Rhodora [electronic resource]

Rhodora

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Vol. 77 March, 1975 No. 809

Che Nem England Botanical Club, Заг.

Botanical Museum, Oxford Street, Cambridge, Mass. 02138

Conducted and published for the Club, by

ALFRED LINN BOGLE, Editor-in-Chief
ROLLA MILTON TRYON
STEPHEN ALAN SPONGBERG
GERALD JOSEPH GASTONY
RICHARD EDWIN WEAVER

Associate Editors

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Rbodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 77 March, 1975 No. 809

ON THE EPIBIOTIC AND PELAGIC
CHLOROPHYCEAE, PHAEOPHYCEAE,
AND RHODOPHYCEAE OF THE
WESTERN SARGASSO SEA

WILLIAM J. WOELKERLING

This paper provides a taxonomic account of the epibiotic
and pelagic Chlorophyceae, Phaeophyceae, and Rhodophy-
ceae collected during six cruises to the Western Sargasso
Sea and follows two previous studies by the author (Woel-
kerling, 1972; 1973) of non-planktonic algae from this re-
gion. Earlier published records (e.g. Collins, 1917; Conover
& Sieburth, 1964; Farlow, 1914; Hentschell, 1921; Pratt,
1935; Winge, 1923) of green and red algae and of brown
algae other than Sargassum from the Sargasso Sea are
few and fragmentary, and, with one.or two exceptions, they
do not include identifications to species level.

Nearly 75 percent of the taxa encountered during this
investigation have not been reported previously from the
Sargasso Sea, and these new records raise the total known
flora of the region to include 10 Chlorophyceae, 25 Phaeo-
phyceae, and 33 Rhodophyceae. Epibiotic Bacillariophyceae
(see Carpenter, 1970) and epibiotic Cyanophyceae (see
Carpenter, 1972; Hentschell, 1921) are not treated in this

paper.

2 Ећодога [Vol. 77

Methods of sampling and processing are outlined else-
where (Woelkerling, 1973) ; voucher material (with speci-
men numbers prefaced by wJw) has been retained in the
author's personal collections, currently housed at WIS.
Other herbarium abbreviations follow Lanjouw & Stafleu
(1964).

Data provided for each taxon includes references to
records of occurrence in adjacent regions and/or of general
taxonomie value, the type locality and reported location of
the type collection (in most cases, location of types has not
been verified), known distribution based on published rec-
ords, and collection data for all specimens examined. This
information usually is followed by ecological and/or taxo-
nomic notes. In cases where specific identification has not
been possible due to fragmentary and/or very young or
small plants, the available data has been summarized briefly
at the generic level. The genus Sargassum presents special
problems regarding species identification, and these are
outlined in the discussion of that taxon.

Enibiotic taxa can be divided conveniently into two eco-
logical groups, namely the permanent element and the in-
vading element. The former includes all taxa epibiotic on
Sargassum fluitans and/or S. natans, the two brown algae
which comprise the vast bulk of Sargasso Sea vegetation,
estimated by Parr (1939) to be in excess of 4 X 10? metric
tons. These two species apparently are endemic to the
Sargasso беа. The invading element includes all other
“macroscopic” taxa (Ascophyllum, Fucus and their asso-
ciated epiphytes (Woelkerling, 1972), species of Sargassum
other than S. fluitans and S. natans, еїс.).

The permanent epibiotic element comprises both taxa
which are normally of small size (less than 1 cm tall) and
diminutive forms of taxa normally of larger stature. Spe-
cies in both groups, however, frequently bear reproductive
structures, in marked contrast to Sargassum fluitans and
S. natans which never have been found with receptacles
(see, however, Parr, 1939, page 49) and reproduce solely
by fragmentation as far as is known.

1975] Sargasso Sea — Woelkerling 3

Sincere thanks are due Mr. Gordon Volkmann of the
Woods Hole Oceanographic Institution for making arrange-
ments for the collection of samples in the Sargasso Sea and
for making passage possible for the author on one of the
cruises. Thanks are also due Dr. Elizabeth M. Gordon for
examining collections of the Ceramiaceae.

DIVISION CHLOROPHYTA
CLASS CHLOROPHYCEAE
ORDER TETRASPORALES
Family Palmellaceae

Genus Pseudotetraspora Wille, 1906
Pseudotetraspora marina Wille 1906:20, Taf. 1, Figs. 32-36.
TYPE LOCALITY : Steinviksholm, Drontheimsfjord, Norway.
TYPE: not located.

