscissed; younger flowers, 3 to fl 5, in bud; pr
420 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
fibrous strands represent the supply to each sepal, a midstrand and two lat-
erals reaching the tip.
There are six to nine traces, situated in a median row, in the base of
each petal (Fic. 16). These may be stelar bundles extending directly into
the petal or they may be branches of a stelar bundle. The number of
traces varies slightly. Morrow (1965) reports three and Gupta (1960)
five from receptacular strands and one from a perianth-stamen bundle. A
single median procambial strand develops first in each petal followed by
three strands at a later stage in ontogeny. The two laterals from this
group of three divide very near the central stele and with the midvein
and one or more small traces from the receptacle form the seven major
strands (Fics. 16, 17). These often produce parallel branches at higher
levels.
Traces to stamens arise in two whorls in the same manner as petal sup-
plies by the branching or direct conversion of a vertical bundle into a
stamen trace. Antipetalous traces may arise as a branch of the same verti-
cal bundle which formed the median petal trace, or as a branch or conver-
sion of an adjacent bundle. Stamen traces are large bundles which divide
in the base of the filament (Fics. 17, 18) into two traces which are oriented
xylem to xylem in the filament with the phloems lateral in position, but
which reunite in the distal part of the filament.
In the receptacle below the gynoecium, about ten large vertical bundles
(bright spots, Fic. 16) are arranged in a central ring with smaller strands
external to them. Slightly higher, all stelar bundles are divided into three
groups, one of which supplies each carpel stalk. Some 14 bundles are
present in a close group in the lower part of each stalk. One of the larger
bundles becomes the dorsal bundle of each carpel and the others form the
lateral and ventral bundles. There are usually four major pairs of lateral
bundles and two ventrals (Fic. 21). The latter may be distinguished by
position and by their extension with the dorsal bundle higher into the
style. Other small bundles are aligned along the ventral face of the locule
and at anthesis extend about one-half the length of the ovary. Branches
of the ventral bundles and the dorsal bundle extend into the style while
lateral bundles and branches of the ventrals and the dorsal vascularize the
ovary wall around the locule (Fics. 21, 22). Two or three small bundles
from the carpellary stele remain in median positions and, with a branch
from one ventral bundle, form the ovular supply. In the funiculus these
bundles are nearly confluent but divide into separate bundles in the cha-
lazal region (Fics. 21, 22).
ORGANOGENY
The value of broadening surveys of floral anatomy to include organ-
ogeny and histogenesis has been emphasized recently (Tepfer, 1953; Esau,
1965; Kaplan, 1968). Gupta (1960) includes a brief description of or-
ganogeny in Nannorrhops, but floral histogenesis has not previously been
done for a palm. The difficulty of obtaining suitable stages for such
' 1969] UHL, NANNORRHOPS RITCHIANA 421
le mc 22. Wash drawings of cleared gynoecia, prepared as for Fic. 20.
. 21, entire gynoeci sho nly bundles of the pram carpel in dorsal
F
esponding lateral bundles omitted for clarity. Detatms: db, dorsal bundle;
ib, lateral bundles; ov, ovule; ov s, ovular supply
studies in most palms is obvious. In Nannorrhops, however, particularly
: age sequences in maturation of both inflorescence branches and
c
that results in mature flowers over a long period of time, necessary material
for ontogenetic a of flowers up to anthesis may be found on
a single inflorescence bran
422 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
2000
ca
16004 st
1200+ pe| FP
pe
pe
es
8004 é
© 400+
Po
E
£
=
D
c
* 100+
: 180 220
Distance from center of axis in microns 23
Fic. 23. Plot of the radial path of a major bundle of the floral axis. DETAILS:
ca, carpel trace; pe, petal trace; se, sepal trace(s); st, stamen trace.
Above the insertion of the bracteole and subtended floral axis, floral
organs arise in acropetal succession on the flanks of the apex. The floral
apex is relatively long and is broadly ovate in outline; the one illustrated
in FiguRE 24 is ca. 50u long and 60 wide. Floral organs are similar 1n
shape in earliest stages and are developed in whorls of three, but each
whorl is actually a low spiral since no three organs are at exactly the same
eve
Sepals are essentially triangular in outline and slightly narrower than
1969 | UHL, NANNORRHOPS RITCHIANA 423
Fics. 24-27. Hrstocenests. Fic. 24, near-median longisection of a floral
apex; Fic. 25, transection showing the ep ordium of a floral se oaed and the pro-
ing it; F
of petal primordia ee. 27, transe tion f an older floral apex showing young
tee of the pa whorl of stamens. All enema to scale, Fic. 24; scale
equals 50u, Detarts: fl p, floral borconpeeietae pe, petal; pr, prophyll; se, ‘sepal:
st 1, stamen of ie whorl; t 1 and t 2, first and sions! tunica layer
other appendages. After initiation, the separate sepal primordia increase
I
size (Fic. 5), closed petals protrude ait from the sepals reaching a
424 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
length of 5 to 6 mm. in late bud. The thickened apical regions of the petals
mature first. Later elongation is by a basal meristematic region (Fic. 37).
Stamen primordia are initiated in two whorls of three and are elliptic
to triangular in outline. FicurE 27 shows the lower whorl in early stages
and Ficure 28 a later stage of the upper whorl. Initial growth is by apical
and marginal meristematic areas. Anther sacs develop in adaxial and
lateral positions (Fic. 30). Development of other aspects of the anther
is similar to but not as regular as that described by Boke (1949) for
Vinca rosea (= Catharanthus roseus). Sporogenous cells are formed by
divisions of primary parietal cells. The tapetum develops later and is
one to two cells wide (Fic. 36). In mature stages the endothecium is a
single layer of large cells (Fic. 19).
The three carpels are separate in origin (Fics. 28, 30) and show the
familiar crescentic shape illustrated for developing carpels by other authors
(Tepfer, 1953; Esau, 1965). In early stages carpels resemble stamens in
size and shape (Fics. 28, 29). Marginal and adaxial growth (Fic. 30)
provide the horseshoe-shaped primordium with a solid base and develop
what has been called an adaxial lip (Tucker, 1959). The ovule primordium
arises ventrally on one side in the base of the shallow cup-like lamina.
Directly above the insertion of the ovule, ventral sutures of the carpels
are open (Fic. 33). The submarginal position of the ovule can be seen in
a young carpel (Fic. 33).
Fusion of the three carpels is ontogenetic and begins in the style. FIc-
URES 32 to 34 are a series of transections of a gynoecium 230, in height.
Only the upper 130, of the styles are connate. Ficure 34 is the first sec-
tion (proceeding distally) which shows connation. Fusion is by meri-
stematic activity along the appressed ventral faces of the carpels. Initially
epidermal cell walls become pointed and interlock (Fic. 35). Subsequent
cell divisions produce a solid zone of tissue with no evidence of epidermal
layers (Fic. 18). This zone closes the ventral suture of each carpel and
joins the three carpels. Fusion progresses gradually toward the base of
the gynoecium so that in the flower at anthesis, stylar and ovarian parts
are connate but stipes are still separate (Fics. 17, 18).
HISTOGENESIS
The floral apex. Esau (1965) states that the amount of zonation of a
floral apex may depend on its “determinateness,” zonation being lost oF
obscured in more determinate apices. This applies well to the floral apex
of Nannorrhops which is relatively indeterminate and shows distinct zona-
tion. The apex (Fic. 24) is zonate with a two-layered tunica, a centra
group of large corpus initials, and a rib meristem, Barnard (1960) states
that two-layered tunicas are relatively common in both floral and vegeta-
tive apices of monocotyledons and lists them in the Gramineae, Cypeta-
ceae, Juncaceae, and Liliaceae. Rohweder (1963) has since demonstrated
two-layered tunicas in the floral apices of Commelinaceae.
1969 | UHL, NANNORRHOPS RITCHIANA 425
Fics. 28-31. cerca yond continued. Fic. 28, transection of a floral apex
) flow
showing carpel primordia; Fic. 29, longisection of a youn er, stamen, and
carpel ptieedia posreinacs Ake equal in length; Fic. 30, transection of a young
nie ristematic activity adaxial in two upper carpels, marginal in lower;
, transection of base of an older flower showing adnate carpels, ee
and petals. All referable to scale Fic. 29; scale equals 50”, DETAILS: ca, :
Pc, procambial strand; pe, petal; se, sepal; st, stamen; st 1, stamen of Tex
horl; st 2, stamen of upper whorl.
hy
a
w
ss
Prophyll and floral primordium. The prophyll is Aen sion otc
and is initiated first (Fic. 24, right). In early stages it is aes in
outline. There appear to be oblique or periclinal divisions in the dermat-
426 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
: Fics, 32-35. DEVELOPMENT OF SYNCARPY, Fics. 32-34. Successive transec-
tions through a young gynoecium ca. 2304 long. Fic. 32, ovarian part of gynoe-
n
2
e . 32,
op of ovules, carpels free, ventral sutures open; Fic. 34, 404 above Fic. 33,
first section showing fusion of carpels; Fic. 35, transection of epidermal layers
Ww 0 carpels in mat i
dermal cells. All referable to scale Fic. 34; scale equals 504. DE
dermis; ov, ovule; unlabeled arrow, Fic. 34, indicates area of fusion of carpel
ogen in the initiation of the tips of the prophyll. This is the only place
where periclinal divisions were observed in the first tunica layer. After
initiation, each segment of the prophyll is extended by marginal growth,
— vs
1969 | UHL, NANNORRHOPS RITCHIANA 427
the two extensions meeting to complete the abaxial sheathing part of the
prophyll. The adaxial part of the sheath is adnate to the axis to a slightly
higher level and apparently develops by intercalary growth.
Floral organs. All floral organs are initiated by periclinal divisions in
the second tunica and usually only one underlying corpus layer. Initiation
of petals is illustrated in Fic. 26, stamens in Fic. 27, and carpels in Fic.
28. Only anticlinal divisions were observed in the first tunica layer dur-
ing the development of floral organs.
Procambium. The difficulties of determining direction of maturation
of procambium are well recognized. In all organs studied for Nannorrhops,
development of the first procambial strands appears to be acropetal. The
first recognizable procambium in a floral stalk is in the form of three central
strands. In all floral organs, a single median strand of procambium de-
velops first. This is present in sepals when they are about 160, high.
Stamens and carpels are about 40, in length when the median strand is
recognizable. A single procambial strand is present in petals when they
are about 250 long and three strands are developed when the petals are
early elongation of these organs to enclose developing stamens and carpels
which achieve more maturity before elongation.
DISCUSSION
The cincinnus. For obvious reasons the sometimes huge inflorescences
of palms have not been readily available for detailed studies. Within the
family much diversity is found in both major axes and ultimate flowering
units. Evolution in the inflorescence of Nannorrhops appears to have re-
sulted in complex patterns of maturation rather than in extreme conden-
Sation and/or fusion. Consequently study of this genus is particularly
helpful in understanding other genera where more reduction is present.
The monopodial systems of major axes are described in a previous paper
(Tomlinson & Moore, 1968). With the initiation of the first flower,
growth in the inflorescence changes abruptly from monopodial to sym-
podial.
Designation of the flowering unit in Nannorrhops as a cincinnus is not
readily evident macroscopically because the five to six flowers within each
cluster are successively younger. Details of anatomy and ontogeny, how-
ever, show that the basic unit in each flower cluster is a single flower bear-
ing a distinctive bracteole on its axis. In the axil of the bracteole, a new
floral primordium is initiated at an angle of approximately 75° on the
alternate and abaxial side of each successive floral stalk. Thus the theo-
retical main axis of the unit is reversed at each primordium and the result
is a short scorpioid cyme or cincinnus (Rickett, 1955).
Comparison of the ultimate units of Nannorrhops and Aristeyera (a
428 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Fics. 36, 37. Fic. 36, transection of part of anther, for magnification refer to
7 q
scale Fic. 34; Fic. 37, near-median longisection of young flower; scale e uals
50u, DETAILs: ca, carpel; pe, petal; sp c, sporogenous cells; st, stamen; ta, tape-
tum.
triad of flowers, Uhl, 1966) suggests that the angle of divergence and
position of the bracteole and its subtended primordium determine the
shape and consequent definition of the flower cluster. In Avisteyera, each
floral primordium is borne on the adaxial side of the axis rather than the
abaxial as in Nannorrhops and the angle of divergence is approximately
25° to 45°. This type of analysis seems to be applicable to many of the
diverse ultimate units in palms which are to be treated in detail elsewhere.
Realization that the bract on the flower may be distinctive in shape and
anatomy is also useful in interpreting other units. In Aristeyera no vas-
cular bundles are present in the bracteole. The second bracteole is bi-
carinate, however, suggesting a prophyll as in Nannorrhops. The signifi-
cance of the prophyll, a bract which is morphologically and anatomically
different from other bracts in the inflorescence, is not apparent at this time.
The flower. Barnard (1955, 1957a,b, 1958) found that in the
Gramineae, Cyperaceae, and Juncaceae, stamens were initiated in deeper
layers of the floral meristems than other floral organs and, therefore, more
closely resembled axial buds. Sharman (1960) also thought stamens
(Gramineae) were cauline since they are more like buds in initiation and
leaves in patterns of initiation and growth. Because of the nature of palm
leaves, developmental patterns are obviously complex and cannot be com-
1969 | UHL, NANNORRHOPS RITCHIANA 429
pared to those of floral organs except in very earliest stages. Leaf pri-
mordia in some palms as described by Periasamy (1962) seem similar
to those of floral organs in Nannorrhops but histogenesis has not been
studied.
Evidence from organogeny and histogenesis in Nannorrhops suggests
that all floral organs are homologous. Stamens and all other floral ap-
pendages arise by periclinal divisions in the T, and one or more corpus
layers. In early stages organs are similar in form and all receive an initial
median procambial strand. Later growth patterns differ according to the
whorl involved. Sepals develop rapidly and enclose other organs, but re-
main separate from other floral whorls; while petals, stamens, and carpels
become briefly adnate showing zonal growth for a short distance at the base
of the flower (Fics. 14, 31). In addition to evidence from histogenesis,
the shape and anatomy of the mature stamens suggest a laminar or foliar
nature. The filaments are very wide at the base (Fic. 5). Further the
large vascular bundle divides near the base of the filament suggesting the
multiple trace condition of foliar stamens (Canright, 1952; Moseley,
1958)
In general the vascular system of the Nannorrhops flower is similar to
that of Rhapis (Uhl, Morrow, & Moore, 1969) and differs from that of
the arecoid palms, Juania, Ravenea, and Ceroxylon (Uhl, in press). Ma-
supply of the carpels. In Juania, Ravenea, and Ceroxylon, carpels are
connate peripherally and ventral sutures are not completely closed at an-
thesis; in Rhapis carpels are separate and those of Nannorrhops are con-
nate by ventral faces. Ventral sutures are closed at anthesis in both the
latter taxa. The ovular supply in the arecoid genera is a single bundle
formed by fusion of a branch from each ventral bundle. In both Rhapis
and Nannorrhops, a branch from one ventral and branches of several
other bundles form the ovular supply.
Within the palms the Nannorrhops flower is relatively unspecialized but
within the Coryphoideae, it is one of a few in which extensive syncarpy
is developed. Ontogenetic development of syncarpy would seem to re-
late N. annorrhops to other Coryphoideae with separate carpels, and stylar
origin of the fusion suggests further connection to a group of coryphoid
genera in which the carpels are connate by the stylar regions only. A sec-
ond type of fusion is seen in the sepals. The connate base arises as a unit
with no evidence of union during ontogeny. Many coryphoid genera have
connate sepals (Morrow, 1965), the significance of connation here is not
understood at the present time.
Much has been written and argued about the basic nature of the angio-
sperm carpel (Eames, 1961; Tucker, 1959). Consideration of many as-
pects of carpel structure in palms is beyond the scope of this paper and
will be considered in a later survey. It is tempting, however, to point out
here that certain features of carpels in the monocotyledons (Rhapis,
Nannorrhops, Juania, Ravenea, Ceroxylon) seem equally and, perhaps,
430 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
more primitive than those of the Ranales (sensu Eames, 1961). In early
stages, carpels of Nannorrhops are separate, stipitate, and conduplicate,
with open ventral sutures. These are features considered primitive in
carpels ( Bailey & Swamy, 1951; Baum, 1961). The ovule, in both Nan-
norrhops and Rhapis, is attached basally and submarginally to one side
of the laminate region. The large vascular supply to the ovule and its
origin in both genera, when considered with the unspecialized form, lead
to the surmise that a single ovule may possibly be primitive in palms.
ACKNOWLEDGMENTS
Acknowledgment is due to Professor Harold E. Moore, Jr., for valuable
help during this study. Thanks are also extended to Mrs. Donald Ferguson
for technical assistance.
LITERATURE CITED
Barney, I. W., & B. G. L. Swamy. 1951. The conduplicate carpel of dicotyle-
dons and its initial trends of specialization, Am. Jour. Bot. 38: 373-379.
Barnarb, C. 1957a. Floral histogenesis in the monocotyledons. I. The Gramin-
eae. Austral. Jour. Bot. 5: 1-20.
. 1957b. Floral histogenesis in the monocotyledons. II. The Cypera-
ceae. Ibid. 115-129.
. 1958. Floral histogenesis in the monocotyledons. III. The Juncaceae.
Ibid. 6: 285-298. is
. 1960. Floral histogenesis in the monocotyledons. IV. The Liliaceae.
Ibid. 8: 213-225.
Baum, H. 1952. Uber die “primitivste” Karpellform. Gsterr. Bot. Zeitschr. 99:
63 4.
Boxe, N. H. 1947. Development of the adult shoot apex and floral initiation in
Vinca rosea L. Am . Jour. Bot. 34: 433-439.
. 1948. Development of the perianth in Vinca rosea L. Ibid. 35: 413-425.
. 1949. Development of the stamens and carpels in Vinca rosea L. Ibid.
36: 535-547.
CanricHt, J. E. 1952. The comparative morphology and relationships of the
Magnoliacese I. Trends of specialization in the stamens. Am. Jour. ot.
39: 484—
Carr, S. G. 146 & D. J. Carr. 1961. The functional significance of syncarPy-
ma 11: 249-256.
EAMES, A. J. 1961. Morphology of the angiosperms. McGraw-Hill Book Co.,
Esau, K. 1965. Plant Anatomy. John Wiley & Sons, Inc., N.Y.
Gupra, S. C. 1960. Organogeny and floral coer of Nannorrhops ritchieana
. Wendl. Jour. Res. Agra. Univ. 9: 103-
KapLan, D.R. 1968. Histogenesis of the era and gynoecium in Downin-
gia bocigalutés Am. Jour. Bot. 55: 933-950.
KAussMANN, B. 1963. Pflanzenanatomie. Gustav Fischer. Jen ane
Morrow, L. O. 1965. Floral morphology and anatomy of seitats Coryphoidea
(Palmae). Ph.D. Thesis, Cornell Univ.
1969 | UHL, NANNORRHOPS RITCHIANA 431
MoseLey, M. F., Jr. 1958. Morphological studies of ns Nymphaeaceae — I.
The nature of the stamens. Phytomorphology
PERIASAMY, K. 1962. Morphological and ontogenetic ieee in Palms — 1. De-
ve lopment of the plicate condition in the palm-leaf. sade ees 12:
54-64.
Rickett, H. W. 1955. Materials for a page 8 of botanical terms — III. In-
florescences. Bull. Torrey Bot. Club 82: 445.
ROHWEDER, Otto. 1963. Anatomische und nacda Untersuchungen an
Laubsprossen und Bliiten der Commelinaceen. Bot. Jahrb. 82: 1-99.
SHARMAN, B. C. 1960. Developmental anatomy of the stamen and carpel pri-
mordia i in oo odoratum, Bot. Gaz. 121: 192-198.
TEPFER, S. S. 1953. Floral anatomy and ontogeny in Aquilegia formosa var.
truncata and Ranunculus repens. Univ. Calif. Publ. Bot. 25: 513-648
ToMLINnson, P. B., & H. E. Moore, Jr. 1968. ass in Nannorrhops
ritchiana (Palmae). Jour. Arnold Arb. 49: 16-
Tucker, S. C. 1959. Ontogeny of the ie hihi oe the flower in Drimys
winteri var. chilensis. Univ. Calif. Publ. Bot. 30: 257-366.
UuL, N. W. 1966. Morphology and anatomy of the inflorescence axis and flow-
ers of a new palm, Aristeyera spicata. Jour. Arnold Arb. 47: 9-22.
. 1969. Floral anatomy of Juania, on and Ceroxylon (Palmae-
Arecoideae). Gent. Herb. 10 (4): in p
, L. O. Morrow, & H. E. Moore, a 1969. sored of the palm Rhapis
excelsa, VII. Flowers. Jour. Arnold Arb. 50: 138-1
ZIMMERMANN, M. H., & P. B. TomLtInson. 1965. retnda of the i. Rhapis
excelsa, I. a ae axis. Jour. Arnold Arb. 46: 160—
L. H. Battey Hortortum
CORNELL UNIVERSITY
IrHaca, NEw Yorx 14850
432 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ASPECTS OF MORPHOLOGY OF AMENTOTAXUS FORMOSANA
WITH A NOTE ON THE TAXONOMIC POSITION OF THE GENUS
Hsuan KENG
THE GENUS Amentotaxus was established by Pilger in 1916 (Bot.
Jahrb. 54: 41), based on the type species, A. argotaenia (Hance) Pilger.
This species, described from sterile material only, was originally desig-
nated as a member of the genus Podocarpus. As soon as its compound
staminate strobilus became known, it was transferred to Cephalotaxus,
and finally to a separate genus, Amentotaxus.
The genus Amentotaxus is endemic to eastern Asia. It was first col-
lected from a small islet near Hongkong and also from southern Kwang-
tung. Subsequently it was reported from southern Formosa, western
Hupeh and Szechuan, and from southern Yunnan and northern Tonkin
(see Fic. 1). Fossil remains have been recorded from Europe and west-
ern America (Sporne, 1965). It was generally considered as a monotypic
genus; Li (1952), however, recognizes that there are at least four dis-
tinct entities (which he considered species) involved, based on color
and relative width of the stomatal bands, and geographic distribution.
Chuang and Hu (1963), on the other hand, point out that the characters
of the stomatal band appear to be less constant, and maintain that there
is only one species, namely A. argotaenia (Hance) Pilger.
It is rather difficult to make a judgment on this controversial issue
without thoroughly examining suitable materials with reproductive struc-
tures, which unfortunately, are not available. For simplicity of nomen-
clature, since all the materials used in this study are from a small locality
in southern Formosa, the binomial Amentotaxus formosana Li is, ac
cordingly, adopted. It would be interesting to have reports on the
strobilate structures based on the materials from other parts of the
geographical range of the genus.
The plants of Amentotaxus are small to medium-sized, dioecious,
evergreen trees. A limited number of them are perhaps in existence, and
they grow in almost inaccessible places. Moreover, they are not repre
sented in any botanical garden or arboretum in the world. Owing to the
The stomatal and ovulate structures were reported by Florin (1931, 1938—
45); his interpretation of the latter, as indicated in a drawing reproduced
in 1951, p. 375, fig. 64, was apparently based on poorly preserved her-
barium material, and is inadequate. Only fragments of the embryonic
1969 | KENG, AMENTOTAXUS FORMOSANA 433
300 mis
Ficure 1. Geographic ie ee of the genus Amentotaxus. 1, Lantao Is-
land, near Hongkong; 2, Mt. Lo-fau-shan, Kovangtung: a, ni and S. Kaoh-
siung, lg (Formosa a); oe isi ae -shan, Hupeh; 5, Mt. Omei-shan, Szechuan;
, , Yunnan; 7, Cha Pa, Tonkin ( res on the ore ar specimens
cited in on 1952 ?.
development were given by Sugihara (1943); and his chromosome num-
er, m = 11, on the basis of counts from the female gametophyte, is in-
correct, as pointed out by Chuang and Hu (1963). The pollen morphology
has been carefully investigated by Erdtman (1957).
MATERIALS AND METHODS
Material preserved in FAA (including leaves, staminate strobili, ovulate
strobili, seeds, and seedlings), and dried material (including young ovulate
strobili, seeds, and seedlings) taken from herbarium specimens, were
received from Professor Ching-en Chang of the Pingtung Agriculture Col-
lege, Taiwan. All the materials were collected by Professor Chang from
near Shin-Huah Farm, Shaw-Jia, Dah-Wu, Taitung, between 1965 and
1968, Clearings of testeatiie were made with 5 percent NaOH at room
temperature. Microtome sections 10 to 12 » thick were stained with a
safranin-fast green combination.
434 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
vas. b. u.epl = pal t. spon. t.
Ficure 2. Transverse section of a leaf, and surface ~e parce views of a
stoma. a Diagram of the transverse section of a leaf; C, portions of
A, enlarged, soar the cellular details; D, sectional view gee the axis)
of a stoma; E, surface view of a stoma "(the broken lines marked guard ae’
: gua
are drawn at a different focus). Nore: aper. = apertures; guard Cc. =
cells; J. epi. = lower epidermis; pal. t. = palisade tissue; res. d. = resin ducts;
Spon. t. = Fn gpg tissue; stom. b. = stomatiferous band; subs. ¢. = subsidiary
cells: u. epi. = upper epidermis; vas, b. = vascular bundle.
Leaves. Foliage leaves are persistent, spirally arranged on the branch-
lets but twisted at the base into two rows in one plane. Internodes are
o 7 mm. long (average). Each leaf consists of a lamina and a Very
short oetiole, Laminae are coriaceous, bifacially flattened, with the adaxial
a
1969 | KENG, AMENTOTAXUS FORMOSANA 435
surface upward. They are linear, often strongly falcate, acute or more
often acuminate at apex and slightly oblique at base, 5—7 cm. long, 0.5—1
cm. broad. Two very prominent stomatiferous bands are present on the
abaxial surface and run parallel to the elevated midrib, one on each side
of it (see Plate I, a & b). Petioles are strongly decurrent on the branch-
lets.
Anatomically, each lamina possesses only one large, median vascular
bundle with a resin canal beneath (see Fic. 2, A & B). The assimilatory
tissues consist of one to two (near and at the midrib) rows of palisade
cells and numerous polygonal, elongate, and dissipated spongy cells. The
upper and lower epidermis are both well defined, the former with slightly
more thickened cuticle layer. Stomata are arranged in longitudinal rows,
their axes oriented more or less parallel to the midrib of the leaf. Each
stoma (see PLATE II, b; Fic. 2, E) is encircled by 7 to 9 subsidiary cells.
Strong papillae of the subsidiary cells surround the orifice of the stomatal
apparatus like a wall, while the guard cells are also heavily cutinized.
Sclereids are abundant, slender, branched or unbranched at one or both
ends, and generally lying between the midrib and leaf-margins and per-
pendicular to them (see PLATE II, b).
Staminate strobilus. The compound staminate strobili are produced
within the large winter bud which is borne on the top of the previous year’s
branchlets. They are short-stalked, usually four (sometimes three, rarely
two or five) together, subtended by four rows of imbricate bud-scales (see
Piate I, a; Fic. 3, A). These scales are leathery, strongly keeled and
more or less pointed. The true terminal bud of these branchlets is gen-
erally in the center and is further protected by small, thin scales (Fic.
, B); it remains dormant and resumes its activity only after expansion
and withering of the surrounding compound staminate strobili.
Each compound staminate strobilus is spike- or catkin-like, from which
is derived the generic name Amentotaxus (Fic. 3, C). When fully ex-
panded, the compound strobilus can reach a length of 2.5 to 3 cm. or
more. It consists of approximately 20 to 30 globular staminate strobili
somewhat decussately arranged (though not quite regular), growing along
the main axis in four rows. These globular staminate strobili are clearly
recognizable especially in the middle portion of the spike, since the distal
Ones are overcrowded and fused, and the lowermost ones are sometimes
adherent to the side (secondary) branches rather than being on the main
axis itself.
The staminate strobilus is globular or ovoid, 2.5 to 3 mm. in diameter
in bud (see Fic. 3, D). It is composed of 9 to 12 closely compacted
microsporangiophores,! which are peltate, with four or five (varying from
f. 20 & 21) and Pseudotaxus (Florin, 1948 a, p. 389, f. 2), microsporangiophore is
perhaps preferable, as a designation, to microsporophyll; although no trace of the
subtending bracts has been found at the base of the stalk in Amentotaxus.
436
[voL, 50
JOURNAL OF THE ARNOLD ARBORETUM
yt Py =
= ry tem Be
a Shes
Y, fi WY Y
Tt ko
ry
Ficure 3. Compound staminate strobilus, staminate strobilus and mic
sporangiophore. A, External view of an un :
of four (one is not seen) compound staminate strobili; B, the same, with
ro-
folded winter bud, showing 2 rage
1969 | KENG, AMENTOTAXUS FORMOSANA 437
two to eight) microsporangia hanging underneath in a semicircle and with
a short stalk near the center (see PLate II, e; Fic. 3, E). The outline of
the peltate microsporangiophores, as seen from the outer surface, varies
from round to deltoid, to more commonly polygonal, due to mutual
compression.
At maturity the thickened outermost layer and one or two (in part)
inner layers of the saerirtg walls are retained (see Plate II, d &
e). The microspores are wingless
Ovulate strobilus. The ovulate strobilus is globular to ovoid, flat-
tened dorsiventrally (see Pirate I, b; Fic. 4, Ay, As, By, B en C2).
These strobili are situated singly in the axils of foliage leaves. “The ovule
is solitary, terminal on the strobilate axis, and subtended below by five
(or six) pairs of opposite and decussate, sterile bracts; three pairs of
which are lateral and prominently keeled, and the other two (or three)
pairs are dorsiventral and only slightly curved (Fic. 4, Ay). The stalk of
the strobilus is slender, about 1-1.5 cm. long, more or less flattened and
narrowly winged.
oung ovules, at the stage of about 3 mm. in length (excluding the
sterile bracts) (Fic. 4, A,), possess an elongate conical nucellus, of
which the upper part is loosely enveloped by a single layer of integument,
the lower half, however, is seemingly associated only with the cupular aril-
lus primordium. The integument (if it is interpreted as confined to the
portion above the arillus only) and the arillus, at this stage, appear to be
completely separated. The vascular supply to the ovule, as seen in cross
section, consists of 8 to 10 normally oriented vascular bundles. They
terminate at the end of the ovule far below the nucellus — neither the
integument nor the arillus primordium is visibly vascularized.
ee the slightly older ovules at the stage of about 5 mm. in length (Fic.
B;), as a result of the enlargement of both the nucellus and the integ-
evidently embedded in the cupular arillus. The vascular supply can be
observed near the base of the ovule.
In the still older ovules, at the stage of about 6.5 cm. in length (Fic. 4,
C3; Fic. 5), the nucellus is enveloped by and fused with the integument
except for the uppermost part which remains free. Nearly two-thirds of
the integument, in turn, is embedded in and completely united with the
arillus. Isolated tracheids may be found in the lower part of the ovule at
a fairly high level in the region where the fusion of aril and integument
occurs,
of the bracts ae compound staminate strobili removed, showing the hidden
terminal bud inside; C, a compound staminate strobilus, showing a number of
ovoid to shies staminate strobili more or less decussately callgge ge on an
axis; D, a globular staminate strobilus (taken from the me ian }
larged; E, and E., two views of a spemnsinieor showing five microsporangia
arranged in a semicircle below; F, and F,, two p scatelie te eal views
of the sporangiophor
438 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
A general outline of the tissues in the largest ovule available is shown
in Ficure 5. The arillus consists of 10 to 15 layers of parenchyma cells
with rows of resinous cells lining the epidermis near the rim. The cuticle
is thin. The lower portion of the integument is composed of 12 to 20
layers of small, partly closely packed, and partly loosely dissipated
parenchyma cells. There is no clear distinction of the arillus from the
integument below the level of fusion. The upper portion of the integu-
ment is heavily cuticularized. The cells near the micropyle are enlarged,
sclerenchymatous, and oriented horizontally. The nucellus is prominently
beaked; the beak is hemispheric, and composed of numerous small polyg-
onal cells with moderately heavy walls rather loosely arranged especially
towards the micropylar end. A large portion of the nucellus at this stage,
is digested and replaced by the multicellular megagametophyte. Isolated
tracheids and short rows of tracheids are observed at the lower part of
the peripheral region where the integument and arillus are merged.
The ovular structure of Amentotaxus in general, as noted by several
authors (e.g. Florin and others), is rather similar to that of Torreya; but
its vasculature is very much simpler. In Amentotaxus, although there are
8 to 10 vascular strands entering the base of the ovule, only the isolated
tracheids are present in the lower part of the ovule, in the region where
the integument and arillus meet. In Torreya, however, there are two vas-
cular strands running up inside the arillus nearly to the apex of the seed,
each of which then sends a branch through a foramen in the stony layer
of the integument; each branch forks, forming a loop which encircles the
seed. Oliver (1903) proposed the “hyposperm theory” to explain this
peculiar vascular structure. According to this theory, all the basal part
(the “hyposperm”) of the ovule is an intercalated growth and phylo-
genetically younger than the extreme tip (the “archisperm’’). The
branching of the integumental vascular bundles inwardly is also reported
in Austrotaxus (Saxton, 1934, p. 419, fig. 18) and Cephalotaxus (Singh,
1961, p. 160, fig. k). In the case of Amentotaxus, no traces of such
branching are present. With the intercalary growth of the lower part of
the integument and arillus concomitantly with the enlargement of the
nucellus, the integument becomes evidently embedded in the arillus and
fused with it. There seems to be no evidence to prove that the lower part
of the Amentotaxus ovule is a “hyposperm,” or is phylogenetically youns-
er than the upper part.
Seed and seedling. The seed is ellipsoid-oblong, drupe-like (PLATE
I, c), 3.2 to 3.6 cm. long, 1 to 1.2 cm. broad, and slightly flattened dorsi-
ventrally. The outer part of the seed coat is completely covered and fuse
with the arillus, except the extreme tip which is exposed (Fic. 6, A & B).
The merged structure is soft-leathery in texture although the outer por-
tion is easily blistered and disintegrated when soaked in water. The
nucellus is almost entirely replaced by the ivory female gametophyte
(“endosperm”) which has an entire rather than ruminate margin (as 7
Torreya). The embryo is linear, lying in the center of the gametophyte.
Ficure 4. External and sectional views of three ovulate strobili. A, and As, two views of a young strobilus; A, and A,, longi-
tudinal and transverse sections of the same strobilus; B, and B., two views of a slightly older strobilus; Bs, longitudinal section
of the same; C, and C., two views of an older strobilus ; C;, longitudinal section of the same; Nore: micr. = micropyle; integ.
= integument; ster. br. = sterile bract; vas. b, = vasc cular bundle.
VNVSOWYOA SAXVLOLNAWNV ‘ONAN [6961
ocr
440 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
In common with other conifers, germination is of epigeal type. In the
one year old seedling examined (Prater I, d), the cotyledons have dropped,
but the two cotyledonary scars are clearly evident. The juvenile leaves
are 3.5 to 4 cm. long, 3.5 to 4 mm. wide, with two glaucous stomatiferous
bands underneath. Fundamentally of spiral arrangement, since the inter-
nodes are of variable length, the juvenile leaves appear subopposite or
rarely subverticillate.
TAXONOMIC POSITION OF AMENTOTAXUS
AND THE CLASSIFICATION OF THE CONIFERALES
Pilger (1926) assigns Amentotaxus, together with Cephalotaxus, to the
Cephalotaxaceae on the basis of the compound nature of their staminate
strobili. Kudo (1931), after seeing the ovulate strobilus, hitherto un-
known, maintains that “it (Amentotaxus) must be included in a new
family Amentotaxaceae, or in a new subfamily or tribe of Taxaceae,
but not in Cephalotaxaceae” (p. 311). As a result, a new family Amento-
taxaceae was proposed by Kudo and Yamamoto (in Kudo, 1931). Koid-
zumi (1932) strongly felt that the new family was not necessary. He,
therefore, established a subfamily Amentotaxoideae (including both Amen-
totaxus and Austrotaxus) within the Taxaceae. Later on, following his
enumeration of various similarities and dissimilarities among the Taxaceae,
Cephalotaxaceae, and Amentotaxus, he (1942), recognized that Amento-
taxus and Cephalotaxus are in fact related, and moreover, suggested that
Taxaceae and Cephalotaxaceae should be merged into one family and both
reduced to subfamilial status. Florin (1948, 1951) emphasized the dif-
ferences of ovulate strobili and stomatal structures between Cephalotaxus
and Amentotaxus, and thus sustained the transference of Amentotaxus
from Cephalotaxaceae to Taxaceae. Chuang and Hu (1965) report the
chromosome number of Amentotaxus argotaenia (Hance) Pilger (or A.
formosana Li) to be x = 7, which is different from those reported from
Taxus (x = 12), Torreya (x = 11), and Cephalotaxus (x = 12). They,
therefore, support Kudo and Yamamoto in maintaining Amentotaxus in
a separate family, the Amentotaxaceae. The present writer is inclined
to think that (1) Amentotaxus is probably better placed in the Taxaceae
than in the Cephalotaxaceae or in a separate family; (2) the Taxaceae
are not isolated, but are likely allied to the Cephalotaxaceae, probably
through Amentotaxus. These two points are elaborated in the following
paragraphs,
Features which distinguish Amentotaxus from other members of the
Taxaceae such as the spicate compound staminate strobili, the peculiar
stomatal structure (with larger number of subsidiary cells, thickened
papillae, etc.), etc., appear to be insufficient to warrant a separate family
status. The difference in chromosome number is probably inadequate to
be cited as a justification for the establishment of the Amentotaxaceae.”
*For example, in a recent report (Hair & Beuzenberg, 1958) on the chromosome
numbers of the Podocarpaceae, the following two closely related genera possess such
1969 | KENG, AMENTOTAXUS FORMOSANA 441
aS integ.
Z Fah
Z EO}
Zee
2» GN nuc. b
ca iS
(a)
ei ON aril
, res. ¢
ese AN
‘\ rely WY
i 6 a
ribet Wie
|
a A
trach.
ee
rte a
()
ot} Ch,
FIGURE . ot ae sa an ovule, details of Figure 4C;. bangs
muc. b. cellus beak; esin cells; muc. = nucellus; @ =
megagametophyte: trach. = nae ids.
On the other hand, the resemblance of Amentotaxus to the Taxaceae, es-
pecially to the genus Torreya, in the general structure of staminate strobili,
ovulate strobili, microsporangiophores, ovules, etc. is overwhelming. There-
fore, Janchen’s (1949) treatment including both Amentotaxus and Torreya
in the Tribe Torreyeae under the Taxaceae appears to be a logical one.
4 Tange of variation: Dacrydium (x = 15, 12, 11, 10, heaped & = 19, 18,
17, 13, 12, 11, 10); whereas other members of the fam o have various feent
basic numbers: Acmopyle (x = 10), a eiveaiami (x = ae iscedaiae (a = 13),
Phyllocladus (x = 9), and Saxagothaea (x = 12).
442 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Ficure 6. External and sectional views of a seed.
A more or less similar view is held by Kudo (1931), Koidzumi (1932),
Li (1952), and others.
Florin in a series of papers (1948, 1951, 1954) strongly advocated sep-
aration of Taxaceae (which includes the following five genera: Taxus,
Amentotaxus, Torreya, Austrotaxus, and Pseudotaxus [ = Nothotaxus])
from the rest of the Coniferales to form a separate order, the Taxales, 4
view originally expressed by Sahni (1920) but modified by Florin with
the exclusion of Cephalotaxus. A quite different scheme proposed by
Buchholz (1934), was summarized by Chamberlain (1935, pp- 229; 230)
as follows: The order Coniferales can be divided into two suborders: one
the Pinineae (as Phanerostrobilares or Pinares) with an obvious cone,
includes the Pinaceae, Taxodiaceae, Cupressaceae and Araucariaceae; the
other, Taxineae (as Aphanostrobilares or Taxares) without such an ob-
vious cone, contains the Podocarpaceae, Taxaceae, and Cephalotaxaceae.
Florin (1951, pp. 363, 364) fully endorsed Wilde’s (1944) postulation
that in Podocarpus, the species with 1-ovulate strobili are independently
derived from those with multiovulate strobili, and represent the ultimate
stage of reduction. In addition, his own interpretation (1951, 1954) of
the ovulate strobilate structures of the modern conifers as possibly
evolved from a much more complicated structure of fossil groups such
1969} KENG, AMENTOTAXUS FORMOSANA 443
as found in the palaeozoic Lebachia, Ernestiodendron, Walchia, and
Pseudovoltzia, has been widely appreciated. Paradoxically, he insists
that the l-ovulate strobilus of the Taxaceae is a primitive rather than
a derived condition; therefore the family Taxaceae is of entirely different
origin from the rest of the other Conifers. This is mainly because of his
emphasis on the finding of 1-ovulate Palaeotaxus in the Triassic and
Taxus jurassica in the Jurassic rocks. “Because of its high geological age”
he noted (1951, p. 349), “Palaeotaxus can hardly derive from any cone-
bearing type.” It seems he does not realize the possible existence of the
exceptionally fast rate of evolution, designated by Simpson (1944) as
tachytelic evolution. Many authors, such as Chamberlain (1935, p. 439),
Pulle (1937), Takhtajan (1953, p. 34), etc. express their notions that the
single ovulate strobilus of taxads is most likely derived from the multi-
ovulate cones. The present writer (Keng, 1963) also points out that the
evolution of the ovulate strobili in the genus PAyllocladus (belonging to
the Podocarpaceae, or according to some authors, the monogeneric family,
Phyllocladaceae) might indicate the possible mode of how the single,
pseudo-terminate ovule of taxads could have been achieved. Incidentally,
Phyllocladus is somewhat intermediate between the Taxaceae and Podo-
carpaceae; on morphological ground it is probably correct for it to be
placed in the Podocarpaceae (Maheshwari, 1962).
Although, as discussed above, Amentotaxus should be better classified
in the Taxaceae rather than Cephalotaxaceae, it does not mean that the
Taxaceae and Cephalotaxaceae are totally unrelated as suggested by Florin.
The present writer agrees with Saxton (1934), Pulle (1937), Koidzumi
(1942), and many others that these two families are in fact related. In
this connection, it is rather interesting to mention the views of Singh
(1961) who has contributed an excellent account on the life history of
Cephalotaxus drupacea Sieb. & Zucc. In his discussion of the relation-
ships of the Cephalotaxaceae and Taxaceae, he pointed out a number of
similarities between these two families and noted that they “resemble each
other in wood structure, pollen structure, and to some extent embryogeny””
(p. 193). He was, unfortunately, dominated by Florin’s misconception
that the Taxaceae are isolated and reached the contradictory conclusion
that “it appears best to regard the Taxaceae and the Cephalotaxaceae as
unrelated” (p. 193).
If we accept the general view that the compound staminate strobilus
is a primitive condition (Wilde, 1944), that the peltate sporangiophore
is more antiquated than the dorsiventral ones (Florin, 1948), and that
the one-ovulate strobilus is derived from a multiovulate strobilus (Pulle,
1937), and also if we assume that the Taxaceae and Cephalotaxaceae are
phylogenetically affiliated, then an ideal ancestral form of Taxus-Amento-
taxus-Cephalotaxus complex would hypothetically possess the following
Synthetic strobilate features. rae
taminate or microsporangiate strobili—a cluster of spike-like com-
pound strobili surrounding a terminal bud and enveloped by numerous
bud-scales (cf. Amentotaxus); each compound strobilus composed of
444 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
many ovoid or globular strobili; each strobilus consisting of many peltate,
spirally arranged sporangiophores with a number of sporangia on the
undersurface around the stalk (cf. Taxus, Pseudotaxus); each peltate
sporangiophore further subtended by a leafy bract (cf. Pseudotaxus, see
Florin, 1948a, p. 389, fig. 2; or Austrotaxus, see Saxton, 1934, p. 423,
figs. 20 & 21).
Ovulate or megasporangiate strobili—a strobilus composed of many
imbricate ovuliferous scales each with several to two (or one) ovules on
its upper surface (cf. Cephalotaxus); ovules erect, with only one integu-
ment and surrounded by a cupular arillus but free from it (cf. Taxus,
Pseudotaxus); the integument supplied by a number of lengthwise vascu-
lar bundles, each of which gives a horizontal branch in the middle of
the integument, toward the inner part of the ovule to supply the nucellus
and gametophyte (cf. Torreya, see Oliver, 1903; Austrotaxus, see Saxton,
1934, p. 419, fig. 18; Cephalotaxus, see Singh, 1961, p. 160, fig. *).
To summarize, firstly, since the resemblance of Amentotaxus to Torreya
(Taxaceae) is so overwhelming, it seems logical to include Amentotaxus
in the Taxaceae; secondly, since the family Taxaceae is intricately affiliated
to the Cephalotaxaceae on the one hand and possibly to the Podocar-
paceae on the other, Buchholz’s scheme of classification of the Coniferales,
therefore, appears to be sound.
ACKNOWLEDGMENTS
I am most grateful to Professor Ching-en Chang, who kindly made
several trips to the southern part of Taitung District, Formosa, to collect
the materials for this study, and to Dr. H. L. Li and Dr. Gloria Lim for
reading the manuscript. My thanks are also due to Professor B. Y. Yang,
Dr. Ding Hou, and Mr. M. C. Kao for supplying some of the literature;
to Dr. C. C. Hsii for translating several paragraphs from the Japanese
literature; and to Mr. D. Teow for taking the photomicrographs.
LITERATURE CITED
Bucuuotz, J. T. The classification of Coniferales. Trans. Illinois Acad. Sct.
25: 112, 113. 1934.
CHAMBERLAIN, C. J. Gymnosperms, structure and evolution. xi + 484 PP»
gs. Univ. of Chicago Press. 1935.
Cuuane, T. L, & W. W. L. Hu. Study of Amentotaxus argotaenia (Hance)
Pilger. Bot. Bull. Acad. Sinica, II. (Taipeh) 4: 10-14. 1963. :
ErptMAN, G. Pollen and spore morphology/plant taxonomy. 147 pp., fronits.,
pls. Almqvist & Wiksell, Stockholm. 1957
FLorin, R. Untersuchungen zur Stammesgeschichte der Coniferales und Cor-
daitales, I. Sv. Vet-akad. Handl. III. 10: 1-588. 58 pls. 1931.
———. Die Koniferen des Oberkarbons und des unteren Perms. -Vill.
Palaeontographica 85B: 1-729. 1938-45. [Cited in Florin, 1951.]
1969 | KENG, AMENTOTAXUS FORMOSANA 445
On the eee and relationships of the Taxaceae. Bot. Gaz. 110:
31-39. 1948
. On Nothotaxus, a new genus of the Taxaceae from eastern China. Acta
Horti Berg. 14: 385-395. 1948a.
- Evolution in Cordaites and conifers. Ibid. 15: 285-388. 1951.
. The female reproductive organs of conifers and taxads. Biol. Rev. 29:
367-389, 954
Harr, J. B., & E. _ BEUZENBERG. rage i evolution in the Podocarpaceae.
Nature 181: 1584-1586, (June) 19
JANCHEN, E. Das System der aie ‘Akad. Wien. Sitz-ber. 158: 155-262.
1949. [Cited in Li, 1952.
Kenc, H. Taxonomic position of Phyllocladus and the classification of Conifers.
Gard. Bull. Singapore 20: 127-130. 1963.
Koizumi, G. Notes on Amentotaxaceae. [In Japanese.] Acta Phytotax. Geo-
bot, i: 185. 1932.
. Further notes on Amentotaxaceae Kudo. [In Japanese.] Jbid. 11:
135, 136. 1942.
Kupo, Y., & Y. Yamamoto. Amentotaxaceae. In: Kupo, Mater. Fl. Formosa
IV. Jour. Soc. Trop. Agric. (Taihoku) 3: 110, 111. 1931.
Li, H. L. The genus Amentotaxus. Jour. Arnold Arb. 33: 192-198. 1952.
MAHESHWARI, P. The overpowering role of morphology in taxonomy. Bull.
Bot. Surv. India 4: 85-94. 1962.
Otiver, F. W. The ovules of the older gymnosperms. Ann, Bot. 17: 451-476.
1903.
PILcer, R. Cephalotaxaceae. IN: ENGLER & PRANTL, Nat. Pflanzenfam. ed. 2.
13: 267-271. 1926.
Putte, A. Remarks nc as system of the Spermatophytes. Med. Bot. Mus.
Utrecht 43: 1-17.
SAHNI, B. On certain ede features in the seed of Taxus baccata, with re-
marks on the antiquity of the Taxineae. Ann. Bot. 34: 117-133. 1920.
Saxton, W. T. Notes on Conifers VIII. The morphology of Austrotaxus spi-
cata Compton. Ann. Bot. 48: 412-427. 1934.
Smupson, G. G. Tempo and mode in evolution. Columbia Univ. Press. New
York. 1944,
SINGH, H. The life history and systematic ag oe oe drupacea
ieb. et Zucc. Phytomorphology 11: 153-
SPoRNE, K. R. The morphology of ioe tia Univ. Library.
London, 1965.
SucrHaRA, Y. Notes on Amentotaxus. [In Japanese.] Bot. Mag. Tokyo 57:
404, 405. 1943.
TAKHTAJAN, A. L. — principles of the system of higher plants. Bot.
Rev. 19; 1-45,
Wipe, M. H. A new gamma of coniferous cones. I. Podocarpaceae
(Podocarpus). Ann. Bot. II. 8: 1-41. 1944.
Yamamoto, Y. Cephalotaxaceae. Suppl. Icon. Pl. Formos. 3: 1-5. 1927.
eae, Amentotaxaceae. Ibid. 5: 7-11. 1932.
DEPARTMENT OF BOTANY
UNIVERSITY OF SINGAPORE
Bukit Trmau Roap
SINGAPORE, 10
446 JOURNAL OF THE ARNOLD ARBORETUM [VoL. 50
EXPLANATION OF PLATES
PLATE I
(Scale in each figure in 1 mm. divisions.)
Amentotaxus formosana Li. a, Cluster of compound staminate strobili from
an unfolded winter bud borne on the tip of a branchlet (cf. Fic. 3A); b, solit pd
ovulate strobilus borne in the axil of a leaf (which has dropped off) (cf.
4, C.); c, one fully mature and three young seeds (cf. Fic. 6A); d, celine
of which the two cotyledons (on the first node) ae dropped off.
PLATE II
(Scales: in a, 150 w; in b and c, eat & in d and e, 200 uw; in f, 2 mm.)
Amentotaxus formosana. a, Transverse section of a leaf showing the midrib
region (cf. Fic. 2B); b, lower (abaxial) surface of a leaf (after ee show-
ing the stomata and sclereids : c, stomata in transverse eo (cf. Fic. 2D);
d, transverse section of a microsporangiophore (ef, Pe. 3, .F,)i e, kon gitudinal
—. of a BP coms aa f, longitudinal section of an ovulate ennai
44.
(cf.
Jour. ARNOLD ArB. VOL. 50 PLATE I
KEenc. AMENTOTAXUS FORMOSANA
Jour. ARNOLD Ars. VOL. 50 PLATE II
KENG, AMENTOTAXUS FORMOSANA
1969} RUDENBERG & GREEN, LONICERA, II 449
A KARYOLOGICAL SURVEY OF LONICERA, II
Lity RUDENBERG AND PETER S. GREEN *
IN THE FIRST PAPER presenting the results of this survey, all the chromo-
some numbers recorded for the genus Lonicera, to that date, were as-
sembled, together with many new counts. Since that time the study of
Lonicera has continued, but to bring the investigation to a conclusion all
the additional counts that have been made using the Arnold Arboretum
collections are presented below (together with three further records that
have appeared in the literature).
Cytological methods, documentation and nomenclature used here fol-
low those of the first paper, to which reference should be made
An attempt was made to note differences in karyotype morphology and,
certainly, differences in the overall size of chromosome complements were
observed between different species. Also, variation in individual chro-
mosomes, their size, centromere position, and the presence and size of
satellites were noted, but considering the relatively large number of spe-
cies in the genus and the few individuals investigated, it has not proved
possible to compare and correlate these differences, and their groupings,
with the infrageneric classification proposed by Rehder (1903).
At metaphase the chromosomes, in many cases, were so contracted
that two satellites were not always visible. Thus, it was not possible to
determine whether or not Lonicera modesta had a satellited chromosome
pair. More details of morphology could be observed at late prophase. In
some cells, pretreatment with oxyquinoline (Tjio & Levan, 1950) caused
a Structural differentiation of the chromosomes by revealing positively and
negatively heteropycnotic segments. Homologues of similar size could
then be identified by the location of the centromere and by the individual
distribution of these segments. A comparable pattern has been observed
in several homologues of different species of Lonicera. Ficures 1 to 10
present examples which were encountered of nuclei in mitosis (most ex-
amples taken from species in different subsections of Rehder’s classifica-
tion).
A few comments may be made. In four cases both diploid and tetra-
ploid plants have been recorded within the same species. In Lonicera
ferdinandii Franch., the earlier undocumented counts and all the plants
at the Arnold Arboretum appear to be diploid, except for one (AA 21595)
which is tetraploid. This particular bush is an old one, raised from seed
of Rock 13519 collected in S.W. Kansu, China, in 1925, yet phenotypical-
* In this survey, the cytological investigations have been eae out by one of us
(L. R.), and the complementary taxonomy by the other (P.S
“Part I was published in Jour. Arnold Arb. 47: 222-247. 1966.
450 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ly it does not appear to differ significantly from the diploid. In L. alpigena
L., Poucques (1949, pp. 129 & 186) has recorded » = 9 and 2n = 18, both
of which numbers were confirmed by counts on a plant in the Arnold Ar-
boretum (AA 91-60) which, unfortunately, died before an authenticating
herbarium specimen was collected. However, in this species, the tetraploid
number, 2” = 36, has been found in two plants of f. mana (Carr.) Zabel
(see below). In L. maximowiczii (Rupr.) Maxim. var. sachalinensis
Fr. Schmidt we can now document a tetraploid (7 = 18 and 2n = 36),
in contrast to the diploid number of 2n = 18 recorded for the species by
Janaki Ammal & Saunders (1952, p. 540). The plant on which their
count was based does not appear to have been documented and it is now
impossible to know which variety may have been involved, or to confirm
its identity. Lastly, in our first paper we recorded a plant of L. modesta
Rehd. var. modesta as diploid (n = 9 and 2n = 18) and of var. lusha-
nensis Rehd. as tetraploid (n = 18 and 2n = 36), both plants having
been raised from seed sent from the Lushan Botanic Gardens in China.
Here, however, there is need for taxonomic reassessment, as we have
pointed out (Riidenberg & Green, 1966, p. 225). Available herbarium
material has proved inadequate to enable one to come to a sound con-
clusion, but it may well prove that two species are involved where
diagnostic distinctions need careful delineation.
It is, perhaps, worth drawing attention to the fact that in the whole
of both subsections TATARICAE and OcHRANTHAE, including many culti-
vars and hybrids, but with one exception, no polyploid plants have been
observed. The exception is Lonicera floribunda Boiss. & Buhse (AA
341-44) which is tetraploid. Within and between these subsections hy-
bridization takes place readily, yet meiosis in most of these diploid hybrids
is, with the exception of some plants with bridges, perfectly normal. A
few of the plants studied at the Arnold Arboretum form bridges at ana-
phase I, especially L. x bella; meiosis was, therefore, checked the next
year to determine its constancy and whether or not the frequency of
these bridges could be correlated with the seasonal variation in climate.
It was found that the number of cells showing bridges was not the same
for the two years. It was smaller after the more normal spring, in contrast
to one with especially cold nights and periods of drought.
LITERATURE CITED
JANAKI Amat, E. K., & B. SauNnpERS. 1952. Chromosome numbers in spe-
cies of Lonicera. Kew Bull. 1952: 539-541.
Love, A. 1968. IOPB Chromosome number report XIX. Taxon 17: 573-577.
Love, A., & D. Loéve. 1966. Cytotaxonomy of the alpine vascular plants of
Mount Washington. Univ. Colorado Studies, Ser. Biol. 24: 1-74
Poucques. M.-L. pe. 1949. Recherches caryologiques sur les Rubiales. Revue
Gén. Bot. 56: 5-27, 74-138, 172-188. [Lonicera pp. 84-95, 129, 186.]
REHDER, A. 1903. Synopsis of the genus Lonicera. Ann. Rep. Missouri Bot.
Gard. 14: 27-232.
1969 | RUDENBERG & GREEN, LONICERA, II 451
| RUDENBERG, L., & P. S. GREEN. 1966. A karyological survey of Lonicera, I.
Jour. Armold Aah 47: 222-247.
| sinks R. L., & G. A. MULLIGAN. 1968. Flora of the Queen Charlotte Islands,
ol ree & A. Levan. 1950. The use of oxyquinoline in chromosome anal-
ysis. Anal. Estac. Exp. Aula Dei 2: 21-64.
|
TABLE. Additional chromosome numbers in Lonicera
csp
DOCUMENTATION GENERAL
SPECIES n 2n AND COLLECTOR DISTRIBUTION
Subgenus Lonicera (Subgen. Chamaecerasus (L.) Rehd.)
Sect. IsoxyLOSTEUM Reh
Subsect. MrcrostyLaE Rehd.
L. angustifolia Wall. ex DC. 9 See Mehra & Gill in Himalayas
1
1291 (PUNJAB), Simla,
W. Himalayas
*D. syringantha Maxim. 18 AA 405-35, Palmer, North & West China
1 June & 26 Aug. 1936
*var. wolfii Rehd. 18 36 AA 4992-2, Allen, West China
1 June 1927, also Dudley
& Dodd, 28 May 1965
*cv. Grandiflora 36 AA 1089-61, agree
18 May 1
Sect. IstKA (Adans.) Rehd.
Subsect. CAERULEAE Rehd.
L. villosa (Mich.) Roem. & Schult. 18 See Love & Love (1966, p.51). Northeastern
Based on Love & Love North America
7496 & 7591, Mt. Wash-
ington, New Hampshire
WOLAAOTAV CIONUYV AHL JO TWNYNOL
OS “10A]
eo -— all 2 y
Subsect. PILEATAE Rehd.
*L. pileata Oliv. 18 AA 151031-B, Dudley & Central and
Dodd, 28 May 1965 western China
9 AA 225-28-E, Gre
4 Nov. 1965 and ae 225-28)
Kobuski & Roush, 14 Sept.
1931
*L. nitida Wils. 18 AA 923-49, Green, Western China
4 Nov. 1965
Subsect. VESICARIAE (Komar.) Rehd.
L. ferdinandii Franch. 18 AA 21595 (Rock 13519, Northern China
Kansu, 1925), Kreps,
25 May 1964
Subsect. BRACTEATAE (Hook. f. & Thoms.) Rehd.
L. altmannii Reg. & Schmalh.
*var. pilosiuscula Rehd. 18 AA 14999, Rehder, Turkestan
5 May 1927
Subsect. DisTeGIar (Raf.) Rehd.
L. involucrata (Richards.) Banks ex Spreng. See Taylor & Mullig, Northern ei
9 18 (1968, p. 109). “ee on CTS and south into
35077 & CT 35434, Rocky om
Graham Is., British
Colombia
Subsect. ge Rehd.
L. alpigen
f. nana ae Zabel 36+ AA 14994-1, Allen, Central and southern
13 August 1927 European Mts.
36 AA 803-35, Green,
26 May 1965
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462 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
NOTES ON WEST INDIAN ORCHIDS, I
LESLIE A. GARAY
DuRING THE couRSE of routine identifications of collections from
various parts of the West Indies, several new species as well as a number
of nomenclatorial changes have been noted. A study of the flora of the
West Indies is currently under way by Dr. Richard A. Howard of Har-
vard University, which will document both the distribution and diversity
of all orchid species known in that floristic region. In the mean time,
notes, similar to this one, will be published seriatim.
Habenaria Dussii Cogn. in Urb. Symb. Antill. 6: 307. 1909.
There is a flower from the holotype of H. Dussii Cogn. given by Pro-
fessor Cogniaux to the collections of the Orchid Herbarium of Oakes
Ames. Since then the type specimen has been destroyed in Berlin dur-
ing World War II. This single flower enabled me to identify the fol-
lowing two collections reported for the first time outside the island of
Guadeloupe.
Puerto Rico: Sierra de Luquillo, open grass-sedge savannah, wet, in cloud
forest along El Toro trail, south side of El Yunque, R. A. Howard & G. Taylor
18701 (AMES).
St. Vincent: St. David Parish, Soufriére Mountain, in tundra-like growth at
elevation of 2800 ft. Entire plant green, G. R. Cooley 8446 (AMES).
Cryptophoranthus erosus Garay, sp. nov. Fic. 1a-d.
Epiphytica, caespitosa, usque ad 3 cm. alta; radicibus crassiusculis,
elongatis, satis profusis, flexuosis, glabris; caulibus secundariis erectis,
atropurpureis; sepalo postico spathulato-rhombeo, valde concavo, 3-
nervio, dorsaliter apicem versus carinato mucronatoque, 14 mm. longo,
6.5 mm. lato; sepalis lateralibus usque ad apicem in synsepalo conniven-
tibus, valde concavis, dorsaliter carinatis, acutis, 15 mm. longis, mises
se 6 mm. latis; petalis carnosis, subfalcato-lanceolatis, acuminatis, ae
nerviis, 4 mm. longis, 1 mm. latis; labello breviter angusteque unguicu-
lato, deinde suborbiculari expanso, margine valde eroso; disco utrinque
carnoso carinato in medio, antice pectinato, 4.5 mm. longo, 3 mm. lato;
—~-
1969] GARAY, WEST INDIAN ORCHIDS, I 463
columna clavata, late alata, clinandrio lacero; ovario cylindrico, verru-
coso, 2 mm. longo
Dominican Republic: in the vicinity of Constanza. Flowers deep purple.
Collected by Rev. Donald Dod and cultivated by him for Bro. Alain H. Liogier
13508 (NY, type!).
This new species vegetatively resembles C. sarcophyllus (Rchb.f.)
Schltr. from Venezuela, but the latter has broader, entire leaves, as well
as dissimilar petals and lip.
Pleurothallis Dodii Garay, nom. nov.
Basionym: Pleurothallis cryptantha Cogn. in Urb. Symb. Antill. 7: 176.
1912, not Barb. Rodr. 1877.
A recent collection by Rev. D. Dod, s.n. (Ny), of this rare species in
the Dominican Republic: Las Abejas, Cabo Rojo, has shown that
the disc of the lip is covered with fine, but sparsely distributed, hairs as
are the margins. This character, although not mentioned in the original
description by Cogniaux, is present on the holotype which I have re-
cently examined in Bruxelles.
Lepanthopsis Dodii Garay, sp. nov. Fic. 2e-f.
Epiphytica, caespitosa, usque ad 8 cm. alta; radicibus filiformibus,
flexuosis, glabris; caulibus secundariis erectis, gracilibus, vaginis satis
distantibus, adpressis, sursum dilatatis hispidulisque omnino obtectis,
usque ad 4 cm. longis; foliis tenuibus, ellipticis, acutis vel obtusiusculis,
margine muricato-denticulatis, usque ad 2 cm. longis, 6 mm. latis; in-
florescentiis capillaribus, subdense multifloris, usque ad 4 cm. longis;
bracteis infundibuliformibus, acuminatis, 1 mm. longis; floribus tenuibus,
diaphanis, patentibus, glabris; sepalo postico ovato-lanceolato, acuminato,
uninervio, 2 mm. longo, 1 mm. lato; sepalis lateralibus inter se usque ad
medium connatis, ovato-lanceolatis, acuminatis, uninervis, 2 mm. longis,
inter se 1.5 mm. latis; petalis ellipticis vel subrhombeis, acutis vel ob-
tusis, 1 mm. longis, 0.5 mm, latis; labello carnoso, triangulari-cordato, 3-
nervio, 1 mm. longo latoque; columna humili, crassa, apoda; ovario pedi-
cellato 2 mm. longo.
Dominican Republic: Polo, epiphytic on trees, Rev. D. Dod 43 (ames, type!).
Lepanthopsis Dodii Garay differs from L. acuminata Ames in having
smaller flowers, proportionately shorter and broader sepals, and dif-
ferently shaped petals.
Since my revision of the genus in The Orchid Journal 2: 467-469.
1953, I have examined the types of all West Indian Pleurothallis species.
Among these species the following need to be transferred to the genus
Lepanthopsis.
464 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Lepanthopsis barahonensis (Cogn.) Garay, comb. nov.
Basionym: Pleurothallis barahonensis Cogn. in Urb. Symb. Antill. 7: 177.
1912.
Lepanthopsis blepharophylla (Griseb.) Garay, comb. nov.
Basionym: Pleurothallis blepharophylla Griseb. Cat. Pl. Cub. 260. 1866.
Lepanthopsis dentifera (L. O. Wms.) Garay, comb. nov.
Basionym: Pleurothallis dentifera L. O. Wms. in Ceiba 1: 227. 1951.
Lepanthopsis Fuertesii (Cogn.) Garay, comb. nov.
Basionym: Pleurothallis Fuertesii Cogn. in Urb. Symb. Antill. 7: 178. 1912.
Brachionidium ciliolatum Garay, sp. nov. Fic. 3g-j.
Epiphytica, parvula, ascendenti, usque ad 7 cm. alta; radicibus fili-
formibus, glabris; rhizomate ascendenti, cauliformi, vaginis scariosis, in-
fundibuliformibus imbricantibusque omnino obtecto; caulibus secun-
dariis vix ullis, monophyllis; foliis pergameneis, oblongo-ellipticis, acutis,
subpetiolatis, usque ad 2 cm. longis, 5 mm. latis; inflorescentiis singulis,
unifloris; pedunculo capillari, in medio univaginato, usque ad 3 cm. longo;
bracteis infundibuliformibus, ovariis pedicellatis aequilongis; floribus pro
genere satis parvulis, ciliolatis; sepalo postico ovato-lanceolato, subacu-
minato, 3-nervato, margine ciliolato, 7 mm. longo, 4 mm. lato; sepalis
lateralibus usque ad apicem connatis, ibi bidentatis, ellipticis, obtusis, 4-
nervatis, margine ciliolatis, 6 mm. longis, 4 mm. latis; petalis ellipticis,
apice subito in apiculo triangulari-subfalcato, acuminato productis, 3-
nervatis, margine ciliolatis, 6 mm. longis, 4 mm. latis; labello carnoso, €
cuneata basi subsigmoideo, antice triangulari, acuto, 3-nervato, margine
valde ciliolato; disco callo pulvinari, antice exciso ornato; toto labello 3
mm. longo, 2.5 mm. lato; columna humili, crassa, vix 1 mm. alta; ovarlo
pedicellato ca. 2 mm. longo.
Puerto Rico: Pico del Oeste, Sierra de Luquillo, 1020 m. alt. Epiphytic or
chid, plants with 3-4 leaves; flowers yellow-green, apparently do not open.
Study trail area. R. A. Howard & L. I. Nevling 16929 (ames, type!).
This new species closely resembles B. parvum Cogn. both in size and
in general appearance. It differs, however, in the shape of the floral
segments which are not caudate. Both B. tetrapetalum (Lehm. & Kral.)
Schltr., and B. simplex Garay, although similar in appearance to B. cilio-
latum Garay, have dissimilar and eciliate lips.
Epidendrum isochilum var. tridens Rchb. f. in Ber. Deutsch. Bot.
Ges. 3: 277. 1885.
Syn.: Epidendrum belvederense Fawc. & Rendle in Jour. Bot. 47: 123. 1909.
1969] GARAY, WEST INDIAN ORCHIDS, I 465
Ficures 1-4, West Indian orchids. Fic. 1, a-d, Crypto phoranthus erosus
Garay; y Fic. 2, e-f, Pig Wass Dodii Garay; Fic. 3, g-j, Brachionidium cilio
latum Gar aray; Fic. 4, k-n, Campylocentrum constanzense Garay. All
greatly magnified.
466 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
There appears to be no distinction between Epidendrum belvederense
Fawc. & Rendle and E. isochilum var. tridens Rchb. f. as a study of the
holotypes indicates. Judging from the number of specimens which I
have examined of this species from the Dominican Republic, this variety
seems to be much more common than the typical variety, which is de-
scribed as having an entire lip.
Epidendrum neoporpax Ames in Bot. Mus. Leafl. Harvard Univ. 2:
Liz, 1934.
Basionym: Epidendrum Porpax Rchb. f. in Flora 48: 278. 1865 not Rchb. f.
Syn: Epidendrum vestitum Ames in Sched. Orch. 4: 51. 1923.
Epidendrum Porpax var. domingensis Cogn. in Urb. Symb. Antill. 7: 181.
1912.
This rather rare Cuban species has been found recently in Costa Rica,
and rediscovered by Mr. Ariza Julia, s.n., in the Dominican Republic:
Sabaneta de Yasica, Puerto Plata Province. An examination of the
type of Epidendrum Porpax var. domingensis Cogn. in the Bruxelles
herbarium convinces me that it is identical with E. neoporpax Ames.
Epidendrum Sintenisii Rchb. f. in Ber. Deutsch. Bot. Ges. 3: 241s
Syn.: Epidendrum monticolum Fawc. & Rendle in Jour. Bot. 47: 124. 1909.
Recently I had the opportunity to examine and to compare the holo-
types of E. Sintenisii Rchb. f. and E. monticolum Fawc. & Rendle. As
a result of this study, I am convinced that they are conspecific. Epi-
dendrum Sintenisii is now recorded from Puerto Rico and Jamaica.
Stellilabium minutiflorum (Krzl.) Garay, comb. nov.
Basionym: Telipogon minutiflorus Krzl. in Ann. Nat. Hist. Mus. Wien 33:
14. 1919.
Syn.: Telipogon Lankesteri Ames Sched. Orch. 3: 23. 1923.
Stellilabium Helleri L. O. Wms. in Brittonia 14: 443. 1962.
This rather rare Costa Rican species has recently been found in the
Dominican Republic: Casalito Bonao by Rev. D. Dod, s.n. (NY):
This is also a new record for the West Indies. Stellilabium Helleri L. O.
Wns., of which I also have studied the holotype, agrees in every respect
with Kraenzlin’s type material which I examined in Vienna. Telipogon
Lankesteri Ames likewise, does not offer any criterion by which it could
be kept separate from S. minutiflorum (Krzl.) Garay.
Polyradicion Garay, gen. nov.
Pfitzer in describing the genus Polyrrhiza stated that it consists of four
West Indian species. Of these four he mentioned only one in making
BIB = een ance
1969] GARAY, WEST INDIAN ORCHIDS, I 467
an Official transfer, namely P. funalis (Sw.) Pfitz. Thus, the genus Poly-
rrhiza is typified by this species. In Flora of Jamaica, Fawcett and Rendle
regard P. funalis (Sw.) Pfitz. to be a synonym of Dendrophylax funalis
(Sw.) Benth., a judgment which I consider to be correct. Since Poly-
rrhiza automatically becomes a synonym of Dendrophylax through this
transfer, it leaves the other species without a validly published generic
name. Since there are only two species involved I reject the idea of con-
servation in favor of a new name which I propose here with the same
etymological meaning as was used by Pfitzer. The genus is, thus, charac-
terized as follows:
Sepala petalaque simillima, aperta, lanceolata; labellum maximum, 3-
lobum, lobi laterales quam lobum intermedium multoties breviores, basi
in calcari valde evolutum producta; columna humilis, crassa, apoda, basi
labellum adnata; clinandrium humile; anthera incumbens, opercularis;
pollinia 2, stipiti nudi, distincti glandulae affixa.
Plantae epiphyticae, aphyllae; radices crassae, valde evolutae; caules
vix ulli; pedunculi laterales, graciles, arcuati, abbreviati, semper uniflori;
flores majusculae.
Species 2, Indiae Occidentalis incolae.
Typus: Angraecum Lindenii Lindl.
Polyradicion Lindenii (Lindl.) Garay, comb. nov.
Basionym: Angraecum Lindenii Lindl. in Gard. Chron. 135. 1846.
Syn.: Aeranthus Lindenii Rchb. f. in Walp. Ann. Bot. Syst. 6: 902. 1864.
Dendrophylax Lindenii Benth. ex Rolfe in Gard. Chron. ser. 3. 4: 533.
1888.
Polyrrhiza Lindenii Cogn. in Urb. Symb. Antill. 6: 680. 1910.
Distribution: Florida, Cuba.
Polyradicion Sallei (Rchb. f.) Garay, comb. nov.
Basionym: Aeranthus Sallei Rchb. f. in Walp. Ann. Bot. Syst. 6: 902. 1864.
Syn.: Dendrophylax Sallei Benth. ex Rolfe in Gard. Chron. ser. 3. 4: 533.
Polyrrhiza Sallei Cogn. in Urb. Symb. Antill. 6: 680. 1910.
Distribution: Dominican Republic, Haiti.
Dendrophylax gracilis (Cogn.) Garay, comb. nov.
Basionym: Polyrrhiza gracilis Cogn. in Urb. Symb. Antill. 6: 679. 1910.
An examination of the holotype, Wright 3300, in the Orchid Herbarium
of Oakes Ames has shown clearly that it is referable to the genus Dendro-
phylax Rchb. f. It is closely allied to D. hymenantha Rchb. f., differing
in its shorter, 1-flowered peduncle and in the size of its flowers which
are twice as large. Dendrophylax hymenantha Rchb. f., however, has been
united with D. varius (Gmel.) Urb., but this decision requires further
study.
468 JOURNAL OF THE ARNOLD ARBORETUM [VvoL. 50
Campylocentrum constanzense Garay, sp. nov. Fic. 4k-n.
Epiphytica, caespitosa, aphylla, usque ad 4 cm. alta; radicibus nu-
merosis fasciculatis, filiformibus, flexuosis, glabris; caulibus nullis vel
vix ullis; inflorescentiis numerosis, fasciculatis, erectis, capillaribus, sim-
plicibus vel dichotome ramosis, supra laxe plurifloris, omnino setaceo-
hirsutis, usque ad 4 cm. longis; bracteis ovato-cucullatis, acutis vel ob-
tusiusculis, extus setaceo-hirsutis, 1 mm. longis; floribus minimis, hyalinis;
sepalo postico ovato, acuto vel obtusiusculo, uninervio, extus sparse setaceo-
hirsuto, 1.5 mm. longo, 1 mm. lato; sepalis lateralibus oblique ovatis,
obtusiusculis, extus setaceo-hirsutis, 3-nerviis, 2 mm. longis, 1 mm. latis;
petalis subfalcato-ovatis, obtusiusculis, 3-nerviis, glabris, 1.5 mm. longis,
1 mm. latis; labello anchoriformi-lobato, antice breviter apiculato, basi
calcarato, calcari cylindrico obtuso, setaceo-hirsuto; disco in medio lon-
gitudinaliter carinato, antice setaceo-hirsuto; toto labello 3 mm. longo,
antice 1.5 mm. lato; columna humili, crassa, vix 1 mm. alta; ovario
cylindrico, muricato-hispidulo, cum pedicello 2 mm. longo.
Dominican Republic: Constanza, epiphytic on trees, Rev. D. Dod 66 (AMES,
type!).
This species is quite unique in the Section DENDRopHYLopsis Cogn. be-
cause of the anchor-shaped lip and a distichously branching, setaceous in-
florescence.
BoraNnicaL MusEuM
HARVARD UNIVERSITY
1969] SMITH, POLLEN OF AFRICAN VERNONIA 469
POLLEN CHARACTERISTICS OF AFRICAN SPECIES OF VERNONIA
C, Ear LE SMITH, JR.
DURING A TAXONOMIC sTUDY of the species of section STENGELIA of
the Composite genus, Vernonia, pollen of a number of species was examined.
Whenever possible, a floret from a specimen of the type collection was
dissected and the anthers macerated in lacto-phenol and methylene blue.
A single grain from each slide was photographed. Size was determined by
measurements of ten grains from each slide, after scanning to ascertain
whether the grains measured fell into more than one size class. Measure-
ments were made to the outside of the reticula, but did not include the
length of spines.
Obviously, not all of the specimens examined belong to section STEN-
GELIA, although all of the species have been assigned here because of thin
terminal appendages on otherwise firm or chartaceous phyllaries. Per-
haps a future student of the genus will find corollary characters on which
the sections of the genus can be more firmly based. The large number
of species involved in Africa alone precludes this in my short-term ex-
amination of the section STENGELIA.
Pollen sizes range from an average of 29.1 » for Vernonia praecox Welw.
ex O. Hoffm. to 69.5 » for grains of V. wittei Hutch. & Burtt (Fic. 6).
The average pollen diameter is 51.9 ». The largest number of species with
a similar pollen size fall into the next highest size class, 53.3 ». A total
of seven species have an average pollen size of 51.7 ». Twenty-six species
fall into larger pollen-size classes and 24 species fall into smaller pollen-
size classes than the 15 species in the median groups. Thus, except for
the few plants having either very large pollen grains or very small pollen
grains in relation to the average pollen size for this group of species, the
species are well clustered with the greatest number falling centrally.
No attempt was made to study the anatomy of the pollen grains of
this group. Morphologically, all of the grains examined are similar. All
are nearly spherical and are evidently tricolporate, although this some-
times is difficult to determine (Fic. 4). In all of the pollen examined, the
outside of the grain is marked by a raised reticulum. T his may be thin,
but in the majority of the species the surface reticulum is moderately to
heavily thickened. Often, the pattern of the reticulum is very regular
with polar alveoli surrounded by a ring from which radiate, at regular
intervals, a series of bars. These are crossed on the sides of the grains
at regular intervals by bars of equal size. On one side of the grain a
longitudinal alveolus extends from pole to pole and one of the pores oc-
curs in this at the equator (Fic. 1). Frequently, the polar rings are
470 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Ficures 1-5, pollen grains of Vernonia species. Fic. Vernoma guineensis
var. poi average pollen meter 51.74; the seed grain, in polar view,
shows the regularity of the reticulation common to many of the upright or shrub-
1969 | SMITH, POLLEN OF AFRICAN VERNONIA 471
broken by this longitudinal opening so that if flattened, the total open
area would resemble a dumbbell. On the sides of the grain, the alveoli do
not appear to be geometrically balanced in any of the species. Where
the reticulation follows a similar pattern in all of the grains examined, I
have called it regular in spite of a lack of an exact geometric pattern. In
many species, polar areas may be defined or not, but the reticulation over
the remainder of the surface of the pollen grain appears to be randomly
placed (Fic. 5). I have called this type of reticulation irregular.
In most of the species, the reticulation is further decorated by spines
arising from the top and sides, The spines all appear to be simple conical
protuberances of the same material from which the reticulum is formed.
V. albo-violacea De Wild. has almost no spines on the reticulum. The
spines on pollen grains of other species vary from small to large, but they
are uniform in size for a species. Only V. bojeri Less., V. gerberiformis
Oliver & Hiern, V. mandrarensis Humbert, and V. prolixa S. Moore lack
spines completely.
The reticulation of V. gerberiformis is unusual among the species ex-
amined. The reticulation is relatively narrow and produced upward from
the surface of the grain in a wing-like projection (Fic. 3). I shall make
no attempt to formulate an adaptive advantage for this deviation from
the usual pattern among the Vernonia pollen grains studied. With an
average pollen diameter of 66.3 » (measured at the outside of the reticu-
lum), this is the second largest grain examined.
The surface of the pollen grains was examined under oil immersion
(485 &) in order to study the surface details. In none of the grains was
a regular pattern or design seen. Some of the reticula and pollen surfaces
are not smooth. The roughness is not readily discernible and does not
appear to be produced in a regular pattern. However, the reticulation on
some of the species (for example, V. adoensis Sch.-Bip. ex Walp., V. ger-
beriformis) is not completely contiguous with the surface of the pollen
grain. When the reticulum is heavy, it frequently stands above the grain
surface on a series of pedestals which may be relatively far apart or
rather close together (Fic. 2). The presence of the pedestals negates the
possibility that this is an artifact created when the surface of the om
shrinks away from the reticulum. In the pollen grains of other species with
an equally heavy reticulum, the elevation of the reticulation above the
surface does not seem to be present.
ll m
median section. Fic. 5, Vernonia castellana,
this shows the
rosette of leaves.
472 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
On the basis of the gross morphological characteristics of specimens
and on a detailed study of the achenes and flowers, the species have been
grouped into clusters which show similarities. Pollen grain data have
been regrouped (TABLE 1) on this basis (excluding species which apparent-
ly do not belong to section SrENGELIA). In general, the characteristics
of the pollen grains seem to support the groupings by overall morphology.
For instance, the first group of three species, V. /asiopus O. Hoffm. in
Engl., V. brownii S. Moore, and V. albo-violacea, varies relatively little
in average pollen grain diameter; the grains have reticula of medium thick-
ness with short or nearly absent spines.
Number 5
of 4
Species 3
54 3; 4 433 Sis 34 64 695
Average Diameter of Pollen Grains
Ficure 6. Graph illustrating the number of species of Vernonia, section
STENGELIA, with pollen falling into each size class. Note that pollen size for
most of the species falls near the median, 51.9z.
The second group of species, V. polyura O. Hofim., V. filigera Oliver &
Hiern, V. longipetiolata Muschler, and V. oxyura O. Hoffm. in Engl., again
agree well in pollen characteristics as well as in overall morphology, €X-
cept for grain size in one species. The pollen grain size of three of the
species ranges from an average diameter of 40.4 » to 46.9 ». The average
pollen grain size of grains of V. filigera is 56.6 uw. It is hardly desirable to
exclude the species from this grouping on this one feature alone, but it
does necessitate another careful look at the specimens to be included here.
The break in size observed in the example cited above is perhaps better
illustrated in the group of species clustered around the type species of the
section, V. adoensis Sch.-Bip. ex Walp. On the basis of their gross
morphology, these species fall readily into a group. An examination of
the details of the achenes and flowers discloses no major discrepancy 10
the pattern. For the most part, the morphology of the pollen grains of
these species supports the grouping. Average pollen diameters for most
species of the group range between 56.6 » and 63.0 ». However, the
average pollen grain diameters of V. shirensis Oliver & Hiern and V.
woodii O. Hoffm. (which are now considered synonymous) are 46.9 p.
1969 | SMITH, POLLEN OF AFRICAN VERNONIA 473
The species of section STENGELIA fall into two distinct groups on the
basis of plant habit. The bulk of the species are rank-growing upright
sub-shrubs from a perennial base, or upright shrubby plants. A few
may become tree-like. The pollen of many of these species has an average
diameter of 50.1 » or more, except for the species grouped around V.
polyura. The reticulum on the grains is generally heavy.
The other group of species is distinguished by a rosette habit with
flowers borne on a, usually, leafless scape. The scape may be unbranched
and support a single head or it may support several heads. Many of the
average pollen grain diameters for this group are less than 50 ». The
reticulation on the grains is often thin and very irregular. However, more
exceptions occur among the species with basal rosettes than among the
species with an upright habit. For example, the pollen of V. gerberiformis,
which was previously described, is very different from the usual pattern
of grain size and morphology. The grains of V. myassae Oliv. in Hook.,
V. pumila Kotschy & Peyr. and V. anandrioides S. Moore have a heavy,
regular reticulum. Furthermore, the grains of the last two have an
average diameter of 53.3 ». The pollen grains of the species with a basal
rosette are more variable in size and morphology than are the pollen
grains of the other species assigned to section STENGELIA.
In only three of the species examined, pollen grains of two size classes
occurred. Both V. calvoana (Hook. f.) Hook. f. and V. insignis (Hook.
f.) Oliver & Hiern have pollen grains similar in size and morphology.
In both, the larger grains appeared to be normal. The smaller pollen
grains appear to have been aborted. Because they were removed from
herbarium specimens, it was impossible to apply germination tests for
viability of the grains to confirm my assumption that the smaller
grains are not functional. About half the pollen grains of V. achyrocepha-
loides Hutch. & Bruce were also smaller than the grains measured, and
appeared to be nonfunctional. So little is yet known about the biology
and genetics of species of section STENGELIA that I can make no as-
sumptions as to the cause of the difference in pollen grain sizes. It is,
perhaps, significant that all of the species have average pollen diameters
near the upper limit of pollen size for this group of 64 species.
SUMMARY
474 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
achenes and floral morphology, pollen morphology and average pollen
diameter confirms the groupings for the most part.
The species traditionally included in section STENGELIA can be divided
into a group with basal rosettes and heads borne on scapes versus a
group of upright sub-shrubs or shrubs (rarely tree-like) with heads
on the sides or ends of branches. In general the first group has smaller
pollen grains with thin, irregular reticula. The second group generally
has larger pollen grains with heavy, more regular reticula. The group
with basal rosettes is less homogeneous than the other in regard to pol-
len size and morpholo
In only three species of the 64 examined were pollen grains of two
size classes found. In all instances, the smaller grains appeared to be
nonfunctional. All three species have pollen diameters in the upper size
range. Insufficient knowledge of the biology and genetics of these species
precludes an explanation.
TABLE 1. Comparison of pollen characters with gross morphology of
species of Vernonia section Stengelia
POLLEN
RETICULATION
SPINES
SPECIES SIZE, THICKNESS PATTERN LENGTH DISTRIBUTION PROVENIENCE
V. albo-violacea $i7 Medium Regular Almost none Bequaert 492, Congo
V. Ob i 56.6 Irregular Short Frequent Brown 2656, Uganda
V. lasiopus Sa.3 ss Regular = Occasional Volkens 444, Tanzania
V. oxyura 40.4 Thin + Regular Medium Numerous Buchanan s.n., Malawi
V. longipetiolata 42.0 Medium Regular 7 Occasional Kassner 2746, Congo
V. p 46.9 Thin Irregular Large si Goetze 866, Tanzania
V. filigera 56.6 Medium . Medium Numerous Schimper 1530, Ethiopia
V. nyassae 48.5 Heavy Regular Small Frequent Thomson s.n., Zambia?
V. swynnertonii 51,7 ae Irregular Short +Numerous Swynnerton 1908, Rhodesia
V. gerberiformis 66.3 Winged = None Schweinfurth 2688, Sudan?
V. wittei 69.5 Medium i Short Frequent de Witte 543, Congo
V. chthonocephala 35.6 Thin a ae Welwitsch 3886, ing
V. subaphylla 38.8 ° is o Few Carson 10, Zam
V. praemorsa 40.4 Medium 2 is Occasional Stolz 104, Hotaeny
V. agricola 43.6 Thin si Medium Numerou Kassner 2136, Zambia
V. castellana 48.5 sg 2 Gossweiler 2883, Angola
V. anandrioides 53.3 Heavy Regular Short Occasional Gossweiler 2132, Angola
V. pumila ae Medium + Regular ‘i Frequent Elliot 7037, Kenya?
V. homilocephala 54.9 + Heavy Irregular ‘s Numerous Elliot 7058, Kenya?
V. eat 50.1 Heavy " Short Ww Homblé 881, Congo
V. pleiotaxo |e si fc : Frequent Quarré 2654, Congo
V. procera 53.4 . ° Numerous Chevalier 7899, Congo?
V. lancibracteata 58.2 6 Regular Ha Frequent Eyles 291, Zambia
V. firma 50.1 Medium Regular Short Frequent Schweinfurth 3153, Sudan?
V. vallicola 58.2 Heavy ‘s Medium a
Gossweiler 3781, Angola
VINONUAA NVOIMAV AO NATIOd “HLIWS 6961
SLY
TABLE 1. Comparison of pollen characters with gross morphology of
species of Vernonia section Stengelia (Continued )
POLLEN RETICULATION SPINES
SPECIES S1zE yp THICKNESS PATTERN LENGTH DISTRIBUTION PROVENIENCE
V. uni 51.7 Heavy Irregular Medium Numerous Braun 1979, Tanzania
V. calvoana var
microce phala S3:3 Medium Regular Short Frequent Lightbody 26259, 2 oC
\3 -
i | | i ANS
Map 2. Distribution of Flindersia pimenteliana F. Muell.
Morwood NGF 6204 (a, BO, BRI, K, L, LAE, NSW 99656, US), McAdam 291 (LAE),
Ross NGF W1001 (a, BO, BRI, L, LAE, NSW 99660), Womersley & Brass N GF
11025 (A, BRI, K, L, LAE); W of Bulolo near Bulolo-Watut Divide, Frodin &
Hill NGF 26355 (x, L, LAE, Nsw 99658), Upper Long Iskand Creek, near Bulo-
lo, Havel & Henry NGF 17024 (a, BO, BRI, CANB, K, L, LAE), Upper Nauwata-
Banda logging area, near Bulolo, Havel & Kairo NGF 11140 (BRI, K, 1, LAE),
NGF 11142 (Bo, K, L, LAE); near Dengalu Village, Womersley NGF 19063 (kK,
L, LAE, NSW 99668); Wau, White NGF 2529 (1, LAE, Nsw 99655); Edie Creek,
1969 | HARTLEY, THE GENUS FLINDERSIA 497
Streimann NGF 17476 (x, LAE); Kauli Creek, 5 miles S of Wau, Hartley 11513
(A, LAE), Henty NGF 14726 (x, L, LAE, NSW 9 9654), Millar NGF 14516 (k, L,
LAE, NSW 99657), van Royen NGF 16302 (B0, L). Papua. CENTRAL DISTRICT:
Lala River, Carr 15805 (cans, L), 15989 (kK, L); Isuarava, Carr 15475 (1),
15569 (L), 15969 (kK, Lt); Mafulu, Brass 5339 (a, Ny, US); Boridi, Carr 13152
(A, K, L, NY), 14408 (A, K, L, NY), 14808 (L), 14910 (A, K, L, NY); Mt. Obree
to Laruni Spur, Lane-Poole 382 (BRI). NORTHERN District: Managalase area,
S side of Hydrographers Range near Siarane, Pullen 6263 (CANB); Bariji-
Managalase area, N side of Sibium Range, S of Toma, Pullen 6358 (CANB, LAE),
6387 (CANB, LAE). Queensland. Cook District: Cape York Peninsula, Mt.
Finnegan, Brass 20322 (a); Great Dividing Range ca. 6 miles S of Mossman
and near “Devil Devil Creek”, Smith 3953 (prt); Mt. Spurgeon, White 10596 (a,
K); Danbulla, Jones 1117 (c ANB); Atherton Tableland, Juara Creek area, near
Danbulla, Smith 3780 (ert); Atherton Tableland, Lake Barrine, Kajewski 1114
(A, BRI, Ny, P); Atherton, Mocatta, February 1913 (prt); Atherton Area, Webb
E
Yungaburra, Dreghorn, December 1935 (A, BRI, K); Gadgarra, Dreghorn 22E
(BRI), January 9, 1934 (A, BRI, NY), White 1566 (A, BRI); Glenallan, Malanda,
Hayes (srt); Paronella Park, on Mena Creek, ca. 14 miles S sel Innisfail, Smith,
August 5, 1948 (srr). NortH KENNEDY District: Koolmoon Creek, ca. 11
miles SSE of Ravenshoe, Smith 4588X (srt); Evelyn, Bailey, oak 8, 1899
(BrI-holotype of Flindersia mazlini F. M. Bailey; x-isotype); Coast Range,
Anonymous, February 1866 (BRI); oe Bay, Dallachy cee Geniece
of Flindersia pimenteliana F. Muell.; , BO, K, NSW 99651), Anonymous (BRI,
MEL); Mt. Macalister, Dallachy, Apri on 1869 (MEL); Mt. Fox, Clemens,
September-December 1949 (GH, MICH
C. T. White (1921) has previously placed Flindersia mazlini in the
Synonymy of F. pimenteliana.
The type collection of Flindersia chrysantha was made at 2300 m. in
West New Guinea and is typical of a number of collections from mountain
rain forests in New Guinea. The leaflets of these collections tend to be of
thicker texture and have more prominent veins and less tapering bases and
apices than typical F. pimenteliana. These are not sharply defined differ-
ences, however, and probably are only environmental modifications. Mer-
rill and Perry noted in their original description of F. chrysantha that the
petals differed from those of F. pimenteliana in color (yellow vs. red) and
pubescence (glabrous vs. pubescent abaxially). With the larger number
of collections that are now available it can be seen that both of these
characters are quite variable in typical F. pimenteliana from lower eleva-
tions in New Guinea. Flower color ranges from red to pink, cream or
white, and the petals range from pubescent to glabrous abaxially.
F aes sex and the distribution of,sexes is extremely variable in this
species. In some collections, such as Moll BW 9745 and Dreghorn, Decem-
ber 1935, all of the flowers appear to be perfect. In others, such as
Dreghorn, January 9, 1934, some appear to be perfect and many appear
to be functionally staminate. A condition where all of the flowers on a
specimen appear to be functionally staminate is found in a number of
collections from New Guinea including Brass 29153 and Carr 14910. Still
other collections, such as Brass 11128 and Hartley 12018 have a majority
498 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
of functionally carpellate flowers mixed with flowers that appear to be
functionally neutral
Insect galls are often formed from the flowers in New Guinea. These
have the appearance of young fruits and were occasionally mistaken for
them by collectors.
Other than Flindersia unifoliolata, which seems to be very closely re-
lated, F. pimenteliana does not appear to have any very close relatives.
As is indicated above in the outline of species relationships, I think its
having exclusively simple trichomes would place it closer to those species
which share that character than to any of the others, but the leaf differ-
ence and especially the difference in hypocotyl position makes any very
close relationship seem unlikely
5. Flindersia unifoliolata Hartley sp. nov.
Arbor usque 15 m. alta; ramulis, rhachidibus et laminis subtus glabris
vel minute et sparse puberulis, pilis simplicibus. Folia opposita vel sub-
opposita, unifoliolata vel interdum imparipinnata et unijuga, 3-9 cm.
longa; petiolulis foliolorum lateralium 1—4 mm. longis, rhachidi ad apicem
extensa usque 1 cm. longa foliolum terminale ferente; petiolis foliorum
unifoliolatorum usque 1.8 cm. longis; laminis subcoriaceis, consperse pellu-
cido-punctatis, ellipticis, equilateris vel parum inaequilateris, 3-8 cm.
longis, 1.4~3.2 cm. latis, basi acuta usque cuneata plerumque aequilatera,
venis primariis utrinque 8-16, apice obtuse acuminata. Capsula secedens
in valvas distinctas maturite, elliptica, 7.8 cm. longa; exocarpio in sicco
atro-rufescente, glabro, muricato, processibus inaequilongis, usque 2 mm.
longis; endocarpio brunnescente et leviter ferrugineo-maculato. Semina 2
inquoque latere dissepimentorum, utrinque alata, 3.5—-4 cm. longa; hypo-
cotylo laterali parum adscendente. Flores non visi. Holotypus: Sayer 136
(MEL).
Queensland. Coox District: Mt. Bellenden Ker, alt. 5200 ft., Sayer 136
(MEL-holotype); Mt. Bartle Frere, in low scrub, 4000-5000 ft., Martin & Hy-
land 1881 (pr).
The localities of the above collections are about 15 miles apart about
40 miles south of Cairns, Queensland. The holotype is a fruiting branch
while the Martin & H yland specimen is sterile.
Closely related to Flindersia pimenteliana which appears to be restricted
to lower elevations in this part of Queensland. The fruits and seeds of the
two species appear roughly identical.
6. Flindersia amboinensis Poir. in Lam. Encycl. Suppl. 4: 650. 1816.
Neotype: DeVriese & Teysmann, Moluccas, Ceram.
Arbor radulifera Poir. in Lam. Encycl. 6: 58. 1804 (provisional name, based
on plate and description by Rumphius, Herb. Amboin. 3: 201. ¢. 129. 1743).
Flindersia earbet nce Spreng. Geschicht. Bot. 2: 76. 1818 se illegit.).
1969 | HARTLEY, THE GENUS FLINDERSIA 499
Flindersia oo Lane-Poole ex White & Francis, Proc. Roy. Soc.
— 38: 232. t. 3. 1927. Type: Lane-Poole 362, Papua, Owen Stanley
ange.
Large trees to 45 m.; outer bark gray to brown, smooth or slightly
roughened; inner bark usually yellow grading to white or yellow-brown
toward the cambium; sapwood white, cream, or yellow; heartwood yellow-
brown; branchlets, leaves and inflorescences glabrous to pubescent with
mostly minute, predominantly stellate trichomes. Leaves alternate, im-
paripinnate or (occasional leaves) paripinnate, 18-57 cm. long; rachis
glabrate to appressed- or soft-pubescent; petiolules of lateral leaflets 2-8
mm. long, terminal leaflet on an extension of the rachis 1-4.3 cm. long;
leaflets 2-4(-5) pairs, chartaceous to subcoriaceous, with or without
scattered pellucid dots, glabrous to appressed- or soft-pubescent below,
glabrous or occasionally finely pubescent along the midrib above, elliptic
to elliptic-lanceolate, usually strongly unequal-sided and_ occasionally
falcate, 8-20 cm. long, 3-9.5 cm. wide, base acute to broadly rounded,
usually oblique, main veins 10—20 on each side of the midrib, apex obtuse
to acuminate. Inflorescence terminal, 18—30 cm. long, usually about as
wide as long, axes and branches densely appressed- to soft-pubescent.
Flowers bisexual, 3.5~5 mm. long; pedicels obsolete to 1.7 mm. long; sepals
densely appressed-pubescent, ciliolate, broadly ovate to suborbicular, 0.7—1
mm. long; petals yellow-brown, cream, or red, glabrous to densely ap-
pressed-pubescent abaxially, glabrous to subvillous at about the middle
adaxially, elliptic-oblong, 3-4.5 mm. long; stamens declinate, 2.2—3.6 mm.
long, filaments pilose subapically or (rarely) glabrate, anthers dorsifixed,
obtuse, 0.7-1 mm. long; staminodes 0.7—1.8 mm. long; disc 0.7—1.3 mm.
high; gynoecium about 2 mm. high and about 1.5 mm. wide, ovules 3 on
each side of the placentae. Capsule separating (or easily separable) into
5 distinct valves at maturity, elliptic to elliptic-oblong, 9-21 cm. long;
exocarp drying light to dark brown, densely and minutely pubescent to
glabrate, muricate, the excrescences thin and flattened laterally or thick
and conical, 5-8 mm. long; endocarp reddish brown or cream. Seeds 3 on
each side of the dissepiments, winged at both ends, 5—-8.5 cm. long;
hypocotyl lateral, horizontal.
ILLustraTion. Rumpuivs, G. E., ibid.
Distr1BuTION. Ceram and Tanimbar Islands in the Moluccas eastward
throughout New Guinea: rain forests from sea level to 1700 meters. See
Map 3
Moluccas. CERAM: without definite locality, DeVriese & Teysmann, 1859-
1860 (L-neotype of Flindersia amboinensis Sa are bb 25869 (L). TANIM-
BAR IsLanps: Otimmer, NJFS bb 24346 (L). t New Guinea (West Irian)
and neighboring islands. AROE ISLANDS: aa Island, Dosimanalaoe,
ste bb 25298 (1), NIFS bb 25322 (1); Trangan Island, Ngaibor, N/JFS bb
3 (L). Watceo IsLaNp: E bank of Majalibit Bay 8 km. NE of Waifor,
begs sig Sal 5165 (CANB, L). SALAWATI IsLAND: Kaloal, Koster BW 1495 (tL).
JAPEN IsLanp: Seroei Aisaoe, Sebosiari, Jwanggin BW 10054 (1); Seroei,
500 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
|
|
Map 3. Distributions of Flindersia amboinensis Poir. (dots) and F. acumt-
nata C, T. White (half-filled circles).