DISTRIBUTION ` apparently known only from the Sargasso
Sea and Norway.

SPECIMENS EXAMINED: Sargasso Sea: 31°N-70°W, 5.vii.1970, Volk-
mann (WJW 2735); 32709'N-64^58' W, 16.v.1970, Woelkerling (WNJW
2670); 34°N-70°W, 30.vi.1970, Volkmann (WJW 2727), 6.vii.1970,
Volkmann (WJW 2748) ; 35°54’N-70°30'W, 3.viii.1970, Moore (WJW
2901); 37°N-70°W, 12.v.1970, Woelkerling (WJW 2625); 37°30’N-
TO°W, 8.vii.1970, Volkmann (WJW 2708); 39*07'/N-70?35'W, 16.viii.
1970, Moore (WIW 2919, 2927) ; 39°30'N-71°W, 6.x.1970, Volkmann
(WJW 2871).

The small, amorphous, gelatinous thalli have been found
as epiphytes on Sargassum natans, Sargassum sp., and
Cladophora.

Howe (1920) described а Pseudotetraspora antillarum
from the Bahamas and separated it from P. marina on the
basis of apparent differences in color, shape of the gela-
tinous mass, and cell size. These criteria require further
consideration since the Sargasso Sea specimens could con-
veniently be placed in either taxon. Howe (1920), for ex-
ample, lists cell diameters of 3-7 пт for P. antillarum while
Wille (1906) gives cell diameter of 4-10 „m for P. marina.
The Sargasso Sea plants have cell diameters of 3-14 um.
A critical comparison of the types and other collections of
the two taxa may well show them to be conspecific.

4 Ећодога [Vol. 77

ORDER CHAETOPHORALES
Family Chaetopeltidaceae

Genus Diplochaete Collins, 1901

Diplochaete solitaria Collins 1901:242. Chapman 1961:69,
Fig. 73. Collins 1909:278, Fig. 99. Collins and Hervey
1917:38. Taylor 1960:53.

TYPE LOCALITY : Kingston, Jamaica,

TYPE: NY.

DISTRIBUTION: Bermuda, Jamaica, Sargasso Sea.
SPECIMENS EXAMINED: Sargasso Sea: 26?57'N-72?58' W, 26.iv.1970,

Moore (WJW 2647, 2651); 28°N-70° W, 4.111.1970, Volkmann (WJW

2435); 31°N-70°W, 5.vii. 1970, Volkmann (WJW 2734); 32°09'N-64°

5877, 16.v.1970, Woelkerling (WJW Wie 2668); 34°N-70° W, 30.vi.

1970, Volkmann (WJW 2726, 2747); 35°54’N-70° 30W, 13. viii. 1970,

Moore (WJW 2900); 36°28’N-70°29’ w 15.viii.1970, Moore (WJW

2908); 37°N-70° W, 12.v.1970, Woelkerling (WJW 2618); 38"22'М-

70°58'’W, 12.x.1970, Volkmann (WJW 2891); 39?07'N-70?35"W, 16.

viii.1970, Moore (WJW 2916); 39730/М-717 W, 6.x.1970, Volkmann

(WJW 2860, 2885).

This species appears to be a rather common component
of the Sargasso Sea flora and has been found, sometimes in
considerable numbers, on both Sargassum fluitans and S.
natans as well as on a variety of red algae, Cladophora
(Chlorophyta), and on hydroids.

Cells in the collections examined bear 1-4 setae that may
be oriented in any direction relative to one another but
generally are directed away from the substrate. As noted
by Collins (1909) the freshwater taxa often referred to
the genus Polychaectophora West and West (1903) may be
congeneric with Diplochaete (see also Printz, 1964; С. S.
West, 1908).

ORDER ULVALES
Family Ulvaceae
Genus Enteromorpha Link, 1820
Enteromorpha sp.
Two collections [31°N-69°29’W, 3.iii.1970, Volkmann
(WJW 2369) and 34°N-70°W, 7.iii.1970, Volkmann (WJW
2897) of Sargassum fluitans contained epiphytic plants

1975] Sargasso беа — Woelkerling 5

of Enteromorpha which appear similar to E. flexuosa (Wul-
fen ex Roth) J. Agardh (see Bliding, 1963, for a detailed
account of E. flexuosa). Since the Sargasso Sea plants were
all stunted (under 3 cm tall), however, specific determina-
tion could not be made with certainty.