Mariattoe, NJFS bb 30424 (L). VocELKop Division: Sorong, van Royen 3505
, BRI, K, L, LAE); Warsamson Valley, E of Sorong, Jwanggin BW 5643 (Lt),
Moll BW 11623 (1, Laz); Kebar Valley, Koster BW 7116 (1), Schram BW
7805 (L); pir Twanggin BW 5780 (L, LAE), Koster BW 4433 (BO, CANB, L,
LAE), BW 4470 (cANB, L, LAE), BW 4474 (1), BW 4497 (cANB, L, LAE), BW
7062 (esx, L, LAE); Wa kd, ca. 50 km. W of Manokwari, Schram BW 7615
(L); i Plain, Koster BW 11059 (1, LAE), Schram BW 1813 (CANB, L),
pilieorg BW 10444 (L); 8 km. NW of Manokwari, Koster BW 4350 (CANB,
L, LAE); Lower Pami River, ca. 5 km. N of Manokwari, Koster BW 4359 (Bo,
=
a
Manokwar Koster BW 11890 (1 L, LAE); Momi, Kostermans 321 (t), 337 (L),
8 (L); Wariap, Kostermans 481 (1); Inanwatan, Moetoeri (Steenkool),
ies bb 32680 (L). Hotranpra Division: Hollandia, Schram BW 1678 (CANB,
SouTHERN Division: Opka, ca. 10 km. NE of Ninati, Subdivision Moejoe,
Kalkman BW 6454 (case, L, LAE). Territory of New Guinea. SEPIK DISTRICT:
Wewak-Angoram Area, 3 miles E of Urimo, Saunders 975 (CANB, LAE); 3
miles N of Angoram, Pullen 1882 (A, L, LAE); without definite locality, Leder-
mann 10406 (L). Mapanc District: Usino, Henty NGF 27500 (K, LAE);
Hoogland 5035 (A, BM, BRI, K, L, LAE, US). MOoROBE DistRICT: Oomsis Creek,
Bulolo, Dobson & Havel NGF 9116 (Lae). Papua. WeEsTERN District: Lake
Daviumbu, Middle Fly River, Brass 7517 (A, BRI, L, LAE); Lower Fly River,
E bank opposite re Island, Brass 8032 (a); Upper Wassi Kussa sig
Brass 8634 (A, BRI, L, LAE); Oriomo River, Hart NGF 5018 (Bo, CANB, L,
Nsw 99661), White . Gray NGF 10366 (A, 80, BRI, K, L, LAE, NSW 09662).
1969 | HARTLEY, THE GENUS FLINDERSIA 501
oa
Brown River, Allan & Jones NGF 2751 (srt, CANB, L, LAE); near Karema,
(A, BO, K, L, LAE, NSW 99663); Owen Stanley Range, Lane-Poole 362 (Brt-
holotype of Flindersia macrocarpa Lane-Poole ex White & F rancis; K-isotype).
NorTHERN District: Yodda River, Carr 13913 (a); Dobodura Area, NGF
2063 (LAE); ca. 1 km. W of Popondetta, Hoogland 3737A (CANB, LAE); Mana-
galase Area, S side of Hydrographers Range near Siurane, Pullen 5584 (CANB).
Cultivated. Java. Botanic Gardens, Bogor, Anonymous, January 1890 (Bo, L).
Not, as the specific epithet implies, known from Ambon Island. In the
original description in Herbarium Amboinense Rumphius gave Ceram as
the locality.
As delimited here, this is probably the most variable species in the genus.
Among the variable features are:
Petals glabrous to densely appressed-pubescent abaxially.
Petals glabrous to subvillous adaxially.
Stamens 2.2--3.6 mm. long.
Filaments glabrate to pilose subapically.
Capsules 9-21 cm. long.
Exocarp of capsule glabrate to minutely pubescent.
Excrescences of exocarp flattened laterally or thick-conical.
Endocarp reddish brown or cream colored.
OPI AKRON
IT have not been able to find sufficient correlations among these characters
to recognize more than a single taxon. The color of the endocarp, for
example, does not always correlate with the pubescence of the exocarp and
neither of these characters consistently correlate with the shape of the
excrescences or the size of the capsule. In the flowers, the only tendency
toward correlation is between petal pubescence and stamen length. Per-
haps some definite correlations could be made if flowering and fruiting
material could be studied for each variant. Where I have been able to do
this, however, studying flowering and fruiting collections that were ob-
viously the same morphologic type, it has seemed unlikely. Finally, it
should be noted that each of the variable features of this species was found
to be variable in one or more of the other species of the genus as well.
Flindersia macrocarpa, originally described as having larger capsules
than had previously been attributed to any other species of the genus, now
grades into typical F. amboinensis.
Obviously very closely related to Flindersia acuminata, which differs
mainly in having smaller leaves, narrower, more acuminate leaflets, and
shorter stamens. Neither of these species appears to be very closely re-
lated to any of the other species of the genus.
502 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Flowers that were apparently insect-stung were noted in several collec-
tions. They are atypical in having enlarged, almost cartilaginous petals,
filaments, staminodes, and ovaries. Remains of insect larvae were found
in the ovaries of some of the affected flowers.
7. Flindersia acuminata C. T. White, Queensl. Dept. Agr. Bot. Bull.
21: 5. ¢. 2. 1919. Type: Mocatta, July 3, 1915, Queensland, Ather-
ton,
Small to rather large trees to 33 m.; outer bark brownish, with shallow
longitudinal fissures; inner bark brownish grading to yellowish brown or
cream toward the cambium; sapwood whitish; heartwood yellowish;
branchlets, leaves, and inflorescences glabrous to pubescent with mostly
minute, predominantly stellate trichomes. Leaves alternate, imparipin-
nate or (occasional leaves) paripinnate, 12.5-35 cm. long; rachis gla-
brous to finely pubescent; petiolules of lateral leaflets 4-10 mm. long;
terminal leaflet on an extension of the rachis 1—3.1 cm. long; leaflets 3—5
pairs, chartaceous to subcoriaceous, sparsely to densely pellucid-dotted,
glabrous to loosely pubescent below, glabrous or short-pubescent along
the midrib above, elliptic to elliptic-lanceolate, usually unequal-sided and
often subfalcate, 5-15 cm. long, 1.3—4.8 cm. wide, base acute to rounded,
often oblique, main veins 10-17 on each side of the midrib, apex narrowly
long-tapering to short and bluntly acuminate. Inflorescence terminal, 7—
14 cm. long, usually about as wide as long, axes and branches short-pubes-
cent. Flowers bisexual, 3-4 mm. long; pedicels 0.7—1 mm. long; sepals
sparsely appressed-pubescent, ciliolate, suborbicular, 1—1.2 mm. long;
petals creamy yellow, sparsely appressed-pubescent abaxially, glabrous or
with a few papillae adaxially, elliptic-oblong, 3-3.2 mm. long; stamens in-
flexed apically, 1.5-2 mm. long, filaments glabrous, anthers basifixed,
broadly rounded apically, about 0.5 mm. long; staminodes about 1
mm. long; disc about 1 mm. high; gynoecium about 1.5 mm. high and
about 1 mm. wide, ovules 3 on each side of the placentae. Capsule sep-
arating (or easily separable) into 5 distinct valves at maturity, elliptic-
oblong, 9-12 cm. long; exocarp drying light to dark brown, densely and
minutely pubescent to glabrate, muricate, the excrescences thick and
conical, to 5 mm. long; endocarp very pale brown flecked with medium
brown. Seeds 3 on each side of the dissepiments, winged at both ends,
5 cm. long; hypocotyl lateral, horizontal.
ILLUSTRATIONS. WuitE, C. T., ibid. Francis, W. D., Australian Rain-
forest Trees 427. 1951
DistrIBUTION. Cook District, Queensland; well-drained rain forests.
See Map 3
Queensland. Cook Disrricr: Kuranda, Crothers, January 1926 (BRI);
Forestry Reserve 607 ca. 10 miles W of Cairns, Smith 10121 (sri), Draper
(BRI); Tinaroo Range ca. 15 miles NE of Atherton, Smith & Webb 3372 (BRI);
1969 | HARTLEY, THE GENUS FLINDERSIA 503
Atherton, Mocatta, July 3, 1915 (srt- holotype; MEL-isotype), Mocatta (srt,
MEL), Jones 1284 (CANB); Gadgarra, Barnard 31 (CAN NB), Dreghorn 11 (prt),
20E (Bri), Smith 10144 (Bri), 10155 (prt), 10424 (pri), Volk 1411 (BRI), Webb
1661 (CANB), White 1567 (rt); Innisfail, Michael 403 (a, prt): head of
Johnstone River, White, January 1918 (srr), eases definite locality: Wood
Technology Dept. Queensland Forestry Service 56 (NY
8. Flindersia schottiana F. Muell. Frag. od el Austral. 3: 25. 1862.
Lectotype: Bidwill 95, Queensland, Wide Bay
Flindersia schottiana F. Muell. var. pubescens F. Muell. Frag. Phytogr.
Austral. 5: 143. 1866. gt ie — Pua saris seca Bay.
Flindersia pubescens F. M. Bailey, Queensl. Agr. Jou . 1898. Type:
F. M. Bailey, ieee oe 1883, eer Sees ae.
Large trees to 50 m.; outer bark pale brown or gray, generally quite
smooth; inner bark white grading to light brown or yellow-brown toward
the cambium: sapwood pale yellow; heartwood light brown; branchlets,
leaves, and inflorescences glabrous to pubescent with predominantly stel-
late, usually rust-colored trichomes. Leaves opposite, imparipinnate or
(occasional leaves) paripinnate, 19-54 cm. long (the leaves of immature
trees generally much larger); rachis glabrate to appressed- or soft-
pubescent; petiolules of lateral leaflets usually obsolete, occasionally kag
2.5 mm. long, terminal leaflet on an extension of the rachis 1.1-2.7 c
long; leaflets (4—)5-8 pairs, chartaceous to subcoriaceous, densely ade
cid-dotted, glabrate to appressed- or soft-pubescent below, glabrous or
short-pubescent along the midrib above, narrowly elliptic to oblong,
usually unequal-sided and often falcate, 8-22 cm. long, 1.6~—6.3 cm . wide,
bases of lateral leaflets narrowly to broadly rounded to cordate oblique,
base of terminal leaflet acute to cuneate, main veins 15-22 on each side
of the midrib, apex gradually tapering to subacuminate. Inflorescence
terminal, to 40 cm. long, generally much wider than long, axes and branches
appressed- to soft-pubescent. Flowers bisexual, 4-6 mm. long; pedicels
obsolete to 1 mm. long; sepals minutely appressed-pubescent, ciliolate,
broadly ovate to suborbicular, 1-1.5 mm. long; petals white, sparsely
appressed-pubescent abaxially, as: in the basal one-third to one-half
adaxially, elliptic-oblong, 4-6 mm. long; stamens declinate, about 3 mm.
long, filaments pilose i fee or (rarely) glabrous, anthers basifixed,
bluntly mucronulate, about 1 mm. long; staminodes 1-1.7 mm. long; disc
about 1.5 mm. high; gynoecium 2-2.8 mm. high, about 1.5 mm. wide,
ovules 3 on each side of the placentae. Capsule separating (or easily
separable) into 5 distinct valves at maturity, elliptic to elliptic-oblong,
8~13 cm. long; exocarp drying brown, glabrate to minutely and densely
pubescent, muricate, the excrescences rather thick and conical, to 5 mm.
long; endocarp reddish brown to light brown. Seeds 3 on each side of the
dissepiments, winged at both ends, 5-6 cm. long; hypocotyl lateral, hori-
zontal.
ILLustratTions. Barey, F. M., Comprehensive Cat. Queens]. Pl. P/.
504 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
IV. 1913 (as Flindersia pubescens). Francis, W. D., Australian Rain-
forest Trees 161, 162, 163. 1929; 179, 180, 181, 427 (as F. og
430 (as F. Sasbescons’. 1951. Mawen, J. H., Forest Fl. New S. Wales 2
. 69 & 70. 1905.
~
Map 4. Distribution of Flindersia schottiana F. Muell.
DistRIBUTION. New Guinea and eastern Australia south to the Hastings
i New South Wales; rain forests to 700 meters. See MAP 4
ew Guinea (West Irian). VocELKop Division: Sorong, Rmoe, Pleyte
703 beget gratin near Kp. Baroe, Pleyte 737 (Bo, L); Mlasoen Hill E of So-
‘€
1969} HARTLEY, THE GENUS FLINDERSIA 505
rong, van Royen 3406 (A, L, LAE); Warsamson Valley E of Sorong, Schram
BW 12354 (L, Lae); Kebar Valley, Koster BW 7121 (L), Moll BW 9531 (1,
LAE), Schram BW 7921 (1); Sidai, Schram BW 1752 (CANB, L), BW 1756 (tL),
BW 7607 (1). Papua. WESTERN District: Lower Fly River, E bank opposite
Sturt Island, Brass 7991 (A, BRI, L, LAE); Oriomo Creek, mouth of Yakup
Creek, 40 miles from sea, Womersley NGF 17729 (4, mO, K, 1, S
99681). NorTHERN DISTRICT: Bariji-Managalase Area, N side of Sibium
Range S of Toma Village, Pullen 6383 (CANB E); 2 miles from Mafo
L
along Ibinamo River toward Mt. Suckling, Darbyshire 1164 (A, L, LAE, NSW
99603). Queensland. Coox Duisrrict: Cairns, Bailey (Nsw 99606), Betche,
August 1901 (Nsw 99609, MEL); Trinity Bay, Hill, 1876 (BRI, MEL); Rocky
Creek, Atherton District, Bailey, June 29, 1899 (BRI); Martintown, Bailey,
June-July 1899 (Bri, MEL); Forest Reserve 185, Juara Creek, near Danbulla,
Fraser 19 (prt), Smith 3781 (srt); Danbulla, Jones & Pedley 669 (CANB);
Kairi, White, January 24, 1918 (prt); Atherton District, Mocatta (BRI, MEL);
Forestry Reserve 191, Barron, Wongabel, Forestry Department 1 (srt), 2
(BRI), 3 (BRI, K). NortH KENNEDY District: Rockingham Bay, Anonymous
MEL); Mount Dryander, Kilner & Fitzalan (BM, MEL); Dalrymple Heights and
vicinity, Clemens, June-November 1947 (A, BRI), September-November 1947
(GH, MEL, MIcH), November 1947 (GH, MICH); Eungella Range, via Mackay,
Francis, October 3-12, 1922 (prt); Cape Hillsborough, ca. 15 miles NW o
Mackay, Bardsley, November 8, 1967 (BRI). Port Curtis DIsTRICT: Kal-
power, Floyd, September 5, 1949 (LAE); Baffle Creek District, White, April
1920 (srt). BurRNeTtr District: Mt. Perry, Forestry Department, October
1921 (BRI). Wipe Bay District: near the Hummock, a few miles E of Bunda-
berg, Smith 4100 (prt), 4101 (Bri); Bingera, ca. 10 miles WSW of Bundaberg,
GH, MICH); Wide Bay, Bidwill 95 (x-lectotype of Flindersia schottiana F.
Muell.); Fraser Island, Petrie 31 (srt); Bauple, Clemens, June 13, 1945 (cH),
June 10-20, 1945 (micn); Amamoor, Moore in Swain 337 (prt); Kin Kin,
Francis, May 1919 (prt); Imbil, Epps, August 1914 (srr), Webb 5019 (CANB);
Brooloo, Webb SN 5339 (cANB), 1635 (caNB). Moreton District: Parish
of Monsidale, Webb SN 5422d (cANB); Eumundi, 5 miles S of Cooroy, Bailey
& Simmonds, November 1894 (rt); Blackall Range, Twine, February 1918
(BRI), White, April 1918 (srt); Sawpid Range, White, April 1918 (srt); Nor-
ueensian n
tt Gan 99600): 5 miles S of Brunswick Heads, Gray & Gray -
(CANB); Potts Point, Robbins 2612 (cANB); Byron Bay Lighthouse Road, Con-
stable 6505 (Nsw 83976); Richmond River, Henderson, November 1886 ae
Nsw 99597); Clarence and Richmond (rivers), Northern Woods aa ou
Wales 24 (x, MEL); Whian Whian, Jones 469 (cANB), 939 (cANB); Casino,
506 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
District Forester, December 1, 1905 (BRI, Nsw 99594), McAuliffe 8338/12
(nsw 99593, us); Tintenbar, Baeuerlen 633 (BM, MICH, NSW 99602); Clarence
River, Anonymous (MEL), Moore 137 (MEL), Northern Woods New South Wales
61 (K, MEL), Selwyn (MEL); Clarence River, Grafton, Northern Woods New
South Wales 63 (Nsw 99604), Squire, November 1937 (Nsw 99595); Mt.
Yarrahappini, Briggs 70.05F (Nsw 99601), Smith 2 (Nsw 99592); lower slopes
of Dorrigo Range, Wheen, August 29, 1945 (Nsw 99596); Taylor’s Arm,
Wilshire, June 2, 1905 (Nsw 99605); Bellinger and McLeay Rivers, Anony-
mous (K); McLeay River, MacDonald 183 (MEL), Woolls (MEL); Kempsey,
MacDonald 212 (met), Rudder, May 17, 1891 (Nsw 99591); Hastings River,
Tozer (Thozet?) (Nsw 99599). Cultivated. Java. Bogor Botanic Gardens,
Anonymous, 1903 (us). Queensland. Cook District: Forestry Reserve 310,
Gadgarra, Smith 10824 (prt). Burke District: Mt. Isa, Pedley 1060 (Rt).
Port Curtis District: Rockhampton Botanic Gardens, Simmons 4 (BRI).
Moreton District: Brisbane Botanic Gardens, Bailey (Brn), Blake 2688 (BRI),
White (srt); Kangaroo Point, Brisbane, Francis (Bo), White, November 1912
(BRI); Wickham Reserve, ae os October 23, 1883 (BRI-holotype of
Flindersia pubescens F. M. iley). w South Wales. Sydney Botanic Gar-
dens, Boorman, December one (P).
Flindersia pubescens is apparently recognized as a distinct species by
many Australian botanists. It is considered to have a rather limited
distribution centering on the Cairns area in northeast Queensland whereas
F. schottiana, sensu stricto, is considered to be wider ranging, extending
from New Guinea and north Queensland south to the Hastings River in
New South Wales. The morphologic differences between the two are
given in the following key from White (1921):
Leaflets on flowering shoots subcoriaceous, somewhat falcate, 6.5-13 cm.
long, 2-3.3 cm. broad, quite glabrous on the rachis and under surface
clothed with very close and dense stellate, velvety tomentum, veins and
Neos F. schottiana
Leaflets on the flowering branches chartaceous, 12.5-23 cm. long, 4.5-6.5 cm
broad, rachis densely clothed with comparatively long golden-brown stel-
late hairs, under surface clothed with numerous but more or less scat-
tered stellate hairs, the veins and veinlets prominent F. pubescens
After studying a large number of collections from both Australia and
New Guinea I am convinced that the differences between these two taxa
all break down to a greater or lesser degree and that they represent
environmental adaptations of sub-taxonomic significance. Their geographic
distributions tend to substantiate this, I think, since both forms occur at
widely disjunct localities in New Guinea. For example, Brass 7991, from
the Fly River and Pleyte 737, from the Vogelkop Peninsula, are good
matches for typical F. schottiana, whereas Darbyshire 1164, from the
Northern District of Papua, and Schram BW 12354, from the Vogelkop
Peninsula, are typical of F. pubescens.
As indicated above, in the outline to species relationships, Flindersia
schottiana, F. bourjotiana, and F. xanthoxyla appear to be more closely
related to one another than to any of the other species of the genus.
Beyond this, however, the three are mutually quite distinct.
1969 | HARTLEY, THE GENUS FLINDERSIA 507
9. Flindersia bourjotiana F. Muell. Frag. Phytogr. Austral. 9: 133.
1875. Type: Dallachy, Queensland, Rockingham Bay.
Flindersia tysoni C. DC. Bull. Herb. Boiss. ser. 2. 6: 986. 1906. Type: Tryon,
August 1901, Queensland, Mossman River.
Large trees to 35 m.; outer bark gray or brown, rather smooth; inner
bark pale yellow-brown; sapwood whitish; heartwood pale yellow-brown;
branchlets, leaves, and inflorescences glabrous to pubescent with mostly
minute, predominantly stellate trichomes. Leaves opposite, imparipinnate
or (occasional leaves) paripinnate, 9-33 cm. long; rachis glabrate to
appressed- or velvety-pubescent; petiolules of lateral leaflets 1.5-4 mm.
long, terminal leaflet on an extension of the rachis 1—2 cm. long; leaflets
(1-)2-4 pairs, chartaceous to subcoriaceous, densely pellucid-dotted,
glabrous to rather densely short-pubescent below, glabrous to sparsely
appressed-pubescent above, elliptic to lanceolate, equal- or only slightly
unequal-sided, 5.5-17 cm. long, 1.5—4.8 cm. wide, base acute to obtuse,
main veins 12-17 on each side of the midrib, apex obtuse to acute or
occasionally bluntly acuminate. Inflorescence terminal, to 20 cm. long,
generally much wider than long, axes and branches sparsely to rather
densely short-pubescent. Flowers bisexual or (a few to many flowers in
an inflorescence) functionally staminate, 6-10 mm. long; pedicels obsolete
to 1 mm. long; sepals glabrate to sparsely appressed-pubescent, ciliolate,
broadly ovate to suborbicular, 1-2 mm. long; petals white or greenish
white, sparsely appressed-pubescent abaxially, glabrous adaxially, elliptic,
5—9.5 mm. long; stamens declinate, 4—5.7 mm. long, filaments sparsely to
rather densely pilose subapically, anthers dorsifixed, mucronate, 1—1.2 mm.
long; staminodes 1-2 mm. long; disc thin in bisexual flowers and com-
paratively thick in functionally staminate flowers, 1-1.5 mm. high;
Synoecium in bisexual flowers 2-3 mm. high, about 1.5 mm. wide, ovules
3 on each side of the placentae; gynoecium in functionally staminate
flowers poorly differentiated, narrowly conical, about 1 mm. high, without
ovules. Capsule separating (or easily separable) into 5 distinct valves at
maturity, elliptic, 7-15 cm. long; exocarp drying blackish-reddish brown,
glabrous, muricate, excrescences slender, often recurved, to 4 mm. long;
endocarp light brown. Seeds 3 on each side of the dissepiments, winged
at both ends, about 5.5 cm. long; hypocotyl lateral, ascending.
ILLUSTRATION. Francis, W. D., Australian Rain-forest Trees 426. 1951.
Distripution. Northeast Queensland, Cook and North Kennedy Dis-
tricts; rain forests to 900 meters. See Map 5.
Queensland. Cook District: Mossman River, Tryon, August 1901 (srI,
NSw-isotypes of Flindersia tysoni C. DC.); near Ayton, Gittons 575 (BRI
35658); Bailey’s Creek Area, Smith 11556 (prt); Mt. Lewis, ca. 10 miles N
of Mt. Malloy, Schodde 3326 (cANB); Kuranda, DuRietz, August 1927 (BRI);
Atherton, Curry 5 (ny), Jones 1289A (CANB), 1289B (cANB), Mitchell, August
1911 (Nsw 99683), Tardent X230 (srt), Webb 2502 (cans); Atherton Table-
508 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
P 5. Distributions of Flindersia bourjotiana F. Muell. (dots) and F.
xanthoxyla (A. Cunn. ex Hook.) Domin (half-filled circles).
land, Tardent (a, BRI); ercnmne! Fraser 22 (BRI), Webb 5131 (cANB); Gad-
garra, Dreghorn 12 (BRI), Kajewski 1140 (a, BM, BRI, NY, US), Trist 33 (NY);
Lake Barrine, Barnard, June 15, 1941 (CANB) ; ’ Glenallan, Malanda, Tardent
188 (BRI); Ravenshoe, Manuell 31 (kK); Russel River, Anonymous (MEL); In-
nisfail, Michael 132 (GH), Petrie (A); Johnstone River, Bancroft, 1885-1886
(BRI, MEL), Michael, May 1916 (prt); Bingil Bay Road, Gittons 575 (BRI
35542); Etty Bay, Webb 905 (cans). NortH KeNNepy District: Herberton,
Mocatta, February 1917 (srr); Kirrama Range W of Kennedy, between So-
ciety Flat and Yuccabine Creek, Smith 3205 (srt); Murray River, Anonymous,
December 12, 1861 (meL); Rockingham Bay, Dallachy (met-holotype of
Flindersia hausjoviana F. Muell.; Bo, MEL-isotypes)
Flindersia tysoni was seneaiiialy placed in the synonymy of F. bour-
jotiana by White (1921).
10. Flindersia nooner ae (A. Cunn. ex Hook.) Domin, Bibliot. Bot.
22 (89): 298. 1927.
sree xanthoxyla A. Cunn. ex Hook. in Hooker’s Bot. Misc. 1: 246. t. 54.
830. Type: Cunningham 117, Queensland, Brisbane River
1969 | HARTLEY, THE GENUS FLINDERSIA 509
Flindersia oxleyana F. Muell. Frag. Phytogr. Austral. 1: 65. 1859 (nomen
illegit., based on Oxleya xanthoxyla A. Cunn. ex Hook.).
Medium to large trees to 40 m.; outer bark gray or gray-brown, fairly
smooth; wood yellow; branchlets, leaves, and inflorescences glabrous to
pubescent with mostly minute, predominantly stellate trichomes. Leaves
opposite, imparipinnate, 15-32 cm. long; rachis glabrous to appressed- or
short-pubescent; petiolules of lateral leaflets obsolete to 6 mm. long,
terminal leaflet on an extension of the rachis 0.8—2.8 cm. long; leaflets
(2—)3-5 pairs, membranaceous to chartaceous, very brittle when dry, with
or without scattered pellucid dots, glabrous to appressed- or soft-pubescent
below, glabrous to sparsely appressed-pubescent above, elliptic to lanceo-
late, usually unequal-sided and often falcate, (2.2-)4-13 cm. long, (0.6-)
1.3-3.2 cm. wide, base obtuse to cuneate, often oblique in lateral leaflets,
main veins 11-15 on each side of the midrib, apex long-tapering, acute.
Inflorescence terminal, to 25 cm. long, usually much wider than long, axes
and branches appressed- to short-pubescent. Flowers bisexual, 4-5 mm.
long; pedicels obsolete to 2 mm. long; sepals glabrous to sparsely ap-
pressed-pubescent basally, ciliate, broadly ovate, about 1 mm. long; petals
pale yellow, glabrous, elliptic-oblong, about 4.3 mm. long; stamens decli-
nate (becoming straight after anthesis), about 3 mm. long, filaments
sparsely pilose subapically, anthers dorsifixed, obtuse apically, about 1
mm. long; staminodes about 1.3 mm. long; disc 0.7-1.2 mm. high; gynoe-
cium 1.5 mm. high, about 1 mm. wide, ovules 3 on each side of the
placentae. Capsule separating (or easily separable) into 5 distinct valves
at maturity, elliptic-oblong, 6.5~11 cm. long; exocarp drying dark brown
to pale gray-brown, densely and minutely pubescent, muricate, excrescences
rather narrowly conical, unequal in length, to 4 mm. long; endocarp
yellow-brown. Seeds (2—)3 on each side of the dissepiments, winged at
both ends, 3.3-5 cm. long; hypocotyl lateral, horizontal.
ILLUSTRATIONS. CUNNINGHAM, A., ibid. Francis, W. D., Australian
Rain-forest Trees 164, 165. 1929 (as Flindersia oxleyana) ; 184, 185. 1951.
Maren, J. H., Forest Fl. New S. Wales 2: t. 73 & 74. 1906 (as F.
oxleyana).
DistRIBUTION. Southeast Queensland and adjacent New South Wales;
rain forests to 500 meters. See Map 5.
Queensland. Burnetr District: Edenvale Hill, near Kingaroy, Michael
3106 (BRI), Wipe Bay District: Mary River Scrub, Gympie, Kenny (BRI);
Imbil, McAdam 83 (prt, NY), 85 (A, BRI), 87 (BRI, NY), Weatherhead, July
1917 (srt). Moreton District: Parish of Monsidale, Webb SN 5422c (CANB);
Yarraman, Clemens, August 4-15, 1944 (A, Ny), Floyd, August 29, 1949 (LAE),
Webb 5143 (cans), SN 5337 (cans); South Pine River near Samford, Hub-
bard 5941 (a, Bri, K); South Pine River near Bald Hills, White 7155 (a, BO,
NY); Samford Range, Shirley & White, April 1918 (BRI); Samford, Tracey in
Webb & Tracey 3392 (cans); Petrie, Blake 3079 (srt); Enoggera, Bailey
(Nsw 99643): Three-mile Scrub, Enoggera Creek, Bancroft (Bri); Brisbane
River below Breakfast Creek Bridge, Bailey (BRI, NSw 96644); Brisbane River,
510 JOURNAL OF THE ARNOLD ARBORETUM [ VoL. 50
Cunningham 109 (GH), 117 (BM-holotype of Oxleya xanthoxyla A. Cunn. ex
Hook.; k-isotype), Hill (MEL); Sherwood, Brisbane River, Hubbard 5942 (x),
Anonymous, December 18, 1930 (BRI); Tamborine Mountain, Longman &
White, February 1917 (pr1). Without definite locality: Wood Technology
Dept. Queensland Forest Service 80 (NY). New South Wales. Acacia Creek via
Killarney, Queensland, Boorman, February 1905 (Nsw 99639), Dunn, No-
vember 1905 (NSW 99642), Dunn 252 (NSW 99640); Tweed River, Anony-
mous 59 (MEL), Moore 14125 (pm); Murwillumbah, Charles, January 9, 1905
(Nsw 96649); Whian Whian, Jones 943 (CANB), Webb 2457 (CANB), 5241 (CANB),
White 12769 (srt); Wollongbar Experimental Farm, Lismore, Johnson &
Constable, June 11, 1957 (Nsw 96650); Tintenbar, Baeuerlen, March 1892
(a); Richmond River, Fawcett (MEL), Henderson (MEL), Watts, 1902 (NSW
99641); Richmond River to the Tweed River, Moore (pm, GH, K); Hastings
River, Boorman, August 1907 (Pp); Sandiland Ranges, Boorman, November
1904 (Nsw 96646). Without definite locality. Leichhardt (K, Nsw 99638).
Cultivated. Queensland. Wipe Bay District: Gympie, Wickham and Channon
Streets, Kenny, January 17, 1907 (BRI). New South Wales. Sydney ot
Gardens, Boorman, February 1907 (NSW 96647), Camfield, January 1894 (N
96648), December 1896 (Ny), January 1898 (MEL, Us).
11. Flindersia bennettiana F. Muell. ex Benth. Fl. Austral. 1: 389.
1863. Lectotype: Bidwill, Queensland, Wide Bay.
Flindersia leichhardtii C. DC. Monogr. Phanerog. 1: 731. 1878. Type:
Leichhardt, 1845, Queensland, Moreton Bay.
Small to large trees to 43 m.; outer bark pale gray, to reddish brown,
quite smooth; inner bark whitish; branchlets, leaves, and inflorescences
glabrous to minutely pubescent with stellate trichomes. Leaves opposite,
imparipinnate, 8.5-36(—45) cm. long; rachis glabrous to appressed- or
rarely short-pubescent; petiolules of lateral leaflets 1-6 mm. long, terminal
leaflet on an extension of the rachis 0.9-3.5 cm. long; leaflets 1-3 (—4)
pairs, subcoriaceous to coriaceous, densely pellucid-dotted, glabrous or
sparsely appressed-pubescent below, glabrous above, ovate to elliptic to
elliptic-oblong, equal- or occasionally unequal-sided and subfalcate, 6—18.5
cm. long, 1.7—6.7 cm. wide, base obtuse to subacute, often slightly oblique,
veins prominent above, main veins 10-30 on each side of the midrib, apex
obtuse or occasionally acute. Inflorescence terminal or occasionally ter-
minal and upper-axillary, to 25 cm. long, as wide or nearly as wide as
long, axes and branches appressed- to short-pubescent. Flowers bisexual
or (a few to many flowers in an inflorescence) functionally staminate, 3-6
mm. long; pedicels 0.3-3.5 mm. long; sepals sparsely to densely short-
pubescent, ciliolate, ovate, 1-1.5 mm. long; petals white, sparsely ap-
pressed-pubescent abaxially, glabrous stile, elliptic-oblong, 2.5—5 mm.
long; stamens declinate, 2.5-4 mm. long, filaments glabrous, ae dor-
sifixed, bluntly mucronulate, about 0.9 mm. long; staminodes 1-1.7 mm.
long; disc about 1 mm. high; gynoecium in bisexual flowers 2—2.8 mm. high,
about 2 mm. wide, ovules 2 on each side of the placentae; gynoecium in
functionally staminate flowers poorly differentiated, conical, about 1 mm.
1969 | HARTLEY, THE GENUS FLINDERSIA 511
high, without ovules. Capsule separating (or easily separable) into 5
distinct valves at maturity, elliptic to elliptic-oblong, 4-7 cm. long; exocarp
drying medium to very dark reddish brown, glabrous, muricate, excres-
cences narrowly conical, unequal in length to 4 mm. long; endocarp
reddish brown. Seeds 2 on each side of the dissepiments, winged at both
ends, 3—4.3 cm. long; hypocoty] lateral, horizontal.
ILLUSTRATIONS. Francis, W. D., Australian Rain-forest Trees 168, 169.
1929; 186, 187. 1951. Maren, J. H., Forest Fl. New S. Wales 3: t. 77 &
78. 1906
DIstRIBUTION. Southeast Queensland and northeast New South Wales;
rain forests to 300 meters.
Queensland. Wipe Bay District: Fraser Island, Epps 30 (ny), Forestry
Dept. 131 (prt), 132 (BRI), 133 (BRI), Hubbard 4403 (x), Petrie 30 (srt),
Trist 6 (NY); Dundowran via Gympie, Tryon, July 1928 (srt); Maryborough
District, Young, September 1916 (prt), Mt. Bauple, Kajewski, September
1922 (a, BRI); Wide Bay, Bidwill (x-lectotype of Flindersia bennettiana F.
Muell. ex Benth.; cH, ny); Gympie, Hamilton-Kenny 145.05 (NSw 99582);
Kin Kin, Francis (prt, Nsw 99586); Forest Survey Camp, Amamoor, Anony-
mous, April 22, 1919 (prt); Imbil, Floyd, September 18, 1949 (LaE), McAdam
90 (A, BRI, NY), Weatherhead in Swain 346 (sri), Webb 5013 (CANB). More-
TON District: Eumundi, Bailey, June 1893 (BRI, Nsw 99588), Simmonds,
June 1895 (a); Candle Mountain, White, May 5, 1918 (prt); Moreton Bay,
Cunningham (pm), Hill (xk, MEL), Leichhardt, 1845 (p-holotype of Flindersia
leichhardtii C. DC.); Brisbane, Boorman, April 1899 (MEL, Nsw 99585), White,
June 3, 1926 (a, BO, BRI); Mount Cotton, Scortechini (BRI, MEL); Tamborine
Mountain, Clemens, March 1947 (prt); Beech Mountain, White 1907 (A, BRI,
NSw 99584); Southport, Simmonds, June 1913 (a, Ny); Meyer’s Ferry, Nerang
River, White, October 20, 1917 (BRI, NSW 99583). Without definite locality:
Bennett (nsw 99587). New South Wales. Tweed River, Anonymous, November
1897 (MEL); Tweed River District, Grime, July 1900 (Nsw 99576); Cudgen,
Murwillumbah District, McKee 9547 (BRI, CANB, NSW 99589); Hastings Point,
Trapnell, June 7, 1960 (BRI); east foothills of Nightcap Range ca. 2-3 miles
Scrub, near Byron Bay, Forsyth, November 1898 (NSW 99575); Whian Whian
State Forest, Jones 937 (cans), Webb 5242 (cans); Richmond River, Baeuer-
len 244 (MeL), Boorman, February 1899 (p), Fawcett (MEL), Henderson 22
(MEL), (BM, BO, MEL, Nsw 99573), Mrs. Hodgk (MEL); Dalwood, Richmond
River, Graham in Watts 37 (MEL, NSW 99579); Richmond and Clarence Dis-
tricts, Bennett (MEL); Minyon Falls, N of Lismore, de Beuzeville, July 16,
1936 (Nsw 99566); Dorroughby, 14 miles NE of Lismore, Byrnes, September
1953 (Nsw 99580); Wollongbar Experimental Farm, Anonymous, November
1897 (Nsw 99570), Manager Experimental Farm, Wollongbar 6 (Nsw 99577);
Lismore, Forest Guard, March 1909 (Nsw 99568), Baeuerlen 350 (A, NSW
99574), 698 (Nsw 99581); Ballina, Baeuerlen 836 (MEL); 8 miles N of W ood-
burn, Williams J43 (Nsw 99571); Clarence River, Beckler (MEL), Wilcox, 1875
(MEL); Iluka, Cameron 93 (Nsw 99567). Cultivated. New South Wales. Syd-
ney Botanic Gardens, Boorman, September 1919 (BRI, NSw 99578, us), Cam-
field, September 1896 (ny), February 1899 (Nsw 99569, us), Mueller (Nsw
512 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
99590); Sydney, Paddington, Vernon, March 11, 1879 (MEL). Victoria. Mel-
bourne Botanic Gardens, Anonymous, October 23, 1934 (MEL)
There is apparently no authentic material of Flindersia leichhardtii in
the Australian herbaria and it has not previously been placed in the
synonymy of F. bennettiana.
As is indicated above, Flindersia bennettiana, F. collina, F. dissosperma,
and F. maculosa comprise a group of related species. They appear to be
related to one another in a linear sequence, beginning with a rain-forest
species, F. bennettiana, and ending with a xerophyte, F. maculosa. The
following outline shows the apparent relationships of these species as
indicated by various morphologic features.
pacer stellate; leaves imparipinnate, 8.5-36(45) a hes the rachis
III sb. des. 2 10 Wd atin anraceto is else re a ae aed . F. benne ae
Trichomes stellate and a leaves imparipinnate or age 5-14
crs TOG, AT YARIS WIE. cod os pow a es Keke de vee dnkas . F. collina.
Trichomes predominantly lepidote; leaves “seeing trifoliolate, 1.5—
G5 crm. Ini, the vachin Wine... 02. kek cen ecu . F. dissosperma.
Trichomes predominantly lepidote; leaves 1-9 cm. long, simple. .........-
Se iene ea te te ee Parsee or aired ore ead Gaia: 14. F. maculosa.
12. Flindersia collina F. M. Bailey, Queensl. Agr. Jour. 3: 354. 1898
(based on Flindersia strzeleckiana F. Muell. var. latifolia F. M.
Bailey). Lectotype: Bailey, Queensland, Moreton District, Main
Range.
Flindersia see F. Muell. var. latifolia F. M. Bailey, Synop. Queensl.
Fl. 1st suppl. 12. 1886.
Medium to large trees to 40 m.; outer bark mottled, green and brown,
exfoliating in thin roundish flakes leaving shallow depressions; inner bark
reddish grading to cream toward the cambium; sapwood yellow to yellow-
brown; heartwood yellow to pale brown; branchlets, leaves, and inflor-
escences glabrous to lepidote or minutely pubescent with predominantly
scale-like and stellate trichomes. Leaves opposite or subopposite, impari-
pinnate or trifoliolate, 5-14 cm. long; rachis glabrous to lepidote or
appressed-pubescent below, glabrous or short-pubescent above, narrowly
to broadly winged laterally, the wings extending 0.3—2.4(—4) mm. on each
side; leaflets sessile, 1-2(—3) pairs, chartaceous to coriaceous, with scat-
tered pellucid dots, glabrous or sparsely lepidote or appressed-pubescent
below, glabrous above, elliptic to obovate to broadly spatulate, 2-9 cm.
long, 1-4.7 cm. wide, base obtuse or (in some terminal leaflets) attenuate,
veins prominent above, main veins (10—)12—16 on each side of the midrib,
apex obtuse to rounded, usually retuse or emarginate. Inflorescence ter-
minal or terminal and axillary, to 17 cm. long, generally about as wide as
long, axes and branches glabrate to short-pubescent. Flowers bisexual or
(a few to many flowers in an inflorescence) functionally staminate, 4.7-5-3
nd
1969 | HARTLEY, THE GENUS FLINDERSIA 513
mm. long; pedicels 0.54.5 mm. long; sepals appressed-pubescent, ciliate,
broadly ovate, about 1 mm. long; petals white, glabrate to appressed-
pubescent abaxially, short-pubescent in the basal half adaxially, elliptic,
4-5 mm. long; stamens declinate, 3.5-4 mm. long, filaments glabrous,
anthers subdorsifixed, subacute apically, about 1 mm. long; staminodes
about 2 mm. long; disc about 0.75 mm. high; gynoecium in bisexual
flowers about 1.75 mm. high, 1.25 mm. wide, ovules 2 on each side of the
placentae; gynoecium in functionally staminate flowers poorly differen-
tiated, conical, about 1 mm. high, without ovules. Capsule separating (or
easily separable) at maturity into 5 distinct valves, elliptic to elliptic-
oblong, 2.8-5 cm. long; exocarp drying dark brown, glabrous or glabrate,
muricate, excrescences 1-2 mm. long; endocarp rather dark brown. Seeds
2 on each side of the dissepiments, winged at both ends, 1.5—2.5 cm. long;
hypocotyl lateral, horizontal.
ILLustrations. Francis, W. D., Australian Rain-forest Trees 170, 171.
1929; 188, 189. 1951. Maren, J. H., Forest Fl. New S. Wales 3: ¢. 81
& 82. 1907
Map 6. oe ae “ oo collina F. M. Bailey (half-filled genie
F. dissosperma (F. Muell.) Domin (dots), and F. maculosa (Lindl.) B
(open circles with eee line
514 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
DistRIBUTION. Southeast Queensland and adjacent New South Wales;
rain forests and rather dry scrubs to 700 meters. See Map 6.
Queensland. LEICHHARDT District: Taroom, Mobsby, October 1912 (BRI).
BurNEtt District: Eidsvold, Bancroft, April 1911 (pri), April 1923 (a),
April 1928 (Nsw 99671); Goodnight Scrub, ca. 40 miles SW of Bundaberg,
Smith 9882 (BRI); West Wooroolin, NW of Kingaroy, Everist, March 23,
1961 (prt); Kingaroy, Smith 3115 (pri, Ny); Nanango, Grove 131 (BRI);
Cooyar and Charlestown, Gympie Forest District, Swain, March 1917 (BRI);
without definite locality (probably Mt. Perry fide L. S. Smith), Keys(?) 800
(BRI). Wipe Bay District: Bundaberg, Boorman, July 1912 (Nsw 96676);
Kepnock, about 2 miles E of Bundaberg, Smith 4163 (prt); Childers, Helms,
January 3, 1899 (BRI), Anonymous (NSW 99670); Bauple, Clemens, June
20, 1945 (GH, MICH); near Imbil, Smith & Webb 3129 (BRI, CANB). MARANOA
District: about 20 miles W of Mitchell, Everist, March 7, 1950 (prt). Dar-
LING Downs District: Chinchilla, Beasley 52 (prt); Bunga Mts., White, Octo-
ber 1919 (srt); Rangemore School Area, Cooyar-Bunya Mountains Road,
Smith 10260 (A, BRI, K, NSW 96677); Toowoomba, Longman, October 1910 (kK,
Nsw 99675). Moreton District: Parish of Monsidale, Webb SN 54226
(CANB); Yarraman, Clemens, August 1944 (A, BRI, NY, US), Floyd, August 28,
1949 (LAE), Webb SN 5338 (cans); Kilcoy, English, October 1919 (A, BRI);
Jimna, near Kilcoy, Webb 5248 (cans); Crow’s Nest, Clemens 43641 (A);
Main Range, Bailey (srt-lectotype of Flindersia collina F. M. Bailey), Pente-
cost 29 (BRI, NSW 99669); Flagstone Creek, ca. 8 miles SE of Toowoomba, 7
miles SW of Helidon, Smith, October 13, 1965 (BRI); Helidon, White 8765 (A);
Fernvale, ca. 13 miles NW of Ipswich, Bevington, 1909 (srt); Rosewood,
Anonymous, October 1908 (BRI); Brisbane | i abhi 164 (pm), Octo-
ber 1828 (kK, oe Moreton Bay, pa 8 (BM, GH, NY), pee (K);
bar Road, Everist & Webb 1413 (srt). Without definite locality: Bowman
(MEL), Trist 19 (Ny). New South Wales. Acacia Creek, near Killarney, Queens-
land, Boorman 15 (Nsw 99673), February 1905 (xk), Dunn, September 1905
(MEL, NSW 99672), October 1905 (Kk), February 1906 (Nsw 99674); Unumgar,
Jones 2371 (CANB). Without definite locality. Hill 53 (MEL).
13. Flindersia dissosperma (F. Muell.) Domin, Bibliot. Bot. 22 (89):
298. 1927.
Strzeleckya dissosperma F. Muell. in Hooker’s Jour. Bot. aig Gard. Misc.
9: 308. 1857. Type. Mueller, Queensland, Burdiken Riv
Flindersia strzeleckiana F. Muell. Frag. Phytogr. Austral. 1: “6S. 1859 (nomen
illegit., based on Strzeleckya dissosperma F. Muell.).
Small trees to 10 m., developing from a divaricately branched shrub
stage; outer bark mottled, dark gray and white, cream or salmon, rough-
scaly on the trunk, smooth above; inner bark reddish grading to cream
toward the cambium; branchlets, leaves, and inflorescences glabrous to
lepidote or minutely pubescent with scale-like and stellate trichomes.
Leaves opposite, imparipinnate or trifoliolate, or (rare occasional leaves)
simple, (0.8—)1.5-6.3 cm. long; rachis sparsely lepidote below, winged
1969 | HARTLEY, THE GENUS FLINDERSIA 515
laterally, the wings extending 0.5—1.5 mm. on each side; leaflets sessile,
1—2 pairs, chartaceous to subcoriaceous, with scattered pellucid dots, sparse-
ly lepidote below, glabrous above, elliptic, spatulate, oblong or sublinear,
0.6-3.7 cm. long, 0.2-0.7 cm. wide, base obtuse, main veins usually indis-
cernible, 8-10 on each side of the midrib, apex rounded to acute, occa-
sionally retuse. Inflorescence terminal or rarely terminal and upper-
axillary, to 8 cm. long, usually nearly as wide as long, axes and branches
sparsely to rather densely lepidote to appressed- or short-pubescent.
Flowers bisexual or (a few to many or all of the flowers in an inflorescence)
functionally staminate, 3-4 mm. long; pedicels 0.7-2 mm. long; sepals
glabrous or glabrate, ciliate, suborbicular, 1-1.3 mm. long; petals white to
cream, glabrous, broadly elliptic, 3-3.5 mm. long; stamens inflexed
apically, about 2.5 mm. long, filaments glabrous, anthers subdorsifixed,
bluntly mucronulate, about 1 mm. long; staminodes 0.5—1 mm. long; disc
about 0.5 mm. high; gynoecium in bisexual flowers about 1.5 mm. high
and 1 mm. wide, ovules 2 on each side of the placentae; gynoecium in
functionally staminate flowers poorly differentiated, pulvinate, about 0.4
mm. high, without ovules. Capsule separating (or easily separable) into
5 distinct valves at maturity, elliptic, 2-3 cm. long; exocarp drying dark
reddish brown, glabrous, muricate, the excrescences 1-2 mm. long; endo-
carp brown. Seeds 2 on each side of the dissepiments, winged at both ends,
1.5—1.8 cm. long; hypocotyl lateral, horizontal.
ILLUSTRATION. Battery, F. M., Comprehensive Cat. Queensl. Pl. t. 73.
1913 (as Flindersia euieciionas.
DistripuTion. East central Queensland; dry scrubs to 300 meters. See
AP 6.
Queensland. NortH KENNEDY District: Maryvale Station, Daintree (MEL);
W of Charters Towers, Blake 14906 (prt); Charters Towers, Michael 1275
(BRI), Stephens North Queensl. Nat. Club 10468 (Nsw 99689); Millchester Hill,
Stephens North Queensl. Nat. Club 9056 (srt); Burdiken, Kennelly 208 oe
Mueller (met-holotype of Strzeleckya dissosperma F. Muell.; K-isotype); H
bert’s Creek, Bowman 70 (MeL); near Bogie Range, lower Burdekin River, ca.
42 miles S of Ayr, Smith 4532 (prt); Cape River, Daintree (met), Fitzalan
(MEL). SourH KENNEDY District: Head of Suttor River, Sutherland (MEL);
Laglan, about 80 miles W of Clermont, Everist, October 12, 1960 (BRI, K); 18
March 1920 (srr). LercHHarpT District: Peak Downs, Mueller (MEL); 9
miles SW of Anakie, Adams 1281 (CANB, MEL, NS 99685): E of Emerald,
Webb 2251 (cans); Emerald, Webb SN 5296 mene Blair Athol, Massey 30
(BRI); ca. 3 miles N of Clermont on road to Charters Towers, Smith 3160 (BRI);
Clermont, Small, September 1912 (BRI, K, NSW 99621); Chirnside, ca. 6 miles
S of Capella, Smith & Webb 3428 (BRI); 2 miles N of Emerald, Bisset E198
(BRI); 4 miles E of Girrah Homestead, 36 miles N of Blackwater Township,
1967 (prt); 7 miles N of Goowarra, Anderson, October 15, 1965 (BRI); about
3 miles E of Goowarra, Johnson 943 (A, BRI); 3 miles E of Parkes Homestead,
516 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Speck 1672 (CANB, NSW 99686); Dingo, O’Shanesy 2013 (MEL); 8.5 miles SW
of Duaringa, Speck 1820 (BRI, CANB, NSW 99687); near Duaringa, Simmons,
May 1938 (RI); between the Barcoo and Springsure, Bailey (MEL). Without
definite locality: Bailey (BRI, NSw 99688), Fitzalan (MEL).
14. Flindersia maculosa (Lindl.) Benth. Fl. Austral. 1: 389. 1863.
Elaeodendron maculosum Lindl. in Mitchell Trop. Austral. 384. 1848. Type:
Mitchell, November 1846, New South Wales, Balonne River.
Flindersia maculata F. Muell. Quart. Jour. Trans. Pharm. Soc. Victoria 2:
44. 1859 (nomen illegit., based on Elaeodendron maculosum Lindl.).
Small to medium trees to 15 m., developing from a divaricately branched
shrub stage; outer bark mottled, dark gray or brown and cream or white,
smooth-scaly on the trunk, smooth above; wood yellow; branchlets, leaves
and inflorescences glabrous to lepidote or minutely pubescent with scale-
like and stellate trichomes. Leaves opposite, simple; petioles lepidote
below, short-pubescent above, 1-12 mm. long; leaf blades chartaceous to
subcoriaceous, with scattered pellucid dots, minutely and sparsely lepidote
and appressed-pubescent below, glabrous or short-pubescent on the midrib
above, narrowly elliptic-oblong, oblanceolate or sublinear, 1-8 cm. long,
0.25-1 cm. wide, base narrowly obtuse to attenuate, main veins usually
indiscernible, 12-17 on each side of the midrib, apex rounded to obtuse,
occasionally retuse. Inflorescence terminal or rarely terminal and upper-
axillary, to 7.5 cm. long, usually about as wide as long, axes and branches
sparsely to densely lepidote to appressed- or short-pubescent. Flowers
bisexual, 4-4.5 mm. long; pedicels obsolete to 2.5 mm. long; sepals
glabrous, ciliate, broadly ovate to orbicular, 1-1.5 mm. long; petals white
to cream, glabrous, obovate, about 4 mm. long; stamens inflexed apically,
about 3 mm. long, filaments glabrous, anthers subdorsifixed, obtuse to
bluntly mucronulate, 1-1.2 mm. long; staminodes about 0.5 mm. long;
disc about 0.75 mm. high; gynoecium about 1.5 mm. high and 1 mm. wide,
ovules 2 on each side of the placentae. Capsule separating (or easily
separable) into 5 distinct valves at maturity, elliptic, 2.3-2.7 cm. long;
exocarp drying dark reddish brown, glabrous, muricate, the excrescences
1—1.5 mm. long; endocarp brown. Seeds 2 on each side of the dissepiments,
winged at both ends, about 1.8 cm. long; hypocotyl lateral, horizontal.
ILLUSTRATIONS. BAILey, F. M., Comprehensive Cat. Queensl. Pl. ¢. 73
bis. 1913. Maten, J. H., Forest Fl. New S. Wales 1: 213. t. 39. 1904.
DIsTRIBUTION. Central Queensland south to southcentral New South
Wales; dry, rather open places. See Map 6
Queensland. BurKE District: 57 miles W of Hughenden, McCray, Septem-
ber 21, 1967 (BRI); Hughenden, Brass & White 63 (A, BO, BRI, K). MITCHELL
Everist & White 134 (srt); Aramac, Paulton (mEL); Barcaldine, Francis,
1969 | HARTLEY, THE GENUS FLINDERSIA Si?
March 1920 (srr); between Emerald and Longreach, Jarvis, October 1913 (srr);
Blackall, Bailey (Nsw 99620), Everist 1562 (a, BRI), White 12387 (A, K, US).
Grecory SoutH District: Thylungra, about 75 miles NW of Quilpie, Bverist
3787 (BRI). WarRrEGO District: about 19 miles N of Thargomindah, Sm
6346 (BRI); about 34 miles N of Charleville on Ward River Road, Smith 341
(A, BRI, K, MEL, NY); between Cunnamulla and Wyandra, Key, October 1940
(CANB); near Cunnamulla, White 11782 (Br1). MARANOA District: Maranoa,
Anonymous (MEL); Bollon Area, Epps, June 26, 1953 (BRI); 10 miles N of Dir-
ranbandi, Key, October 17, 1937 (cANB); Buckinbah, near St. George, Jones
215 (BRI): St. George, W edd, December 1893 (BRI); Nindigully District, Anon-
ymous, November 1938 (cans); Warrie, Nindigully, Allen A547 (CANB), Roe
R10 (cANB), October 1937 (cANB). DARLING Downs District: 36 miles W of
Goondiwindi on Talwood Road, Webb SN 5319a (cans), SN 5319b (CANB),
SN 5319c (cANB), SN 5319d (cans); Goondiwindi District, Jones C50 (CANB),
Webb 1611 (CANB), 2498 (CANB). New South Wales. Yantara Lake, Anonymous
385 (MEL); E of Broken Hill, Pidgeon & Vickery, August 1939 (Nsw 99617);
Koonenberry Mountains, ca. 62 miles SSE of Milparinka, Constable 4609
(Nsw 64979), V. E. Expedition (MEL); Mount Hope Station, 3 miles N of
White Cliffs, Constable 4595 (Nsw 67368); Wonominta River, ae Jan-
uary 1887 (MEL): Wilcannia, Johnson 547/90 (Nsw 5181), Kennedy, 1886
(MEL); about 12 miles S of Wilcannia, Hogan, October 1955 (MEL); between
Wilcannia and Cobar, Campbell 109 (cans); 30 miles E of Wilcannia, Riek &
Common 322 (cANB); above Morinda (Menindee) and Mt. Murchinson
(Murchiston), Anonymous (MEL); Menindee District, Neila-garri Station,
Constable, November 20, 1947 (Nsw 4879); Mt. Murchinson, Anonymous (MEL),
Dallachy (Bm, GH, K, NY), Dallachy & Goodwin (MEL); Darling River, Anony-
mous (MEL), Dallachy (pm), Kennedy, 1866 (MEL); about 1 mile E of Ivan-
hoe, Johnson, May 6, 1955 (Nsw 99615); Mossgiel, Bruckner, October 1885
(mex), M ueller (GH); between the Lachlan River and Darling River, Bruckner,
1885 (MEL), Mueller (p); Lachlan River, Tucker (met); Upper Lachlan River,
Curran 21 (MEL); 11 miles from Ivanhoe on Paddington Road, Whaite 1388
(Nsw 99616); between Kirriby and Lauradale, Warego River to Paroo River,
Boorman, October 1912 (srt, NSW 99619, Us); near Morton Boolka, Morris
766 (Nsw 99618); Barringun, Foyster, 1884 (MEL); Bourke, Betche, Novem-
ber 1887 (Nsw 99623), McDougall, 1901 (Nsw 99613), Wuerfel, 1884 (MEL);
Clover Creek, Bourke, Mackay 112 (MEL); 20 miles SE of Bourke, Riek &
N
99632); Dunlop Station, ca. 8 miles S of Louth, Etheridge 28 (Nsw 99622);
Cobar, 4 mile, Wilcannia Road, Forestry Officer 24 (Nsw 99629); between the
Bogan and Darling Rivers, Morton 77 (met); Bogan River, Anonymous (MEL);
Bogan, Morton, 1880 (MEL); W of Nevertire, Clarke, June 14, 1944 (CANB);
Jackson, October 1911 (Nsw 99628); Balonne River, St. George’s Bridge (ca.
25 miles W of Mungindi), Mitchell, November 1846 (k-holotype of Elaeoden-
dron maculosum Lindl.: Nsw 99612-isotype); Brewarrina, Boorman, Novem-
ber 1903 (GH, NSW 99630, ny, us); Yarrawin Station, Barwon River, 30 miles
SE of Brewarrina, Froggatt 16 (nsw 99627); Carinda, ca. 40 miles SW of
Walgett, Bucknell & Lowe, August 1965 (Nsw 99631); Pilliga, Rupp 13 —
99625); Pilliga Scrub, Anonymous, November 1932 (MEL); between Pilliga an
Wee Waa, Bassett, November 14, 1947 (Nsw 99626); plains near Baradine
Forsyth, October 1899 (NSW 99624); Ellenborough Falls(?), Boorman, 1904
(A, P, us); Murrumbidgee River, Bennett 3 (mEL); Cobar to Riverina Dis-
518 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
tricts, Anonymous (MEL); Riverina, Morton, 1880 (MEL). Without definite
locality. Bidwill 74 (k), Anonymous (NSW 99614).
15. Flindersia ifflaiana F. Muell. Frag. Phytogr. Austral. 10: 94. 1877.
Type: Hill, Queensland, Trinity Bay.
Flindersia brachycarpa Merr. & Perry, Jour. Arnold Arb, 20: 332. 1939. Type:
Brass 8389, Papua, Wassi Kussa River.
Medium trees to 35 m.; outer bark gray-brown, thick, deeply longitudi-
nally fissured; inner bark red, grading to cream toward the cambium;
sapwood yellow or yellow-brown; heartwood yellow-brown or brown;
branchlets, inflorescences, and capsules glabrous to pubescent with mostly
minute, predominantly stellate trichomes. Leaves opposite, paripinnate,
15-34 cm. long; rachis glabrous; petiolules 4-9 mm. long; leaflets 3-6
pairs, chartaceous to subcoriaceous, densely pellucid-dotted, glabrous,
ovate-elliptic, elliptic, elliptic-lanceolate, or occasionally lanceolate, usually
unequal-sided and often subfalcate, 6-13.5 cm. long, 2.8-5.2 cm. wide,
base rounded to subcuneate, often slightly oblique, main veins 14-22 on
each side of the midrib, apex rounded to narrowly obtuse, rarely subacu-
minate. Inflorescence terminal or terminal and upper-axillary, 14-25 cm.
long, usually as wide as long or somewhat wider, axes and branches
glabrate to appressed- or short-pubescent. Flowers bisexual or (a few to
many flowers in an inflorescence) functionally staminate, 2.5-3 mm. long;
pedicels 0.5-2.5 mm. long; sepals sparsely appressed-pubescent, ciliolate,
ovate-triangular, 1—1.2 mm. long; petals white, sparsely appressed-pubes-
cent abaxially, glabrous or with a few papillae adaxially, elliptic to
elliptic-oblong, 2—2.5 mm. long; stamens inflexed apically, about 1 mm.
long, filaments glabrous, anthers subdorsifixed, bluntly mucronulate, about
0.5 mm. long; staminodes 0.5 mm. long; disc about 0.7 mm. high; gynoe-
cium in bisexual flowers about 1.5 mm. high, about 1 mm. wide, ovules
2 on each side of the placentae; gynoecium in functionally staminate
flowers poorly differentiated, turbinate, about 0.5 mm. high, without ovules.
Capsule comparatively woody and heavy, separating to one-half or more
of the length but not completely, rounded short-cylindric, 3.2—-5.5 cm.
long; exocarp drying blackish brown to reddish brown, densely and
minutely pubescent, often cracked with age, muricate, the excrescences
rather widely spaced, to 2 mm. long; endocarp reddish brown. Seeds 2 on
each side of the dissepiments, winged at the apical end only, 2.7-3.3 cm.
long; hypocotyl lateral, ascending.
ILLUSTRATION. FRANcis, W. D., Australian Rain-forest Trees 426. 1951.
DistRIBUTION. Southern Papua south to the Atherton Tableland,
Queensland; rain forests to 400 meters. See M .
Papua. WESTERN District: Tarara, Wassi rege oe 8389 (a-holo-
type of Flindersia brachycarpa Merr. & Perry; BRI, L, LAE-is s); Oriomo
Creek, mouth of Yakup Creek, 40 miles from sea, yy ie ON GF 17728 (A,
Map 7. Distributions of Flindersia ifflaiana F. Muell. (dots) and F. australis
R. Br. (half-filled circles).
BO, K, L, LAE, NSW 99667). Queensland. Cook District: Cape York Peninsula,
Cape Grenville, Y oung 64 (prt); Endeavour River, Parsieh(?) (MEL); moun-
tains near Mossman, Rosenstrom (prt); Great Dividing Range ca. 6 miles S
Of Mossman, Smith 3964 (prt); Mt. Molloy, Crothers, 1934 (A, BO, BRI, NY);
Barron River, Anonymous 46 (MEL); Forestry Reserve 1073, N of Kuranda,
Dansie 1995 (srt, K), 1996 (BRI), 2008 (srt); Timber Reserve 315, Smith-
field, Kuranda, Doggrell A36 (sri); Black Mountain near Kuranda, Jones 800
(CANB), 1511 (cANB), Webb 5189 (cANB); a few miles N of Kuranda, Smith
520 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
5309 (BRI); Trinity Bay, Hill (met-holotype of Flindersia ifflaiana F. Muell.;
K-isotype); Cairns, Bailey (Nsw 99684); Freshwater Creek, Bailey (BRI);
Kamerunga, Cowley 43 (prt); Edge Hill, near Cairns, Morris North Queensl.
Nat. Club 4084 (srt); Redlynch, near Cairns, White 12813 (A, Ny); Atherton
Area, Webb 2547 (cANB), 5208 (CANB). Without definite locality: Anonymous
10 (rt), Wood Technology Dept. Queensland Forest Service 91 (A, NY). Cul-
tivated. Queensland: Cook District, Cairns, on esplanade, White, February
1918 (BRI).
In the original description of Flindersia brachycarpa, Merrill and Perry
stated that it strongly resembled F. ifflaiana but differed “.... . in
having larger leaves with the leaflets strongly oblique at the base and
more acute at the apex, and a little larger fruits.” These have proven to
be quite variable characters in many species of Flindersia and now, with
additional collections available, it is evident that they do not serve to
distinguish F. brachycarpa from F. ifflaiana.
As is pointed out above, Flindersia ifflaiana is apparently more closely
related to F. australis than to any of the other species of the genus. The
two do not appear to be particularly closely related, however. The leaves
of F. iffaiana are opposite and paripinnate whereas those of F. australis
are usually alternate and basically imparipinnate. Additional differences
are given in the key to species.
16. si Seg australis R. Br. in Flinders’ Voyage 2: 595. t. 1. 1814.
Type: Brown, September, 1802, Queensland, Broad Sound.
Medium trees to 25 m.; outer bark gray to brown, smooth or with
shallow longitudinal fissures, often exfoliating in flakes; inner bark reddish;
branchlets, leaves, inflorescences, and capsules glabrous to pubescent with
mostly minute, predominantly stellate trichomes. Leaves alternate, sub-
opposite or opposite, in mature growth closely crowded at the branchlet
apices, imparipinnate or (occasional leaves) paripinnate, (5.5—)9-34 cm.
long; rachis glabrous to densely appressed-pubescent, near the base often
narrowly crisped-winged laterally; petiolules of lateral leaflets obsolete to
3(—5) mm. long, terminal leaflet sessile or on an extension of the rachis
to 3.2 cm. long; leaflets 2-4(—6) pairs, chartaceous to subcoriaceous, with
scattered pellucid dots, glabrous to densely appressed-pubescent below,
glabrous or sparsely short-pubescent on the midrib above, broadly to
narrowly elliptic to occasionally lanceolate, equal- or slightly unequal- sided,
(2.4-)3-12 cm. long, (0.8—)1.5-4.3 cm. wide, bases of lateral leaflets
obtuse to cuneate, base of terminal leaflet sbiuke to attenuate, main veins
8-18 on each side of the midrib, apex obtuse to acute or subacuminate,
occasionally rounded. Inflorescence terminal or terminal and upper-axil-
lary, often on leafless branchlets, to 15 cm. long, about as wide as long,
axes and branches rather sparsely to densely pubescent. Flowers bisexual
or (a few to many flowers in an inflorescence) functionally staminate,
6.5-7.5 mm. long; pedicels 0.2-1 mm. long; sepals densely pees
pubescent, pee ciliate, ovate-triangular, 2.2-2.5 mm. long; petals
1969] HARTLEY, THE GENUS FLINDERSIA 521
white to cream, densely appressed-pubescent (except for the margins)
abaxially, sparsely short-pubescent in the basal one-half to two-thirds
adaxially, elliptic-oblong, 6-7 mm. long; stamens declinate, 3.5—4.5 mm.
long, filaments glabrous or with a few papillae and/or crisped trichomes
subapically, anthers subdorsifixed, mucronate, 1—1.3 mm. long; staminodes
-5 mm. long; disc in bisexual flowers rather thin, about 1.5 mm. high;
disc in functionally staminate flowers comparatively thick, about 1 mm.
high; gynoecium in bisexual flowers about 3 mm. high, and 1.4 mm_ wide,
ovules 2 on each side of the placentae; gynoecium in functionally stami-
nate flowers poorly differentiated, conical, about 0.6 mm. high, without
ovules. Capsule comparatively woody and heavy, separating to one-half
or more of its length but not completely, rounded short-cylindric, 4.6—9
cm. long; surface of exocarp drying blackish, reddish brown, or pale
brown, densely and minutely pubescent, muricate, the excrescences rather
densely crowded, often recurved, to 10 mm. long; inner part of exocarp
very rough where exposed by dehiscence; endocarp cream, yellowish, or
pale brown. Seeds 2 on each side of the dissepiments, winged at the apical
end only, 3.4~5 cm. long; hypocotyl lateral, horizontal.
ILLUSTRATIONS. Brown, R., ibid. FRANcis, W. D., Australian Rain-
forest Trees 158, 159. 1929; 176, 177. 1951. Mawen, J. H., Forest FI.
New S. Wales 2: t. 67 & 68. 1905.
DistripuTion. Eastcentral Queensland south to northeastern New
South Wales; rain forests and rather dry thickets. See Map
Queensland. SoutH KENNEDY District: Pinevale via Mackay, Webb &
Tracey 3319 (prt), LetcHHARDT District: Lake Elphinstone, Dietrich (MEL);
Morambah Homestead, 43 miles SW of Nebo Township, Story & Yapp 121
(BRI, K, MEL); 4.6 miles SW of Duaringa, Speck 1670 (cANB); Coomooboolaroo,
Berney 1919 (prt); Expedition Range, Byerly (MEL); 36 miles WSW of Theo-
dore Township, Sean 6928 (BRI, K); 18 miles N of Taroom, Speck 1861
(BRI, CANB, K). Port CurTIs weiagse Broad Sound, Brown, September 1802
(k-holotype: BM, MEL, P-isotypes); r Ogmore, ca. 75 miles NW of Rock-
hampton, Smith, ‘October 18, 1951 as hea ae Thozet (MEL). BURNETT
District: Eidsvold, Bancroft, April 1912 (prt). Wipe Bay District: Kolan
River, Smith’s Crossing, ca. 14 miles WNW of Bundaberg, Smith 4159 (BRI);
Maryborough, Clemens, October 27, 1948 (BRI, GH, MIcH); Mt. Bauple, Clem-
ens, June 10-20, 1945 (micH); Imbil, McAdam 84 (A, BRI); Cooroy, Douglas,
November 2, 1962 (BRI). DARLING sina District: Chinchilla, Beasley 51
— Moreton Disreict: Yarraman, nig August 29, 1949 ens Coal
ford, Meebold 7999 (nv); Brisbane River, (cae ee 60 (K), Mueller, July,
1855 hoor dowd November 1917 4 NSW 99565); tema Creek, near ue
572 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Law (MEL); Unumgar, Jones 2367 (cANB); Murwillumbah, Charles 458.04
(nsw 99562); Whian Whian State Forest, Webb & Tracey 378 (CANB), 1953-
1958 (BRI); Rous, near Lismore, Cheel, September 1926 (Nsw 99560); Rich-
mond River, Cameron (MEL), Fawcett (MEL), Henderson (MEL), Hodgkington
(mEL); Clarence River, Moore (met), Beckler (MEL); Kempsey, MacDonald,
October 3, 1895 (MEL). Cultivated. Queensland: Brisbane Botanic Gardens,
Bailey, November 11, 1884 (srt), Blake 2797 (prt), Hubbard 4745 (x), White
2442 (kK). New South Wales: Sydney Botanic Gardens, Boorman, October 1920
(Ny, us), November 1920 (Nsw 99563).
EXCLUDED NAMES
FLINDERSIA GREAVESII C. Moore, Cat. Nat. Industr. Prod. New S. Wales
53. 1861
This name appeared in the “indigenous woods” section of a catalogue
that accompanied an exhibit. Several other timber species were listed, each
represented, apparently, by a wood sample. The description follows:
A magnificent tree, the monarch of the northern forests, attaining a height of
150 feet, 3 to 6 feet in diameter, distinguishable from every other species of
the genus by its dark brown and rough scaly bark, as well as by other charac-
ters; timber used for house building purposes. Mountain brushes on the Clar-
ence [River, New South Wales].
The wood sample has apparently been lost, but an herbarium specimen
at Sydney (Nsw 99604), labelled Flindersia greavesii by Moore and
bearing the number 63, which corresponds to the number of that species
in the catalogue, could be designated as the type collection.
The only really diagnostic statement in Moore’s description is that per-
taining to the bark, which would almost certainly apply to Flindersia
australis. The specimen mentioned above, however, clearly matches F.
schottiana (published a year later), a species with relatively smooth bark.
Maiden (Forest Fl. New S. Wales 2: 152. 1905) listed Flindersia
greavesit as a synonym of F. australis and also designated it as a nomen
nudum. He concluded, as I have, that the description referred to FP. aus-
tralis and that the type collection referred to F. schottiana. In addition,
he pointed out that a tree in the Sydney Botanic Gardens, labelled by
Moore as F. greavesii, is really F. australis and that Mr. W. A. B. Greaves,
after whom the tree was named, gave him a fruit which he stated was that
of F. greavesii and that it actually was F. australis.
Moore must have realized his error, too, since he did not mention Flin-
dersia greavesii in his Census of the plants of New South Wales (1884)
or in his Handbook of the flora of New South Wales (1893). Flindersia
australis and F. schottiana were both mentioned in these publications.
Although I do not think Flindersia greavesii can be discounted as a
nomen nudum, I think this name should be excluded since the description
and the type collection clearly refer to two different species.
1969} HARTLEY, THE GENUS FLINDERSIA 523
FLINDERSIA PAPUANA F. Muell. Descript. Notes Papuan Pl. 1(5): 84.
1877. Type: D’Albertis, Fly River, Papua (not seen).
Mueller named this species from a single immature fruit and stated that
it was “. . . just a temporary name for the Papuan plant to place it on
the record until foliage and flowers can be obtained.” Thus it is a pro-
visional name and is not validly published.
LITERATURE CITED
Atry-SHaw, H. K. Diagnoses of new families, new names, etc. for the seventh
edition of Willis’ “Dictionary.” Kew Bull. 18: 249-273. 1965.
BalLey, F. M. Meliaceae. Queensl. Fl. part 1. 225-243. 1899.
BenTHAM, G., & F. Muetter. Meliaceae. Fl. Austral. 1: 378-390. 1863.
CANDOLLE, C. DE. Meliaceae. Monogr. Phanerog. 1: 399-752. 1878.
ENGLER, A. Rutaceae. Nat. Pflanzenfam. III. 4: 95-201. 1896.
. Rutaceae. Nat. Pflanzenfam. ed. 2. 19a: 187~358. 1931.
ErDTMAN, G. Pollen morphology and plant taxonomy. Angiosperms. xii +
539 pp. Chronica Botanica, Waltham, Massachusetts. 1952.
Francis, W. D. Australian Rain-forest Trees. xi + 347 pp. Commonwealth of
Australia. 1929
Australian os forest Trees. xvi + 469 pp. Commonwealth of Aus-
tralia. 1951.
Harrar, E. S. Notes on the genus Flindersia R. Br. and the systematic anat-
om e important flindersian timbers indigenous to Queensland. Jour.
Elisha Mitchell Soc. 53: 282-293. 1937
Hartiey, T. G. A revision of the Malesian species of Zanthoxylum (Ruta-
ceae). Jour. Arnold Arb. 47: 171-221. 1966
. A revision of the genus Lunasia (Rutaceae). Jour. Arnold Arb. 48:
460-475. 1967
Mercatre, C. R., & L. CHALK. Anatomy of the Dicotyledons 1: Ixiv + 724
pp. Clarendon Press, Oxford.
Price, J. R. The distribution of alkaloids in the Rutaceae. In: Chemical
Plant Taxonomy, T. Swatn, Ed. 429-452. Academic Press, London and
New York. ,
Ritcuir, E. oe of Flindersia species. Rev. Pure and Appl. Chem. 14:
47-56.
SMITH- a : Chromosome numbers in the Boronieae (Rutaceae) and
their bearing on the evolutionary development the tribe in the Aus-
tralian flora. Austral. Jour. Bot. 2: 287-303.
WELsH, M. a A ctsinsintae “maple” (Flindersia ah Sone, Woods No. 25:
18-23. 19
Waite, C. T. tes on the genus Flindersia (Rutaceae). Proc. Linn. Soc.
New S. Wales 46: 324-329. 1921.
Wittaman, J. J., & B. G. Scuusert. Alkaloid-bearing plants and their con-
ta ined alkaloids. US. Dept. Agr. Tech. Bull. No. 1234. 247 pp. 1961.
524
JOURNAL OF THE ARNOLD ARBORETUM
[voL. 50
INDEX TO EXSICCATAE
text
Adams, 1281, 1317 (13)
Allan & Jones NGF 2751 (6)
Allen A547 (14)
Baeuerlen 244, 350 (11); 633 (8);
698, - (11)
Bailey 10 (8)
Balansa 163, ag (1)
Barnard 31
Beasley 51 eae ge (it)
Beckler 7620 (13 & 14)
Bennett 3
Bidwill 74 oe 95 (8)
Bisset E198
Blake 2688 (8) 2797 (16); 3079
(10); 14906 (13)
Boorman 15 (12)
Bowman 70
Brass 5339 (4); 5565, 7517 (6); 7991
(8); 8032 (6); 8389 (15); 8495,
8542 (2b): 8634 (6); 20322 (4);
28881, 28906 (2b); 29153 (4)
Brass & — i 12535 (4)
Brass & White 6.
Briggs 75.05F @)
Cameron 93 (11)
Campbell 0109 (14)
Carr 13152 (4); 13913 (6); 14408,
14808, 14910, 15475, 15569, 15805,
15969, 15989 (4)
Charles 458.04 (16)
Clemens 4739, 5086, 6825 (4): 43213,
43373 (3); 43641 (12)
Collins W966 (4)
Constable 4595, 4609 (14); 6505 (8)
Cowley 43 (15
unningham 18 (12); 60 (16); 109,
117 (10); 164 (12)
Dansie 1995, 1996, 2008 (15)
Darbyshire 1164 (8
obson & Havel NGF 9116 (6)
Docters van Leeuwen 10478, 10618,
10682 (4)
The numbers in parentheses refer to the corresponding species and varieties in the
Doggrell A35 (3); A36 (15)
Dreghorn 11 Ai i2 (9); 13, 14 G@):
20E (7); )
Dunn 252 a
Eddowes NGF 13086 (2b)
Epps 30 (11); 31 (8)
Etheridge 28 (14)
Everist 1562, 5787 (14)
Everist & Webb 1413 (12)
Everist & White 134 (14)
Floyd NGF 7472, NGF 7527 (4)
Floyd & Morwood NGF 6204 (4)
Forbes 421 (6)
Forestry Department 1, 2, 3 (8);
237, 132, 433 (it)
Forestry Officer 24 (14)
Fournier & Sebert 22 (1)
Franc 1738, =. 1956, The ‘comparative anatomy of the Flagellariaceae. Kew
ull. 11: 491-501.
SOLEREDER, H., & F ._ J. Mever. 1929. Systematische anatomie der mo
donen. Vol. 4. Farinosae. Gerbriider Borntraeger, Berlin. of, ee
Wawra, H. 1875. Beitrag zur Flora der Hawai’schen Inseln. Flora
nN
5
oD hy
77
o
4
DEPARTMENT OF BOTANY
ARIZONA STATE UNIVERSITY
TEMPE, ARIZONA 85281
556 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
THE ECOLOGY OF AN ELFIN FOREST IN PUERTO RICO, 9.
CHEMICAL STUDIES OF COLORED LEAVES
RICHARD J. WAGNER, ANSTISS B. WAGNER, AND RICHARD A. HOWARD
ONE OF THE SPECTACULAR FEATURES of tropical vegetation is the fre-
quent occurrence of brightly colored leaves. The color may be found
in young foliage, produced either regularly or seasonally, or in the ma-
stroyed. Red color may also be a pathological symptom frequently
associated with phosphorous deficiency. Macmillan (1952) stated ‘‘antho-
cyanin may appear temporarily in the young leaves, and, if abundant be-
fore the chlorophyll is largely developed, a bright red immature foliage
results. This is very evident in many tropical trees, e.g. Mesua ferred,
a species of Calophyllum, Eugenia, Cinnamomum etc. The coloration is
at times so vivid that from a distance such trees appear to be in flower.”
Anthocyanins are usually red in acid solution and may become purplish
to blue as the pH is increased. A previous paper (Howard, 1969) has
shown that the pH of the plant sap of the component species of the elfin
forest on Pico del Oeste ranges from 2.5 to 6.5. Anthocyanins are often
associated as well with the occurrence of sugars in the plant cells and, in
temperate areas, with the occurrence of frost or low temperatures. Crock-
er (1938) stated “anthocyanins appear in many plant cells mainly in
the early spring and in autumn at times of low temperatures; under these
conditions soluble sugars are also abundant in plant organs. Arthur finds
that low temperature favors the development of anthocyanin in the apple
without a change in sugar content. He also points out that the small
amount of pigment found in cells calls for relatively little sugar as a build-
ing material and concludes that temperature probably acts directly rather
than through sugar accumulation.” More recent work has shown an as-
sociation of color due to anthocyanin with a shortened photoperiod and
suggests that it may be regulated by a phytochrome system. Finally it
is evident to anyone familiar with a fall season in New England that po-
tential for the development of color is also inherent in certain plants.
The elfin forest on Pico del Oeste displays a localized brilliance in the
vivid colors of young leaves and in the flush growth of many of the
woody components. The development of color with age was noted only
1969 | WAGNER, WAGNER, & HOWARD, ELFIN FOREST, 9 557
in Miconia pachyphylla, Mecranium amygdalinum, and Calycogonium
squamulosum. Perhaps the greatest year-round color, however, is found
in many plants of the bromeliad genus Vriesea.
We were not able to attempt tests of the ratio of red to far-red light
as a factor in the leaf color we observed in the various plants of the
cloud-dominated environment. We were able, however, to establish a
small laboratory through the courtesy of Mr. Joseph Martinson, in which
we attempted simple tests to examine the chemical bases previously pro-
posed for the color we observed. Tests were run to determine the sugar
content of young and old leaves, and of the red- and green-leaved forms
of Vrieseas, and that of the water soluble phosphorus.
The epiphyte Vriesea sintenisii occurs throughout the forest: on the
branches of isolated trees, on upper branches of trees forming the canopy,
with many young plants on the horizontal branches or, occasionally, on
the ground in cut-over areas. The plants that are exposed to the sky
exhibit a brilliant red color. Within the forest, on shaded branches, and
frequently on the ground other plants of the same species lack the red
color and are pale green in appearance. Although the color difference is
intense, the red plants may have the leaf bases green within the rosette
but very few plants could be truly called intermediate, that is, partly
green and partly red. The principal variation is in the intensity of red.
tips while the protected or shaded seedlings are all green. The intensity
of color does not appear to vary throughout the year or to suggest a
photoperiod variation. However, the maximum variation in daylength
during the year in Puerto Rico is only 2 hours and 18 minutes.
Gleason and Cook (1927) do not mention Vriesea sintenisi in their
description of elfin forest types, and previous workers on the family or
on the flora of Puerto Rico have not described the color variations or
suggested any taxonomic value for them. Our initial encounter with
these two color forms of Vriesea sintenisii suggested that two ecotypes
were present.
The brilliance of the red form of Vriesea sintenisii suggested that the
plants might make attractive ornamentals. However, when bright red
plants were taken to a lower elevation within the Luquillo Mountains
and placed in an area with less cloud cover, the plants died even when
supplied with water daily. Red color forms which were returned to Bos-
ton and kept in the greenhouse under high humidity, without any ad-
justment of the natural photoperiod of the Boston area, retained their
red color but failed to flourish, remaining in a vegetative state long after
plants of comparable size on Pico del Oeste had flowered, shed seeds, and
reproduced vegetatively. Red-colored plants taken from the exposed
tree tops and placed on the forest floor in the shade gradually lost their
red color and by the time of flowering were almost completely green.
The green-colored plants were also subjected to transplant experiments.
These, too, failed to survive when transferred to a lower location, while
558 JOURNAL OF THE ARNOLD ARBORETUM [vor. 50
those transported to the greenhouse in Boston grew well in the new loca-
tion and flowered on schedule when compared with plants on Pico del
Oeste. Green-colored plants within the elfin forest were also transferred
from their protected positions to exposed positions by strapping branches
holding these plants to upper branches of the forest canopy. These green
plants died in the exposed positions. Unfortunately we were unable to
attempt a gradual transfer of these plants from one position to another.
Clearly the green plants were physiologically adapted to shaded location
and could not survive an abrupt although seemingly slight change in ex-
posure to greater light. The red plants became adapted to the shaded
location with the apparent loss or masking of the red pigment.
A second bromeliad found in the elfin forest was Guzmania berteroniana.
Plants of Guzmania occurred primarily on the trunks of Prestoea mon-
tana or on the ground in protected areas on the lee slopes of the trail.
All plants of Guzmania were green and no red forms were seen. Trans-
plant experiments produced results nearly comparable to those for green
forms of Vriesea. Plants taken to lower elevations died; those taken to
Boston have persisted, but the rate of maturation was slower; plants which
were transferred to exposed locations in the canopy died even more
quickly than did the green forms of Vriesea.
When initial sugar tests suggested a higher sugar level in red-leaved
plants of Vriesea, we continued a comparative study through two years,
making analyses of plants in various stages of development.
In each test an average of eight plants was collected in the Pico del
Oeste forest and taken to the laboratory. The leaves were all separated
and washed thoroughly in running water and hand dried with towels.
Roots and rhizomes were discarded. After the fresh weight of the leaves
was obtained for each plant they were oven dried at 70°—75° for two days.
When the dry weight was obtained the leaves were finely cut, and after
a thorough mixing of the fragments of the individual plants, 1 gm. of
dry leaves was placed in an Erlenmeyer flask; 100 cc. of distilled water
was added; and the mixture was simmered for 30 minutes. When cooled
to air temperature, distilled water was added to regain the original
volume of 100 cc. The solution was allowed to mix for about two hours,
then the sugar content was estimated quantitatively, following the
colorimetrical method of Folin and Wu (1920). In a second Folin-Wu
tube 0.1 cc. of 1:10 diluted HCl was added and the tube submerged for
5 minutes in a boiling water bath to hydrolize the higher sugars to a
hexose before a second colorimetric sugar determination was made. The
results are expressed in the following tables as percentages of sugar per
gram of dry weight of the plant, and are the average of the eight plants
of each sample category. There was no significant variation between the
eight samples.
Soluble phosphorus determinations were made from 2 grams of the
dried material which was diluted with 100 cc. of distilled water and sim-
mered at 90°C. for 30 minutes; this was cooled to air temperature and
distilled water was added to regain the original weight. The soluble
1969] WAGNER, WAGNER, & HOWARD, ELFIN FOREST, 9 559
phosphorus was determined according to the procedure of Benedict
and Theis (1924): 10 cc. of the extract and 10 cc. of a standard were
mixed with 1 cc. of a 5% hypochlorate solution to decolorize the brown-
ish mixture; after 24 hours the solution was filtered and 5 cc. of the
standard and the unknown were measured in test tubes; the results are
expressed as mg.% of dried plant material.
1966 RED-COLORED VRIESEA
COLUMN NO. 1 Z 3 4 5 MEAN
WATER 83% 83% 83% 81% 80% 82%
HEXOSE 1.92% 1.88% 1.88% 1.44% 1.51% 1.73%
TOTAL SUGAR 2.27% 1.96% 2.05% 1.72% 1.96% 1.99%
1966 GREEN-COLORED VRIESEA
WATER 86% 85% 86% 85% 82% 84.8%
HEXOSE 1.27% 1.32% 1.29% 1.09% 1.36% 1.26%
TOTAL SUGAR 1.56% 1.82% 2.36% 1.15% 2.06% 1.79%
MNS: 1. Young plants in vegetative rosettes. 2. Inflorescence present, basal
flowers open. 3. Inflorescence mature, basal flower in young fruit stage. 4. Fruit
forming, seeds turning black. 5. Fruiting stage, seeds mature.
1967 RED-COLORED VRIESEA
COLUMN No. 1 2 3 MEAN
WATER 80% 85% 80% 82%
HEXOSE 1.5% 1.9% 1.8% 1.7%
TOTAL SUGAR 1.8% 2.1% 2.0% 2.0%
SOLUBLE PHOSPHORUS 23mg% 30mg% 21mg% 24mg%
1967 GREEN-COLORED VRIESEA
WATER 83% 86% 83% 84%
HEXOSE 1.2% 1.5% 1.4% 1.4%
TOTAL * SUGAR 1.4% 1.7% 1.6% 1.6%
SOLUBLE PHOSPHORUS 26mg% 24mg% 25mg% 25mg%
Corumns: 1. Young plants. 2. Inflorescence mature. 3. Fruit mature.
GUZMANIA BERTERONIANA
CoLUMN No. 1 z 3 : MEAN
WATER 86% 85% 87% 88% 86%
HEXOSE 1.0% 1.1% 2.0% 1.2% 1.3%
Tor 1.3% 1.3% 2.2% 1.2% 1.5%
SOLUBLE PHOSPHORUS 24mg% 25mg% 25mg% n.e. 25mg%
Cotumns: 1. Young vegetative rosettes. 2. Mature vegetative plants. 3. Inflores-
cence mature. 4. Fruits mature.
The water content of the leaf tissue of the components of the elfin
forest has been given in a previous paper (Howard, 1969). Tissues tested
contained from 93 percent water to 44 percent water. Vriesea ranked in
560 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
the lower portion of the upper third of all the plants tested in water
content. All the plants tested which revealed a higher water content
were understory plants. The red color-form of Vriesea, being primarily
a canopy epiphyte had a slightly lower water content than did the green-
colored Vriesea which occurred in protected locations. The water content
of the plant tended to diminish as the plant matured and was lowest when
the plant was in fruiting condition. The total sugar in the green-colored
form increased to the flowering period, decreased during the development
of the fruit, but increased as the fruit matured
Hexose and total sugar content proved to average higher in red-
colored Vrviesea and the total sugar content was approximately 40 per-
cent higher in the red than in the green forms during the growing season.
In Guzmania the water content averaged higher than in Vriesea, in-
creasing slightly in the life cycle to the fruiting stage. Hexose and total
sugars increased in Guzmania during the development of the inflorescence
and was 100 percent higher in the flowering cycle, exceeding the red
form of Vriesea.
The amount of soluble phosphorus averaged about the same in the red
and green Vrieseas. However, it seemed to increase in the red Vriesea
during the flowering period and decrease in the green forms during the
same period.
The amount of soluble phosphorus is evidently not a limiting factor in
the coloration found in the red form of Vriesea sintenisii.
In Guzmania, soluble phosphorus levels remain fairly constant through
the life cycle and were approximately the same as those of Vriesea. Since
both bromeliads would receive inorganic materials as wind-blown debris
collected in the leaf bases directly, it appears the supply or utilization
of phosphorus is equally available to the plants no matter what their
position in the forest structure.
The distinctive coloration of young developing leaves in tropical forests
is mentioned in many previous studies. An earlier study (Howard, 1969)
indicated red, orange, pink, yellow, and bronze tones in the young leaves
of specific plants within the elfin forest of Pico del Oeste. Many taxa
which did not produce colored leaves when young did show a lighter
green or a yellow-green color to the young foliage. Tests for hexose
sugars and for total sugars were made on selected leaves from individual
taxa. One to four brightly colored leaves were taken, as available, from
a flush of growth and tests on these were compared with material from
older leaves of the same branch. The following results were obtained:
TAXON Color of young Percent Percent Percent
leaf water hexose sugar all sugars
yglf oldlf yglf oldlf yglf oldlf
Calycogonium
squamulosum pale red 75 74 4.3 2.6 a ee
Calyptranthes
krugit yellow-green 61 48 5.7 4.45 7.85 4.40
1969 | WAGNER, WAGNER, & HOWARD, ELFIN F OREST, 9 561
Cleyera
albopunctata yellow 70 «(57 Bos 875 8.35 8,90
Eugenia
borinquensis deep red 70 3a 562 530 5.70 8.05
Gonocalyx
portoricensis pink 89 75 2.8 z3 —_ —_
Hillia
parasitica light green 83 81 2.86 4.45 3.57 5.00
Hornemannia
racemosa orange-pink of ae 4.96 8.50 6.08 8.88
6
sintenisii yellow-green 70 63 7.05 7.30 9.55 8.00
Mecranium
amygdalinum yellow-green 69 70 coagulates copper reagent
Marcgravia
Sintenisit
juvenile leaves red 82 78 7.6 ; — —
86 = 81 7.20 8.84 7.90 8.86
adult leaves red 84 69 3.4 3.8 _— —
89 69 7.10 10.00 7.35 10.10
Miconia
pachyphylla red-purple 62 56 coagulates copper reagent
Miconia
foveolata red-pink 71 68 tek ° 373 782 67535
Miconia
pycnoneura red-purple 60 $3 coagulates copper-reagent
Ocotea
Spathulata bronze-red 62 68 7.7 6.7 ~ —
Tabebuia
rigida red-purple 83 70 6.4 3.7 — —
Torralbasia
cuneifolia yellow-green 71 63 8.80 9.55 8.84 10.89
Wallenia ;
yunquensis wine-red 75 63 9.80 7.85 10.00 8.90
With the exception of one plant of Marcgravia sintenisii and those of
Eugenia and Hornemania, all taxa tested which had red or orange, brightly
colored leaves had a higher hexose sugar content in juvenile foliage than
in mature foliage of the same plant. In the taxa which produced nage!
or pale green juvenile foliage, the hexose sugar content was lower
juvenile leaves than in mature leaves, with the exception of C, aie
562 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
krugii. For total sugars in yellow-green Wain leaves, the same ex-
ception occurs, with the addition of lex sinten
It is of interest to record that the plant saci obtained during the
month of April from four taxa of the Melastomataceae all contain some
chemical substances which form a gelatinous coagulate in the Folin and
Wu copper reagent. One sample of Calycogonium squamulosum, of the
same family, formed the coagulate in March but a second test in April
from the same plant did not. A similar reaction was experienced for Mz-
conia foveolata.
The contrast between the higher hexose and total sugar content in the
brightly colored juvenile leaves and that of the mature green foliage in
the woody plants is similar to that condition found in Vriesea sintenisit
when the anthocyanin-dominant plants were compared with the green
color-form.
VARIATIONS IN pH OF CELL SAP
The pH of liquid expressed from leaf tissues was mentioned in relation
to vulnerability to insect damage in an earlier paper in this series ( How-
ard, 1969). Where possible liquid was obtained from fully expanded
leaves and leaves normally green but which had not completely hard-
ened. The leaves were pressed between clean microscope slides, a drop
or two of expressed fluid collected, and the pH checked with a Beckman
pH meter. The amount of liquid and the ease with which it could be ob-
tained varied considerably between the plants of the elfin forest of Pico
del Oeste, and also varied with the time of year for individual plants.
The liquids varied in their color and consistency, and in the rate of color
changes upon exposure to air. The results of this survey are presented in
the following table:
Variations in the liquid extract of crushed leaves and in the pH of plant sap
Color or consis-
tency of sap Feb. July Nov.
CYATHEACEAE
Cyathea pubescens clear a 4.6 cat
Prestoea montana yellow-green 5.1 6.2 5.4
ACAE
Anthurium dominicense clear 5.0 5.5 5.8
BROMELIACEAE
Guzmania berteroniana rusty —_ 4.0 —
Vriesea sintenisii clear — 5.1 aS
DIOSCOREACEAE
Rajania cordata — 4.9
1969 | WAGNER, WAGNER, & HOWARD, ELFIN FOREST, 9 563
ZINGIBERACEAE
Renealmia antillarum _ 5.2 5 4.8
PIPERACEAE
Peperomia emarginella — 5H) 5.0 5.3
Peperomia hernandiifolia a 5.0 4.6 5.1
CHLORANTHACEAE
Hedyosmum arborescens orange-brown 4.5 5.3 5.6
MoracEAE :
Cecropia peltata — uk a we
URTICACEAE
Pilea krugii lilac Sa 5.1 6.0
Pilea yunquensis _— Sad 5.9 6.4
LAURACEAE j
Ocotea spathulata gelatinous ie | 4.9 4.9
ELIACEAE
Trichilia pallida dirty green 5.6 Pe 5.4
AQUIFOLIACEAE :
Ilex sintenisii — 5.3 Suh 5.1
CELASTRACEAE
Torralbasia cuneifolia — 5.0 4.8 4.7
OCHNACEAE es
Sauvagesia erecta yellow-green 4.9 4.9 ‘
MARCGRAVIACEAE 48
Marcgravia sintenisii _ 4.6 5.4 .
THEACEAE .
Cleyera albopunctata — 3.8 4.1 ;
GUTTIFERAE ‘i
Clusia grisebachiana yellow-orange 3.9 3.9 ;
BEGONIACEAE : -
Begonia decandra cherry-red 2.5 2.5 :
MYRTACEAE
Calyptranthes krugii — — Hs : -
Eugenia borinquensis dirty brown 4.8 4.
MELASTOMATACEAE F 4
Calycogonium squamulosum lavender 3.4 - 4)
Mecranium amygdalinum rose-purple g2 re 7
Miconia foveolata red-purple 3.9 =
Miconia pachyphylla rose 3.7 Bh oy
Miconia pycnoneura rose 3.3 ; ¢
ERICACEAE
Gonocalyx portoricensis — 3.3 .
Hornemannia racemosa — 3.9 .
564 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
MyrsINACEAE
Ardisia luquillensis dirty lavender 4.4 4.4 4.4
Grammadenia sintenisii } — 4.1 4.5 4.1
Wallenia yunquensis — 4.2 4.1 3.5
SAPOTACEAE
Micropholis garciniaefolia straw color 4.2 4.1 4.8
SYMPLOCACEAE
Symplocos micrantha blue-purple 4.0 4.2 4.8
OLEACEAE
Haenianthus salicifolius — | See 5-1
var. obovatus
CONVOLVULACEAE
Ipomoea repanda — 4.5 6.2 5.9
BIGNONIACEAE
Tabebuia rigida a 5.0 Sue 53
GESNERIACE
Alloplectus aes — 5.1 4.6 55
Gesneria sintenisii rust-brown 53 Sa 5.6
ACANTHACEAE
Justicia martinsoniana slimy 5.5 4.1 6.5
RUBIACEAE
Hillia parasitica lime-green 4.9 4.7 4.9
Psychotria berteriana — 5.0 a4 57
Psychotria guadalupensis brick-red 4.9 5.0 4.9
CAMPANULACEAE
Lobelia portoricensis lime-green 5.0 4.8 4.7
COMPOSITAE
Mikania pachyphylla _ 5 5.9 5.8
The leaves from which the liquid was expressed to obtain the records
in the preceding table were a normal green color. Where the color is
not indicated the liquid was pale green. Accessory pigments, however,
were present in several plants and were usually seen in the petiole; yet
there was no consistent correlation with the color of the plant liquid ob-
tained. The petioles and lower leaf surface of Peperomia hernandiifolia,
Wallenia yunquensis, and Tabebuia rigida were red-purple in appearance.
Miconia foveolata had red pubescence and Miconia pachyphylla ap-
peared to have an underlying tone of red-purple to the leaf blade. The
veins of Begonia decandra appeared bright red in fresh condition, and
the cherry red color of the expressed liquid suggests this pigmentation
was released in crushing. Psychotria guadalupensis exhibited a red pig-
*In a recent paper Lundell has established a new genus based on this taxon and
has made the combination Cybianthopsis sintenisii (Urb.) Lundell, "Wrightia 4: 68.
8.
1969] WAGNER, WAGNER, & HOWARD, ELFIN FOREST, 9 565
ment in the petioles and inflorescence axis but not in the leaf blade. The
brick red color of the liquid extracted was darker than the pigmentation
observed in the whole plant.
It was not possible to extract liquid by the hand pressure method used,
from any member of the Gramineae, Cyperaceae, or Orchidaceae.
The pH of the extracted liquid ranged from 2.5 in Begonia decandra
to 6.5 in Justicia martinsoniana. In three taxa tested in February, July,
and November, the pH was identical. For 12 taxa two identical readings
were obtained in the three unit test. The pH was highest in November
for 20 taxa, with high readings in July in 10 taxa, and in February for
6. In 16 taxa the lowest pH reading was obtained in February, in 9 taxa
during July, and for 4 taxa in November. In all cases except the her-
baceous species the material examined was taken from a single marked
plant for the tests. In a general observation the greatest number of
plants would be in flower in July, in fruit in November, and in dormant
condition during February.
The average pH of the expressed fluid of 11 taxa, in which the juvenile
leaves are predominantly red, is 4.6 in a range of 3.4 to 5.2. The com-
parable pH of 6 taxa, in which the young leaves are yellow or yellow-green,
is 4.7 with a range of 3.5 to 5.2. The average pH of the remaining 23
taxa of dicotyledonous plants tested, in which the young leaves are not
noticeably different in color from the mature foliage, is 4.9 in a range of
2.7 to 6.0. The color of the young leaves and the associated pH ap-
pears to be characteristic of the individual species, with no direct correla-
tion evident between the color of the leaf or of the expressed pigment,
the pH, or the sugar content.
LITERATURE CITED
Benepict, S.R., & R.C. THets. A modification of the molybdic method for
the determination of inorganic phosphorus in serum. Jour. Biol. Chem.
61(1): 63-66. 1924.
Crocker, W. Growth of Plants. pp. 320, 321. Reinhold Co. 1948.
Fo.tn, O., & H. Wu. A system of blood analysis. Supplement 1. A simplified
and improved method for determination of sugar. Jour. Biol. Chem.
41(3): 367-374. 1920.
Gieason, H. A., & M. T. Coox. Plant Ecology - Porto Rico. Pts. 1 and 2.
Sci. Surv: Porto Rico Virgin Is. 7: 1-173. a7.