Genus Monostroma Thuret, 1854

Monostroma pulchrum Farlow 1881:41. Collins 1909:211.
Collins, Holden, and Setchell 1900:658. Taylor 1957 :72.
TYPE LOCALITY: Watch Hill, Connecticut.

ТҮРЕ: PH.

DISTRIBUTION : Connecticut to Nova Scotia, Sargasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 39°05’N-69°48’W, 10.v.1970,
Woelkerling (WJW 2572); 39°11'N-69°24'’W, 10.v.1970, "Woelkerling
(WJW 2550).

Two small plants of Monostroma pulchrum were found
as epiphytes on Fucus vesiculosus L., which had apparently
drifted out into the Northwestern fringes of the Sargasso
Sea (see Woelkerling, 1972). Critical studies are needed
to determine whether M. pulchrum is really specifically
distinct from the more widely distributed and better known
M. oxyspermum (Kuetzing) Doty (see Bliding, 1968, p. 585,
under Ulvaria oxysperma).

Genus Percursaria Bory, 1823
Percursaria percursa (C. Agardh) Rosenvinge 1893:963.

Bliding 1963:20, Figs. 5-6. Collins 1909:197. Kylin

1949:16, Fig. 9. Taylor 1957:61; 1960:54.

Enteromorpha percursa (C. Agardh) J. Agardh Chap-

man 1961 :66, Fig. 70.

TYPE LOCALITY: Adriatic Sea.

TYPE: LD.

DISTRIBUTION: widely distributed.

SPECIMENS EXAMINED: Sargasso Sea: 34°N-70°W, 13.v.1970, Woel-
kerling (WJW 2612); 37°N-70°W, 12.v.1970, Woelkerling (WJW
2627).

In both cases the host plant was Sargassum natans. The
main axes and laterals were pleuriseriate while the smaller
laterals were uniseriate in the specimens examined.

6 Ећодога [Vol. 77

Genus Ulva Linneaus, 1753
Ulva sp.

One young plant [34°N-70°W, 7.111.1970, Volkmann
(wJw 2412) of Ulva was found as an epiphyte on a frag-
ment of the grass Spartina. Specific determination was not
possible. The Spartina fragment apparently had drifted
out from the U. S. coast.

ORDER CLADOPHORALES
Family Cladophoraceae

Genus Cladophora Kuetzing, 1843

As noted by Taylor (1960, p. 78), the genus Cladophora
has been difficult to cope with in the American tropics and
critical studies are badly needed. The recent monographs
of Soderstrom (1963) and van den Hoek (1963), which
will probably provide a basis for such studies, have been
used in making specific determinations during this investi-
gation.

Cladophora dalmatica Kuetzing 1843:263, van den Hoek

1963 :186, Figs. 601-35.

C. oblitterata Soderstrom 1963 :47, Figs. 38-54A.

TYPE LOCALITY: Split (Spalato), Yugoslavia.

ТҮРЕ: L (No. 937/281/406).

DISTRIBUTION: probably widespread.

SPECIMENS EXAMINED: Sargasso Sea: 26°57 N-72°58'W, 26.iv.1970,
Moore (WJW 2646); 31°N-69°29'W, 3.11.1970, Volkmann (WJW
2374); 34?N-70^W, 10.1.1970, Volkmann (WJW 2205), 13.v.19770,
Woelkerling (WJW 2614) ; 37°N-70°W, 12.v.1970, Woelkerling (WJW
2620); 37*30'N-70^W, 8.vii.1970, Уобтатп (WJW 2704); 38722'N-
70°58’W, 12.x.1970, Volkmann (WJW 2886); 39°07 N-70°35'W, 16.
viii.1970, Moore (WJW 2918).

Specimens up to 3 cm tall have been found both on Sar-
gassum fluitans and S. natans. In all cases the apical cells
were under 20 „m in diameter and the branch systems
showed a distinct acropetal organization.

Cladophora laetevirens (Dillwyn) Kuetzing 1843:267. van

den Hoek 1963 :128, Figs. 409-429, 433, 440.

1975] Sargasso Sea — Woelkerling 7

TYPE LOCALITY: England.

NEOTYPE: BM (H4351/60/6); see van den Hoek 1963;
p. 128.

DISTRIBUTION: probably widespread.