Howarp, R. A. The ecology of an elfin forest in ita Rico. 8. Studies of
stem growth and form and of leaf structure. Jour. Arnold Arb. 50: 225—
262. 1969. ;
Macmitian, J. F. Tropical Planting and Gardening. p. 14. Macmillan & Co.
5th ed. 1952
ARNOLD ARBORETU
HARVARD ce
566 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
THE GENERA OF PORTULACACEAE AND BASELLACEAE
IN THE SOUTHEASTERN UNITED STATES *
A. LINN BocLe
PORTULACACEAE A. L. de Jussieu, Gen. Pl. 312. 1789, “Portulaceae,”
nom. cons. (PURSLANE FAMILY)
Fleshy or succulent herbs or subshrubs [shrubs or rarely small trees],
with erect to procumbent branches, reproducing vegetatively by means of
rhizomes, stolons, or axillary bulblets. Roots fleshy, basal or adventi-
tious, fibrous, tuberous, cormose, or plant with a simple to branched
taproot. Leaves alternate, opposite, or in basal rosettes, the blades en-
tire, cylindrical (or nearly so) to flat; stipules scarious, fimbriate, of
tufted hairs, or absent. Inflorescence terminal or lateral, basically cymose
but often appearing racemose or paniculiform, or flowers solitary in the
axils. Flowers perfect, regular, inconspicuous or often showy, erect,
spreading or nodding. Perianth biseriate (or uniseriate). Sepals (in-
volucral bracts?) 2 [4-8], imbricate, + equal, free or basally connate,
deciduous or persistent. Petals (tepals?) 4-6 (sometimes 2 or 3),
or basally connate, hypogynous or perigynous, often ephemeral. Stamens
few to many, alternate with and/or opposite the petals, free or inserted
on the corolla base; filaments filiform; anthers 2-loculate, dehiscing
longitudinally and introrsely. Gynoecium 2—9-carpellate, syncarpous;
styles as many as the carpels, + united, rarely simple, with linear to
capitate stigmas; ovary superior to half-inferior, or inferior, unilocular;
ovules many to few [—1], sometimes on long, ascending funiculi, ana-
tropous or amphitropous; placentation free-central or basal, the placenta
2—9-parted or 1. Fruit a pyxis or capsule [rarely an achene], dehiscence
*Prepared for a generic flora of the oe United States, a project of the
Arnold Arboretum and the Gray Herbarium of Harvard University made possible
through the support of the National Stiaas gomnernta (Grant GB-6459X, principal
investigator Carroll E. Wood, Jr.). This treatment follows the format established
in the first paper in the series (Jour. Arnold Arb. 39: 296-346. 1958) and continued
through those in volumes 40-SO (1959-1969). The area covered includes North and
South Carolina, nea Florida, Tennessee, Alabama, Mississippi, Arkansas, and
Louisiana. The escriptions apply primarily to the plants of this area, with supple-
mentary sageiageerie in brackets. References which the author has not seen are
marke d by isk.
I wish t eek Carroll E. Wood, Jr., for reading and editing the acne and
i N. Avery
nre
to R. B. Channell and S. Ware for living plants of Talinum calcaricum. The illus-
trations are the work of Sydney B. Devore, Diane C. Johnson, and Virginia B.
Savage.
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 567
circumscissile, or by 3 longitudinal valves. Seeds one to many, + round-
reniform, flattened. Seed coat often crustaceous, sometimes arillate,
smooth or variously sculptured (rugulate, granulate, muricate, tuber-
culate). Embryo curved to annular, enclosing the abundant mealy endo-
sperm; cotyledons (1—)2 [rarely 4]. Type Genus: Portulaca L.
A small family of 15 to 31 genera with centers of distribution in west-
ern North America, southern South America, and South Africa, but ex-
tending to eastern Siberia, Australia, New Zealand, and Madagascar.
Many of the genera are small or monotypic and restricted in distribution.
Only a few have more than about a dozen species (Anacampseros, Ca-
landrinia, Claytonia, Montia sensu lato, Portulaca, Talinum). Inter-
esting patterns of distribution are evident in several genera: Portulaca
and Talinum have attained wide distributions in the tropics and sub-
tropics of both hemispheres; Calandrinia has centers of development in
western North America, South America (Chile), and Australia; and
Montia, which is principally boreal in range, is represented in far flung
areas of the Southern Hemisphere.
De Candolle and Bentham & Hooker viewed the family in a narrow
sense, essentially as it is regarded today, including 12-15 genera but
no’ subfamilial categories, while Fenzl (followed by Endlicher) took a
much broader view of the family, recognizing seven tribes and including
many genera now placed in such related families as Basellaceae, Aizoa-
ceae, and Molluginaceae. Similarly Baillon established three series with-
in his Portulacaceae: Portulaceae, Aizoideae, and Mollugineae.
The system of the family established by Franz, which was based on
a wide range of morphological and anatomical characteristics, has been
followed with modification by Pax & Hoffman and by Eckardt. The
genera of Basellaceae originally included by Franz were removed, as were
some of doubtful affinity (Hectorella Hook. f., and Lyallia Hook. f.
which have recently been established as a family Hectorellaceae). Of
the two subfamilies now recognized in the Portulacaceae, the smaller
Montioideae Franz includes only Montia L., Claytonia L., and the mono-
typic Chilean Wangerinia Franz. Rydberg and, more recently, Nilsson
have split eight segregate genera out of the Claytonia-Montia complex,
but these are not universally recognized. The large subfamily Portula-
coideae consists of about 13 genera arranged in two tribes, each with
two subtribes. Of primary interest in our area are tribe Portulaceae,
subtribe Portulacinae, containing only Portulaca L., and tribe Calan-
drinieae, subtribe Calandriniinae, containing Talinum Juss. and four
related genera.
The Portulacaceae share several features with other Centrospermae,
including the presence of betacyanins (betalains) in place of anthocyanins,
curved to annular peripheral embryos, and basal placentation in a com-
und ovary. The family is considered closely related and possibly an-
cestral to the Basellaceae through such features as intraxylary phloem
(weakly developed in some Montioideae), uniovulate ovaries in ad-
568 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
vanced genera, a tendency toward unisexual flowers, and similar floral
plans. Relationships to the Caryophyllaceae, Chenopodiaceae, and Ama-
ranthaceae are seen in the morphology and anatomy of the seeds, and
to the Aizoaceae (through Sesuvium), from which the Portulacaceae
differ principally in characteristics of floral organization and the lack of
well-developed anomalous secondary growth. A relationship to the
Primulaceae, advanced particularly on embryological evidence (Gui-
gnard, cf. Talinum), has found little support.
Embryological studies in a few of the better known genera reveal
anatropous to amphitropous, crassinucellar ovules in which the inner
of the two integuments forms the micropyle. A Polygonum-type embryo
sac is formed from the chalazal cell of a linear tetrad. Endosperm for-
mation is initially nuclear, later becoming entirely cellular (or only
are tetrasporangiate, and pollen grains are generally three celled (rarely
two celled) when shed. Embryos with only one cotyledon are reported
in Claytonia virginica L., while four cotyledons have been found in Ana-
campseros lanceolata (Haw.) Sweet.
The pollen morphology of the family is diverse, ranging from tricolpate
to rugate (with several to many colpi distributed evenly over the sur-
face), to forate (with many evenly distributed round pores). The sur-
faces of the grains usually bear small spines (Erdtman, Franz).
On the basis of studies of floral development and anatomy the peri-
anth and androecium in the family have been variously interpreted as
biseriate or uniseriate, while the pentamerous floral plan evident in many
genera has been considered as either basic, or derived from a basically
trimerous (Payer, Eichler) or bimerous (Sharma) pattern. Thus the two
(to several) sepals are considered by some as involucral bracts, and the
petals as tepals (cf. Pax & Hoffman, 237, 238). In the light of con-
flicting interpretations, as well as for taxonomic convenience, the tra-
ditional terms sepal (calyx) and petal (corolla) are used here
The ovary has been shown in several genera (Anacampseros, Calan-
drinia, Montia, Portulaca) to be septate early in ontogeny. The septa
are lost as the ovary develops, resulting in free-central or basal placen-
tation, with the ovules arranged in one to several groups, each group
presumably representing a lost locule. Anatomical data for the family
are fragmentary and not particularly distinctive. The intraxylary phloem
reported to occur in Montia and its allies is not always well defined. A
petiolar vascular supply of a single bundle has been reported in species
of Portulaca and illustrated in the monotypic genera Mona O. Nilss. and
Neopaxia QO. Nilss.
Chromosome counts available for some of the better known genera
(Calandrinia, Claytonia, Lewisia, Montia, Oreobroma, Portulaca, Tali-
num) show diploid numbers ranging from 2” = 8 to ca. 191, in either
dysploid or euploid series, suggesting a rather complex evolution through
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 569
the development of polyploidy followed by the formation of aneuploids.
The family is of only minor economic importance. Species of Anacamp-
seros, Calandrinia, Lewisia, Montia, Portulaca, and Talinum are grown
as arnanentals, A few species are used as vegetable or salad “greens”
or in folk medicine for various internal complaints. Species of Portulaca
and Calandrinia have proven poisonous to livestock in the southwestern
United States and Australia due to high concentrations of oxalic acid.
REFERENCES:
Barton, H. Portulacacées. Hist. Pl. 9: 54~80. 1886.
Becker, C. Beitrag zur ee Anatomie der Portulacaceen. [Diss.]
F clad. Alenentievs Universitat. 41 pp., 9 figs. Munich. 1895.
BENTHAM, G., & J. D. Hooker. se Gen. Pl. 1: 155-159. 1862.
BRANDEGEE, K. Studies in Portulacaceae. Proc. Calif. Acad. Sci. 4: 86-91.
1894.
CANDOLLE, A. P. pe. Portulaceae. Prodr. 3: 351-364. 1828.
Davis, G. L. Systematic embryology of the angiosperms. 528 pp. New York.
1966. [Portulacaceae, 11, 14, 219.]
D’Hupert, E. Recherches sur le sac embryonnaire des plantes grasses. Ann.
Sci. Nat. Bot. VIII. 2: 37-128. pls. 1-3. 1896.
Eckarpt, T. Portulacaceae. Jn: H. Metcutor, Engler’s Syllabus der Pflan-
zenfamilien. ed. 12. 2: 90-92. 64.
EICHLER, A. W. mirage cane 2: 125-129. fig. 47. 1878. [Claytomia,
125; Portulaca, 125; Talinu
ENDLICHER, a Portulaceae. ay 2 946-955. 1840.
RDTMAN, CG. 1952. Pollen morphology and plant taxonomy: Angiosperms.
539 pp. Waltham. 1952. Pponidatetean 336, 337
Evertst, S. L. A review of the poisonous plants of Queensland. Proc. Roy.
sai Queensland 74: 1-20, 1964. [Calandrinia, Portulaca, incl. P. olera-
ee E. Monographie der Mollugineen und Steudelieen. 1. Supplement.
Abh. Ann. Wiener Mus. Naturges. 1-2: 279-307.
Franz, E. Beitrage zum Kenntnis der Portulacaceen und Basellaceen. Bot. Jahrb.
42(Beibl. 97): 1-48. 1908.
Gray, A. Portulacaceae. Gen. Pl. U.S. 1: 221-230. pis. 97-100. 1848. [Clay-
tonia, Talinum, Portulaca, Sesuvium.
Howet., T. A rearrangement of American Portulaceae. Erythea 1: 29-41.
1893.
JoHANSEN, D. A. Plant embryology. 305 pp. Waltham. 1950. [Portulacaceae,
LS 7.
Kowat, T. Morphology and anatomy of the seeds in gpm ae Rchb, (In
Polish: English summary.) Monogr. Bot. 12: 3-47. 1961.
LuBBockK, J. A contribution to our knowledge of seedlings. vol, 1. London.
1892. [Portulacaceae, 224-229.]
Martin, A. C. The comparative internal morphology of seeds. Am. Midl. Nat.
36: 513-660. 1946. [Portulacaceae, 558.
Nitsson, 6. Studies in Montia L. and Claytonia L. ~ allied genera. 3. Pol-
len morphology. Grana Palynol. 7: 279-363.
Pax, F., & K. HorrMann. Portulacaceae. Nat. Pfla Lee ed. 2. 16c: 234-
262. 1934. [Portulaca, 246; Talinum, 248; Claytonia, 257.]
570 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Paver, J. B. Traité d’organogénie comparée de la fleur. vii + 748 pp., 145
pls. Paris. 1857. [Portulacaceae, 325-335. pls. 68, 70.
Puiiipson, W. R., & J. P. SKipworRTH. Hectorellaceae: a new family of di-
cotyledons. Trans. Roy. Soc. New Zealand Bot. 1: 32. 1961. [Incl. Hec-
torella, Lyallia. |
Raprorp, A. E., H. E. Antes, & C. R. Bett. Manual of the vascular flora of
the Carolinas. 1183 pp. "Chapel Hill. 1968. [Claytonia, 432; Portulaca,
433; Talinum,
RIcKETT, H. W. Wild flowers of a United States. Vol. 2. The Southeastern
States. Part. 1. New York. 1967. cpa 295, pl. 106; Portulaca,
296, pl. 106, 107; Talinum, 295, 296, pl. 106.]
Rypserc, FP, Pothilscacess. N. Am. Fl. 21: 279-336. 1932. [Portulaca,
Talinum, by P. Wilson. ]
Scumutz, E. M., B. N. Freeman, & R. E. ReEeEp. Livestock-poisoning plants
of Arizona. 176 pp. Tucson. 1968. Rapiiaatene, 145.
ScHouTE, J. On the perianth aestivation in the Portulacaceae and the
Basellaceae, Rec. Trav. Bot. Néerl. 32: 395-405. 1935.
SHARMA, A. & . BHATTACHARYYA, Cytogenetics of some members of
Portulacaceae and related families. Caryologia 8: 257-274. 1956.
SHARMA, H. P. Studies in the order Centrospermales. 1. Vascular anatomy of
the flower of certain species of the Portulacaceae. Jour. Indian Bot. Soc.
33: 98-111. 1954. [Portulaca, Talinum
sai J. P. The taxonomic position a Hectorella caespitosa Hook. f.
Trans. Roy. Soc. New Zealand. Bot. 1: 17-30. 1961. [Hectorella, Lyal-
lia
Upuor, J. C. Tu. Dictionary of economic plants. ed. 2. 591 pp. Lehre. 1968.
[ Claytonia, 136; Portulaca, 426; Talinum, 509.]
Woutpart, A., 3 Masry, The distribution and phylogenetic significance
of the betalains with respect to the Centrospermae. Taxon 17: 148-152.
1968. [Claytonia, Portulaca.|
KEY TO THE GENERA OF PORTULACACEAE
General characters: low, succulent to suffrutescent herbs with simple, —
flat to terete, alternate, alana or basal leaves; stipules scarious, or of tu
hairs, or absent: stems erect to procumbent; inflorescence axillary, a or
scapose, diffuse to ie cad basically cymose but often appearing racemose
or paniculate; flowers complete, small to large, regular, hypogynous to epigy-
nous; sepals 2; petals usually 5 (4-6); stamens 5-100, alternate or opposite the
petals, free or fasciculate; ovary 2-9-carpellate, unilocular, with basal placenta-
tion; ovules few to numerous, ascending; fruit a 3- to many-seeded capsule,
circumscissile or valvate
oa
; ae — scarious or laciniate, or of tufts of hair; inflorescence a
terminal head or congested helicoid cyme, usually surrounded by a whorl of
sane ovary half-inferior to inferior; stamens few to many, perigynous;
fruit a circumscissile capsule; ovules and ar nbc om * eee
>
. Stipules absent; ovary superior; fruit a 3-valved capsule.
B. Leaves cauline or basally tufted on stems from tuberous roots or rhi-
zomes; inflorescence a terminal peduncled cyme, or flowers axillary, soli-
tary; stamens 5-100, alternate with or opposite the petals; fag a
seeds numerous. . Talinu
2 Se 0 eS eS Ree ee we we Be ee wm ee Re el ee
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 571
B. Leaves few to many from a deep-seated corm; flowering stem scapose,
bearing a single pair of opposite leaves; inflorescence a loose raceme;
stamens 5, opposite the petals; ovules andl seeds (3-)6. .. 3. Claytonia.
Subfamily PORTULACOIDEAE [Franz]
Tribe PorTULACEAE [ Franz |
Subtribe Portulacinae | Franz |
1. Portulaca Linnaeus, Sp. Pl. 1: 445. 1753; Gen. Pl. ed. 5. 204. 1754.
Low, annual or perennial herbs with erect, ascending or procumbent,
fleshy or suffrutescent, sometimes reddish stems from fleshy or fibrous
roots. Leaves alternate to subopposite, terete, subterete, or flat, entire,
often congested in an involucre about the flowers. Stipules scarious or
reduced to tufts of hairs, rarely absent. Flowers erect, sessile to sub-
sessile, solitary and axillary or few in terminal heads or compact helicoid
cymes. Sepals 2, opposite, the abaxial larger than the adaxial, united
below. Petals +2, free or basally connate, gelatinous-deliquescent after
flowering. Stamens 6-40 [4-100], inserted perigynously; filaments
usually pubescent below; pollen polyrugate. Carpels as many as the
style branches; styles short, 2—9-parted [rarely simple]; ovary semi-
inferior to inferior, globose to obovate, plurilocular below to unilocular
above; ovules numerous, amphitropous, on a simple or branched free-
central placenta. Capsule membranaceous, chartaceous, circumscissile.
Seeds numerous, reniform to cochleate; seed coat smooth or variously
sculptured, in ours granular to stellate-tubercular or -echinate, brown to
black or gray. Embryo peripheral, annular, surrounding the endosperm.
(Including Portulacca Haw.). Lectotype species: P. oleracea L.; see
Britton & Brown, Illus. Fl. No. U.S. ed. 2. 2: 39. 1913. (Name probably
derived from Latin, portula, a small gate or door, in reference to the
calyptra of the capsule.) — PURSLANE.
A genus of about 100-125 species widely distributed in tropical, sub-
tropical, and temperate regions of the world. Eight to ten species occur
in our area.
In the early literature the North American species were generally
grouped in two or three subgeneric categories of undesignated rank, based
on either leaf shape (flat or cylindrical) or the presence or absence of
pubescence. Wilson (N. Am. FI.) recognized 23 species, but no sub-
generic groupings. In Poellnitz’s provisional monograph of the genus
(1934), 104 species were disposed in two subgenera, one with two sec-
tions and eight subsections. For lack of comparable material, Poellnitz
could not key all of the species to their appropriate subsections, but each
is described and discussed in detail. The American species have since
been monographed by Legrand, who recognized six subgenera, of which
three, containing about 62 species, occur in the Western Hemisphere. All
Fic. 1. Portulaca. a—h, P. oleracea: a, flowering and fruiting branch, 1/2;
b, flower in semidiagrammatic vertical section to show Ww perigynous floral organs
and basal oe X 6; c, nearly mature fruit inclosed above by accrescent
sepals — e bracteole at base of ovary, X 3; d, same, with perianth =
to show top of 6 ovary with constricted apex and persistent style, X 5; e,
sa
j, flower bud just before anthesis to show sepals connate at base, X 8; k, flower
with forward petals depressed to show stamens and style, X 4; 1, withered peri-
anth (sepals and je stamens not visible) adhering to upper part of ¢
cumscissile perica 10; m, base of fruit after dehiscence, fale fou J
ascending basal placentae, one bearing a seed, x 10.
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 573
and ranges. Our species are discussed here under their older, more fa-
miliar names, followed by their disposition in Legrand’s system.
Four flat-leaved species, with carinate to subcarinate sepals, and usual-
ly with yellow flowers, have been described in our area. Portulaca olera-
cea L. (pusley, pursley, purslane, pigweed), 2n = 18, 45, 52, and 54, is
a common procumbent weed of waste places, fields, and cultivated areas,
with almost world-wide distribution. Its flat, fleshy, obovate-cuneate to
spatulate leaves; strongly keeled sepals; yellowish, apically notched
petals; and black, granulate seeds are characteristic. Birds have been ob-
served to eat the seeds and may have played a role in the widespread
distribution, but its origins are lost in antiquity (see discussion in De
Candolle). ‘West of the Mississippi River in Arkansas, Kansas, and
Missouri, the place of P. oleracea may be taken by P. neglecta Mack. &
ush, which differs in its upright ascending habit; larger size; larger,
broader, and thinner leaves; more numerous stamens; muricate seeds;
and different flowering time. Where the two occur together, P. oleracea
flowers open at about 9:30 a.m., while those of P. neglecta open at about
7:40 a.m. Poellnitz suggested that P. neglecta may be only a variety of
P. oleracea. The closely related P. retusa Engelm., ranging from Arkansas
and Missouri to Arizona and Utah, is similar to P. oleracea in its pro-
cumbent habit, but differs from both of the above in its thinner, retuse
leaves; more slender habit; smaller flowers; and echinate-tuberculate
seeds. All three species were recognized by Poellnitz, but Legrand re-
duced P. neglecta and P. retusa to synonymy under P. oleracea.
The fourth flat-leaved species, Portulaca coronata Small (including P
lanceolata Engelm.,2 not Haw.), inhabits granite outcrops and sandy
soils in South Carolina (very rare), Georgia, and Mississippi. In the
southwestern United States it ranges from the granitic region of western
Texas to lower California. The species is especially marked by the flaring,
corona-like rim surrounding the ovary just below the line of dehiscence,
as well as by its subcarinate sepals, lanceolate leaves, yellow (to orange
or red-tinged) petals with acute tips, and gray seeds. According to McVaugh
it is often found growing with P. Smallii on the granitic flat-rocks
of the Southeast, but in slightly more shaded conditions. In Legrand’s
system, P. coronata becomes a synonym of P. umbraticola HBK., which
ranges to Cuba, Central America, and much of South America.
The six remaining species of the area have more or less cylindrical
leaves, dorsally rounded sepals, and, for the most part, characteristic
pubescence of the leaf axils and inflorescences. Portulaca pilosa L..,
* According to Fosberg, P. coronata Small is the only name available for P. lan-
ceolata Engelm., since FP. lance olata Haw. (1803), antedating Engelmann’s name by
tet
°
£,
~
=
3
5 aed
a
Ss
S
=
&
a
e
eo
f
Engelm. Wilson (1932) ‘considered P. coronata a sy
Poellnitz (1934) treated the two as different species with very different seeds.
574 JOURNAL OF THE ARNOLD ARBORETUM [vot. 50
16, has conspicuous stipules of tufted whitish hair in its leaf axils, and
whitish to brownish hairs around the flowers. Its small, obovate, notched
petals are usually purplish pink, with a small mucronate tip at the base
of the notch. It is reported on sandhills and in dry, sandy soils of woods,
roadsides, and cultivated grounds from the Carolinas to Florida, and
westward to Texas, as well as in the West Indies, Mexico, and Central and
South America.
Portulaca Smallii P. Wils., 2n = 16, has only slightly tufted axils and
pale brown hairs in the inflorescence surrounding inconspicuous flowers
with deep lavender to pure white petals. The species is abundant in the
shallow soils of surface depressions and particularly of the marginal eco-
tone of granite flat-rocks in North Carolina and in Georgia to DeKalb
and Pike counties (but not in South Carolina or Alabama). This species
is very closely related to, and possibly derived from, P. pilosa, the two
being distinguished chiefly by the endemism of the former and the
greater overall size and larger number of stamens in the latter (Cotter &
Platt). Portulaca suffrutescens Engelm. is a linear-cylindric-leaved suf-
frutescent perennial with copper or yellow-brown (buff), notched petals,
and black, rounded-tuberculate seeds. It ranges from Arkansas to Ari-
zona and Mexico. Portulaca grandiflora Hook. (rose-moss, moss-rose),
2n = 18, 36, a native of South America, has escaped cultivation and
become widely naturalized in North America and Europe. Its flowers,
2-5 cm. in diameter, are larger than those of any of the native species
and have 40 or more stamens. Flower color ranges from white to pink,
red, orange, salmon, or yellow. Its cylindric leaves also have hair-tufted
axils
Portulaca parvula Gray, with ascending branches, copiously hairy nodes,
nearly cylindrical leaves, yellow, orange, or copper petals (red, accord-
ing to Wilson and Poellnitz), and stipitate capsules, occurs from western
Missouri and Arkansas westward to Colorado, California, and south-
ward into Mexico. This species was redefined by Johnston, who segre-
gated the purple-flowered plants included in it by Gray as P. mundula
I. M. Johnston (Mexico to Oklahoma, Kansas, and Missouri). Such
plants have also been included under P. pilosa L., but Johnston considered
that species attributable to specimens from Curacao, and not conspecific
with the plants of Mexico and the southwestern United States. Legrand,
however, reduced P. mundula to varietal status under P. pilosa and the
remainder of P. parvula to synonymy under P. halimoides L.
Portulaca phaeosperma Urb. has small yellow flowers surrounded by
brownish to whitish hairs and nearly cylindrical leaves with incon-
spicuously tufted axils. It is found in sand-dunes, scrub, and shell-mounds
of southern peninsular Florida and the Keys, southward throughout the
West Indies to Yucatan and Curacao. According to Legrand, P. phaeo-
sperma is synonymous with P. rubricaulis HBK
Chromosome counts indicate that basic numbers in Portulaca may
= 4, 5, 8, and 9. At least two chromosome series are present,
with 2m =8, 16, and 48 in one, and 2n = 18, 36, 45, 54, and 108 in the
other. The Japanese ‘Jewel’ strain of P. grandiflora is reported to have
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE s73
2n = 10, while P. quadrifida L. has 2n = 50 (Bouharmont). These
data indicate that chromosomal duplications or deletions, concurrent with
polyploidy, have played an important part in the evolution of the genus.
There are no records of natural interspecific hybridization, and attempt-
ed reciprocal crosses (involving P. grandiflora and P. marginata HBK.
in one case, P. grandiflora and P. grandiflora ‘Jewel’ in the other) have
failed. Bouharmont reports very irregular meiosis in an artificial hybrid
between the African P. centrali-africana R. E. Fries, 2n = 108, and P.
Kermesiana N. E. Br., 2n = 108. He also found chromosomes to be much
larger in P. foliosa and P. grandiflora than in other species.
Ikeno’s efforts to explain color variation in Portulaca grandiflora in
terms of simple Mendelian principles was only partially successful.
Blakeslee attributed various changes and reversions in habit and in petal
coloration (sectorial and periclinal chimeras, stripes and spots) to spon-
taneous vegetative mutations. Other cytological peculiarities include the
presence of polyploid cells (2” = 36, 72) in adventitious roots of P.
grandiflora and P. foliosa (Bouharmont) and the formation of up to 12
secondary cells of varying size within the primary epidermal cells of P.
grandiflora (Czeika). The presence of betacyanins has been confirmed in
P. grandiflora, P. oleracea, and P. pilosa (Wohlpart & Mabry).
The opening and closing of flowers in response to light intensity and
temperature have been studied by several workers. Iwanami e¢ al. con-
cluded that temperature controls flowering time in Portulaca grandiflora,
while Cotter & Platt suggest that light is the controlling factor in P.
Smallii, Lebrun found that three vegetative races of P. quadrifida from
different sources show a divergence of behavior under the same conditions
appreciably corresponding to the difference in local times of their places
of origin. Similarly, offspring of an artificial hybrid between P. centrali-
africana and P. Kermesiana were intermediate in their behavior to those
of the two parents.
On the basis of floral vascular anatomy Sharma suggests that the pen-
tamerous floral condition in Portulaca is derived from a basically bimerous
plan, with a biseriate (rather than uniseriate) perianth, and a modified
androecium consisting basically of two cycles. The ovary in Portulaca
is generally considered to be semi-inferior to inferior, but Soetiarto &
Ball have demonstrated that in P. grandiflora all the floral organs arise
on the flanks of the floral apex as in a hypogynous flower, and that all
except the carpels are then lifted on the lip of the floral cup. They con-
clude that perigyny is an ontogenetically late development in P. grandi-
flora and may be interpreted as a derived condition.
The anthers of Portulaca tuberosa and P. grandiflora have binucleate
tapetal cells and binucleate pollen grains at anthesis, but P. oleracea has
multinucleate tapetal cells and trinucleate grains. Seed coat morphology
may be an important specific character, but is not necessarily consistent in
all species. Kowal states that in both P. oleracea and P. sativa L. seeds
occur with sculpturing of two forms and that a particular specimen may
produce seeds of one or both types.
Several species of the genus are grown as garden ornamentals. Por-
576 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
tulaca oleracea is reported in Jamaican folk-lore to be of value in the
treatment of cardiovascular diseases (possibly due to the presence of
(— )-noradrenaline in relatively high amounts in the fresh plant tissues),
as a diuretic, and as an antiscorbutic. It is also widely cultivated and
used as a vegetable or salad-green, or as fodder for pigs. Portulaca pilosa
is sometimes used as a diuretic, an emmenagogue, and as a stomach tonic.
High oxalate or nitrate content in plants of P. oleracea and other species
may prove toxic to livestock.
REFERENCES:
Under family references see Davis, EICHLER, ErpTMAN, EveERIST, GRAY,
HowELL, JOHANSEN, Kowa, Luspock, Martin, Raprorp et al., RICKETT,
ScHMuTz, SHARMA & BuHaTTAcHARYyA, H. P. SHARMA, UpHor, WoHLPaRT &
Masry.
BLAKESLEE, A. F. A dwarf mutation in Portulaca showing vegetative reversions.
Genetics 5: 419-433. 1920. [P. grandiflora. |
. Inheritance of germinal peculiarities. Portulaca, Carnegie Inst. Wash.
Year Book 19: reise 1921.
, Jr. A vegetative reversion in Portulaca. (Abstr.)
Brooklyn Bot. Gard Mem. 1: 18. 1918. [P. grandiflora
BouHARMONT, J. Note sur la cytologie de quelques espéces de Portulaca. Bull.
Soc. Bot. Belg. 98: 175-188. 5.
CaNDOLLE, A. P. pe. Origin of cultivated plants. Reprint of ed. 2 (in English).
Stechert-Hafner, N.Y. 1886 (1959). [P. oleracea, 87-89.]|
a E. Contributions to the histogenesis of the Caryophyllales, I.
Tra . Microscop. Soc. 20: 97-164. pls. 8-25. 1889. [P. oleracea,
pls. "12(5), 22(15).]
Connarp, M. H., & P. W. ZmmMeERMAN. The origin of adventitious roots in
cuttings of Portulaca oleracea L. Contr. Boyce Thompson Inst. 3: 337-
346. 1931
Portulace oleracea. Am. Jour. Bot. 22: 453-4
. Macrosporogenesis and embryology of ae oleracea, Am. Jour.
Bot. 27: 326-33 40.
Costa, A. S., & A. M. B. CARVALHO. Common purslane (Portulaca ret agha
a heccgges of the tomato spotted wilt virus. (In Portugese; English s
ry.) ows 19(1, nota 6): XXI-XXV. 60.*
Baca °”D. Jj, &@ KB. Parr. Studies on the ecological life history of Portu-
laca Smallii. ae 40: 650-668. 59.
Cze1ka, G, Uber das spontane Auftreten von Mitosen und inaqualen Cytoki-
nesen in der endopolyploiden Epidermis von Portulaca grandiflora. Planta
51: 566-574. 1958.
Epwarbs, S. Portulaca pilosa. Bot. Reg. 10: pl. 792. 1824.
Enomoto, N. Studies on an ever-segregating race in Portulaca grandiflora.
Jap. Jour. Bot. 1: 137-151. 3
Fence, P. C., L. J. Haynes, & K. E. Macnus. High concentration of (—)-nora-
drenaline in Portulaca oleracea L. Nature 191: 1108. 1961.
FosBerc, F, R. Notes on North American plants. IV. Am. Midl. Nat. 29: 785,
786. 1943. [P. coronata, P. lanceolata.|
Cooper, D. C. Microsporogenesis and the — of the male gametes in
3
~s
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 577
Fow ter, H. S. The two-faced plant. Cornell Plantations 7: 26-28. 1950-1951.
[P. oleracea, notes and instructions on use.
Gray, A. Plantae Lindheimerianae, te Boston Jour. Nat. Hist. 6: 154, 155.
1850. [{Portulaca(ceae) by A. Engelmann.
Contributions to American botany. 1. Revision of some polypetalous
genera and orders precursory to the Flora of North America. Proc, Am.
Acad. Arts Sci. 22: 270-314. [Portulaca, 274.
Hooker, W. J. oe grandiflora Hook. Bot. Mag. 56: pl. 2885. 1829;
58: pl. _. 183
Huane, T. C. Tt mg — “ Formosan plants, (2). Taiwania 13: 15-101.
1967. [P. pilosa, 75; fig. 5
IKENO, S. Studies on the ae of flower-colors in Portulaca grandiflora.
Jour. Coll. Agr. Univ. Tokyo 8: 93-133. 1921.
Iwanami, Y. The movements of the stamens of Portulaca — I. Bot.
Mag. Tokyo 75: 133-139; II. 75: 289-295; III. 75: 331-3 1962.
. Hosutno. The opening and closing movement of the flower of Por-
tulaca eg II. bid. 76: 108-114.
camienseani The opening and closing movements of the flower of Por-
on candi, I. (in Japanese; English summary.) Jbid. 75: 443-
1962
nay i. ‘ie Hon MA, & S. TAKANO. Interspecific crosses with Portulaca. 1.
(Abstract; in Japanese .) Jap. Jour. Breeding 8: 193. 1958.
LeBRUN, J. Le mouvement d’ouverture et de fermeture des fleurs chez diverses
te a Oa “Bull. Soc. Bot. Belg. 99: 19-35.
Lecranp, C. D. Revisando tipos de Portulaca. Communic. Bot. Mus t.
Nat. Montevideo 2(24): 1-10. pls. 1, 2. 1952. [Species fe Humboldt, oa
pland, & Kunth
; especies americanas de Portulaca. Anal. Mus. Hist. Nat. Monte-
video 7(3): 1-147. pls. 1-29. 1962. [Monograph of American spp. ]
MacKenzir, K. K., & B. F. Busn. New plants from Missouri. Trans. Acad.
Sci. St. Louis 12: 79-89. pls. 12-17. 1902. [P. neglecta, n. sp.]
McMurray, N. The blooming of Portulaca. Am. Bot. 25: 133, 134. 1919.
McVaucH, R. The vegetation of the granitic flat-rocks. Ecol. Monogr. 13: 119-
166. 1943. [Distribution map of P. coronata in southern U.S., 141; in-
cludes P. coronata, P. Smallii.| ;
Murpy, W. H. Granite rock outcrop endemics from the standpoint of spe-
ciation. (Abstr.) ASB Bull. 15: 72. 1968. [P. Smallit.]
PoELLNITz, K. von. Versuch einer Monographie der Gattung Portulaca L.
Repert. Sp. Nov. 37: 240-320. 1934.
. Die Portulaca-Arten Westindiens. Jbid. 50: 89-103. 1941.
Racuavan, T. S., & A. R. SRINIVASAN. Cyto-morphological features of Por-
tulaca tuberosa Roxb. Proc. Indian Acad. Sci. B. 14: 472-488. 1941,
[Embryology, including review of previous work on P. oleracea, P. grandi-
ora.
Snow . i“ The purslane-worm. (Copidryas Gloveri Grote). Science 10: 204.
1887. [P. oleracea, P. retusa.
bape S. R., & E. Baty. Ontogenetical and experimental studies of the
floral apex of Portulaca grandiflora. 1. Histology of transformation of the
shoot apex into the floral apex. Canad. Jour. Bot. 47: 133-140. pls. 1-8.
1969. ;
Sovices, R. Embryogénie des Portulacacées. Développement de l’embryon
578 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
chez le Portulaca oleracea L. Compt. Rend. Acad. Sci. Paris 207: 763-
770. 1938.
STEINER, E. Cytogenetic studies on Talinum and Portulaca. Bot. Gaz. 105:
3 79. 1944.
Syakupo, K., S. Kawapata, & A. UstHara. On the plant having » = = 5 chromo-
somes in Portulaca. Jap. Jour. Genetics 35: 107-109. 1960.
Witson, P. Portulaca. In: P. A. Rydberg, Portulacaceae. N. Am. Fl. 21: 279-
336. 1932. [ Portulaca, 328-336.]
Tribe CALANDRINIEAE Fenzl
Subtribe Calandriniinae | Franz |
2. Talinum A. L. de Jussieu, Gen. Pl. 312. 1789, nom. cons. prop.
Annual or perennial, glabrous, succulent herbs, subshrubs [or shrubs],
with simple or branched, short to elongate stems with fleshy, tuberous
roots or rhizomes. Stems frequently terminating in wiry, short or long
peduncles. Leaves alternate to subopposite or basally tufted, exstipulate,
cylindrical or flattened, entire, fleshy. Inflorescence a terminal or axillary,
wiry-peduncled cyme, or flowers sometimes solitary on short pedicels in
the leaf axils. Sepals 2, deciduous or persistent. Petals 5, rarely more,
free or connate at the base, often showy, purplish to red, pink, yellow, or
white, ephemeral. Stamens 5-100, alternate with or opposite the petals or
in fascicles opposite and basally affixed to the petals; filaments slender,
sometimes colored. Style elongate, deeply 3-parted to subcapitately lobed
or capitate, equaling or exceeding the stamens; ovary superior, unilocular
(at least above); ovules numerous, amphitropous, placentation free cen-
tral. Fruit a unilocular many-seeded, chartaceous capsule, dehiscing locu-
licidally from the apex to the base by 3 valves. Seeds compressed, round-
reniform, dull gray or shiny brown to black, smooth, striate, or tuberculate,
distinctly to indistinctly arillate. Embryo peripheral, incompletely an-
nular; endosperm starchy. (Talinum Adans., 1763, nom. superfl.; Heli-
anthemoides Medic., Phemeranthus Raf., Litanum Nieuwl.) LECTOTYPE
species: T. patens A. L. Juss., nom. illegit. = T. paniculatum (Jacq.)
Gaertn., typ. cons. prop.; see J. E. Dandy, Taxon 18: 465. 1969. (Origin
of name obscure, said to be derived from the aboriginal name of an African
species.) — FAME FLOWER.
A genus of about 50 species distributed in the tropical, subtropical, and
temperate regions of both hemispheres, but best developed in North
America (30-35 species), and particularly in Mexico (Rose & Standley).
The genus was last monographed by Poellnitz (1934) who recognized
47 species but found no basis for maintaining the three sections estab-
lished by De Candolle (§ Phemeranthus Raf., with cylindrical leaves;
§ Talinastrum DC., with flat leaves; and § Talinellum DC., since trans-
ferred to Calandriei HBK.).
Eight species are now recognized in our area, six wholly or partially
east of, and two mostly west of the Mississippi River. Two flat-leaved
species frequent the lowlands, forming erect, suffrutescent plants ranging
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE
Fic. 2. Talinum. a-l, T. calcaricum: a. flowering plant, X 1/2; b, flower bud,
dusts before anthesis to show sepals, X 3; c, flower, X 3; d, stamen before
anthesis, X 20; e, f, two views of stamen fees anthesis, x 20; g, gynoecium
ith one — of ovary removed to show free-central placentation, X 10; h,
Cross section of ovary showing placentation a € f is, X 10; i,
lateral view of ovule ns time of anthesis showing ginning to grow from
funiculus, x 40; j, just before dehiscence, < 5; k, <— Nas thin, tightly
fru
1 f . develo d (the two sides not m midline of
deed a oF grees 4 x sec a the curved embryo Frain Te endosperm
(black stipe Genecasisiviass a ES.
580 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
in height from about 6 dm. to more than 1 m. Talinum paniculatum
(Jacq.) Gaertn. (including “7. patens Willd.” and T. reflexum Cav.), 2n
= 24, is sparingly branched with terminal, paniculate inflorescences,
terete peduncles, and red or pink to yellowish flowers with about 15-20
stamens. The species occurs in both the New and Old Worlds, ranging
in North America from Florida to Arizona, southward through the West
Indies and Mexico to Central and South America. It favors sandy and
cultivated soils, and is seen only rarely in waste places in the Carolinas
as an escape from cultivation (Radford et al.). Several authors recognize
var. sarmentosum (Engelm.) Poell., with procumbent branches, in Texas.
In contrast with T. paniculatum, T. triangulare (Jacq.) Willd. is much
branched and bears corymbiform, racemose or cymose inflorescences with
three-angled peduncles and purplish to pink or yellow flowers with about
30 stamens. It occurs from southern peninsular Florida and the Keys,
where it inhabits hammocks, pinelands, and waste places, to the West
Indies, Mexico, and Central and South America.
The cylindric-leaved species of the southeastern United States are
herbaceous “rock plants,” particularly in the Appalachian highlands,
favoring exposed, arid sites in the shallow soil or debris on the margins
or surfaces of granite, limestone, or serpentine outcrops. They are dis-
tinguished primarily by floral characteristics, including the number of
stamens, length of style relative to stamens, persistence of sepals, and
seed color. (See Ware for a key to these species in the Southeast.)
The best-known and most widespread Talinum of our area is T. tereti-
folium Pursh, 2n = 24, 48, which ranges from the serpentine barrens of
southeastern Pennsylvania, southward in acid soils to the granite out-
crops of the Piedmont of the Carolinas and Georgia, and to the Altamaha
Grit region of Georgia. Its pink flowers have 15—20 stamens, a three-
lobed style as long as the stamens, small deciduous sepals, and small, shiny,
black seeds. Three species with more restricted distributions have been
segregated from T. teretifolium. Talinum calcaricum Ware, confined to
shallow soil at the margins of limestone exposures in the cedar glades of
middle Tennessee and northern Alabama, has purple-pink petals, per-
sistent sepals, 25-45 stamens shorter than the three-lobed style, and
large, shiny, dark-brown seeds marked by longitudinal rows of low,
broad, raised cells. Talinum Mengesii Wolf, 2n = 24, occurring on sand-
stone or granite in Alabama, Georgia, and Tennessee, has deciduous-
sepaled pink flowers with 50-80 (40-100) stamens shorter than the sub-
capitate style, and small, shiny, black seeds. Wright finds that “7. tere-
tifolium is replacing T. Mengesii on granite outcrops in central Georgia
where their distribution ranges overlap, and where their niche require-
ment appears to be very similar.” Talinum appalachianum Wolf, 2n =
24, is confined to granite outcrops along the Coosa River in two counties
of Alabama. Its flowers have five stamens which alternate with the
petals, and there are sometimes one or two extra stamens opposite the
petals. The three-lobed style is longer than the stamens, and the seeds
at maturity are smooth and brown
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 581
West of the Mississippi two more or less cylindric-leaved species enter
our area in Arkansas. Talinum calycinum Engelm., 2n = 24, with red
petals and 30-45 stamens, occurs in southernmost Illinois and from Mis-
souri and Arkansas to Montana, Nebraska, Kansas, Oklahoma, Texas,
New Mexico, and Mexico (Poellnitz). Talinum parviflorum Nutt., 2n
= 48, with white to rose-colored petals and 4-8 stamens, also occurs in
southern Illinois and ranges northward from Arkansas to Minnesota and
North Dakota, westward to Colorado, Texas, Arizona, and possibly Mexi-
co. Talinum rugospermum Holz. may approach our area to the northwest,
for it occurs from northwestern Indiana through Illinois to northeastern
Iowa, Wisconsin, and eastern Minnesota. It is distinguished by its deeply
trifid style, 12-25 stamens with subglobose anthers, and_ strongly
wrinkled seeds.
The flowers of all the species are very delicate and ephemeral. Several
authors have pointed out the need for studying living plants, in either
the field or cultivation, since critical characters, such as the color, size,
shape, and persistence of the floral organs, are damaged or lost in pressing
and drying. The flowers usually open for only a short period in full
sunshine, the time and duration of daily flowering differing among the
species, or even among different populations of the same species, but
appearing to be more or less characteristic in a given geographical area.
Wolf observed the usual flowering time in T. Mengesii to extend from
about 11 a.m. to 5 p.M., while in T. appalachianum the flowers open be-
tween 3 and 4 p.m. and close between 5 and 6 p.m. Ware reports for T.
calcaricum flowering times of 1 to 6 P.M. in Tennessee, but 3:30 to 6:00
P.M. in Franklin County, Alabama. It is not known, however, whether
anthesis is controlled by temperature, light intensity, or both. The sig-
nificance of flowering times and of the relative lengths of styles and
stamens in the genus has not been investigated. Harshberger, the only
reporter of pollinators, observed two species of the hymenopteran Cal-
liopis on flowers of 7. teretifolium.
Ecological studies by Ware have shown that Talinum calcaricum and
T. Mengesii are restricted to their narrow ecological niches through in-
ability to compete with other species in more favorable sites. Adaptations
of species of Talinum which favor survival under the conditions of ex-
treme drought which often prevail in their habitat include succulence of
leaf and stem, cylindrical leaves with low surface-to-volume ratio, few
sunken stomata, pronounced cuticle, profusely branched root system, and
the ability to flower even when a water deficit exists in the leaves (Gup-
till, Harshberger, Ware).
Steiner found no record of interspecific hybridization in Talinum, and
his attempts to cross 7. teretifolium with T. parviflorum and T. parvi-
florum with T. aurantiacum Engelm., 2n = 48, of the southwestern
United States were unsuccessful. Both his chromosome counts (eight
species) and those of others indicate a polyploid series with 2n = 24, 48,
and 72. He also reported polyploid cortical cells with 21 = 96 in the
root tips of T. parviflorum. Ware tried the crosses T. calcaricum
582 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
calycinum, T. calcaricum *« Mengesii, and T. Mengesii calcaricum,
but the resulting low seed yields led him to conclude that partial genetic
barriers exist between these morphologically similar taxa.
n the basis of seed morphology and anatomy, Kowal proposed that
“Talinum patens Willd.” and T. reflexum Cav., which have been treated as
conspecific with 7. paniculatum since early in this century, are distinct
species. In our specimens the development of the aril varies considerably
in different species. At maturity the seeds of T. calcaricum, T. Mengesti,
and T. teretifolium are enclosed in a fragile, membranaceous, although
tightly investing, aril, and all appear dull gray. Disintegration or removal
of the aril reveals a shiny dark brown seed in T. calcaricum, and shiny
black seeds in the other two species.
Guignard found the embryology of 7. paniculatum to be the same as
that of Portulaca oleracea and some species of Calandrinia (i.e., Polyg-
onum-type embryo sac, solanad-type embryogeny). Anatomical details
of the genus include the presence of unicellular epidermal papillae and
drusiform calcium-oxalate crystals in the leaves. Incomplete septa have
frequently been described in the unilocular ovary. The pollen of T. tereti-
folium is 12-forate (i.e., with + circular apertures distributed evenly
over the surface of the grain), with three of the fora larger than the rest
(Franz, Erdtman), while that of T. triangulare var. crassifolium Hort. are
about 18-rugate (i.e., with elongate apertures distributed evenly over the
surface), with a thin, psilate exine (Huang). Sharma describes the floral
vascular anatomy of T. paniculatum and interprets the androecium as
consisting of two alternate cycles of stamens. Saponins are reported in
the leaves of T. paniculatum.
The genus is of little economic importance. Talinum paniculatum
(Jewels of Opar) and T. triangulare, as well as several western American
species are grown as border or rock-garden plants for their foliage, and
red, white, or yellowish flowers. The roots of 7. aurantiacum Engelm.
are cooked and eaten by Indians of the southwestern United States, while
the leaves of T. portulacifolium (Forsk.) Aschers., of Africa and of
triangulare in the Americas, are sometimes cultivated as a leafy vegetable.
Talinum cuneifolium (Vahl) Willd. is used in Tanganyika as an aphro-
isiac.
REFERENCES:
Under family references see BRANDEGEE, EICHLER, ERDTMAN, Gray, HoweLL,
Kowat, Martin, Paver, Raprorp eft al., Rickett, H. P. SHarma, and UPHOF.
ALEXANDER, E. J. Talinum stig Addisonia 21: 61, 62. pl. 703. 1942.
Epwarps, S. Talinum teretifolium. Bot. Reg. 29: pl. 1. 1843.
FAIRCHILD, D. Talinum, a summer vegetable for Florida. Fla. State Hort. Soc.
Proc. 56: git 1943.
Fassett, N. C. Notes from the herbarium of the University of Wisconsin. 3.
Rhodora 30: 205-207. 1928. [Distinguishes T. rugospermum Holz. from
T. teretifolium Pursh.]|
FREEMAN, O. M. Notes on some plant associations in Greenville and Pickens
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 583
ea South Carolina. Castanea 23: 46-48. 1958. [T. teretifolium, 47.]
GuIcnarp, J. L. Embryogénie des Portulacacées. Développement de l’embryon
chez le Talinum patens Willd. Compt. Rend. Acad. Sci. Paris. 261: 5599~
5601. 1965. [7. paniculatum. |
GupTILL, P. L. The morphology and life cycle - oT teretifolium Pursh.
M. S, Thesis, Emory University Library.
HARSHBERGER, J. W. An ecological study of a ie Talinum with descrip-
tions of two species. Bull. Torrey Bot. Club. 24: 178-188. pl. 299. 1897.
17. nig T. napiforme DC., T. Greenmanii, sp. nov., illustrated. ]
Hoxzincer, J. M. Talinum rugospermum, n. sp. Asa Gray Bull. 7: 115~117.
1899, [Segregated from 7. teretifolium.
———.. The geographical distribution of the teretifolium group of Talinum.
Ibid. 8: 36-39. 1900.
Huanc, T. C. Pollen grains of Formosan Plants (2). Taiwania 13: 15-110.
1967. [T. triangulare var. ere Sem 75, fie. 54.
Linpsay, G. The giant Talinum: Talinu guadalupense Dudley. Cact. & Suc-
cul. Jour. 23: 35-39. 1951. rIllustrated.)
McVaucH, R. The vegetation of the granitic flat-rocks of the southeastern
United States. Ecol. Monogr. 13: 121-166. [T. teretifolium, T. Mengesii,
renee K. von. Zur Kenntnis der Gattung 7 — Adans. (Portula-
caceae). Ber. Deutsch. Bot. Ges. 51: 112-127.
oe der Gattung Talinum Adans. 7% Sp. Nov. 35: 1-~
Rose, J. © = 4 F.C. ernge a genus Zalinum in Mexico. Contrib. U.S.
Sims, J. Times reflexum ay oe Mag. 37: be 1543. 1813. [Yellow-flow-
ered farm fom South America, = 7. paniculatum.]
STEINER, E. Cytogenetic studies on - Talinum oa Paha Bot. Gaz. 105:
374-379. 1944. [Chromosome eo for 8 spp.]
SUBRAMANIAM, T. V., & S. N. CHANDRASEKHARAN. Talinum triangulare Willd.
(Portulacaceae), a little known pai e pot-herb. Madras Agr. Jour. 40:
Thy Se 1953>
TEoporo, N. G. The Talinum: its culture er me. Philipp. Jour. Agr. 9: 395~
399. 4 pls. 1938. (Farmers Circular 4
Ware, S. A new Talinum (Portulacaceae) eae the cedar ‘em of middle
Tennessee. Rhodora 69: 466-475. 1967. [T. calcaricu
. On the ecology of Talinum Mengesii ee cetilaraces\: Bull. Torrey
Bot. Club 96: 4-10.
. Ecological role of Talinum (Portulacaceae) in cedar glade vegetation.
Ibid. 96: 163-175.
& UARTERMAN. Seed germination in two species wei Talinum (Por-
tulacaceae). (Abstr.) ASB Bull. 14: 44. 1967. [T. calcaricum, T. Mengesit;
require cold treatment (6 weeks), light, alternating “panes ale s. |
Wuerry, E. T. General notes: Talinum in Virginia. Claytonia 1: 55, 56. 1935.
(T. teretifolium.]
Witson, P. Talinum. N. Am. Fl. 21: 280-289. [31 spp. recognized. ]
Wo tr, W. Notes on Alabama Plants. Am. Midl. Nat. 6: 151-158. 1920. [T.
Mengesii.]
The status of Talinum in Alabama. /bid. 22: 315-332. pls. 1, 2. 1939.
Wricur, V. K. Competition on granite outcrops between two species of Talinum.
(Abstr.) ASB Bull. 16: 72. 1969. [T. teretifolium, T. Mengesii.]
584 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Subfamily MONTIOIDEAE Franz
3. Claytonia Linnaeus, Sp. Pl. 1: 204. 1753; Gen. Pl. ed. 5. 96. 1754.
Small perennial [or annual] unbranched herbs from deep-seated corms,
[branched or unbranched tap-roots, rhizomes or stolons], producing 1 to
many succulent, scapose flowering stems. Leaves basal, 1 to many in a
rosette, petiolate [or sessile], + fleshy, glabrous, exstipulate, narrowly
linear to lanceolate, oblanceolate, elliptic, spatulate, narrowly ovate, or
cellate flowers, the pedicels often spreading or recurving in fruit. Sepals
2, free, ovate, rounded, obtuse or acute at apex, persistent. Petals 5 [or
fewer | slightly united basally, broadly oblong to obovate, exceeding the
sepals (or exceeded by the stamens and sepals in C. virginica {. micro-
petala Fern.), apex rounded to truncate or slightly emarginate, pink to
white, with pink veining, rarely yellow; convolute in the bud, drying in
place. Stamens 5, opposite the petals; filament adnate to the petal claw;
anthers pink; pollen mostly 3-colpate. Gynoecium 3-carpellate; style 1,
with 3-cleft stigma; ovary superior, often 3-angled, unilocular usually with
6 (3-6) ovules on a basal placenta. Fruit a membranaceous, ovoid cap-
sule dehiscing by 3 valves, the valves often inrolling after dehiscence.
Seeds shiny, brown to black, smooth to alveolate, orbicular to lenticular.
Embryo with 1 or 2 cotyledons, peripheral, curved around the endosperm.
LECTOTYPE SPECIES: C. virginica L.; see Britton & Brown, Illus. Fl. No.
Us. ed. 2.52: 37. 1913, (Named for John Clayton, 1686(?)-1773,
English doctor and botanist, resident of eastern Virginia from 1705, who
contributed material to Gronovius for the Flora Virginica.) — SPRING
BEAUTY.
A distinctly North American genus of about 32 species, when con-
sidered in the broad sense of Gray or Poellnitz, or of about 20 species
when interpreted in the narrower sense of Greene, Robinson, Rydberg,
Pax & Hoffman, Swanson, and Nilsson. Two complex species (C. virginica
L. and C. caroliniana Michx.) of sect. CLayTonta occur in eastern North
America. The remaining species, including C. lanceolata Pursh, C. tube-
rosa Pallas ex Willd., and C. umbellata S. Wats., closely related to ours,
are distributed from the Rocky Mountains, westward to the Pacific
Ocean, northward to Alaska, and westward to eastern Siberia and the
Kamchatka Peninsula.
Both species of our area are cormose perennials inhabiting shady wood-
lands, grassy banks, wet meadows, and bog habitats or in association
with bald granite exposures (in South Carolina, plants of this habitat are
more robust than those of adjacent habitats). The two differ mainly in
leaf shape and size. Claytonia virginica (spring beauty, rose elf, grass
flower), with two varieties and two forms, has linear to linear-oblanceolate
——— 5
_
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 585
leaf blades more than eight times longer than broad (Davis, Voss) and
gradually narrowed to the petiole. It occurs from Nova Scotia and New
Brunswick, westward across southern Quebec and Ontario to Minnesota,
and southward to Texas and Georgia (Davis). The more northern C.
caroliniana is generally smaller than C. virginica, with possibly darker
flowers, and has spatulate leaves less than eight times longer than broad,
with the blade narrowed abruptly to the petiole. It ranges from south-
western Newfoundland, westward through southern Quebec and Ontario
and the northern parts of the border states to eastern Minnesota, south-
ward through Nova Scotia to New England and along the Appalachians to
western North Carolina and northern Georgia, and westward through
Kentucky and Tennessee to northwestern _Arkansa as. Davis says that
C. virginica blooms earlier.” Voss, however, describes sites in central
Michigan in which populations of the two species are thoroughly inter-
mixed. Here their flowering periods overlap, but C. caroliniana reaches
its peak of flowering slightly earlier than C. virginica. No obvious evi-
dence of hybridization was seen, however. Uttal reports a hybrid swarm
between the two species at a location in Virginia where highly disturbed
conditions have broken down the ecological barriers usually separating
them there.
Narrow-leaved and broad-leaved taxa are recognized within Claytonia
virginica by Davis, Fernald, Gleason & Cronquist, and Lewis et al. This
variation is apparently correlated with chromosome numbers. At least
45 different chromosomal races are known in C. virginica, with numbers
ranging from 2m = 12 to about 191. Rothwell, and Lewis et al., have
worked out the chromosomal evolution of C. virginica throughout its
range. Lewis proposes that “C. virginica has evolved from an ancestral
narrow-leafed race having » = 6 from which the widespread m = 12 +
and southern m = 7 races arose. From the latter was derived an n = 14
race also common in the south. These races and at least some of their
higher polyploid derivatives make up the narrow-leafed var. acutiflora
(= var. Simsii of Davis). From continuing autoaneuploidy at the diploid
level evolved the n =~ 8 race where, we believe, particular chromosomal
redundancies with certain genetic combinations or duplications lead to
the expression of broad leaves. Such plants became widespread in the
north as did the morphologically similar m = 16 + race. These races and
their derivatives, largely aneutetraploids, represent the more id
evolved broad-leafed var. virginica. We agree with Davis . . . that
other gross morphological features correlate with leaf width.”
Similar lines of evolution in chromosome number, although less exten-
sive, are also known in Claytonia caroliniana L., in which reported
chromosome numbers range from 2” = 16 to 38, and in the western C.
lanceolata, 2n = 16 to 72.
The generic and subgeneric classification of Claytonia is chaotic. Two
to ten infrageneric categories, designated or incorrectly cited as sub-
genera and sections, have been recognized. The nomenclature is con-
586 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
fusingly intertwined with that of the closely related Montia L. and various
segregate genera. The boundary between the two is indistinct, prompting
numerous transfers of species and sections. The Claytonia-Montia com-
plex has been interpreted in three divergent ways, all of which have found
some support in recent literature.
Claytonia has been distinguished from Montia either on meristic
grounds (e.g., by Linnaeus, Gray, Poellnitz) or on the basis of habit
the former case Claytonia is said to have
5 more or less equal petals slightly connate basally, usually 5 stamens,
and capsules with several seeds (up to 6), while Montia is distinguished
by distinctly unequal basally connate petals, 3 stamens, and 2 or 3
ovules and seeds per capsule. In this “broad” sense Gray (1887) recog-
nized twenty species of Claytonia, but only one of Montia (M. fontana
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Torrey & Gray (1838) recognized four sections in Claytonia (Clay-
tonia, Limnia (Haw.) Torr. & Gray, Alsinastrum Torr. & Gray, Nato-
crene Torr. & Gray). Much later (1887) Gray divided the species of the
genus among two subgenera or sections (it is not clear which) and 7 sub-
groups based on habit (‘‘section” Euclaytonia with subgroups Cormosae,
Caudicosae, and Rhizomatosae; “section” Limnia (Haw.) Torr. & Gray,
with subgroups Limnia, Alsinastrum Torr. & Gray, Naiocrene Torr. & Gray,
and Montiastrum Gray).
In the subsequent literature these subgroups are referred to as sections,
although Gray apparently did not designate them as such. More recently,
Poellnitz placed 32 species of Claytonia in 10 sections and recognized only
five species of Montia. This restricted view of Montia finds support in
recent papers by Moore and by Walters who recognize only M. fontana
L., with four subspecies, distinguished by characters of the seed coat, in
Europe, North America, and Australia.
A second approach to the complex is to transfer certain sections of
Claytonia to Montia. Swanson, in pointing out transitional floral types
in both genera, notes that all Montioideae are basically pentamerous and
draws generic and sectional lines primarily on plant habit (as did Greene
and others before him), rather than on a meristic basis. In Swanson’s nar-
rower circumscription Claytonia, with about 20 species in four sections
(Caudicosae (Gray) Poell., Rhkizomatosae (Gray) Poell., Claytonia, and
Limnia (Haw.) Torr. & Gray), becomes limited to those perennial and an-
nual species forming an unbranched basal rosette from a tuber, tap
root, or rhizome, and producing upright, axillary, scapose flowering
branches bearing a single pair of opposite leaves subtending the generally
simple inflorescence (“section” Euclaytonia and “section” Limnia-Lim-
nia of Gray). The remaining species (subgroups Naiocrene, Alsinastrum,
and Montiastrum of Gray) are transferred to Montia, which is then
characterized by plants of varying habit which are branched above the
base, with cauline leaves and axillary or terminal inflorescence. Swanson
recognizes eight species of Montia, in four sections (Montia, Limnalsine
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 587
(Rydb.) Pax. & Hoffm., Naiocrene (Torr. & ah Pax & Hoffm., Mon-
tiastrum (Gray) Pax & Hoffm.), in North Americ
The third approach to the complex is that of Rydberg (followed by
Pax & Hoffmann) who also restricted Claytonia, recognizing only two sub-
genera (Euclaytonia and Belia (Steller) Rydb., treated as sections by
Pax & Hoffmann). In contrast to previous workers, however, Rydberg
established segregate genera for the sections or species excluded from
Claytonia (Crunocallis Rydb., Limnalsine Rydb., Limnia L., Montiastrum
(Gray) Rydb., and Naiocrene (Torr. & Gray) Rydb.). Such splitting has
been supported and furthered by Nilsson, who segregated three new
monotypic genera primarily on the basis of pollen morphology (Maxia O.
Nils., Mona ©. Nils., Neopaxia ©. Nils.), raising the number of genera
in the complex to ten.
Swanson considers the members of sect. CAUDICOSAE, with heavy tap-
Phology (decreasing size, increasing asymmetry, increase in satellite
number) support Swanson’s sequence. Their data suggest at least two
lines of evolution in Claytonia, “from sect. Caudicosae through sect.
Rhizomatosae to sect. Claytonia, and from a taxon similar to C. sibirica
(2n = 12, 24, 36, 48; in sect. Caudicosae) to sect. Limnia.”
Flower color in our species ranges from white to dark pink, with pink
to rose veining. In Claytonia virginica a yellowish blotch often occurs
above the claw of the petal. Orange-yellow-flowered variants are re-
ported in C. virginica (f. lutea R. J. Davis) from Maryland and Pennsyl-
vania, as well as in the western C. Janceolata from Idaho and Washington.
Flowering occurs in the spring for two to three weeks, beginning in late
March in Kentucky, with a new flower opening every day and lasting
for 2-3 days. Anthesis is apparently controlled by temperature, with
not more than two flowers open at the same time on an inflorescence
(Wood). The plants are apparently outbreeding, adapted to insect pol-
lination by Dipterans and Hymenopterans (Lovell), but self-pollination
occurs if the flowers fail to open due to poor weather conditions. Flowers
of C. virginica are proterandrous, being functionally staminate the first
day, pistillate the second and third days. The life cycle of Andrena
erigeniae (Hymenoptera), a common short-tongued bee pollinator that
appears and disappears with the flowers of C. virginica, may be keyed to
that of the plant species. Seed dispersal through forcible ejection has
been observed in C. alsinoides Sims and C. sibirica L., with the seeds
shooting out 1—1.5 meters.
Pollen of Claytonia (sensu Greene and Swanson) is mostly tricolpate,
with some grains 6-rugate, as opposed to polyrugate or dodecacolpate in
those former species of Claytonia which are now placed in Montia or the
various segregate genera. Pollen-grain size in C. virginica has been found
to be slightly larger in tetraploids than in diploids.
588 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Anatomical and developmental aspects of the embryo sac, embryo,
seed, and young sporophyte of Claytonia virginica are reported in detail
by several workers. This species differs from other members of the genus
in usually having only one developed cotyledon in the embryo. In the
seedling, formation of the corm begins in the first year through abnormal
development of pericyclic cells of the primary root. The corm soon be-
comes covered with a protective layer of cork.
The genus is of no particular economic significance. In parts of West
Virginia Claytonia caroliniana is known as “tangle-gut.” The leaves are
eaten as ‘“spring-greens” after being steeped in hot grease. Leaves of C.
perfoliata Donn ex Willd. (2n = 12, 24, 36) are eaten like spinach. A
tea was made for use as a diuretic from leaves of C. sibirica L. The starchy
tubers or taproots of C. virginica, C. caroliniana, and C. acutifolia Pallas
are edible
REFERENCES:
Under family references see BRANDEGEE, DAvis, EICHLER, ErpDTMAN, GRAY,
HoweELt, JoHANSEN, Kowa, LuppocK, Martin, NiLsson, RapForp et @l.,
RIcKETT, ScHMUTZ et al., UpHor, and Wonvpart & MApry
AtiosHina, A. L. 1963. Morphology of pollen grains in the genus Claytomia
Gronoy. and allied genera. (In Russian). Bot. Zhur. 48: 1191-1196. 1963.
Coox, M. T. The development of the embryo-sac and embryo of Claytonia
virginica. Ohio Nat. 3: 349-353. 1903.
Davis, R. J. The North American perennial species of Claytonia. Brittonia
18: 285-303. 1966. [Only recent treatment of our perennial species. ]
& R. G. Bowmer. Chromosome numbers in Claytonia. Brittonia 18:
37, 38. 1966.
FENZL, E. Monographie der oa yom Raggy ao Abhandl. Ann.
Wiener Mus. Naturges. 1 & 2: 245-3 a
FREEMAN, O. M. Notes on some plant peak ae in “Greenil and Pickens
counties, South Carolina. Castanea 23: 46-48. 1958. [C. virginica.]
Graves, J. A. Does Claytonia develop during the hen months? Asa Gray
Bull. 5: 17. 1894.
Gray, A. A revision of some polypetalous genera and orders. Proc. Am. Acad.
Arts Sci. 22: 270-314. 1887. [Claytonia, 273, 278-284
GREENE, E. L. Flora Franciscana. 1-2: 177-180. 1891.
———. Miscellaneous specific types —1. Leaflets 2: 45-48. ee [Orange-
flowered form, C. chrysantha from Mt. Baker, Wash., 45
Haccius, B. Embryologische und _histogenetische studie en an 5 Panakeesia
Dikotylen. ” I. Claytonia virginica L. Osterr. Bot. Zeitschr. 101: 285-
303.
Hoim, T. Claytonia Gronov., a morphological and anatomical study. Mem.
Natl. Acad. Sci. 10: 25-37. 1905.
Lepesour, C. F. Fl. Ross. 2: 146-151. 1844-46. [Claytonia, by E. Fenz
pe W. H. Aneusomaty in aneuploid populations of Claytonia virginica.
Am. Jour. Bot. 49: 918-928. 1962. [Map.
——. Bo ionge sou evolution in plants. Bot. Rev. 33: 105-115. 1967.
mmary discussion of chromosome numbers and races and their geo-
paler es distribution in species of Claytonia. |
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 589
———=, KK. L. Orrver & ¥. Suna, cig areas a . a virginica and
its allies. Ann. Missouri Bot. Gard. 54: 153-1
—— & Y. Supa. Karyotypes in relation to Ce Bees and phylogeny in
Claytonia. Ibid. 55: 64-67. 1968.
. SupA, & B. MacBrype. Chromosome numbers of aia vir-
ginica in the St. Louis, Missouri, area. /bid. 54: 147-152.
Lovett, H. B. The life story of three spring wild flowers. Wild Le 19: 61-
64. pls. 10, 11. 1942. [C. virginica, Mertensia virginica, Jeffersonia
Marroguin, A. S., & H. A. WinTER. Un estudio morfologico de la plantula
de Claytonia virginica L. (English summary). Ann. Escuela Nac. Cienc
Biol. Mex. 2: 191-215. 1940. [See Biol. Abstr. 16: no. 18827. 1942.]
Moore, D. M. The subspecies of Montia fontana L. Bot. Not. 116: 16-30.
1963.
Nixsson, 6. Studies in Montia L. and Claytonia L. and allied genera, 1. Tw
new genera, Mona and Paxia. Bot. Not. 119: 265~285. 1966; Scie
and additions. Ibid. 469; 2. Some chromosome numbers. Ibid : 464-468;
3. Pollen morphology. Grana Palynol. 7: 279-363. 1967. [Maxia O. Nils.,
359, 360.
Oswatp, F. W. An abnormal form of spring-beauty. Phytologia 5: 50, 51.
1954
POELLNITZ, K. von. Claytonia Gronov. and Montia Mich. Repert. Sp. Nov.
30: 279-325. 1932. [Monograph. |
Raymon, M. Le Claytonia virginica L. dans le Québec. Nat. Canad. 76: 201-
204. 1949, [Distribution and ecological notes on C. virginica and C. caro-
liniana in Quebec. ]
ReEep, R. M. “Tangle-Gut.” Castanea 28: 177. 1963. [C. caroliniana.]
Ronson, . — Claytonia. In: A. Gray et al. Syn. Fl. N. Am. 1(1): 270-
272. :
ROLLINS, = ic Orange-yellow-flowered Claytonia virginica. Rhodora 60: 258,
259. 19
ROTHWELL, x, V. Aneuploidy in Claytonia virginica, Am. Jour. Bot. 46: 353-
360
: fe Kump. Chromosome numbers in populations of Claytonia vir-
ginica from the New York metropolitan area. Am. Jour. Bot. 52: 403-407.
1965.
RypgerG, P. Studies in the Rocky Mountain flora, 16. Bull. Torrey Bot. Club
3: 137-161. 1906. [Separates Limnia L., Crunocallis Rydb., and Naio-
crene (Torr. & Gray) Rydb., from Claytonia a. |
Flora of the Rocky Mountains and adjacent plains. xii + 1110 pp.
New York. 1917. [Separates Montiastrum (Gray) Rydb. from Claytonia. |
Sus, J. Claytonia — Bot. Mag. 24: pl. 941. 1806.
Somes, M. P. An variety of plant Iowa Nat. 2: 67, 68. 1909.
Sovéces, R. Sei gautne des Portulacacées. Développement de l’embryon
chez le Claytonia perfoliata Donn. Compt. Rend. Acad. Sci. Paris 221:
111-113. 45.
Star, A. E. Leo ins induced meiotic chromosome breakage in Clay-
tonia virginica L. . Abstr. B. 28(5): 1811B. 1967
Swanson, J. R. A panne of relationships in Montioideae a
Brittonia 18: 229-241. 1966. [Redefinition of generic limits and sub
generic categories in Claytonia and Montia.
590 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Torrey, J., & A. Gray. Fl. N. Am. 1(1): 198-202; Suppl. 676, 677. 1838.
{Species grouped in four “sections”: Claytonia, Limnia, Alsinastrum,
Naiocrene. |
Urrat, L. J. A hybrid population of Claytonia in Virginia. Rhodora 66: 136-
139. 1964.
Voss, E. G. The spring beauties (Claytonia) in Michigan. Mich. Bot. 7: 77-
93. 1968. [Excellent account of distribution and variation in Michigan. |
Watters, S. M. Montia fontana L. Watsonia 3: 1-6. 1953. [M. fontana with
four subspecies based on seed-coat morphology, in northwestern Europe. |
Woopcock, E. F. Morphology of the seed in Claytonia virginica. Pap. Mich.
Acad. Sci. 5: 195-200. pls. 19, 20. 1926.
Wutts, J. C. The distribution of the seed in Claytonia. Ann. Bot. 6: 382,
383. 1892.
BASELLACEAE Moquin-Tandon, Chenopod. Monogr. Enum. x. 1840, nom. cons.
( MADEIRA-VINE FAMILY)
A small family of herbaceous, somewhat succulent, glabrous, ( ?dextrorse-
ly) twining vines, lax herbs [or subshrubs?] producing annual shoots
from perennial, fleshy rhizomes or tubers; leaves fleshy, alternate, ex-
stipulate, sessile or petioled; inflorescence an axillary or terminal raceme,
spike, or panicle of numerous small flowers, each subtended by a small
bract; bracts of the pedicel 2, small, opposite, membranaceous or fleshy,
caducous or persistent; flowers regular, bisexual [or unisexual]; sepals
(involucral bracts?) 2, free or basally connate; petals (tepals?) 5, mem-
branaceous or slightly fleshy, basally connate to form a shallow floral cup,
aestivation quincuncial or imbricate; stamens 5, inserted on lip of floral
cup or on bases of the petals; filaments [erect or] recurved in bud; an-
thers 4-locular, insertion of filament basal or versatile; gynoecium of 3
united carpels; ovary superior, unilocular; ovule 1, basal, campylotropous
to anacampylotropous; styles 3, basally united, stigmas slender, + bifid
[or capitate to clavate, or style 1 with capitate stigma entire to 3-lobed];
fruit a utricle [or berry], included in the perianth; seed 1, with copious
endosperm; embryo annular [or spirally twisted]. Tyee cenus: Basella
L
A family of four or five genera containing 15—20 species, most native
to the New World tropics or the Andean regions of South America. Ba-
sella, including about five species and thought to have originated in the
tropics of the Old World, has probably achieved its present pantropic
distribution through cultivation. Of the New World genera, Tournonia
Mog. and Ullucus Loz. are monotypic, Anredera Juss. (sensu stricto) is
generally considered to be monotypic, and Boussingaultia HBK. consists
of 10-15 species. Amredera and Boussingaultia were united by Baillon,
and more recently by van Steenis, under Anredera. The family is rep-
resented in the southeastern United States only by a single species of
Anredera (A. leptostachys), of the tribe Boussingaultieae Benth. & Hook.
The genera of Basellaceae fall into two natural groups: those with
spirally twisted embryos and stamen filaments erect in the bud (Basedla,
Tournonia, Ullucus) and those with annular embryos and filaments out-
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 591
wardly reflexed in the bud (Anredera, Boussingaultia). These two groups
have consistently been given systematic recognition as tribes, subtribes,
or series, when included in the Chenopodiaceae (Endlicher, Bentham &
Hooker, Baillon), or as subfamilies or tribes of the Basellaceae (Moquin-
Tandon, Engler, Ulbrich, Eckardt). Franz treated the Baselleae as a
tribe of subfam. Montioideae in the Portulacaceae and included the five
genera of Basellaceae and the transitional genus Portulacaria Jacq.
Anatomically the Basellaceae differ from Chenopodiaceae in the ab-
sence of anomalous secondary growth and in the presence of bicollateral
stelar bundles resulting from the tardy development of internal phloem.
Bicollateral bundles, and similar features of floral ontogeny and mor-
phology, are shared with some Montioideae (Portulacaceae).
Observations on Basella rubra L. and Anredera vesicaria (Lam.) Gaertn.
f. indicate that the Basellaceae share with other Centrospermae possession
of betacyanin pigments (here basellain-r, basellain-v) in place of antho-
cyanins. Saponins have been found in the seeds of this species, while
calcium oxalate occurs in the form of druses and single crystals. Slime
cells occur in parenchymatous tissues of the various taxa, and stomata of
the rubiaceous type occur on both surfaces of the leaves.
As in the Portulacaceae, the perianth is interpreted as either biseriate,
with two sepals and five petals, or uniseriate, with a single cycle of five
sepals (or tepals) subtended by two large involucral bracts. The posi-
tion of the five stamens in opposition to the five ‘“‘sepals’” suggests the
loss of either the corolla or an intervening cycle of stamens, most probably
the latter. According to Payer, and Eichler, the floral plan in Basella
rubra is basically trimerous, the present pentamerous cycles each arising
as successive whorls of three, one member of the outer perianth and androe-
cial cycles aborting. Sharma considers the basal placentation of B. rubra
to represent an extreme reduction from an originally axile placentation.
Embryological characteristics of the family include bitegmic ovules in
which the inner integument forms the micropyle. Within a bulky nu-
cellus the chalazal megaspore of a linear tetrad develops into Polygonum-
type embryo sac. The endosperm is initially nuclear, later becoming
cellular. Anthers of the family are tetrasporangiate and produce a
glandular tapetum of multinucleate cells. Pollen grains are three celled
when shed. ;
The pollen of the family is polymorphic. Grains are spheroidal in all
genera except Basella, in which elaboration of the exine in the inter-
apertural areas has produced a cubical shape. Aperture configuration
varies from a basic pattern of six furrows arranged as on the sides of
a cube (Basella, some species of Boussingaultia), through reduction in
size and increase in number of apertures (as in “Boussingaultia lepto-
stachys,” Ullucus tuberosus), to many pores distributed evenly over the
surface of the grain (Amredera vesicaria). Similar patterns occur in Por-
tulacaceae and other families of Centrospermae.
Knuth states that pollination in Basella rubra is largely cleistogamous,
with a few chasmogamous flowers pollinated by small, short-tongued in-
592 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
sects. Observations on the floral biology of American taxa are lacking.
Structures resembling berry-like fruits found in an inflorescence of Anre-
dera leptostachys from southern Texas were originally mistaken for fruits,
but proved on dissection to be the much enlarged, fleshy petals enclosing
an insect larva. The . attack apparently stimulates this aberration
of growth in the host flow
Chromosome counts a species of three genera suggest that the base
number of the family is 12, with 2n = 24, 48, and 60 in Basella, 24 and
28 in Boussingaultia, and 24 or 36 in Ullucus.
The family is of economic importance in that several species are culti-
vated for their fleshy leaves, which are used as a substitute for spinach
(Basella rubra), or for their fleshy, starchy rhizomes or tubers (U/lucus,
Anredera vesicaria, species of Boussingaultia). The tubers of Ullucus
tuberosus are an ancient and important food crop in Bolivia, Chile, Colom-
bia, Peru. Basella rubra and some species of Anredera and Boussingaul-
tia which are cultivated as ornamentals have become naturalized in many
places. The reddish pigments of Basella rubra fruits are used in Asia as
food coloring and may serve as a substitute dye for carmine.
The dextrorsely twining habit often cited in descriptions of the Basel-
laceae is open to question. Nevling notes that this characteristic is sel-
dom constant among climbers, and Sloane’s illustration of Anredera
vesicaria shows leftward twining stems.
REFERENCES:
Acostis Sorts, M. Ullucus tuberosus. Estudio botanico-morfolégico, micro-
grafico, farmacognésico, quimico y aplicative del melloco. Anal. Univ
Ecuador 57: 263-276. 1936.*
Barton, H. Chenopodiacées. Hist. Pl. 9: 130-217. 1888. [Series Baselleae,
145-148, 197, 198
BeILteE, M. L. Organogénie florale du Boussingaultia baselloides. Actes Soc.
Linn. Bordeaux 56: CLVI. 1901. [Includes vascular anatomy.
BentHam, G., & J. D. Hooker. Chenopodiaceae. Gen. Pl. 3: 43-78. 1880.
[Subfam. Baselleae, 48; tribe Eubaselleae, tribe Boussingaulteae, 76-78.]
Branpt, W. Ein Beitrag zur vergleichenden vos bnahiaaaa der Centrospermen.
Festschrift A. Tschirch, 13-22. Leipzig.
Davis, G, L. Systematic embryology of the angiosperms. 528 pp. New York.
1966. [Basellaceae, 55.]
DecatsNE, J. Ullucus tuberosus. Revue Hort. III. 2: — 1848. [Color
plate, description, discussion of cultivation and distribution. |
Diers, L. Der Anteil an Polyploiden in den Verdsteaspictats der West Kor-
dillere Perus. Zeitschr. Bot. 49: 437-488. 1961. [Diploid and tetraploid
chromosome numbers in Boussingaultia sp., B. diffusa, Basella alba, 449,
452, 453, 455.]
ECKARDT, T. Basellaceae. Jn: H. Metcutor, Engler’s Syllabus der Pflanzen-
familien. ed. 12. 2: 92. 1964.
ErcHier, A. W. Bliithendiagramme 2: 128, 129. fig. 48. 1878. [Basellaceae;
mainly on Basella rubra.|
ENGLER, A. a der Pflanzenfamilien. ed. 5. 248 pp. Berlin. 1907. [Basel-
laceae, 124.]
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 593
Franz, E. Beitrage zum Kenntnis der Portulaceen und Basellaceen. Bot. Jahrb.
42(Beibl. 97): 1-48. [1908] 1909. [Morphology of inflorescence, flower,
pollen; vegetative anatomy. |
HEGNAUER, R. Chemotaxonomie der Pflanzen. Band 3. Dicotyledoneae:
Acanthacese- Cyrillaceae. 473 pp. Basel & Stuttgart. 1964. [Basellaceae,
Hopce, W. H. Three neglected Andean tubers. Jour. N.Y. Bot. Gard. 47:
214-224. 1946. [Ullucus tuberosus and other tuber-bearing crop plants of
the Andean Indians
KnutH, P. Handbuch “der Bliitenbiologie 3(1): 280, 281. Leipzig. 1904.
[ Basella rubra. |
MacBripg, J. F. Flora of Peru. Pt. 2(2). Field Mus. Publ. Bot. 13(2). 1936.
[Basellaceae, 573-577; notes on cultivation of various species by the Andean
Indians, incl. Basella, "Ullucus, Boussingaultia, Anredera
Moourn-Tanpo DON, A. Chenop udearent ta monographica enumeratio, xi + 1 p.
Paris. 1840. [Establishes the family Basellaceae (p. x), eeaciies it
from Chenopodiaceae, Amaranthaceae, Portulacaceae; gives a conspectus of
three genera.
————.. Basellaceae. DC. Prodr. 13: 220-230. 1849. [Recognizes two subfam-
ilies and six genera
Moror, Note sur + Panauateit des Basellacées. Bull. Soc. Bot. France 31:
104-107. 1884, [Ontogeny of bicollateral bundles in Basella rubra, Bous-
singaultia baselloides, Ullucus tuberosus.
Nakal, T. Notulae ad Plantas Asiae Orientalis (18). Jour. Jap. Bot. 18: 91-
120. 1942. [Establishes a new suborder Baselliineae (Chenopodiales), con-
taining Basellaceae Mogq., and new segregate family Ullucaceae Nak. for
Ullucus Loz
NESTERENKO, P. A. Pigment extracted from berries of Basella rubra as a
i 1936.
NeEvutno, L. I., Jr. Some ways plants climb. Arnoldia 28: 53-67. 1968, [An-
redera cordifolia, 54.]
Payer, J. B. Traité d’organogénie comparée de la fleur. vii + 748 pp., 145 pls.
Paris. 1857. [Basellaceae, 313-316, pl. 75; Basella rubra.
ScHouTe, J. C. On corolla aestivation and phyllotaxis of floral phyllomes.
Verh. Akad. Wet. Amsterdam Afd. Natuurk, 1. 34(4): 1-77. 1935a. [Ba-
sellaceae, 8.]
. On the perianth aestivation in the Portulacaceae and the Basellaceae.
Rec. Trav. Bot. Néerl. 32: 396-405. 1935b.
SHarMA, H. P. Contributions to the morphology and anatomy of Basella rubra
Linn. Bull. Bot. Soc. Bengal. 15: 43-48. 1961. [Detailed study of vege-
tative and floral anatomy. |
SLOANE, H. Nat. Hist. Jamaica 1: 138. pl. 90, fig. 1. 1707. [Fegopyrum scan-
ens = Anredera vesicaria.
STEENIS, C. G. G. J. van. Basellaceae. Fl. Males. I. 5: 300-304. 1957. [Ba-
sella, Anredera incl. Boussingaultia.
Uxsricu, E. Basellaceae. Nat. Pflanzenfam. ed. 2. 16c: 263-271. 1934.
VOLKENS, G. Basellaceae. Nat. Pflanzenfam. III. la: 124-128. 1893. Ergan-
zungsheft II: Nachtrage III zum Teil II-IV:105. 1908
Witson, P. Basellaceae. N. Am. Fl. 21: 337-339. 1932
Woutpart, A., & T. J. Masry. The distribution and phylogenetic significance
of the betalains with respect to the Centrospermae. Taxon 17: 148-152.
1968.
594 JOURNAL OF THE ARNOLD ARBORETUM [vor. 50
1. Anredera A. L. de Jussieu, Gen. Pl. 84. 1789.8
Twining or scrambling, herbaceous vines with slender, much branched,
somewhat fleshy, glabrous and sometimes reddish stems produced an-
nually from fleshy rhizomes or tubers. Leaves alternate, exstipulate,
petioled [or sessile], slightly fleshy, entire margined, blades suborbicular
to elliptic, ovate [or cordate], with apex acute to acuminate [or obtuse],
and base gradually or abruptly narrowed [to truncate or cordate]. In-
florescence axillary or terminal, a simple nodding raceme, racemose spike,
or panicle, with numerous small, sessile or pedicellate flowers in axils of
small bracts. Bracts of the pedicel 2, small, opposite, free or deciduous
[or basally connate and persistent at level of pedicel articulation]. Flow-
wale
ash
pik
-
separa
gor
wee
‘
Fic. 3. Anredera. a-d, A. leptostachys: a, part of stem with inflorescences,
X 1/2; b, flower, X 12; c, partial vertical section of gynoecium to show ovule,
xX 24; d, teratological flower, the tepals accrescent from insect attack — note
thickened pedicel,
cording to Rickett (1960), the investigation of a proposal to conserve the name
A Mis Juss. (1789) over — Adans. (1763) has shown that “the actual type
of Anredera (Sloane’s specimen j
th . genus is roan not identical with Fall
even taxonomic synonyms. Anredera therefore stands without conservation, so far as
7 ”
Fallopia is concerned.
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 595
ers fragrant, perfect [or imperfect]. Sepals 2, + adnate to the floral cup,
nearly flat to boat-shaped [keeled or narrowly to broadly winged along
the back]; slightly shorter than [to slightly exceeding| the petals at
anthesis. Petals 5, small, white or greenish [turning purple at maturity in
some species] thin or somewhat fleshy, united basally to form a short floral
tube, more or less spreading at maturity. Stamens 5, opposite and inserted
on the petal bases. Filament filiform or subulate, recurved in the bud. An-
thers oblong [to ovate], versatile, pollen rugate or polyforate. Ovary
small, superior, ovoid [or slightly compressed] 1-locular. Styles 3, free,
with bifid, papillose stigmas [or styles variously fused below, with
capitate to clavate stigmas, or style 1, with capitate, 3-lobed stigma].
Ovule 1, basal, subsessile [or sessile]. Fruit a utricle with fleshy or
parchment-like pericarp, enclosed by the perianth. Seed erect, lenticular,
with crustaceous [or coriaceous] seed-coat. Embryo semiannular [to
annular]. Cotyledons plano-convex [or subclavate]. Type species: An-
redera spicata J. F. Gmel. = Anredera vesicaria (Lam.) Gaertn. f.; see
Gmel. Linn. Syst. Nat. 2: 454. 1791, and P. Wilson, N. Am. FI. 21: 337.
1932. (Derivation of generic name unknown.) — MADEIRA-VINE.
A New World genus of 10-15 species, inhabiting tropical regions from
southern Florida and southernmost Texas, through Central America
(about three species), to northern South America (about 13 species) ; rep-
resented in our area only by Anredera leptostachys (Moq.) Steenis in
southern Florida, and in southern Texas by both A. leptostachys and A.
vesicaria (Lam.) Gaertn. Anredera leptostachys is distinguished by its
three bifid styles and wingless sepals, while 4. vesicaria, better but er-
roneously known as A. scandens (L.) Mogq., has broadly winged sepals
and three undivided styles. Both species belong to sect. ANREDERA (see
below) and may be most closely related to the South American A. cordi-
folia (Tenore) Steenis and Boussingaultia floribunda Mogq.
The distribution of Anredera leptostachys in our area is not well de-
fined, but it is known from Dade, Monroe, and Collier counties, Florida.
The best known location appears to be Key West, where the species has
been collected in hammocks and where it is said to occur in vacant lots
and fence rows.
In his original circumscription of the Basellaceae Moquin-Tandon
(1840) recognized two genera in his subfamily Anredereae Endl.: Anre-
dera Juss., characterized by winged sepals (“perigone”) and Boussin-
gaultia HBK. with wingless sepals. He later (1849) segregated a third
genus, Tandonia Mogq., to accommodate those species of Boussingaultia
distinguished primarily by ovaries bearing a single style with capitate
stigma. Bentham and Hooker, however, reunited the two genera, and
Tandonia has since been accorded only sectional rank.
Volkens divided Boussingaultia into two sections: TANDONIA (Mogq.)
Volk., containing the single-styled species, and EvUBoUSSINGAULTIEAE
Volk., containing the three-styled species. In the latter section Volkens
included the type of the genus, B. baselloides HBK. Hauman, how-
596 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
ever, in the only recent revision of Boussingaultia, pointed out that the
original description of B. baselloides mentions a single style, with a
capitate, three-lobed stigma, and that the flower erroneously figured by
Volkens for the type species is one of B. leptostachys Moq., with three
bifid styles. The type species thus belongs to Volkens’s section TAn-
DONIA. Hauman replaced the name EUBOUSSINGAULTIEAE with sect.
MoguINELLtA Haum. He further clarified some areas of confusion in
identification and nomenclature, and provided a ‘“‘provisional’”’ enumera-
ion of specimens and a key to thirteen species but did not give detailed
descriptions for them. Ulbrich accepted Hauman’s treatment of the
genus.
Baillon, and more recently van Steenis, reduced Boussingaultia to
synonymy under the older Anredera. Van Steenis states that winged
sepals occur in some species of Boussingaultia and that the difference
between the two genera is only a matter of degree in the development of
the wings. In his expanded genus Amredera van Steenis recognizes two
teristics formerly used. Section TANpDoNIA, with bracts basally connate
and persistent on the pedicel, is not changed in composition. Section
ANREDERA, with bracts free and caducous, is formed by the addition
of Anredera vesicaria (as A. scandens) to the former sect. MOQUINELLA
Haum. of Boussingaultia. The systematics of the genus is currently in
a state of flux, since van Steenis transferred only a few species of Bous-
singaultia to Anredera. The remaining species must be cited under Bous-
singaultia until formally transferred. Soukup attempted to transfer
several South American species, but his combinations are invalid for lack
of proper basionym citation. The genus is badly in need of critical study
and revision.
A tendency toward imperfect or functionally imperfect flowers may
exist within the genus. Van Steenis concluded that the flowers of the
type specimens of Boussingaultia baselloides HBK. “appear . . . female
with small barren anthers.” He also states that fruits of Anredera cordi-
folia have never been found and that A. scandens (= A. vesicaria) does
not produce seed in Malaysia. Furthermore, Hauman describes sexually
dimorphic flowers of Boussingaultia ramosa (Moq.) Hemsley, and Heim-
merl describes a collection of “B. gracilis” from southern Brazil in which
“all flowers are female.” The isch and extent of dioecism among the
species of Amredera should be investigated.
Beille described the floral vascular anatomy of “Boussingaultia basel-
loides” (cultivated). He found five vascular bundles in the pedicellar
stele, each bundle supplying a single trace to a “sepal” and its opposing
stamen, and three of the bundles each supplying a single trace to one
of the three carpels. There are no vestigial vascular traces in the in-
tervals between the “sepals,” from which he concluded that the abortion
of the ‘‘corolla” is complete.
1969 | BOGLE, PORTULACACEAE AND BASELLACEAE 597
Chromosome counts for the genus indicate that both diploid and tetra-
ploid species exist: » = 12 for Anredera cordifolia, and n = 24 in Bous-
Singaultia diffusa (Mogq.) Volk.
Anredera leptostachys, A. cordifolia, and A. vesicaria are widely grown
as Ornamental vines for their foliage and fragrant flowers. They can be
propagated vegetatively from their fleshy rhizomes or tubers, or in A.
cordifolia from small tubercles which form in the leaf axils. Webb re-
ports that plants of “Boussingaultia baselloides,’ naturalized in New
South Wales, Australia, are suspected of causing death of cows (symp-
toms of irritant poisoning). Hot-water extracts of these plants proved
A tangle of misidentification has developed around plants of Boussin-
gaultia collected in South America and those cultivated and naturalized
in various areas. Many of these have been identified as B. baselloides
HBK., but they are specimens of Anredera cordifolia (including B. gra-
cilis Miers, B. gracilis {. pseudo-baselloides Haum., and B. baselloides
sensu Hook. Bot. Mag. p/. 3620). Hauman states that very few of the
many specimens labeled B. baselloides which he examined were correctly
identified and that most were attributable to B. gracilis. The latter was
introduced into horticulture during the 18th century and has become
widespread. For this reason references in the literature to B. baselloides
HBK., such as those of Webb, are especially subject to question.
REFERENCES:
Under family references see BAILLON, BENTHAM & Hooker, BEILLE, DIERs,
< T, EICHLER, FRANZ, HEGNAUER, MAcBripe, Moquin-Tanpon, Moror,
ScHoute (1935b), SLOANE, STEENIS, ULBRICH, VOLKENS, WILSON, WOHLPART
BAKHUIZEN VAN DEN BRINK, JrR., R. C., & C. G. G. J. VAN — Nomina
conservanda proposita. Anredera Juss. Taxon 5: 198. 1956.
GAERTNER, J. Fruct. Sem. Pl. 3: 176, 177. pl. 213, fig. " ued 1805.
[ A. vesicaria. }
Hauman, L. Notes sur le genre Boussingaultia HBK. Mus. Nac. Buenos
Aires. Bot. 33: 347-359. 1925. [Revision.]
Hemmmert, A. Basellaceae. In: WETTSTEIN, R. V., & V. SCHIFFNER. Ergebn.
Bot. Exp. Sud-Brasil 233, 234. 1908. [Collection of B. gracilis with all
flowers female. |
Hooker, W. J. Boussingaultia baselloides. Bot. Mag. 64: pl. 3620. 1837. [A.
cordifolia, not B. ceeseeniggeh HBK
Humpotpt, A., A. BoNPLAN 73 Kun Nov. Gen. Sp. Pl. 2: 190. 1817
te quart. [ Anredera aoe y ‘Ibid. 7: 164-196, pl. 645 bis. 1825 [B. basel-
ta : ‘B., C. M. Rowe t, Jr., & M. C. Jounston. Flowering Plants and
Ferns of the Texas Coastal Aas Communities. 1961. [A. vesicaria, B.
Morton, J. F., & R. B. Leprn. 400 Plants of South Florida. 134 pp. Coral
Gables. 1952. [B. leptostachys, 28.]
NEvLING, L. I., Jx. Jn: I. O. P. B. chromosome number reports VI. Taxon 15:
117-128. 1966. [A. cordifolia, e = 327]
598 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
RickeTtT, H. W. Report of the Committee for Spermatophyta. Conservation
of generic names II. Taxon 9: 14-17. 1960. [2428. Anredera Juss., 14;
conservation unnecessary. |
Seppon, H. R., & W. L. HinpMarsu. Boussingaultia baselloides, “\ambs tails,”
a reputed poisonous climber. Jour. Counc. Sci. Industr. Res. Australia 2:
53, 54. 1929.*
Souxup, J. El genero Boussingaultia HBK, fue reducido a sinonomia de Anr
dera Juss. Biota 6: 158, 160. 1966. [Includes five invalid pct la
under Anredera. |
Wess, L. J. Guide to the medicinal and poisonous plants of Queensland. Counc.
Sci. Industr. Res. Australia. Bull. 232. 1948. [B. baselloides HBK., 24.]
THE ARNOLD ARBORETUM
OF
HARVARD UNIVERSITY
1969 } WEAVER, FOTHERGILLA 599
STUDIES IN THE NORTH AMERICAN GENUS FOTHERGILLA
(HAMAMELIDACEAE)
RicHarp E. WEAVER, Jr. !
FOTHERGILLA is a small genus of spring- blooming shrubs in the Ha-
mamelidaceae endemic to the southeastern United States. The first speci-
mens of the genus were collected by Dr. Alexander Garden of Charleston,
South Carolina, and the material was sent by him to Linnaeus. The genus
was named in oe of Dr. John Fothergill, a London physician and pa-
tron of early American botanists. As they usually occur in very localized
clumps, none of the species of Fothergilia is abundant. Hence the genus
is relatively poorly represented in most herbaria.
The genus Fothergilla is one of a considerable number of plant genera
(e.g. Bartonia, Hudsonia, Leiophyllum, Cleistes, Clethra) which show a
disjunct distitbutional pattern in the SGuthenstens United States, i.e., the
genus is well represented by one or more species in both the Atlantic
Coastal Plain and the Appalachian Mountains but is absent or very rare
in the intervening Piedmont. The coastal plain and montane populations
of Fothergilla have long been recognized as being CEE distinct;
however, the morphological variation in the genus is great, and as a result
as many as four taxa, two in the mountains and two in the coastal plain,
have been recognized by the various authors.
HISTORY OF THE GENUS
As indicated by the very short list of “excluded names” at the end of
this paper, the generic limits of Fothergilla appear to be fairly well marked.
*Part of a thesis racer Sa in partial fulfillment “ the requirements for = degree
of Master of Arts at Duke University. Much of the e field wo ork w
valuable in the execution of the project; and Mrs. A
North Carolina, who secured pickled buds for hau investigation. Special thanks
are due Dr. oll E. Wood, Jr., of the Arnold Arboretum, Harvard University, with-
out whose interest, en ncouragement, and material aid in the form of pickled buds from
Fothergilla plants at the Arnold Arboretum the effectiveness of this study would have
it whose
nS were essential to this study: A, DUKE, FSU, GA, GH, MO, NCU, NY, TENN, US,
600 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
However, Parrotiopsis jacquemontiana (Decne.) Rehder, a plant native
to Kashmir and Afghanistan, has been referred to Fothergilla by several
authors. Falconer (Proc. Linn. Soc. London 1: 18. 1839) originally re-
ported the plant, unaccompanied by a description, as Fothergilla involu-
crata. Niedenzu (1891) treated the plant as the sole member of his sub-
genus Parrotiopsis of Fothergilla, but remarked that perhaps it should
be treated as a separate monotypic genus. Decaisne (in Jacquemont, Voy-
age dans l’Inde 6: 73. 1844), Clarke (in Hooker f., Fl. Brit. India 2:
426. 1879), and Hooker (Bot. Mag. 122: f. 7501. 1896) considered the
plant in question to be a species of Parrotia. According to Rehder (Jour.
Arnold Arb. 1: 256. 1920) Parrotiopsis differs from Fothergilla in its capi-
tate rather than spicate inflorescence, subtended at the base by large bracts,
and in its less numerous stamens with linear rather than clavate filaments.
The genus Parrotiopsis was proposed by Schneider (Ill. Handb. Laub-
holzk. 1: 429. 1905), and, to my knowledge, has been accepted by all
subsequent authors.
The number of taxa meriting specific rank in the genus Fothergilla has
been the subject of considerable debate for nearly 70 years. During much
of this period four species have been variously recognized as occurring in
the southeastern United States: (1) F. major Lodd., ranging along the
mountains and inner Piedmont from North Carolina to Alabama; (2) F.
monticola Ashe, occupying the same range as the preceding species; (3)
F. gardenii Murr., occurring along the Atlantic and Gulf Coastal Plains
from North Carolina to Alabama; and (4) F. parvifolia Kearney, occupy-
ing approximately the same range as F. gardenii. It can be seen, therefore,
that, although the range of the montane element does not overlap that of
the coastal plain element, the ranges of the commonly accepted montane
taxa are entirely sympatric as are those of the supposed coastal plain taxa.
To my knowledge all investigators during the last 100 years, with the
exception of Sargent (Garden and Forest 8: 446. 1895) have agreed that
the mountain and coastal plain elements of Fothergilla are so distinct that
the recognition of at least two taxa of specific rank is required.
However, the morphological features distinguishing Fothergilla monti-
cola from F. major are less than convincing, and as a result a number of
investigators have questioned the status of F. monticola as a taxon specifi-
cally, or even varietally, distinct from F. major. The primary morpho-
logical character used to distinguish the two is the presence or absence of
a waxy bloom on the undersides of the leaves, F. major being glaucous
and F. monticola nonglaucous. Differences in habit, dentition, and pubes-
cence of the leaves, and color of the inside of the capsule have been
reported by the various authors. More recently, reports of differing
chromosome numbers (Anderson & Sax, 1935; Thomas, in Ernst, 1963)
have added a new dimension to the problem.