SPECIMEN EXAMINED: Sargasso Sea: 31°N-69°29’W, 8.11.1970,
Volkmann (WJW 2359).

The only specimen (about 2.5 cm tall) encountered during
this study occurred as an epiphyte on Sargassum fluitans.
Cladophora socialis Kuetzing 1849:416, 1854:15, pl. 71,

Fig. 1. van den Hoek 1963:43, Figs. 79-91.

TYPE LOCALITY: Tahiti.

TYPE: 1, (937/253/440).

DISTRIBUTION: Europe, Tropical Oceania, Sargasso Sea.

SPECIMEN EXAMINED: Sargasso Sea: 26°57'N-72°58’W, 26.iv.1970,
Moore (ХЈУ 2650).

The plants, up to 2 cm tall, occurred as epiphytes on Sar-
gaassum natans.

Genus Spongomorpha Kuetzing, 1843

Spongomorpha arcta (Dillwyn) Kuetzing 1849:417. Collins
1909 :359. Taylor 1957:90, pl. 6, Figs. 5-6.
Cladophora arcta (Dillwyn) Kuetzing 1843:263. Collins,
Holden and Setchell 1896 :224; 1901 :815.
TYPE LOCALITY: England.
TYPE: presumably in NMW.

DISTRIBUTION: colder waters of Europe and North Amer-

ica.

SPECIMENS EXAMINED: Sargasso Sea: 38°53’N-69°39’W, 10.v.1970,
Woelkerling (WJW 2561, 2567); 39^05'N-69^48' W, 10.v.1970, Woel-
kerling (WJW 2571).

Specimens up to 2 cm tall were found attached to plants
of Ascophyllum nodosum and Fucus vesiculosus which had
drifted out to the Northwestern fringes of the Sargasso
Sea (see Woelkerling 1972). Spongomorpha arcta is prob-
ably not a permanent component of the Sargasso Sea flora.

8 Ећодога [Vol. 77

DIVISION CHROMOPHYTA
CLASS PHAEOPHYCEAE
ORDER ECTOCARPALES
Family Ectocarpaceae

Genus Ectocarpus Lyngbye, 1819

Ectocarpus elachistaeformis Heydrich 1892:470, pl. XXV,
Fig. 14. Boergesen 1914:18, Fig. 11; 1920:435. Collins
and Hervey 1917:70. Earle 1969:132, Fig. 28. Taylor
1928 :107, pl. 14, Fig. 12; 1960:202, pl. 29, Fig. 9.

TYPE LOCALITY: New Guinea.

TYPE: probably destroyed.

DISTRIBUTION: Caribbean Islands, Gulf of Mexico, New
Guinea, Sargasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 31^N-69?29'W, 3.111.1970,
Volkmann (WJW 2382, 2384); 84°N-70°W, 10.1.1970, Volkmann
(WJW 2211).

Plants up to 5 mm tall occurred as epiphytes on Sargas-
sum natans and on an unidentified Sargassum sp. Pluri-
locular organs are relatively abundant but are not as elon-
gate-lanceolate as described by Boergesen (1914).

Genus Feldmannia, Hamel, 1989
Feldmannia irregularis (Kuetzing) Hamel 1931-1939:XVII,

Fig. 61F. Cardinal 1964:54, Fig. 29. Kuckuck 1963 :371,

Fig. 6.

Ectocarpus irregularis Kuetzing. Boergesen 1926:25,
Figs. 12-14. Chapman 1963:11. Rosenvinge et Lund
1941:50, Figs. 23-24. Sauvageau 1933 :101, Figs. 24-27.

TYPE LOCALITY: Adriatic Sea.

TYPE: L.

DISTRIBUTION: northern Europe, Canary Islands, Sar-
gasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 28°N-70°W, 4.111.1970, Volk-
mann (ХЈУ 2431); 31°N-60°29'W, 3.111.1970, Volkmann (WJW
2366); 34°N-70°W, 7.11.1970, Volkmann (WJW 2413).

1975] Sargasso беа — Woelkerling 9

The Sargasso Sea specimens occur as epiphytes on Sar-
gassum fluitans and S. natans and agree well with the ac-
count of Feldmannia irregularis given by Cardinal (1964).
Plurilocular sporangia are common. Chapman (1963) re-
cords this taxon (as Ectocarpus irregularis) from Jamaica
and, following Boergesen (1941), regards E. rallsiae (=
Giffordia rallsiae (Vickers) Taylor (1960, p. 208), a widely
distributed taxon in the American tropics) as conspecific.
Earle (1969), however, maintains G. rallsiae as a distinct
species. The precise relationships of G. rallsiae and Feld-
mannia irregularis thus remain uncertain and further criti-
cal studies of the type and other collections of the two taxa
are needed.