Britton (1905) treated F. monticola as a synonym of F. major. Small
(1903, 1913, 1933) and Radford et al. (1964) made no mention of F.
monticola, and since the plant has been reported as occurring in the South-
east, including the Carolinas, it is assumed that they also rejected it. In-
~b
1969] WEAVER, FOTHERGILLA 601
vestigators who have accepted F. monticola as a distinct species include
Hesse (1909), Rehder (1910), Bailey (1949), Harms (1930), Anderson
and Sax (1935) and Ernst (1963). However, Anderson and Sax (1935)
and Rehder (1910) pointed out that F. monticola might better be treated
as a variety of F. major rather than as a species distinct from it. Ernst
(1963) noted that the two are sympatric and are nearly indistinguishable,
and that the relationship between them is in need of study. It is perhaps
of significance that Ashe (1897), in describing F. monticola, considered it
to be the only species of Fothergilla in the southern Appalachians. In-
deed he states “the only published name that could possibly apply to this
species [F. monticola] is F. major Lodd.” It was Ashe’s opinion, however,
that F. major Lodd. was merely a robust specimen of the coastal plain
species.
The status of Fothergilla parvifolia as a taxon distinct from F. gardenii
is also open to question. The supposed coastal plain taxa reportedly dif-
fer in the shape and dentition of the leaves and in the length of the cap-
sules.
Authors who have accepted Fothergilla parvifolia as a distinct species
include Small (1903, 1913, 1933), Britton (1905), and Harms (1930).
More recent authors, including Ernst (1963) and Radford et al. (1964)
have treated F. parvifolia as a synonym of F. gardenii.
The present paper is an attempt to evaluate, both morphologically and
cytologically, the taxonomy of the genus Fothergilla.
PHENOLOGY
The flowering period of the populations of Fothergilla in the coastal
plain begins in late March in northern Florida and southern Alabama and
continues until mid-May in North Carolina. The peak of flowering in
North Carolina is during the second and third weeks of April. F. alnifolia
(= F. gardenii ) 8 serotina, originally described by Sims (1810) and
taken up by DeCandolle (1830) and Harms (1930) reportedly blooms in
August. I have found no evidence, either from herbarium specimens or
from field observations, of a Fothergilla blooming in August. F. alnifolia
8 serotina was based on a cultivated plant, and its blooming in August
was probably an abnormal occurrence.
The flowering period of the montane Fothergillas is affected by altitude
as well as latitude but generally extends from late March until mid-May
in various parts of their range.
The appearance of the flowers in relation to that of the leaves has been
used by several authors (Small, 1903, 1913, 1933; Britton, 1905) asa
key character for distinguishing the species of Fothergilla. It has been
reported that the flowers of the coastal plants appear before the leaves,
while the flowers of F. major appear as the leaves are unfolding. Observa-
tion of the plants in the field, as well as examination of herbarium speci-
mens, indicates that this is generally true. Although exceptions have been
noted, the distinction remains a fairly usable diagnostic character.
602 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Van Dersal (U.S.D.A. Miscell. Publ. #303, pp. 128, 129. 1938) found
that the seeds do not germinate until the second year, evidently requiring
a period of low temperatures to break the dormancy of the embryo.
Flowers are initiated during the summer growing season and complete
inflorescences are formed before the leaf drop in the fall. Meiosis, how-
ever, does not take place until the following spring.
MORPHOLOGY
Habit. Plants of the genus Fothergilla, generally low shrubs with erect
or strongly ascending aérial stems, spread profusely by means of woody
underground stems, frequently forming dense clumps. The aérial stems are
typically unbranched for one-half to one-third their height, and in the case
of the coastal plain populations may be completely unbranched. The
coastal plain plants are seldom more than one meter tall, although excep-
tional specimens up to 2.6 meters in height have been seen; flowering
specimens of the montane plants are generally between one and three me-
ters in height, but in extreme cases may be nearly 6.5 meters tall.
Hesse (1909), Rehder (1910), and Bailey (1949) have used supposed
differences in habit to distinguish F. major from F. monticola, The
glaucous F. major is reportedly an erect, pyramidal shrub, while the non-
glaucous F. monticola is less tall and more spreading. Examination of
several populations of Fothergilla, consisting of both glaucous and non-
glaucous plants, in the mountains of western North Carolina has given
no indication of such a difference in habit. Most plants of both types are
rather low, spreading shrubs. Occasionally, especially when growing in
moister situations, the plants of both types become taller and more erect.
The tallest and most nearly erect plant examined was nonglaucous.
On the other hand, examination of Fothergilla plants cultivated at the
Arnold Arboretum of Harvard University has shown that several of the
shrubs in the collection are indeed erect with a more or less pyramidal
shape, totally different from any of the numerous plants which were studied
in the field. Other plants at the Arboretum, more typical of the wild type,
are low and spreading. While all of the erect plants are glaucous, the
spreading plants are of both types. The significance of this difference in
habit is unknown, but since it is not consistently correlated with the pres-
ence or absence of a waxy bloom on the undersides of the leaves, it should
be ruled out as a character substantiating the existence of two taxa of
Fothergilla in the mountains of the southeastern United States.
Vestiture. The undersides of the leaves of both the montane and
coastal plain forms of Fothergilla are occasionally covered with a fine,
waxy bloom. As mentioned earlier, the presence or absence of this bloom
has been used as the primary morphological character for distinguishing
F. major from F. monticola.
Since the waxy bloom generally disappears when specimens are heat
dried, and since most collectors regrettably have failed to mention whether
1969 | WEAVER, FOTHERGILLA 603
or not the plants were glaucous, herbarium specimens are useless for the
study of this character. Therefore field studies were necessary. Populations
were studied in the following localities:
PIEDMONT
1. Near Hillsborough, Orange County, North Carolina.
2. Hanging Rock State Park, Stokes County, North Carolina.
MounNTAINS
3. Gorge of the Thompson River, Transylvania County, North Carolina.
4. Gorge of the Horsepasture River, Transylvania County, North Caro-
lina.
The Piedmont populations are composed entirely of nonglaucous indi-
viduals; the mountain populations are composed of approximately equal
numbers of glaucous and nonglaucous individuals. There appears to be
little variation in the amount of bloom among those plants which are
glaucous at all; the bloom when present is very striking and in all proba-
bility is genetically controlled. Aside from the presence or absence of a waxy
bloom the plants are indistinguishable as far as leaf, capsule, and habital
characters are concerned. In view of this fact and since glaucous and non-
glaucous individuals grow side by side in the same population, this single
character seems to offer insufficient grounds for the recognition of two taxa
of any rank.
A single population of Fothergilla in the coastal plain of North Carolina
(Scotland County) was found to consist of glaucous and nonglaucous indi-
viduals. As in the case of the montane populations both types grow side
by side and are identical in other respects.
The aérial organs of Fothergilla, both floral and vegetative, with the
exception of the styles and stamens, are clothed to varying degrees with
a soft, downy pubescence. The individual trichomes are stellate in form,
composed of six to ten unicellular segments radiating from a single modi-
fied epidermal cell. The color of the trichomes varies from dark brown
on the stipules and floral bracts to a pale yellow on the remaining parts.
Interspersed among the stellate trichomes on the apex of the ovary and
persisting on the mature fruit are stiff, yellow, simple trichomes which are
apparently the only simple ones to be found on the plant.
In general the coastal plain plants are pubescent to a greater degree
than are the montane plants; however, in both groups the amount of
pubescence varies among the respective organs, as well as on the same
organ of different plants. The trichomes are generally dense and over-
lapping on the petioles, stipules, calyx, floral bracts, and peduncles. The
aérial stems of the montane plants are generally pubescent on the distal
portions, especially at the bases of the spikes and the leaves where the
pubescence may be considerable, and on the present year’s growth, but
become completely glabrous in the proximal portions; those of the coastal
plain plants are frequently pubescent, at least sparingly so, to the base.
604 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The leaves are pubescent on both surfaces, but generally more densely
so on the abaxial surface; the trichomes are scattered to varying degrees
over the lamina but are often concentrated on the principal veins or in
their axils. The lamina is frequently glabrous in the montane plants but
is always pubescent to some degree in the coastal plain plants examined.
Rehder (1910) reported that the leaves of F. major varied from scattered
to rather densely pubescent beneath, while those of F. monticola were
pubescent chiefly on the veins, the lamina often being nearly glabrous.
Ernst (1963) also noted that F. monticola was pubescent to a lesser de-
gree than F. major. Bailey (1949) was vague on the matter, having de-
scribed the leaves of F. major as pubescent below, at least on the veins, as
opposed to scattered pubescent below in F. monticola.
Examination of herbarium specimens as well as observations of the
plants in the field has indicated that the degree of pubescence of the leaves
on an individual plant of both the glaucous and nonglaucous types is quite
variable. For example, the pubescence on the undersides of the twenty-
five mature leaves taken at random from plants of both the glaucous and
nonglaucous types in each of two populations in Transylvania County,
North Carolina, demonstrated that the range of variation in the degree
of pubescence is the same for both types: in each of the plants studied
the variation ranges from leaves in which the lamina is glabrous to leaves
in which the lamina is covered with a nearly continuous indumentum.
In order to quantify these impressions herbarium specimens prepared
from two populations of Fothergilla (one from Burke County and one
from Transylvania County, North Carolina) were studied intensively,
and the number of trichomes present on four square millimeters of Jaminal
surface at comparable positions on the undersides of the leaves of both
the glaucous and nonglaucous types was counted. The results are sum-
marized in TABLE 1.
TABLE 1. Variation in the Number of Trichomes on the Leaves in
Two Populations of the Montane Fothergilla
RANGE
NUMBER OF MEAN
LECTOR OF VARIATION # OF
LocaLIry § AND NUMBER Form Leaves In # or Harrs HaIrs
Burke Co. Wilbur 7040 glaucous 40 0-40 5.09
Burke Co. Wilbur 7038 non-
glaucous 40 0-30 5.98
Transyl-
vaniaCo. Weaver 300 glaucous 20 7-35 16.65
Transyl-
vania Co. Weaver 301 non-
glaucous 20 4-29 13.55
1969} WEAVER, FOTHERGILLA 605
From the data in Tas.e 1 the following conclusions may be drawn as
far as the populations studied are concerned: (1) the variation in degree
of pubescence of each form is great within one population as well as be-
tween different populations; (2) the variation in the degree of pubescence
of each form is approximately equal in each population.
Even though the samples were small the results indicate that there is
no consistent difference in the degree of pubescence between glaucous and
nonglaucous individuals.
Leaves. The leaves of Fothergilla are simple, alternate, deciduous and
somewhat coriaceous in the coastal plain populations, while they are
thinner and membranaceous in the montane populations. Variation in
shape and type of dentition is great; it is not at all unusual to find on a
single plant leaves with (1) shape ranging from elliptic through ovate
and obovate to suborbicular; (2) bases cordate, truncate, and rounded;
(3) apices acute, obtuse, and emarginate; and (4) margins varying from
coarsely serrate-dentate to entire. Extent of the dentition, however, is
more nearly constant than the type of dentition; in general among the
coastal plain populations the dentition when present is restricted to the
upper half of the leaf, while among the montane populations it almost
invariably extends below the middle of the leaf.
Leaf size has traditionally been used to distinguish the montane and
coastal plain forms of Fothergilla, and it remains a usable diagnostic
character. The leaves of the coastal plain plants are typically and usually
conspicuously smaller than those of the montane plants. Although there
is a considerable overlap in absolute leaf lengths and width between the
two forms, the largest leaf on any given specimen examined from the
coastal plain never exceeded 5.2 cm. in width, while the largest leaf on
a specimen from the mountains was never less than 5.1 cm. wide.
Fothergilla parvifolia, described by Kearney (in Small, 1903), has
usually been distinguished from F. gardenii by the shape and dentition
of the leaves. Britton (1905) and Small (1933) agreed that the leaves
are different: F. parvifolia with leaves broadly ovate, oval, or suborbicular,
toothed mainly from below the middle, and cordate at the base; and F
gardenii with leaves oblong, oblong-ovate, or ovate-orbicular, toothed above
the middle, and narrowed at the base, according to Britton, and elliptic,
elliptic-ovate, or elliptic-orbicular, toothed only near the apex, and cuneate
or rounded at the base, according to Small. Earlier, however, Small (1903,
1913) had reported that the leaf margins of F. parvifolia were coarsely cre-
nate above the middle, while those of F. gardenii were undulate or coarsely
toothed near the apex. he
As may be inferred from these conflicting descriptions the distinctions,
at least as regards leaf shape and dentition, between F. parvifolia and
F. gardenii are hazy at best. In view of the variation among the leaves
of an individual plant as described above, it would seem that the delimita-
tion of taxa of Fothergilla on the basis of leaf shape and dentition is un-
warranted.
606 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Flowers. The perianth of Fothergilla is greatly reduced, consisting of a
gamosepalous calyx, the 5—7 lobes of which are reduced to minute, ir-
regularly shaped teeth. Petals are lacking. The base of the androecium
is adnate to the calyx, forming a shallowly campanulate hypanthium. The
stamens vary in number from 12—32, and are arranged in a single series
on the rim of the hypanthium. The coastal plain populations tend to have
fewer stamens than the montane populations; in the material studied the
number ranged from 12 to 24 in the coastal plain plants and from 18 to
32 in the montane plants. The filaments, easily the most conspicuous parts
of the flower, vary greatly in length with no apparent pattern in a single
flower. The longer ones are conspicuously thickened distally while the
shorter ones tend to be nearly filiform. The pistil is made up of two
carpels fused below but divergent near the apex of the ovary into separate
filiform styles.
Most previous authors have considered the ovary of Fothergilla to be
weakly perigynous; Ernst (1963) described the ovary as being “slightly
recessed in the receptacle” but considered the fruit to be “partly inferior.”
Macroscopic examination of the ovary indicates that it is partially fused
to the receptacle and is therefore semi-inferior.
In order to determine the histological relationship between the ovary
and the hypanthium, flowers of Fothergilla major collected in Transylvania
County, North Carolina, and preserved in 70% ethyl alcohol were dehy-
drated using the tertiary butyl alcohol series suggested by Johansen (1940),
and infiltrated with paraffin. The material was sectioned on a rotary micro-
tome at 12 microns and stained in 1% safranin in 95% ethyl alcohol and
1% fast green in 95% ethyl alcohol. The investigations have shown that
the hypanthium is actually adnate to the ovary to a point slightly above
the base of the ovary, i.e., at least one-third the total length of the latter
(Fic. 1). Therefore the ovary is, indeed, somewhat semi-inferior.
Serial sections of the pistil have given further insight into the morphol-
ogy of that organ. Macroscopically the pistil in cross section appears to
be 2-locular; to my knowledge all previous authors have described the
pistil accordingly.
Actually, as shown in Ficures 2 and 3, the closure of the individual
carpels is not complete from a point immediately below the attachment
point of the ovules to the apex of the ovary. As a result the ovary 4p-
pears to be bilocular below the attachment point of the ovules but uni-
locular above it. In actuality, then, the ovary is characterized by having
a single deeply lobed locule. Horne (1914) reported that a similar situa-
tion, but in reverse, i.e. unilocular below and bilocular above, was found
in Rhodoleia, a member of the subfamily Bucklandioideae of the Hama-
melidaceae native of eastern Asia; he also stated, however, that the
Hamamelidoideae, of which Fothergilla is a member, possessed septa, ren-
dering them bilocular.
Fruits and seeds. The fruit of Fothergilla, a grayish or brownish
loculicidal capsule, is rendered partly inferior by the fusion of the per-
1969] WEAVER, FOTHERGILLA 607
Ficurrs 1~3, Cross section of the flower of Fothergilla major, X 60. Fic.
1, Fusion of the ovary to the hypanthium. Ovary at attachment point
of the ovules. Fic. 3. Ovary below attachment point of the ovules.
sistent hypanthium to the ripened ovary. Each of the carpels dehisces
along a median dorsal suture, and, as a result, the capsule is 4-beaked at
maturity, The capsules of the montane and coastal plain forms are similar
in shape, but those of the montane populations are decidedly larger in
size. Although there is a considerable overlap in total capsule length in
the material at hand, the fruiting hypanthium of the coastal plain form
608 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
varies from 3 to 4.5 mm. in length while that of the montane form varies
in length from 4 to 9.2 mm.
Small (1903, 1913, 1933) and Britton (1905) i that the cap-
sules of Fothergilla parvifolia varied from 6 to 8 mm. in length, while
those of F. gardenii varied from 8 to 10 mm. Examination of specimens
from Jesup, Georgia, the type locality of F. parvifolia, including a speci-
men identified as F. parvifolia by Kearney (Kearney s.n., 1893 [Ny]|) has
shown that the capsules of these plants actually vary from 7 to 10 mm. in
length. In addition numerous capsules of typical F. gardenii measuring
less than 8 mm. have been
Rehder (1947) and Bailey (1949) reported that the capsules of Fother-
gilla major were light brown inside with red markings on the inner sutures,
while those of F. monticola were darker inside without the red markings.
Examination, in the field, of the capsules of the glaucous and nonglaucous
forms of the montane Fothergilla shows that the color of the inside of the
capsules often varies from light to dark brown on the same plants. The
red markings reported by Rehder and Bailey are indistinct at best and
are not constant even among the capsules of a single inflorescence.
The seeds of Fothergilla, two per capsule at maturity, are ellipsoid or
narrowly ovoid with a very hard, shiny, reddish-brown seed coat.
whitish area, decurrent on opposite sides of the seed and including the
sub-apical hilum, is present at the micropylar end. In the material ex-
amined, the seeds of the montane populations vary in length from 6.2 to
7.8 mm., while those of the coastal plain populations vary from 4.8 to
6.3 mm. in length.
CYTOLOGY
The first account known to me of the chromosome number of any of
the species of Fothergilla is that of Anderson and Sax (1935) in their
survey of the chromosome numbers in the Hamamelidaceae. They re-
ported the haploid number to be m = 36 for F. major and n = 24 for F.
monticola, The counts were made from aceto-carmine smears of pollen
mother cells from plants growing at the Arnold Arboretum of Harvar
University. Anderson and Sax pointed out, however, that the “species”
are so similar that it is doubtful whether F. ‘monticola deserves more than
varietal rank. It was their opinion, from the cytological evidence, that “F.
monticola is merely a “presarens! variety which arose spontaneously from
the hexaploid species, F. majo
Thomas (in Ernst, 1963) ats reported the chromosome numbers of
Fothergilla, the counts again having been made from plants growing at
the Arnold Arboretum. His results were as follows:
ARNOLD ARBORETUM
SPECIES ACCESSION NUMBER HapLow NUMBER
F. monticola AA # 4163-A nm = 24
F. major AA # 694-34 n = 36
F. gardenii AA # 684-50 s = 36
1969 | WEAVER, FOTHERGILLA 609
In the course of this study the chromosomes of Fothergilla were again
recounted. Inflorescence buds were originally collected during April, 1966.
In addition buds of F. gardenit were obtained from Mrs. Anne B. McCrary,
of Wilmington, N.C. All buds were fixed in modified Carnoy’s Solution
(3 parts absolute ethyl alcohol: 1 part glacial acetic acid) and stored under
refrigeration in 70% ethyl alcohol.
| dn Be
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* but has several scattered populations in the
pinelands of subtropical Florida. These two species are placed in dif-
ferent sections because the central florets of C. tomentosa are functionally
more easily separated by their achenes (long beaked in C. dentata and
beakless in C. tomentosa), by the length of the ligulate corollas, and by
the position of the heads before and after anthesis (see illustration).
dominance of bilabiate “ray” florets in Gerbera, the vestiges of an inner lip in many
species of Chaptalia, and the tridentate ligule in corollas lacking an inner lip, all indi-
cate that in these two genera, the ligulate ray florets are derived from bilabiate forms
by loss of the inner lip.
*Burkart recorded only Chaptalia alsophila Greene, C. dentata (L.) Cass., C.
olak. as occurring in the United States, but Watson
and Kearney & Peebles (Ariz. Fl. ed. 2. 957. 1960) listed
C. leucocephala Greene as occ urring from Arizona to Mexico and added New Mexico
See
i heal pi oingeon Bot. 3(3): 441. 1930) mentioned that Chaptalia cag
was cultivated at two localities in Yucatan. Burkart, however, referring the
original reference (Millspaugh & Chase, Fieldiana Bot. 3(2): 148. 1904) parry that
the species involved was actually the closely related C. obovata Wright. Burkart did
cite, ea one specimen of C. dentata from Huasteca, Veracruz, Mexi
622 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
head on a cloudy da
i, floret of “second series, x 4 (cf. c); j, bilabiate pateal ‘floret, X 4; k, ti p of
, X 1; g, same on a sunny day, ; h, outer floret, Es 4;
developing fruit, X 1; n, mature achene with pappus, X 2. Note most pappus
ss % hep mitted ol pcg with only enough included to show appropriate
an
|
1969 | VUILLEUMIER, COMPOSITAE TRIBE MUTISIEAE 623
The phenomenon of trimorphic florets found in this genus is not rare
in the Compositae (Uexkill-Gyllenband), but inaccurate observations
on the condition in Chaptalia have led to misinterpretations of both the
origin of the ligulate florets (see footnote 3) and the relationships of the
genus.
Most workers (Cassini, Bentham & Hooker, Hoffmann, Burkart) have
allied Chaptalia with Gerbera L. (Africa and southeastern Asia) and
Trichocline Cass. (Andean with a species in Australia). All three are
similar in habit, having basal rosettes, scapose flowering stems, and the
same type of white tomentum. Palynologically, however, Chaptalia is
distinct from both Gerbera and Trichocline and is similar to Lycoseris
Cass.,° another genus of the Gerberinae (cf. Bentham & Hooker). Wode-
house postulated on the basis of pollen similarities a relationship between
Chaptalia and Lycoseris which he supported with evidence from floral
morphology, suggesting that both have outer florets without (or with
reduced) inner lips and “staminate” florets with undivided styles. Ger-
bera and Trichocline, he maintained, have, in contrast, monomorphic
perfect florets with bilabiate corollas. Examination of specimens, how-
ever, shows the styles of Chaptalia always to be bipartite, even in the
functionally staminate central florets of certain species (presumably the
“staminate” florets of Wodehouse), and the ray corollas of several species
of Chaptalia are distinctly bilabiate. The floral morphology of Gerbera,
like that of Chaptalia, appears to be much more variable than Wodehouse
thought. Monomorphic bilabiate corollas are not a constant character
throughout Gerbera, and eight or nine species (of 30 to 35 species ) have
heads with dimorphic florets. In addition, some individuals of Chaptalia
lack one of the three floret types. In fact, when all the species of both
Chaptalia and Gerbera are carefully examined, there is complete transi-
tion between the two genera. Baillon went so far as to combine them,
and both Bentham and Burkart thought it necessary to state specifically
that, despite the transitions, there are enough characters in combination
to justify maintaining the two as distinct genera.
The occurrence of cleistogamous (always closed) as well as chasmog-
amous (open and radiate) heads in Chaptalia is of special interest, for
cleistogamy is rare in the Compositae, although it has been known in
one species of Gerbera since the time of Linnaeus. Burkart found that
both kinds of heads occur in three Argentine species of Chaptalia (c
exscapa (Pers.) Baker, C. piloselloides (Vahl) Baker, and C. runcinata
HBK.., the last two in the same section as our C. dentata). In contrast
to the species of Gerbera which produce both types of head simulta-
neously on the same plant, species of Chaptalia apparently produce only
one type at a given time on an individual plant (e.g., on a cultivated
plant of C. runcinata cleistogamous heads in May, chasmogamous ones in
i is i i tricolporate, and has
°The pollen of both Chaptalia and Lycoseris is spherical, tricol and
vestiges of spines. Gerbera and Trichocline have pollen which is slightly ellipsoidal,
tricolporate, almost smooth, and which has the “remarkable character” (cf. Wode-
house) of intercolpar thickenings (ie., between the furrows).
624 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
winter (July), and cleistogamous again the next February). Burkart
interpreted 7. seasonal variation as an adaptation to life on the Ar-
gentine pampa
Although Solbrig mentioned the possibility of apomixis in Chaptalia,
various experiments suggest that this does not occur. Burkart emas-
culated and bagged the heads of five South American species but found
no indication of seed production. On the other hand, nonemasculated,
bagged heads produced abundant viable seeds, showing that autogamy
is probable. That self-pollination does occur in nature is suggested both
by the presence of cleistogamy and by the reports that some species
have involucral bracts that curl inward and force the stigmas against
the pollen-producing florets. Recent experiments with C. dentata have
shown that the plants are self-compatible. All of the full, mature achenes
from a head which was not accessible to pollinators germinated. How-
ever, about 50 per cent of the achenes never matured, indicating that
self-pollination is not completely effective, but, once it has occurred,
there are no self-incompatibility barriers (Vuilleumier).
Artificial crosses made by Burkart between three South American
species all produced F; hybrids which were completely, or nearly, sterile.
However, natural hybrids between two other species not investigated are
found in southern Brazil and northern Argentina. Chromosome numbers
have been reported only for Chaptalia piloselloides (a species with cleis-
pagide oy in Burkart’s experiments not apomictic), with 2” = 49- 54,
54, 51-54, 60; C. nutans, 2n = 48; and C. integerrima (Vell.) Burkart
cs C, ape os (Cass.) Baker), 2n = 48.
The genus has no economic importance, but Chaptalia nutans is ap-
parently used by Argentine Indians for infections (the leaves being ap-
plied with a little oil) and for respiratory ailments. Chaptalia tomentosa
has been cultivated to a limited extent (cf. Bailey, Cyclop. Am. Hort. 1:
288. 1900, and 1: 734. 1928, as well as Roy. Hort. Soc. Dict. Gard. ed. 2.
1: 450. 1956).
REFERENCES:
BaLpwin, J. T., Jr., & B. M. Speese. Chaptalia nutans and C. integrifolia:
their chromosomes. Bull. Torrey Bot. Club 74: 283-286. 1947.
BENTHAM, G. Notes on the classification, history, and geographical distribu-
tion of Compositae. Jour. Linn. Soc. Bot. 13: 335-461. 1873. :
& . Hooker. Compositae. Gen. Pl. 2: 163-533. 1873. [Mutzsiaceae,
214-219, 484-504; Chaptalia, 498.
BurkartT, A. Estudio ‘del género de compuestas Chaptalia con especial refer-
encia a las —- argentinas. Darwiniana 6: 505-594. pls. 1-10. 1944.
Cassini, H. Ebauche de la synanthérologie. Opuscules Phytologiques 2: 1-281.
Paris. 1826. [Mutisieas 95-128.
Horrmann, O. Compositae. Nat. Pflanzenfam, IV. 5: 87-387. 1893. [Muti-
sieae 330-350; Cicsecik 345.]
Kocu, M. F. Studies in the anatomy and morphology of the composite flower
II. The corollas of the Heliantheae and Mutisieae. Am. Jour. Bot. 17: 9 995-
1010.
|
1969 | VUILLEUMIER, COMPOSITAE TRIBE MUTISIEAE 625
KNUTH, a ee Handb. Bliitenbiol. 3(2): 213-237. 1905. [ Gerbera,
236,
RICKETT, - Wild Flowers of the United States. Vol. 2. a Southeastern
States. Part. 2, New York. 1967.’ {Chaptalia, 599, pl. 224.)
Sims, J. Chaptalia tomentosa. Bot. Mag. 48: pl. 2257. 1821.
UEXKULL-GYLLENBAND, M. von. Phylogenie der Bliitenformen und der
celta eerste bei den Compositen. Bibliot. Bot. 10(52): 1-81.
2 pls. 1
WODEHOUSE, 4 P. Pollen in the ap aere and oe of
plants. IV. The Mutisieae. Am. Jour. Bot. 16: 297-313. pls. 23, 24.
1929. erg en of ibeieudlee of the genera; C. dentata
pollen, fig. 14.]
Present address:
THE ARNOLD ARBORETUM THE GrAy HERBARIUM
OF OF
HARVARD UNIVERSITY HARVARD UNIVERSITY
626 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
A NEW SPECIES OF ARENARIA FROM THE BHUTAN HIMALAYA
N.C. MAJUMDAR AND C. R. BABU
THE NEW speEctes described in this paper was collected from Bhutan,
a small mountainous country in the Eastern Himalayas, situated between
26° 40’ and 28° 0’ N. latitude and between 88° 10’ and 91° 45’ E.
longitude, and lying between Tibet and India. It is composed of lofty
and rugged mountains which vary in elevation from 300 meters to 7500
meters above sea level, and are separated by deep valleys. The climate
of Bhutan varies according to elevation; thus, the lower southern valleys
are saturated with moisture, hot and steamy; the central valleys enjoy
a temperate coolness; and the extreme northern higher region has the
rigors of frost and ice.
The vegetation of Bhutan, in general, is composed of tropical, sub-
tropical, temperate, and alpine elements. Chila, the locality from which
this interesting taxon has come, is a mountainous ridge situated between
3630 meters and 4120 meters in altitude, in the central tract. It is char-
acterized by alpine vegetation which is composed of herbaceous plants
such as Ranunculus, Gentiana, Primula, Potentilla, Gaultheria, Arenaria,
Cerastium, Stellaria, and Swertia, in addition to shrubs like Symplocos,
Eurya, and Pentapanax.
Arenaria bhutanica Majumdar & Babu spec. nov. Fic. 1.
Pertinet ad subgenus ARENARIA, affinisque nulli speciei huius generis
adhuc usque notae; valde distincta habitu annuo, floribus solitariis, caule
uno, pedicellis duplici linea pilorum ornatis, sepalis subacutis quam pe-
talis brevioribus, seminibus 3—6 in una capsula.
Herba annua, gracilis, ramosa, 5-10 cm. alta; caulis quadrangularis,
prostratus et glaber infra, ascendens vel suberectus et una linea longitu-
dinali pilorum sursum; folia sessilia, opposita quidem aequalia, ad basin
connata, lineari-lanceolata, carnosiuscula, integra, margine haud incras-
sato, ad apicem acuta, pungentia, brunneola, glabra, utrinque punctata,
punctis elevatis tubercularibus circularibus, uninervia, 5-10 % 1.5-2 mm.,;
flores solitarii, axillares et terminales, longe pedicellati, albi, 5-8 mm.
diametro; pedicello gracili, quadrangulari, fructifero paulum recurvato,
linea duplici longitudinali reflexorum pilorum ornato, 1-2.5(-3) cm.
longo; bracteae foliosae, saepe minores foliis inferioribus; sepala 5, usque
ad basin libera, lanceolata, subacuta, marginibus late scariosis, obscure
uninervia, glabra, 3.8-4 0.8-1(—1.2) mm.; petala alba, breviter un-
guiculata, oblongo-spathulata, integra, obtuso-rotundata, sepalis longiora,
5 mm. longa; stamina 10, uniseriata, 3-3.5 mm. longa; filamenta linearia,
1969 | MAJUMDAR & BABU, ARENARIA 627
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RE 1. Arenaria bhutanica, a-h. a, habit; b, sepal; c, petal; d, stamens
with ne without glands at base; e, pistil ; yi capsule: g and h, seeds.
ad ipsam basin connata, eorum singula alterna basi glandulifera, ovoideae
brunneolae insidentia, 0.2 mm. longa; antherae minutae, ovoideae, atro-
purpureae, basifixae, 0.2 mm. longae; ovarium subsessile, ovoideum,
glabrum, tricarpellatum, 1.1~-1.3(—1.4) mm. longum; styli terni, lineares,
papillosi; capsula ovoideo-subglobosa, breviter stipitata, dehiscens ad
apicem in dentes 6 obtusos, 2.8-3 mm. longos; semina (immatura) 3-
va subreniformia, compressa, rubro-brunnea, levia? 0.8—1 mm. lata.
: Chila, on way to Paro, alt. 3630-4125 m., 24 August 1963, G. Sen
FS 721 (holotype and isotype, CAL). Small prostrate, delicate herb with
white flowers; stamens blue
The very slender annual habit, the linear one-nerved leaves, and the
obscurely nerved subacute sepals which are shorter than the petals of this
remarkable species may bring it under sect. OccIDENTALES of the sub-
genus ARENARIA; but apart from the fact that all the eleven species in-
cluded in this section by McNeill (in Notes Roy. Bot. Gard. Edinburgh
24: 115. 1962) are either Spanish or North African in distribution, they
differ from this species in having dichasial cymose inflorescences. Among
628 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
the Indo-Tibetan species, this distinctive species with uncertain affinity
shows superficial resemblance to Arenaria monosperma Williams in its
slender habit, linear one-nerved sharply acute leaves, and solitary axil-
lary and terminal flowers; but it differs in its annual growth, presence
of a single line of hairs on the stems, two lines of hairs on the pedicels,
petals longer than the subacute sepals, and in the 3—6-seeded capsule.
ACKNOWLEDGMENT
The authors are grateful to Rev. Dr. H. Santapau, S.J., Department
of Botany, St. Xavier’s College, Bombay, for kindly making a few cor-
rections in the Latin description.
CENTRAL NATIONAL HERBARIUM
BOTANICAL SURVEY OF INDIA
CaLcuTtA, INDIA
1969 | THE DIRECTOR’S REPORT 629
THE DIRECTOR’S REPORT
THE ARNOLD ARBORETUM DuRING THE FiscaL YEAR ENDED
JUNE 30, 1969
IT SEEMS AN APPROPRIATE TIME, while preparing the annual Director’s
Report, to reflect on the pleasures and difficulties of implementing the
original purpose of the Arboretum “to grow all of the trees, shrubs, and
herbaceous plants hardy in the vicinity of West Roxbury” and the con-
comitant goal of increasing our knowledge of these plants, their relatives,
and the vegetation associated with them in their natural areas. How well
this charge is being accomplished the record will show.
Again in the first months of 1969, New England suffered characteristic
diversity of climate. A severe ice storm in January was localized in the
suburban area, causing extensive damage on the Case Estates in Weston,
but none in Jamaica Plain. Two storms in February and one in March
produced accumulations of wet heavy snow that damaged plants se-
verely, primarily in Jamaica Plain, almost equalling the destruction of
past hurricanes. The weather bureau reported a record of 47.6 inches
of snow in Boston for February alone (annual average is 41.7 inches),
including probably the longest recorded period of uninterrupted snow fall,
78 hours. The accumulation was even greater at the Arboretum than
that officially reported at the airport. The extent of damage is almost
immediately apparent to visitors because of the nearly complete de-
struction of the Magnolia stellata plantings near the Administration
Building as well as in the loss of some entire trees, and many branches,
in the cluster of 40-year old specimens of Prunus sargentii. Elsewhere
on the grounds the damage was equally severe, as in the species apples;
the oaks, where a single large tree well over 200 years old was toppled;
in the conifers; and in the Carpinus and Ostrya collections. The grounds
crew used chain saws and brush chippers to handle the damaged branches
in an immediate effort to clear the grounds, later returning the wood as
mulch to the area. Proper pruning and repair of the damaged trees will
take the rest of the year. Replacements for many specimen trees must
be propagated so that clones of known lineage will be retained. There is
still injury to be assessed in the shrub collection where many specimens
which were bent to the ground, their branches twisted or fractured, may
not live through the season. However, even though the damage of the
winter of 1968-69 will long be evident, the spring season which followed
was again without a late frost and the display of forsythias, lilacs, crab
apples, tree peonies, and rhododendrons was superb. The spring was,
in fact, one of the finest in many years.
630 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Staff :
The President and Fellows of Harvard College approved the promotion
of Associate Curators Dr. Lorin I. Nevling, Dr. Bernice G. Schubert,
and Dr. Carroll E. Wood to be Curators, effective July 1, 1969.
Dr. Alfred Linn Bogle, a graduate of the University of Minnesota was
appointed Assistant Curator at the beginning of this fiscal year; Dr.
Thomas S. Elias, who recently received his degree from St. Louis Uni-
versity, was also appointed an Assistant Curator, effective July 1, 1969.
Both men will work with Dr. Wood in continuing the preparation of the
Generic Flora of the Southeastern United States.
Dr. Beryl S. Vuilleumier concluded her term of work on the South-
eastern Flora project and has taken up research problems on her own
recent South American collections.
Mr. Robert S. Hebb, a recent graduate of the Gardening program at
the Royal Botanical Gardens, Kew, is serving as Assistant Horticulturist.
Mrs. Winifred P. Hebb is an assistant in the herbarium and library.
The President and Fellows approved the appointment of Mrs. Ara R.
Derderian as Honorary Curator of the Bonsai Collection, effective June
1, 1969.
Honors from outside the University were received by two members
of the staff. Dr. Wyman was awarded the Veitch Memorial Medal, in
gold, of the Royal Horticultural Society with the citation “By this
Veitch Medal we today pay tribute to Dr. Wyman’s contribution to the
science, to the practice, and to the literature of horticulture.” Since the
medal is rarely awarded to persons outside of England, we are happy
to record that Dr. Wyman is the third member of our staff to receive this
prestigious honor.
Dr. Bernice Schubert, together with Dr. Lyman B. Smith, of the Smith-
sonian Institution, received the Eva Kenworthy Gray award of the Ameri-
can Begonia Society in recognition of their joint contribution of original
material which aided members of the Society in the study of begonias.
These botanists have published a series of papers on the classification
and distribution of Begonia in Central and South America.
Dr. Wyman was elected a Vice-President of the Massachusetts Horti-
cultural Society, and Dr. DeWolf was asked to serve on the Library Com-
mittee of the Society. Dr. Schubert was elected a member of the Council
of the Society for Economic Botany.
Horticulture:
The development of the Weld-Walter Street tract of land for expansion
of the living collection was begun during the winter months of the cur-
rent fiscal year. This tract of 15 acres is held by Harvard University
for the purposes of the Arnold Arboretum. Our own crew repaired and
reconditioned the stone wall surrounding a large part of the land. After
competitive bidding a contract was let to place units of 4- and 6-foot
chain-link fence around the area. Two driving gates and three pedestrian
gates will control access to the property. A second contract, for land
1969 | THE DIRECTOR’S REPORT 6
ABOVE: Development of the Weld-W alter Street tract of the Arnold Arbore-
tum Se with the cutting of a roadwa
v: View from the summit of the hill showing Route 1 and the Hebrew
Rehabilitation ren Buildings.
632 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
movement, has established 1,840 feet of roadway 24 feet wide which
sweeps gracefully from Walter Street to the highest point, where a turn-
gravel and an oil seal for the present. The land was treated with lime
and fertilizer in the fall, following recommendations of the Soil Conser-
vation Service of the U.S. Department of Agriculture. Planting of species
which will tolerate the dry hillside conditions will be gradual, and the
area will be opened to the public in 1972.
To improve the appearance of the largest of the three ponds along
the meadow road, which had become filled with aquatic weeds during
recent years, it was dredged. Late fall rains and winter snow filled it,
and a program of planting around the pond has be
Income available from the Isabella P. Shaw fund helped provide six
r
dendrons in 1968 and permitted rapid replacement of storm-damaged
plants of mountain laurel along the base of Hemlock Hill.
a
“Pond along the ea said within in Perens which was icon pan
ing the summer of
The growth control chemical Casoron is proving helpful in restricting
grass and weeds near special plantings. Its use has reduced mowing time
required to maintain the appearance of the grounds and has nearly
eliminated the problem of injury to the base of tree trunks, caused by
the use of mowing machines
The genetic dwarf conifer collection established in terraced plantings
near the greenhouses in Jamaica Plain has done well in that location
where it attracts much popular interest and where it has survived the
winter without special protection. Limited to specimens of known origin
1969] THE DIRECTOR’S REPORT 633
and unquestioned identification, this collection has great reference value.
A gift of 21 previously unrepresented taxa from Mr. Joel Spingarn, of
Baldwin, New York, and collections of native plants from locations along
the coast of Maine, made by Mr. Fordham, have increased the variety
of the dwarf conifer collection.
Mrs. Ara R. Derderian has accepted responsibility for curating the
Larz Anderson collection of bonsai, a famous and popular display which
has needed competent care for several years. With the help of members
of the staff she has carefully pruned and repotted most of the plants dur-
ing the winter months. Many of the specimens are thought to be imperial
bonsai because of their age and special character.
For the protection of lawns, nursery area, and roads, special steel edg-
ing is being used in the vicinity of the Dana Greenhouses, and, for more
efficient work procedures, steel storage bins, shelving, and work benches
have been installed in the greenhouse. A new well provided continuous
water pressure and flow for irrigation in the nursery area during the dry
summer months. The emergency electric generator worked well during
electric power failures resulting from the February and March snow
storms. Without it the oil-fueled furnaces would have been inoperative
and the greenhouse plantings might not have survived two periods of
more than six hours without heat.
During the year a total of 672 specimens were planted on the grounds
as replacements or additions. Cuttings or grafts have been prepared for
287 taxa which may need replacement. A total of 152 additional taxa
have been prepared for distribution to Cooperating Nurserymen, to other
arboreta, or to the Friends of the Arnold Arboretum for testing. Staff
research required propagation of 78 taxa.
During the year 11 recent staff introductions were distributed to bo-
tanical gardens and Cooperating Nurserymen. In addition, 169 shipments
of plant materials representing 918 species and varieties were made to
gardens and individuals in the United States and 14 other countries.
Thirty-seven lots of seeds, including 182 taxa, were sent in response to
specific requests from correspondents in the United States and 24 other
countries. The 88 shipments of plant materials received during the year
included 287 taxa, and 72 lots of seeds represented 182 taxa. Plants not
needed by the Arboretum were offered to the Department of Buildings
and Grounds of Harvard University and to other colleges and universities,
as well as to botanic gardens.
Dr. Wyman cooperated with representatives of the U.S. Bureau of
Public Roads and several Roadside Development engineers of the Massa-
chusetts Department of Highways, Bureau of Public Works, in advising
and by supplying some plant materials for programs of highway beautifi-
cation. The Massachusetts Department of Highways also accepted four
truck-loads of trees and shrubs for demonstration plantings along the
new Blue Star Memorial Highway in eastern Massachusetts.
A collection of 36 plants was donated to Channel 2, Boston's educa-
tional television station, for “sale’’ at its benefit auction in early June.
634 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
As the application of electronic data-processing equipment and tech-
niques to collection records has become of great national and international
concern, our staff has collaborated with others on projects under way at
other institutions. Several types of projects are in a trial period but
may be able to incorporate data from our collections in the future. Dr.
Howard serves as chairman of the Plant Records Center of the American
Horticultural Society. The Plant Records Center, operating under a
grant from the Longwood Foundation, has been devising a method for
placing the records of the living and herbarium collections of the Long-
wood Gardens in a retrieval system. The goal of the Plant Records
Center is to establish a central data bank of sources of plant materials
by recording, eventually, the accession records of all botanical gardens
and arboreta in the country. The Arnold Arboretum has the largest col-
lection of woody plants and probably the best records on the origin and
behavior of plant species under cultivation, and its records should be a
valuable addition to those of the Plant Records Center. Other projects
involving electronic data-processing systems are mentioned under the
section on the herbarium.
Case Estates:
The Case Estates in Weston, Massachusetts, are the location for
the nursery area for the Arnold Arboretum, as well as for special display
plantings, demonstration plots for comparison of mulching materials and
pruning techniques, ground covers, street trees, and perennial beds; areas
of natural woodland, and various materials which cannot be accommodated
in Jamaica Plain.
Additions were made to the already established wild flower garden
this year and a small collection of rock garden plants was established
for trial and demonstration.
The diversity and educational nature of the plantings has made the
Case Estates increasingly popular with school classes, colleges, and gar-
den clubs. All of the Weston Schools sent classes for talks or tours dur-
ing the year, while guided tours were held at the request of many
groups from Massachusetts and other states. To facilitate special lec-
tures and the regular popular classes for adults of the surrounding subur-
ban communities, one of the buildings was redesigned as a class room and
was equipped with carpeting, shades for darkening the room (for show-
ing slides), and new chairs. The building occupied by the superintendent
of the grounds was also reconditioned.
After a severe ice storm in January, a special study session was of-
fered for the Friends of the Arnold Arboretum. Forty-four people spent
a cold, sunny day observing the damage and discussing methods of repair
and later care for the injured plants. Afterward the trees were pruned
by the staff, or where necessary, removed.
Dr. Wyman was requested to aid the town of Weston in planning a
small park at the junction of Newton Street. and South Avenue, for
1969 | THE DIRECTOR’S REPORT 635
which the Arboretum donated plants. The town expressed its deep ap-
preciation in a vote of thanks. The Arboretum also made a gift of fiery
red crabapple trees of the cultivar BarBARA ANN for planting near the
new fire station.
Herbarium:
The herbarium collections of the Arnold Arboretum are divided in two
parts. The portion housed in Jamaica Plain is composed of cultivated
plants and serves as a reference collection for the identification of and
distribution studies on plants in cultivation. It now comprises over
136,000 specimens and has particularly representative collections of woody
ornamental trees and shrubs, especially those which form the large part
of our living collections. As a result of the special effort devoted to in-
creasing representation in the cultivated herbarium, additions of some size
came this year from Massachusetts, Pennsylvania, Florida, Puerto Rico,
Mexico, Venezuela, Brazil, and South Africa.
A new collection of historical interest to us consists of herbarium speci-
mens prepared by F. L. Olmsted who worked with Charles Sargent in
planning the Arboretum plantings and who was responsible for the develop-
ment of the park system in Boston and of Central Park in New York.
Given by the Olmsted Associates, this unmounted collection will require
much work before it is fully available. It includes specimens collected
in the Arnold Arboretum in 1875, the oldest material from our collec-
tions and probably some of the first specimens taken from early Arbore-
tum introductions, as well as material of the same period from Central
Park.
The major portion of the Arboretum herbarium, housed together with
that of the Gray Herbarium in the Harvard University Herbaria in Cam-
bridge, is composed of native plants of the floras of the world. It is most
monographic and floristic studies, work in plant a
and palynology, and is used for general identifications or to answer special
questions.
The several research projects of the staff center on a number of geo-
graphic areas. Dr. Wood and his associates continue their studies toward
a Generic Flora of the Southeastern United States, which actually has
implications affecting a much larger area of the country. During the
year Dr. Wood treated the families Betulaceae and Aristolochiaceae for
this project; Dr. Vuilleumier worked on some tribes of the Compositae;
Dr. Bogle, on the families of the Centrospermae; Dr. Long and Sister
Victoria Hayden, as Mercer Fellows, studied the Acanthaceae and Rubia-
ceae, respectively; Dr. Sorensen is studying the Phytolaccaceae, and Dr.
David Bates, of Cornell University, has agreed to continue the work on
the Malvaceae started by the late Dr. Brizicky. Other areas in the United
States were involved in Dr. Schubert’s work on species of the genus Des-
modium for the Manual of the Flora of Texas and Dr. Howard’s descrip-
636 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Students and staff of the 1968 Tropical Botany Seminar held at the Fair-
ci alee Sg Garden and the University of Miami, Dr. Howard was one of the
tive aha of the plants of the Isles of Shoals (near New Hampshire
and Maine).
Dr. , evling is conducting a cooperative program with scientists of the
Universidad Nacional Auténoma de México on the environments and
plant resources of the state of Veracruz. The native plants of the region
are being studied by him and his associates, or by specialists on particu-
lar groups, with emphasis on the ecology and biolog gy of the vegetation,
in addition to purely floristic studies. Special data handling techniques
are being employed in certain aspects of this project. Plants cultivated
in the area are also being studied and will serve as a valuable addition
to the cultivated collections at Jamaica Plain. The Arboretum has helpe
to support, in part, two collectors in Veracruz, Marino Rosas R. an
Guadeloupe Martinez Calderén.
Dr. Schubert has special research interest in some ‘igor of the genus
Dioscorea in Mexico which have very small stature but occur in various
sections of the genus, not being related by their ce habit. She
is also concerned with studies of species of Desmodium occurring through-
out the Americas. Dr. Sorensen has a particular interest in the genus
Dahlia, the national flower of Mexico, and his studies in that country
have increased his understanding of the distribution and growth patterns
of the group
The Caribbean islands are the floristic area of intensive research by
Dr. Howard and Miss Powell. Attention has been focused most recently
on biological studies of elfin forests in Puerto Rico and St. Kitts. Large
—————
1969 | THE DIRECTOR’S REPORT 637
general collections have been studied by them from the Bahamas, Puerto
Rico, Guadeloupe, Martinique, and St. Lucia.
Dr. DeWolf is preparing studies of the family Moraceae or of the genus
Ficus for the floras of Surinam and Venezuela being published in those
countries. Other staff work on South America includes that on Begonia
in Colombia by Dr. Schubert and that on Schoenobiblus, of the Thyme-
laeaceae, by Dr. Nevling, who is also preparing a treatment of the entire
family for the flora of Venezuela.
African members of the Moraceae and the genus Ficus are being studied
by Dr. DeWolf for Uganda and for the Flora of East Tropical Africa;
and African Desmodium is being treated by Dr. Schubert for the latter
flora also.
Several of our botanists are working on aspects of the flora of Asia,
which has for a long time been of much interest to the staff of the Arbore-
tum. Dr. Hu has spent most of the past year in Hong Kong. Dr. Perry
is working with our collections from Papua and New Guinea, concentrat-
ing on the family Myrtaceae. Dr. Hartley has completed the work of
identifying his general collections from New Guinea, of which the dupli-
cate series are being prepared for distribution. His special research con-
cerns the family Rutaceae in tropical Asia and his monograph of the
genus Flindersia is completed. Dr. S. Kazmi, a Mercer Fellow from
Pakistan, is undertaking a revision of the family Boraginaceae from West
Pakistan and Kashmir.
The largest herbarium project during the year was the rehousing of
the fruit and seed collection. This move was necessitated by the appoint-
ment of a senior member to the Gray Herbarium staff. The collection is
again available for consultation in the Cambridge building. The use of
self-sealing polyethylene bags for fruit storage is in the experimental
stage and, hopefully, this technique will permit significant growth with-
out requiring additional floor space.
The importance of the herbarium as a scientific tool cannot be under-
estimated. The scientific needs which use of the herbarium fulfills have
been partly demonstrated already. Each year, however, because of new
techniques or new methods in which old techniques are employed, along
with the growth of the collection, the demands on it become greater and
its overall usefulness is extended. In addition to its use by the resident
staff, many parts of the collection are studied by scholars from other
institutions, usually through a system of inter-institutional loans. Dur-
ing the year just ended loans to other institutions continued at a very
high level: 154 loans to 23 foreign and 39 domestic institutions, the total
of specimens loaned being 19,351. On the other side of the ledger, 12,300
specimens (110 loans from 31 foreign and 27 domestic institutions) were
borrowed for study by our staff. ;
Added to the herbarium this year were 26,985 sheets, increasing the
collection to 908,925 sheets. Of the total collection 136,556 are deposited
in the herbarium of cultivated plants in Jamaica Plain, the remainder in
the collection in Cambridge. While herbarium growth is at a satisfactory
638 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
level, the problem of adequate space to house the collection properly in-
creases proportionately. Several areas are seriously overcrowded and
some emergency measures have already had to be taken. We hope for
adequate expansion space before the need becomes desperate.
The Department of Botany of the Smithsonian Institution is recording
data on type collections in its herbarium. Test cards on a few selected
genera have been sent to other herbaria and data from our herbarium
have been supplied for this project. Through cooperation we learn the
kind of information required, the time and effort needed to record it, the
methods of recovery, and most important, we gain further knowledge on
the accessibility of information in our own collections.
The cooperative project with the Universidad Nacional Autonoma de
México is utilizing the computer facilities of that university to store
and process the data acquired under the project entitled “Environments
and Plant Resources of Veracruz.” At present, data processing tech-
niques are used in the preparation of herbarium labels and for the storage
and recovery of the label data. Bibliographic materials for Veracruz are
also being processed for retrieval by automatic means.
Library:
The use and the size of our important botanical library continued to
increase during the year, necessitating some thought about the amount
and kind of space needed to house the collections in the near future. To-
tal accessions this year were nearly double the number indicated in re-
cent reports. While the number of books purchased was up 40 percent,
binding of periodicals showed a 150 percent increase. The acquisition
of 859 bound volumes brought the total to 55,126, and 138 pamphlets
were added to the collection, which now numbers 21,236. The growth
of the library may be realized from the new total of 76,362 catalogued
items,
Four reels of microfilm and 2141 microfiche cards from various her-
baria were purchased jointly with the Gray Herbarium, to keep current
these important tools of botanical research. A total of 9175 microfiches
are now available for study.
Regular issues of the Gray Herbarium Index of American Plants, the
Index Nominum Genericorum, and the Torrey Index of Botanical Lit-
erature were incorporated to maintain the regular sequences.
We were pleased to receive a large number of single volume gifts dur-
ing the year, in addition to a special gift of volumes from the Olmsted
Associates of Boston. The library of the late Harold H. Knowlton was
also presented to the Arnold Arboretum. A bookplate was prepared with
the Knowlton family seal and the inscription “From the library of Harold
W. Knowlton, presented in loving memory by his family to the Arnold
Arboretum of Harvard University.” The Knowlton library is particularly
strong in volumes on iris, daylilies, and other horticultural groups.
Improvements have been made in the forestry collection housed in
Jamaica Plain. Additional library help has made it possible to complete
1969 } THE DIRECTOR’S REPORT 639
e: ee (scene,
Jnusual accumulations of snow marked the = months of 1969 when sec-
tions of the Arboretum could not be visited on foo
catalogue changes for the books returned from the Harvard Forest library
two years ago. This year the Arboretum accepted the transfer to Jamaica
Plain of Widener Library’s books on forestry which will be recatalogued
later.
A major rearrangement of the periodicals in Jamaica Plain was com-
pleted during the winter to provide space for growth. The American
periodicals now occupy the main library room, with the British, French,
and German language periodicals each in separate alcoves. The reprint
files and the collections of nursery catalogues and pamphlets dealing with
botanical gardens and arboreta were also reorganized.
Systematic Plant Anatomy:
At the time that the fruit and seed collection was moved to provide
additional laboratory space in the Harvard University Herbaria building,
he wood collection was consolidated. This procedure offered an oppor-
tunity to re-examine the collection of dry and preserved wood specimens
and slides and to begin some needed curatorial work. The collection
was increased this year by 400 microscope slides of woods of North
American trees, prepared in a cooperative program with the North
Carolina State University at Raleigh. As usual, specimens of wood sam-
ples and slides were sent on loan as requested.
640 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The Arboretum received as a gift the wood collection of the late Ralph
F. Perry presented by his family through Mrs. Lyman C. Morrill. This
collection of display woods, housed in a special cabinet and maintained
in Jamaica Plain, is a valuable teaching aid which includes both polished
samples and bark sections
Our collections are being used in several active research programs,
including studies of the Ulmaceae by Dr. William Stern and his students
at the University of Maryland, who have used wood samples supple-
mented by material from the living collections. Dr. Bogle is completing
his study of the floral morphology and vascular anatomy of the genera
of the Hamamelidaceae. With the assistance of Mrs. Roca-Garcia, Dr.
Howard has begun an pape sree study of the floral nectaries of the
Puerto Rican species of the genus Marcgravia. Work on nodal and
petiolar anatomy of the bate of dicotyledons continues as a major
program. Special collections of preserved material from Hong Kong
and Macau, supplied by Dr. Hu, have added four families and 40 new
genera to the study. A special project was initiated to prepare a key
to the plants of Barro Colorado Island based on the structural charac-
teristics of the node and petiole. The material was supplied for this
project by Dr. Thomas Croat, in cooperation with the staff of the Missouri
Botanical Garden, as part of their study of the Flora of Panama.
Education:
Two formal courses in the Department of Biology were offered by
members of the Arboretum staff. Dr. Howard taught an advanced one-
semester course in plant systematics, Biology 209, “Phylogeny of the
Flowering Plants,” and Dr. Wood gave Biology 103, “The Taxonomy of
Vascular Plants.” Dr. Hartley taught the Harvard University Extension
Course in general botany throughout the year.
The program of luncheon seminars in systematics held at the Harvard
University Herbaria building was conducted by Dr. Wood during the
fall semester. Several staff members presented lectures in this series
through the year. Drs. Howard, Nevling, and Schubert offered “300”
or research courses for graduate students during the year.
At Harvard, as at other universities throughout the world, there was
student “unrest” this spring. Several senior staff members, who are
members of the Faculty of Arts and Sciences, spent long hours in spe-
cial meetings of the Faculty and of the Department of Biology during
the crisis. They also carried on extended conversations with students
as a small contribution toward better communication and improved un-
derstanding
This year Dr. Howard served as one of three teachers in a Tropical
Botany Seminar sponsored jointly by the Fairchild Tropical Garden and
the University of Miami. The seminar, attended by 12 students from as
many colleges, was financed by the National Science Foundation and
offered at the Fairchild Tropical Garden. He also took part in symposia
in New York, sponsored by the Herb Society of America, and in Geneva,
_—_—_—
1969 | THE DIRECTOR’S REPORT 641
Switzerland, under the auspices of the Jardin Botanique. He presented
lectures on the Baldwin Wallace and Franklin Pierce campuses and was
sponsored by the American Institute of Biological Sciences at St. An-
selm’s College. In addition, he spoke in the series of the Torrey Botanical
Club lectures and that of the Worcester Horticultural Society.
Mr. Fordham prepared special lectures and demonstrations for visit-
ing classes from the Universities of Massachusetts, Rhode Island, and
Connecticut, and for groups from Tufts, Wheaton, and Pine Manor Junior
College. He attended the national convention of the American Rhodo-
dendron Society at Pine Mountain, Georgia, where he gave a talk on
Rhododendron Propagation. Dr. Sorensen was speaker for a seminar
series at the University of New Hampshire. Dr. Nevling gave a seminar
on the nature and diversity of climbing plants at the Universidad Na-
cional Aut6noma de México. Mr. Pride talked about the Case Estates
and about native plants of New England in a two-day lectures series
on Nantucket, in addition to speaking for the Worcester and the Massa-
chusetts Horticultural Societies. Dr. Wood participated in the sym-
posium at the Virginia Polytechnic Institute on the distributional history
of the biota of the Southern Appalachians. His lecture was on “Some
Floristic Relationships Between the Southern Appalachians and Western
North America.”’ In addition, nearly all staff members filled one or more
lecture engagements with garden clubs.
Educational displays utilizing materials from the Arboretum collections
were prepared by the staff for the Fall Harvest Show of the Massachu-
setts Horticultural Society and an Iris exhibit for the Worcester County
Horticultural Show. Mr. Fordham and Mr. Williams presented lectures in
special programs held at the Spring Flower Show of the Massachusetts
Horticultural Society.
The Arnold Arboretum Achievement Award for Botanical or Horti-
cultural Excellence was established last year through a special gift for
that purpose. The award is made to an outstanding student in one of the
high schools in the vicinity of the Arnold Arboretum. This year the
award, a choice of books and plant specimens, was made to Mr. Stephen
Grace, of Jamaica Plain High School, who plans to continue his education
at Salem State College.
Travel and Exploration:
Members of the staff travelled widely during the past year. Dr. Au
returned to Hong Kong where she continued her work towards a revised
flora of Hong Kong and the New Territories. She again taught a class at
Chung Chi College in exchange for facilities for collecting and drying
herbarium specimens. Several new records of plant distribution were ob-
tained for the islands.
Mr. Pride joined a trip to India and Nepal, conducted by Dr. Oleg
Polunin, during which he visited montane areas around Darjeeling and
Katmandu. On the return trip he visited Wisley; Wageningen, and the
642 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Belmonte Arboretum in Holland; and the Floralie Internationale in Paris.
He collected some herbarium specimens and obtained some seeds for
trial at the Arnold Arboretum.
Dr. DeWolf, with the support of a grant from the National Science
Foundation, made field studies of the species of Ficus in Venezuela. He
was able to locate and study 97 trees, or populations of 15 taxa, to obtain
ecological and morphological data from living plants. Several species
which were considered rare or uncommon on the basis of herbarium rec-
ords proved to be abundant in restricted locations. He was also able to
make 38 collections of fig insects for the cooperative study of Mr. J. T.
Wiebes, of the Rijksherbarium, Leiden. Herbarium specimens in sets
of several duplicates were prepared, dried, and returned with the coopera-
tion of the staff of the Instituto Botanico in Caracas. The aid of many
people who assisted Dr. DeWolf is acknowledged with gratitude, among
them, Drs. Tobias Lasser and Leandro Aristeguieta, of the Instituto
Botanico; Dr. José Rafael Garcia, of the Ministerio de Agricultura; Lic.
José de Jesus San José, of the Sociedad Venezolana de Ciencias Naturales;
and Dr. Argirmiro Bracemonte, of the Universidad de la Region Centro-
Occidental.
Dr. Nevling made a trip to Mexico to continue his collaboration with
Dr. Arturo Gémez-Pompa and the staff of the Universidad Nacional
Autonoma de México and the Jardin Botanico in their investigations of
the flora of the state of Veracruz.
Dr. Howard had completed the biological study of the elfin forest on
Pico del Oeste in Puerto Rico as proposed in the original N.S.F. grant,
when an unfortunate airplane crash opened a new area of study on a
ridge less than a mile from the Pico del Oeste study site, where the elfin
forest vegetation was removed for a distance of 300 yards. The exposed
site was visited and marked for subsequent studies of regrowth and the
development of adventitious shoots, invasion of new species, or the re-
placement by taxa of the present vegetation, and for erosion under the
heavy rainfall conditions of the area.
Members of the staff attended professional meetings in their areas of
interest, most of which have been cited in other sections of this report.
Dr. Howard was an invited speaker on the occasion of the 150th an-
niversary of the founding of the Jardin Botanique in Geneva, Switzer-
land. A brief vacation following the assembly permitted a visit to the
tundra areas of Norway to obtain kodachromes useful in teaching.
Dr. DeWolf, again with the aid of a grant from the National Science
Foundation, was able to visit and study at herbaria in England, Germany,
and the Netherlands in connection with his research on the species of
Ficus in Africa and Dorstenia in the New World.
Gifts and Grants:
By permission of the President and Fellows of Harvard University and
with the aid of members of the Committee to Visit the Arnold Arboretum,
invitations were sent this year to solicit membership as Friends of the
1969] THE DIRECTOR’S REPORT 643
Many of the conifers lost
An mple of snow damage to a hemlock tree.
their he while others were stripped of lower branches.
The Friends are an informal group of contributors,
the general work or special col-
embers have participated in the
Arnold Arboretum.
some of whom have been supporting
lections for over twenty years. Its m
open houses and the popular classes in horticulture and botany offered
by the staff and have shared in a program of plant distribution and hardi-
The Friends supplied active political support in ceria
Id
— testing.
a ski tow in the
a bill filed in the Massachusetts legislature to bui
rian Arboretum, and again this year in opposing a swimming ate and
recreational area on the grounds. Over 300 new Friends have joined to
help support the Arboretum during the past year, and it is our hope that
many more will participate as the Arboretum approaches its Centennial
ear,
644 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
A Centennial Fund has been established by the Treasurer of the
University for gifts to be used during 1972 or in anticipation of it. Part
of the development of the Weld-Walter tract has been made possible by
this extra financial aid. We are particularly grateful for the generous
but currently anonymous gifts to be capitalized until 1972 which are to
support field work, plant introduction, and work in special areas of hor-
ticulture.
Work on the Generic Flora continued with support from a grant by
the National Science Foundation, as has Dr. DeWolf’s work on Ficus.
Throughout the year the Arnold Arboretum receives many gifts of
living plants, books, herbarium specimens, and articles of scientific or
historical value which are acknowledged individually by the staff and by
the University.
Publications:
The Arnold Arboretum publishes regularly a scientific quarterly, the
Journal of the Arnold Arboretum, and a popular bulletin, Arnoldia, is-
sued in twelve single or combined numbers. The Journal gives priority
to technical papers by members of the staff but accepts papers from other
authors when the subject matter concerns our collections or involves
topics of particular relevance to the work of the Arboretum. Under the
editorial direction of Dr. Bernice Schubert, 29 articles by 40 authors
were published during the past fiscal year for a total of 582 pages. Sub-
scriptions have increased substantially in the last two years, particularly
since the first forty-five volumes have become available in a reprint edi-
tion. Circulation is the responsibility of Miss Dulcie Powell, who has
recently completed a reorganization of the files and records.
Arnoldia was edited, and largely prepared, this year by Dr. Wyman.
A total of ten numbers containing 119 pages was published. 1 he largest
single issue was an article by Dr. DeWolf entitled ‘“Notes on making an
herbarium.” This is a modern presentation of the techniques of prepar-
ing specimens and contains an excellent bibliography of special articles
on the subject. It will replace an article by the late Ivan M. Johnston
which is long out of print. Up to now this special number has been re-
quested by twenty-two colleges.
Two special publications were issued during the year. The booklet
“Through the Arnold Arboretum,” with photographs by Mary Rosen-
feld, text by Stephanne Sutton, and art work by Pamela Bruns, is a
popular guide to the living collections of the Arboretum. Some of Mrs.
Rosenfeld’s fine photographs were reproduced in the Harvard Alumni
Bulletin of March 17, 1969, an issue devoted to Harvard’s botanical
collections. The Arboretum story, entitled “Harvard is Green,” is avail-
able for distribution as reprints.
The second special publication, Flowers of Star Island, is a study of
the vegetation on the Isles of Shoals, a conference center located off the
coast of Portsmouth, New Hampshire. Dr. Howard prepared the text
and Helen Roca-Garcia the line drawings and silhouettes.
1969] THE DIRECTOR’S REPORT 645
Mercer Fellows:
A portion of the income from the bequest of Mrs. Martha Dana Mer-
cer is used annually as “Mercer Research Fellowships.’ In most cases
the fellowships permit the holder to live in Cambridge or Jamaica Plain
while using the collections of the Arnold Arboretum for his special re-
search studies. A few fellowships have been awarded to individuals who
wished to work with members of the Arboretum staff in order to learn
a particular technique or to become experienced in the operation of
various units within the Arboretum. This year for the first time a fel-
lowship was awarded to a graduate student to pursue an academic pro-
gram leading to a degree, under the direction of a member of the staff.
Mr. Mario Sousa-Sanchez from Mexico, who was admitted to the Gradu-
ate School of Arts and Sciences, will be engaged in studies toward a Ph.D.
degree. His research interest and thesis project concern the genus Loncho-
carpus, an arborescent member of the Leguminosae, as it occurs in
tropical America.
Four scholars were appointed Mercer Research Fellows for all or part
of the year. Arthur Charles Gibson, of Miami University, for work with
the cultivated plants of New England; Sister Mary Victoria Hayden, of
Catherine Spalding College, for work on the family Rubiaceae; Syed Mo-
hammed Anward Kazmi, of Peshawar University, Pakistan, for work on
the Boraginaceae of West Pakistan and Kashmir; and Robert William
Long, University of South Florida, for work on the family Acanthaceae.
Bibliography of the Published Writings of the Staff
July 1, 1968 — June 30, 1969
BarANov, A. I. Fruits of Canarium album (Lour.) Raeusch — Source of little
known foods and medicines of the Chinese. Quart. Jour. Taiwan Mus.
20: 367-374. 1967.
Bocte, A. L. Evidence for the hybrid origin of Petasites warren and P.
vitifolius. Rhodora 70: 533-551. 1968.
+Brizicxy, G. K. A proposal for conservation of the generic name 5015. Kos-
teletzkya Presl, 1835, against Thorntonia Reichenbach, 1828. Taxon 17:
Sal, 3352.
Sorbus anil the problem of generic typification. Jour. Arnold Arb.
49: 502-508. 1968.
05, a proposed starting point for the nomenclature of subgeneric
taxa of vascular plants. Taxon 17: 659, 660. 1968.
W. L. Stern. Notes on the distribution and habitat of Columellia.
Jour. Arnold Arb. 50: 76-79. 1969.
_ W. L. Stern, & R. H. Eype. Comparative anatomy and relationships
of Columelliaceae. Jbid. 50: 36-75. 1969.
DeWotr, Gorpon, P., Jr. Notes on making an herbarium. Arnoldia 28: 69-
111. 1968.
——. Dr. Donald Wyman. /bid. 29: (Suppl.) 3 pp. March 7, 1969.
—. The introduction of our hardy Stewartias. /bid. 29: 41-48. 1969.
646 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
Forpuam, A. J. Hastening germination of some woody plant seeds with im-
permeable seed coats. Internat. Pl. Prop. Soc. Comb. Proc. 17: 223-227.
1967.
Germination of woody legume seeds with impermeable seed coats. The
British Association of Seed Analysts — Bulletin 8: 10-16. 1968.
. Vegetative propagation of Albizia. Am. Nurseryman 128: 7, 63. 1968.
. Dwarf conifers from witches’-brooms, I. Bonsai Bull. 6: 6-11. Fall
1968.
. Dwarf conifers from witches’-brooms, II. /bid. 6: 9-14. Winter 1968-
9.
. Elliottia racemosa and its propagation. Arnoldia 29: 17-20. 1969.
Howarp, R. A. Polygonaceae. In: Flora of the Netherlands Antilles, ed. A.
L. STOFFERS. 2: 88-96. 1966.
The Director’s Report. The Arnold Arboretum during the fiscal year
ended pee 30, 1968. Jour. Arnold Arb. 49: 525-544.
—— eze-dry technique for the plant collector. Rhodora 70: 410-419.
1968.
. Flowers of Star Island, The Isles of Shoals. 106 pp. Arnold Arb.
Spec. Publ. 1968.
. The botanical garden—an unexploited source of information. Bois-
siera 14: 109-117. 1969.
———. The ecology of an elfin forest in Puerto Rico, 1. Introduction and
sig gt studies. Jour. Arnold Arb. 49: 381-418. 1968.
ecology of an elfin forest in Puerto Rico, 8. Studies in stem
growth 3 form and of leaf structure. /bid. 50: 225-262. 1969.
Hu, S.-Y. ae Compositae of China VI. Quart. Jour. Taiwan Mus. 20: 283-
339.
" oa ne of China VII. /bid. 21: 1-52.
. Some interesting and useful plants of Hong Kong. as Chi Bull.
44: 10-20. 1968. Illustrated.
Neviinc, L. I., Jk. Some ways plants climb. Arnoldia 28: 53-67. 1968.
. The ecology of an elfin forest in Puerto Rico, 5. Chromosome numbers
of some flowering plants. Jour. Arnold Arb. 50: 99-103. 1969
& P. R. Retrz. Timeleaceas. Flora Illustrada Catarinense. 1-21. 1968.
O’Connor, J. Arnold Arboretum —a ‘tree trial’ garden. Gard. Chron. 164:
18-20. 1968.
. The Arnold Arboretum. Quart. Jour. Forestry 62: 71-73. 1968.
———. The Arnold Arboretum. Nurseryman and Garden Centre 146: 438,
439. 1968.
The Hunnewell Arboretum. Park Administration 1968: 30, 37. 1968.
PRIDE, G. H. Tupper Hill. Mass. Audubon Mag. 53: 14-17.
Sax, K. Possible mutagenic hazards of some food oo beverages, and in-
secticides. Jap. Jour. Genetics 43: 89-94.
SCHUBERT, B. G. Begonias for the herbarium. ke 36: ais 1969.
. Report of the Standing Committee on Stabilization: report no. 1. Reg-
num Vegetabile 60: 108-114. 1969. (Appendix E, Synopsis of rose
on botanical nomenclature, = 1969.)
SORENSEN, P. D. Discovery of a factor for reproductive self-compatibility in
Dahlia scapigera. Am. Bates Bull. 42(2): 10-14. 1969
. Revision of the genus Dahlia. Rhodora 71: 309-365. 1969.
Sutton, S. B. Through the Arnold Arboretum, text. np. Arnold Arb. publ.
1968
1969 | THE DIRECTOR’S REPORT 647
———. National Historic Landmark: Harvard’s Arnold Arboretum. Arb. Bot.
Gard. Bull. 3: 42-44. 1969.
. The Arnold Arboretum. Horticulture 47: 20, 21, 40, 41, 50. 19
VUILLEUMIER, B. S. The genera of Senecioneae in the southeastern United
States. Jour. Arnold Arb. 50: 104-123. 1969.
. E. Woo R. The typification of Cacalia (Compositae-Senecio-
neae). ak 50: 268-273. 1969.
Woop, C. E., , & G. L. WesstEr. Tautonyms and confusion in the Inter-
national Code: Taxon 17: 645-651. 1968.
Wyman, D. Metasequoia after twenty years in cultivation. Arnoldia 28: 113-
. 1968.
. Potentilla fruticosa varieties in the Arnold Arboretum. Jbid, 125-131.
9
; Plant registrations. bid. 29: 1-8. 1969.
. Some dyeing new plants worthy of trial. /bid, 9-16. 1969.
Casoron—a new weed killer to protect woody plants. bid. 21-23.
1969,
. Tree Peonies. /bid. 25-32. 1969.
. Seven gue? years of growing Rhododendrons in the Arnold Arbore-
tum. /bid. 33-4 69.
The best dis vines. Am, Nurseryman 128(7): 7-9, 70-74. 1968.
The best plants for espaliers. bid. 128(9): 10, 11, 88-94. 1968.
The best fastigiate trees. bid. 128(11): 10, 11, 90. 1968,
The best plants for hedges. Jbid. 129(1): 9, 107-114. 1969.
The best of the ground covers. /bid, 129(8): 12, 13, 92-104. 1969.
. The best plants for screening. /bid. 129(10): 11, 72-78. 1969.
. Botanic gardens and arboreta in North America. In: Supplement to
the Dictionary of Gardening, ed. 2. 201-204. 1969. Clarendon Press, Ox-
ford.
I]
Ricuarp A. Howarp, Director
648 JOURNAL OF THE ARNOLD ARBORETUM [voL. 50
The Board of Overseers of Harvard College
Committee to Visit the Arnold Arboretum
GEORGE Putnam, Chairman, Boston, Massachusetts.
GEORGE R. CLarK, Vice-Chairman, Philadelphia, Pennsylvania.
Mrs. GeorGcE L. BATCHELDER, JR., Beverly, Massachusetts.
Mrs. RALPH BRADLEY, Canton, Massachusetts.
FREDERICK D. Brown, Webster, Massachusetts.
Mrs. Paut C. Casot, Needham, Massachusetts.
WILLIAM FLemerR, III, Princeton, New Jersey.
Mrs. JULIAN W. Hitt, Wilmington, Delaware.
Henry B. Hosmer, Boston, Massachusetts.
RusseELt E. Larson, University Park, Pennsylvania.
MiItrorp R. LAWRENCE, Falmouth, Massachusetts.
Mrs. JoHNn E. Locxwoop, Bedford, New York.
Harop E. Moore, Jr., Ithaca, New York.
R. HENRY Norwes, Jr., Mentor, Ohio.
AuGuSTIN H. Parker, Dover, Massachusetts.
Mrs. RicHaArp W. Pratt, Chestnut Hill, Massachusetts.
FRANCIS W. SARGENT, Boston, Massachusetts.
Mrs. Homer N. Sweet, Boston, Massachusetts.
GEORGE TAYLOR, Kew, England.
Joun D. Warner, Commissioner, City of Boston, Department of Parks and
Recreation.
Mrs. CuHarLeEs D. WEBSTER, Islip, New York.
RIcHARD P. WHITE, Washington, D.C.
NATHANIEL WHITTIER, Medfield, Massachusetts.
Mrs. JoHN G. WILLIAMS, Gladwyne, Pennsylvania.
iy.
1969] THE DIRECTOR’S REPORT 649
Staff of the Arnold Arboretum
1968~1969
RiIcHARD ALDEN Howarp, Ph.D., Arnold Professor of Botany, Professor of
Dendrology, and Director.
Kart Sax, S.D., Professor of Botany, Emeritus.
ALFRED LINN BOocLeE, Ph.D., Assistant Curator.
PAMELA ANNE Bruns, B.A., Artist.
Micuaet AntHony Canoso, M.S., Senior Curatorial Assistant.*
CONSTANCE ELIZABETH DERDERIAN, Honorary Curator of the Bonsai Collection.
GorDON PARKER DEWOLF, Jr., Ph.D., Horticultural Taxonomist.
ALFRED JAMES ForDHAM, Propagator.
Wit1am Ep Grime, B.A., Curatorial Assistant.*
THOMAS GORDON HartLey, Ph.D., Associate Curator of Pacific Botany.
Rosert STEPHEN Hess, S.B., Assistant Horticulturist.
WINIFRED PARKER Hess, S.B., Herbarium Assistant.
HEMAN ARTHUR Howard, Assistant Horticulturist.
Suivu-Yinc Hv, Ph.D., Botanist.
THomaAs MatTrHEw KINAHAN, Superintendent, Case Estates.
Victor FERENC Marx, M.Libr., Librarian.*
Lortn Ives Nevitne, JR., Ph.D., Associate Curator and Supervisor of the
erbaria.*
Dutcie Aricta PowELL, M.A., Botanist.
GrorGE Howarp Prive, M.A., Associate Horticulturist.
HELEN Roca-GarciA, A.M., Research Assistant.
BERNICE GipUz SCHUBERT, Ph.D., Associate Curator and Editor.
PAUL DavIpsEN SorRENSEN, Ph.D., Assistant Horticultural Taxonomist.
STEPHANNE Barry SuTTON, A.B., Archivist.
Rogsert Gerow WILLIAMS, B.S., Superintendent.
CARROLL Emory Woop, Jr., Ph.D., Associate Curator
DonaLD Wymaw, Ph.D., Horticulturist.
* Appointed jointly with the Gray Herbarium.
650 JOURNAL OF THE ARNOLD ARBORETUM
INDEX
Acmopyle, 276, 337, 441
— 38
—uniflorum, 395
— ane
Adenosty]
earl ede 467
— sallei, 467
Aérial Roo
The Ecology of an Elfin
Forest in Puerto Rico, 6., 197
African Species of Hamas, Pollen Char-
469
Aletris ee 370, 382
Alisma,
enotes ambiguus, 103, 254, 257, 260
564
7
,4
Amentotaxus formosana, Aspects of Mor-
tad of, with a Note on the Tax-
nomic Position of the Genus, 432
| atonal 567
Anatomy and Ontogeny of the Cincinni
and Flowers in Nannorrhops ritchiana
(Palmae), 411
Anatomy and peerage of Columel-
ceae, Comparat
Anatomy of the Node and Vasculariza-
i Leaf. Comparative Mor-
phological Studies in Dilleniaceae, IV.,
Anatomy of he Palm Rhapis excelsa, VII.
]
owers
Anatomy, oie of Foiled ari
with Secondary Growth— An Intro-
duction, 7
Angiosperm Pollen, Cretaceous, of the
Atlantic Coastal viegs® and its Evo-
lutionary Significance,
Angraecum lindenii, 467
Anredera, 590, 594-598
, 10.
—dominicense, 101, 103, 246, 251, 261,
i — vulgaris, 6
Appendicisporites
eae roma 20, 27
rocon lin ae). 5,14
Arbor radulifera, 498
Ardisia,
— luquillensis, i 253, 256, 259, 564
Arenaria from the Bhutan Himalaya, A
— sect. Occidentales, 627
— subg. Arenaria, 626
Bi lec 626-628
— monosperma, 628
Aristea, 160
Aristeyera, 427, 428
recon 10
— g. Chamissonis,
Arnoglossum, 272,
— plant eum, 273
Arthrost tylidium sarmentosum, 251, 261
Arundel Formations, Patuxent and, 5-9
Ascarina, 6, 8
Aspects of Morphology of Amentotaxus
formosana with a Note on the Taxo-
nomic Position of the Genus, 432
Aspects of Reproduction in Saurauia, 180
Aspects of the Complex Nodal Anatomy
of the a ean 124
Asteridae,
Asteropo. ‘es
Atlantic Coastal Plain, Cretaceous An-
giosperm Pollen of the, and its Evo-
ce,
Austrotaxus,
Axis of Dracaena fragrans (Agavaceae),
The Vascular System in the. I. Dis-
tribution and Development of Primary
Strands, 370
[voL. 50
1969] INDEX 651
Avensu, Epwarp S. Aspects of the Com-
plex eo Anatomy of the Dioscor-
eaceae,
Basu, C. R., and N. C. Mayumpar. A
New Species of Arenaria from the
Bhutan Himalaya, 626
Basella, 590
Basellaceae, 590-598
so and Portulacaceae, The Gen-
of, in the Southeastern United
ui, 566
Baynton, Harotp W. The Ecology of an
3. Hill-
top
croclimate of Pico del Oeste, 80
Beaucarnia,
recu rvata,
Be panel 207, 253, 256, 259, 563
aa
Betulaceae, 16, 24,
Bhutan Himalaya, “a New Species of
Arenaria from the, 626
Bocte, A. Linn. The Genera of Portu-
lacaceae and Basellaceae in the South-
eas astern ee States, 566
Bolivar
cahpar eon eer
Brachionidium silat, 464, 465
— parvum, 252,
= an gt
Brizicky, Grorce K., and WIL L.
STERN. Notes on ihe Distribution ee
Habitat of conga
BrizIcky, RGE s Niel L.
STERN, and ue arp H. Eype. Com-
parative Anatomy and Relationships
of Columelliaceae, 36
Browntera, 27
Cacalia Jaeger -Senecioneae) ,
Le De son a of,
— sect. Cacalia, 116-118, 272
— sect. Conophora, 117, 118, 273
— sect. Eucacalia, 273
— alpina, 271
—anteuphorbium, 271
oe atriplicifolia, 272,273
hifolia, 272
— suaveolens, 272, 273
Cacalia tuberosa, 273
Calandrinia, 567, 569
Calycogonium, 242
—squamulosum, 199, 241, 253, 256, 259,
560
Calyptranthes, 2
rugii, 199, oy 253, 256, 259, 560,
ae
Camarozonosporites, 16
Campylocentrum constanzense, 465, 468
Caytoniales, 5;
Cheirolepidium, 5
chectial Sindies of Colored Leaves. The
Ecology of an Elfin Forest in Puerto
rs
_ Plants. The re of an El-
5
incinni and Flowers in Nannorrhops
ritchiana (Palmae), Anatomy and
otundus
Charts: oe 570, 584-590
sect. Caudicosae, 586, 587
myer Claytonia, 584, 586, 587
sect. Rhizomatosae, 586, 587
Cleveta 242
— albopunctata, 102, 253, 256, 259, 561
“
a, a 199, 202, 241, 247
253, 256; 259,
Columellia Notes on the Distribution and
Habitat of, 76
652 JOURNAL OF THE ARNOLD ARBORETUM
Columellia, 36-71, 76-79
— lucida, 44, 48, 77, 78
— oblonga, 76-78
— — oblonga, 40, 44, 76, 77
— — sericea, 44, 52, 76, 77
5
Columelliaceae, Comparative Anatomy
nd Relationships of, 36
Columelliaceae, 36-71
Comparative Anatom
ships of Columelliaceae,
el Morphological on in
Dilleniaceae, IV. tomy of the Node
and Vesrotabetiow: o the Leaf, 384
rar 14-18, 2
Compositae tribe Mutisieae, 620-625
— tribe Senecioneae, 104
— subtribe Gerberinae, 620-625
— subtribe e Senecioninae, 104
Concavissimisporites, 5
Conifers, Rainforest, A Revision of the
Malesian and Pacific, I. Podocarpaceae,
in part, 274-314; 315-369
i Relation-
fic
Pactra craves, 462, 465
Cupressaceae, 5, 14
Curatella americana, 395
Cyathea pubescens, 562
Cyatheaceae, 5, 14
Cycadales, 5
Dacrycarpus, 276, 315
Dacrydium, 276, 282-308, 411
— araucarioides, 284, 296-298
[voL. 50
Dacrydium oo 296
296
— gibbsiae, 285, 306
sare sy 285, 306
junghuhnii, 285
—Hscopodioke, 284, 298, 299
, 299
— — araucarioides, 284, sei 294
—— nidulum , 292, 294
novo- -guineense, "283, "286
— — panche eri,
Conifers, I. Podocarpaceae, in part,
274-314; 315-369
1969 | INDEX 653
Dendrophylax gracilis, 467 Dioscoreaceae, Aspects of the Complex
— lind Nodal Anatomy of, ee
i, 467
DeWo tr, Gorpnon P., Jr. A ey Species
bella, 450, 455, 459
— caprifolium, 458
— chrysantha, 456, 459
——— a, 456
—etrusca, 459
— ferdinandii, 449, 453
— flori , 45
— involucrata, 453, 4
ca eo ae 458, 459
a, 457
—— repens, 458
— koehneana, 456
ckii, 457
INDEX
657
Lonicera maackii podocarpa, 457
— maximowiczii sadhalignasts, 450, 454
59
—— modesta, 450
— morrowii, 455, 4
— muendeniensis, 456
— xanthocarpa, 456
— nitida, 45
— notha, 455
— orientalis, 454
— longifolia, 454
—_ eileen
— XX pse ale. chryantha, 456
— quinquelocularis eg eetiase: 457
— syringantha, 45
oo ae 452
— — wolfii, 452
— tartarica, 454
— ‘Albo-Rosea’, 455
— s ieeiaaeedls «
— X xy osteo, 455
— xylosteum
ER H. The Ecology of an
Elfin Forest in Puerto Rico, 7.
at t, and Earthworm Relationships,
AEA 26,
Magothy Formation, 18-21
racial N. C., and C. R. Basu. A
New Species of Arenaria from the
Bhutan Himalaya, 626
Pacific Rainforest Coni-
fers, A Revision of the Podocar-
paceae, in 274-314; 315-369
Marcgravia,
— sintenisii, re 205, 253, 256, 259, 561,
563
Mecranium, 103
ai 102, 199, 253, 256, 259,
$6,
enispemaca, 26
Menodor
ea td ie 273
— atriplicifolia, 273
Miconia, 10.
—foveolata, 102, 199, 247, 253, 256, 259,
563
658 JOURNAL OF THE ARNOLD ARBORETUM
segue mirabilis,
pachyphylla, ce ve 242, 246, 247,
vee 256, 259, 561
— pycnoneura, 241, es 256, 259, 561,
5
Microcachrys, 276, 441
Microclimate of Pica x Oeste, Hilltop
and Forest Influences on. The Eco ology
of an Elfin Forest in Puerto Rico, 3.,
80
Micropholis, 2
Bani aes Atay 199, 246, 254, 257, 260,
564
Microstrobus,
ae pach 103, 205, 254, 257,
Minot, 19
Mir - and K. SusramMan
Glye cos mis pentaphylla ala “
Related Indian Taxa,
Monocotyledons with ons Growth
vs wegee Anatomy of — An Introduc-
?
ak. 567, 569
Moor seltel » JR., N. W. Unt, and L. O.
hee Anatomy of the Palm
Rhapis ‘exces VII. Flowers, 138
Moraceae,
reer Pe Studies in Dilleniaceae,
Comparative. IV. Ana atomy of he
Pa and Vascularization of the Leaf,
384
Morphology of Amentotaxus formosana,
Aspects of, with a Note on the Taxo-
Mutisieae (Com tae) in the South-
eastern United eae The Tribe, 620
Myrica nagi, 357
rile edie ng 27
Myristica
My dalteraite ose, 14
Nageia, 340
[voL. 50
Nageia ovata, 357
— ritchiana, 411-430
phengsear ritchiana (Palmae), Anato-
my Onto ps5 ny of the Cincinni and
4 Jr. The Ecology of
n Forest in Puerto Rico,
Chromosome Numbers of Some Flow-
I a Species of oe from the Bhu-
n Himalaya, A, 626
New Species of eet from Suriname, A,
478
NEWELL, THom Study of the
Genus Joinvillea ( Whee :
7
Nivenia, 160
Nod of the ne
Aspects of ved Complex, 1
Node, Anatomy of the, and enh te
on of the Leaf. Com arative Mor-
phological Studies in Dilleniaceae, IV.,
Nolina,
No ea oie. 14-16, 24-26
North American Genus Fothergilla
(Hamamelidaceae), Studies in the,
Notes on the Distribution and Habitat
of Columellia, 76
Notes on West Indian Orchids, I., 462
Nothotaxus, 442
Ocotea, 103, 242
— spa thulata, 95, 102, 199, 229, 246,252,
255, 258, 561, 563
Ontogeny of fe Cincinni and Flowers
in iota ritchiana (Palmae),
atomy and, 411
Orchids, Pa on West Indian, I., 462
Oxleya
Been etm 483, 508
phar papas 397
— junceu
Pacific es Conifers, A Revisi
ne the Malesian and, I. ore
Nn part, 274-314; 315-369
Pagiophyitum, 5
Palaeotaxus, 443
@
1969 |
Palm Rhapis erie Anatomy of the.
. Flowers,
Palm subfam. ees 138
Patapsco Formation, 9-13
Patuxent and Ayia Formations, 5-9
Pecakipollis, 1
gay omia sears 205; 246, 252, 255,
ae nando 247, 252, 258, 563
Peromonolites, 7
eemannciany reticulatus, 8
Phemeranthus
Pherosphaera
Phyllocladus, a 16, 276-282, 441
— asplenifolius,
—-hepookelixs, i
— — protracta, 278
— major, i
— protractus, 278
Pico del Cests, Hilltop and Forest Influ-
ences on the Microclimate of. The Ecol-
ogy of an Elfin Forest in Puerto Rico,
Pilea,
a Pe 74D, 252.255) 2985 563
= ahee, 207,
— yunquensis, 102, 207, 252, 255, 258, 563
— reflexa, 162, 3
— dentifera, 464
— fuertesii, 464
Plicapollis, 17-20
Podocarpaceae, 5, 440
oak oe of the Male-
Podocarpaceae
sian an Pa cific Rainforest Conifers,
in part 274-314; 315-
— aspleniifolius, 277
— beccarii, 352
— blumei, 349
— caesius, 357
INDEX
Podocarpus cinctus, 332
33
— ewe 359
— japonica, 357
— kawaii, 320
— koshunensis, 357
ice siti 349
—— — ternatis, 349
— ee a 323
— mannii,
— minor, 346
— motleyi, 352
— — rotundifolia, 357
— nagi, 357
—— angustifolia, S57
Pollen, eous Angiosperm, of th
Atlantic Coastal Plain and its Evolu-
tionary Significance, 1
660 JOURNAL OF THE ARNOLD ARBORETUM
Polyradicion, 466
i ee 467
— sallei, 46
Polyrrhiza gracilis, 467
7
Populus, 22
Porocolpollenites, 17, 18
Porophyllum ela 271
Heroes
ubg. Portu ne sl S73
Portlacacen and Basellaceae, The Genera
f he Southeastern United States,
Portulacaceae, 566—
—subfam. Montioideae, 567, 584-590
— subtribe a a 578-583
— ser. Puitilacene, 567
Prestoea, 242
— montana, 199, 251, 261, 562
Primary Strands, Distribution and Devel-
opment of The Vascular System in the
oa of Dracaena fragrans (Agavaceae),
370
=i tae 380, 382
— berteriana, 200, 242, 254, 257, 260, 564
— guadalupensis, 103, 205, 254 257, 260,
564
Pterocarya, 8
Punctatricolporites, 1
Puerto Rico, The “Seg ogy of an Elfin
Forest in, 3. Hi p and Forest Influ-
ences on the ounce of Pico del
te, 80
4. Transpiration Rates and Tempera-
tures of Leaves in Cool Humid En-
vironment, 93
5: romosome Numbers of Some
Flowering sais 99
6. Aérial Roots,
197
7. Soil, Root, and Earthworm Relation-
ships, 210
8. Studies of Stem Growth and Form
and of Leaf Structure, 225
[voL. 50
9. Chemical Studies of Colored Leaves,
556
Rainforest Conifers, A Revision of the
Malesian and Pacific, I. Podocarpaceae,
in part, 274-314; 315-369
Rajania cordata, 101, 205, 247, 251, 261,
562
Ranunculales, 26
Raritan Formation, 14-18
Ravenea, 429
Relationships of Columelliaceae, Compara-
ive Anatomy and,
Renealmia antillarum, 251, 261, 563
Reproduction in Saurauia, — of, 180
Retitricolpites sige
evision of the Genus F —— (Ruta-
“4
Malesian and Pacific
Rainforest Conifers, A, I. Podocarpa-
ceae, in part, 274-314; 315-369
Rhapis, 138-152, 492
— excelsa, 376-378, 380, 382
hapis_ excelsa, Pies of the Palm,
VI 138
II. Flowers
Rhoipteleaceae 16, 24, 27
Rogersia, 8
“eo and Earthworm Relationships, Soil.
The Ecology ee 4 Elfin Forest in
Puerto Rico, 7.,
Roots, Aérial, The cae of an Elfin
orest in Puerto Rico, 6., 197
RUpDENBERG, Lity, and Peter S. GRE
A Karyological Survey of Lonicera, 1
Rugubivesiculites, 14, 16
Salicaceae, 26
22
Saurauia, Aspects of Reproduction in, 180
Saurouls, Lind
t tee 184, 188
rn ESZ, 183, 188
— chiliantha
— excelsa, 182, 184, 188
— humboldtiana, 182, 184, 188
— micayensis, 1
= omichiphi, 182, 188, 191
os peduncularis 187
—ursina, 182, 188
fe
es |
1969]
Saurauia vasicae, 186
Sauvagesia, 10.
— erecta, 102, ee 252, 255,.259, 563
+9,
Sebanincheris ington 397
— castaneifolia,
Scleria secans, ae 261, 262
— sagittatus, 272
Senecioneae in the sara United
States, The Genera of, 10
SmirH, C. EarLe, JR. ad Characteris-
tics of African je ge i“ Pager 469
SOEJARTO, Dya of Repro-
duction in See a
Soil, Root, and Earthworm Relationships.
The Ecology eo - Elfin Forest in
Puerto Rico, 7
Southeastern Un et States, The Gen
of Portulacaceae and Basellaceae in ihe.
566
Southeastern United States, The Genera of
Senecioneae in the,
Southeastern United States, —o oo
Mutisieae (Compositae) in the, 6
msporites, 1
Stellilabium helleri, 466
and Form and of Leaf
Sisittate, ‘Suu of. The Ecology of
est in Puerto Rico, 8., 225
6
RN, Witt1am L., and GEO K.
Brizicky. Notes on “te Distribution
and Habitat of Columellia, 7
STERN, aati L., GEORGE K. BrIzIckKY,
and R mparative
Anatomy and Relationships of Columel-
lia
Straseckya, 484
ones Genus
Le 9 |
°
co
or
oO
au
p
q
ry
3
rx}
°,.8
©
iy
18°
aS)
a5
oO
Study of the fgg Joi tnvillea “(Flagellari-
aceae), A, 527
Studies of Stem Growth and Form and of
af Structure. The Ecology of an
Elfin Forest in Puerto Rico, 8., 225
INDEX
661
SUBRAMANYAM, K.,
Glycosmis pentaphyl (Rutaceae) and
Related Indian Taxa, 15
Suriname, A New oe. of Ficus from,
—
seg —" 199, 241, 246, 254, 257,
o 563
Synosma, 116, 272, 273
— suaveolens, 273
es 242
rigida, 95, 100, 101, 200, 242, 254, 257,
ib 561, 564
Talinum, 571, 578-583
Taxaceae, 440
—subfam. 1: i Secale 440
Taxodiaceae, 5,
ee a 39
a ca,
a, ap ie 466
Telipogon | 466
ig Se ie of Leaves in Cool Humid
mk Transpiration Rates and.
The "Ecolo ogy of an Elfin Forest in
Puerto Rico, 4., 93
ar UeRry 26
Tetracentron, 11
7
— korthalsii subrotunda, 397
— leiocarpa, 397
—macrophy lla, 397
— masuiana, 397
ees nordtiana, 397
agent lia, 39
gg vk B. and M. H. “ey sed
cular my of Mon
as Gis Secondary Growth, 159
Tomurson, P. B., and M. H. Zowmer-
MANN. The Vascular System in the
Axis of Dracaena fragrans (Agavaceae),
662 JOURNAL OF THE ARNOLD ARBORETUM
1. soba ma i Development of
ts ands, 3
Torralasi eer ay 199, 252, 255, 258,
561,
lenin an 441
Tournonia
Transpiration Rates and Temperatures of
Trichilia pallida, 102, 199, 246, 252, 255,
Trichocline,
Tricolpites iat
Tricolpopollenites hee 14-16
Trigon obalanus doichangensis, 27
— cacalia, 271
Unt, Natatie, W. Anatomy and Onto-
geny of the Cincinni and Flowers in
Nannorrhops ritchiana (Palmae), 411
Unt, Natarie, W., L. O. Morrow, and
. E. Moors, Jr. Anatomy of the Pan
Rhapis excels VII. Flowers, 138
CR
Ulmac
United ig Southeastern, The Gen
: Portulacaceae and Basellaceae in the.
eas States, Southeastern, The Genera
of Senecioneae in the, re
United States, Southeas tern, The Tribe
Mutisieae (Compositeae) in the, 620
Urticales, 16, 27
Vacuopollis, 1
Vascular pied of Monocotyledons
with Seco co Growth — An Intro-
duction, 1
Vascular System in the Axis of Dracaena
— rans Reseatiesy. The. 1. Distribu-
velopment of Primary
as
Vascularization of the Leaf, Anatomy of
the Node and. Comparative Morpho-
logical Studies in Dilleniaceae, IV., 384
Vernonia, Pollen Characteristics of Afri-
can ies of, 469
Vernonia sect. Stengelia, 469, 472-474
— abyssinica, 477
pear ON 477
Vernonia adoensis, 476
— chevalieri, 477
_ eet te 477
— clavoana, 476
— — microcephala, 476
— crataegifolia, 477
— denudata, 477
— filigera, 475
— firma, 475
— gerberiformis, 475
— grantii, 476
— guineensis cameroonica, 477
7
— hymenolepis, 4
— incompta, 477
— insignis, 476
— iodocalyx, 476
kotschyan
— lancibracteata, 475
— lasiopus, 4
475
a Sins soe em 475
Soe lata, 475
mandrarensis, 477
— nyassae, 475
— oxyura, 475
IEE ery 475
— polymorpha adoensis, 476
6
[voL. 50
4
1969 |
Vernonia a ai 477
alli
— wittei, ie
— woodii, 476
Vitreisporites, 5, 14
Vriesea,
_ainkenieli, 251,261,
pecan ER, BERYL ie The Gen-
era of Senecioneae in the Southeastern
United States, 104
VuILLEUMIER, BeryL Stmpson. The Tribe
Mutisieae (Compositae) in the South-
eastern United States, 620
VUILLEUMIER, Bervt S., and C. E. Woon,
ectotypification of Cacalia L.
(Compositae- Senecioneae), 268
—
Wacn_er, AnstTiss B., RICHARD J. WAGNER,
and Ric =e . Howano, The Ecology
of an E est in Puerto Rico, 9.
Chemical is of Colored Leaves, 556
W.
AGNER p J., ANsTIss B. WAGNER,
and R rp A. Howarp. The Ecology
of an Elfin Forest in Puerto Rico, 9.
Chemical Studies of Colored Leaves, 556
Walchia, 443
Wallenia, 103, 242
— yunquensis, 102, 199, 227, 246, 247, 253,
257, 259, 561, 564
INDEX
663
Lag ny 567
Weaver, Ricuarp E., Jr. Studies in the
North American Genus Fothergilla
(Hamamelidaceae) , 599
West Indian Orchids, Notes on, 1., 462
Jr., and Beryt S. VuIt-
LEUM MIER. Lectotypification of Cacalia
L. (Compositae-Senecioneae), 26
Xanthorrhoea, 160
— quadrangulata, 162
Yongsonia, 612
Yucca,
— aloifolia, 162
sroeramer Mane M. H., and P. B. ToMtin-
SON ascular pane my of ana aN
ies with Secondary Growth—A
Introduction, 159
MANN, M. H., and P. B. ToMuIn-
tribution and Development of Primary
Stran 7
Zonalapollenites, 8