Genus Giffordia Batters, 1893
Giffordia conifera (Boergesen) Taylor 1960:207. Earle
1969:135, Fig. 21.
Ectocarpus coniferus Boergesen 1914:8, Figs. 5, 6. Col-
lins and Hervey 1917 :69.
TYPE LOCALITY: U.S. Virgin Islands.

TYPE: 6.
DISTRIBUTION: Sargasso Sea and tropical western Atlan-
tic.

SPECIMENS EXAMINED: Sargasso Sea: 28°N-70°W, 4.iii.1970, Volk-
mann (WIW 2430); 34°N-70°W, 7.11.1970, Volkmann (WJW 2416).

The Sargasso Sea specimens occur epiphytically on Sar-
gassum fluitans and S. natans and agree with the original
account of Boergesen (1914). Only plurilocular sporangia
are present. Hamel (1931-39) considers this taxon to be
conspecific with Feldmannia irregularis, but Earle (1969)
maintains it as а distinct species. The status of the taxon
will remain questionable until a critical study of all the
ectocarpoid algae of the western tropical Atlantic is un-
dertaken.

Giffordia mitchelliae (Harvey) Hamel 1939:XIV, Fig. 61c, d.

Cardinal 1964:45, Fig. 23. Earle 1969:138, Fig. 24.
Taylor 1960:206, pl. 29. Figs. 1-2.

10 Ећодога [Vol. 77

Ectocarpus mitchelliae Harvey 1852, p. 142, pl. 12 g.
Boergesen 1914:6, Figs. 3-4. 1941:7, Figs. 1-5. Collins
and Hervey 1917:69.

TYPE LOCALITY: Nantucket Island, Massachusetts.

TYPE: TCD.

DISTRIBUTION ` widespread in tropical and temperate seas.
SPECIMENS EXAMINED: Sargasso Sea: 31^N-69^29'W, 23.iii.1970,
Volkmann (WJW 2361, 2381, 2383); 32°09'N-64°58’W, 16.у.1970,
Woelkerling (WJW 2666); 34^N-70^W, 10.1.1970, Volkmann (WJW

2206, 2218); 39°07’N-70°35'W, 16.viii.1970, Moore (WJW 2925).
Specimens up to 5 em tall occur as epiphytes on Sargas-

sum. fluitans, S. natans, and on an unidentified species of

Sargassum. In all cases plurilocular sporangia are abun-

dant.

Giffordia sandriana (Zanardini in Kuetzing) Hamel 1939:
XIV. Cardinal 1964:37, Fig. 18. Kylin 1947 :10, Fig. 3.
Taylor 1960 :207.

Ectocarpus sandrianus Zanardini in Kuetzing 1849 :451.
Rosenvinge et Lund 1941:44, Fig. 18.

TYPE LOCALITY: Adriatic Sea.

TYPE: L.

DISTRIBUTION: Bermuda, Europe, Sargasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 34°N-70°W, 7.111.1970, Volk-
mann (WJW 2414).

Plants up to 5 cm tall occurred as epiphytes on Sargas-
sum fluitans and bore plurilocular sporangia.
Genus Pylaiella Bory, 1823

Pylaiella littoralis (L.) Kjellman. Cardinal 1964:11, Fig. 1.
Rosenvinge et Lund 1941:5. Taylor 1957 :102, pl. 9, Figs.
1-3. Woelkerling 1972:298.

TYPE LOCALITY: Europe.
TYPE: LINN.
DISTRIBUTION: widespread.

SPECIMENS EXAMINED: Sargasso Sea: 38^553'N-69*39'W, 10.v.1970,
Woelkerling (WJW 2559); 239*11'N-69*24"W, 10.v.1970, Woelkerling
(WJW 2555).

1975] Sargasso Sea — Woelkerling 11

The plants were found as epiphytes on Fucus vesiculosus
along the northwest fringes of the Sargasso Sea and prob-
ably should not be considered as a permanent component of
the Sargasso Sea flora (Woelkerling, 1972).

ORDER SPHACELARIALES
Family Sphacelariaceae

Genus Sphacelaria Lyngbye, 1819

Sphacelaria fucigera Kuetzing. Sauvageau 1901:145, Fig.
35. Taylor 1960:210, pl. 29, Fig. 5. Womersley 1967 :199.
TYPE LOCALITY: Karak Island, Persian Gulf.

TYPE: І, (987/171/412).

DISTRIBUTION: cosmopolitan in tropical and temperate

waters.

SPECIMENS EXAMINED: Sargasso Sea: 34^N-70^W, 10.1.1970, Volk-
mann (WJW 2202); 38°22’N-70°58’W, 12.x.1970, Volkmann (WJW
2896); 39*07'N-70^35'W, 16.viii.1970, Moore (WJW 2921).

The plants were epiphytic on Sargassum natans and an
unidentified Sargassum and bore numerous propagula. One
collection (WJW 2921) also had numerous multicellular
hairs.

ORDER DICTYOTALES
Family Dictyotaceae

Genus Dictyota Lamouroux, 1809
Dictyota sp.

Two collections [32°09’N-64°58’W, 16.v.1970, Woelker-
ling (WJW 2658) and 39930'N-71?W, 6.x.1970, Volkmann
(WJW 2877)] of small plants referrable to Dictyota have
been made during this study. The former, which contained
both male and female reproductive structures and was about
5 сш tall, was found growing on a fragment of Colpomenia,
which probably became detached and drifted out from the
shores of Bermuda. It apparently shares a number of fea-
tures with D. dichotoma, but definite specific determination
was not considered possible.

12 Rhodora [Vol. 77

The latter plants were attached to a fertile piece of Sar-
gassum (origin uncertain) and were very young (less than
2 ст tall); specific determination likewise could not be
made. It seems more probable that Dictyota is an invader
rather than a permanent component of the Sargasso Sea
flora considering that both host plants themselves appear
to be invaders.

Prat (1935, p. 128) makes mention of a Dictyota cervi-
cornis from this region, but no specimens referrable to that
species have been encountered during the present study.

Genus Padina Adanson, 1763
Padina sp.

A very young Padina plant [32°09’N-64°58’W, 16.v.1970,
Woelkerling (WW 2657) ] occurred on a piece of Colpo-
menia which apparently had drifted out from Bermuda.
Specific determination was not possible and it seems prob-
able that Padina is an invader rather than a permanent
component of the Sargasso Sea flora.

ORDER CHORDARIALES
Family Elachisteaceae

Genus Elachistea Duby, 1830

Elachistea lubrica Ruprecht, Collins, Holden, and Setchell
1898 :480. Taylor 1957:140. Woelkerling 1972 :297.
TYPE LOCALITY: Okhotsk Sea.

TYPE: LE,

DISTRIBUTION: reported from eastern North America,
Greenland, and the Okhotsk Sea.

SPECIMENS EXAMINED: Sargasso Sea: 38°53'N-69°39'W, 10.v.1970,

Woelkerling (WJW 2564); 39*05'N-69?48'W, 10.v.1970, Woelkerling

(WJW 2569); 39°11’N-69°24’W, 10.v.1970, Woelkerling (WJW 2547;
2556).

An invader species attached to drifting Ascophyllum and
Fucus (see Woelkerling, 1972). The specimens have been

referred to Elachistea lubrica because of the apparent ab-
sence of moniliform paraphyses (see Taylor, 1957, p. 139),

1975] Sargasso беа — Woelkerling 18

but critical study is needed to determine whether this is a
reliable character of specific distinction. Lund (1959) and
Rosenvinge (1893) have regarded E. lubrica as a variety
of E. fucicola.

Family Chordariaceae

Genus Chordaria C. Agardh, 1817
Chordaria flagelliformis (Mueller) C. Agardh. Kylin 1947:

59, Figs. 51A, D. Lund 1959:121, Figs. 26, 27. Taylor

1957 :148, pl. 12, Fig. 6; pl. 14, Fig. 4.

TYPE LOCALITY: Denmark.

TYPE: not located.

DISTRIBUTION: cooler waters of North America and Eu-

rope.

SPECIMENS EXAMINED: Sargasso Sea: 39?05'/N-69?48'W, 10.v.1970,
Woelkerling (WJW 2576); 39°11'N-69°24’W, 10.v.1970, Woelkerling
(WJW 2554).

The plants were epiphytes on invading specimens of
Fucus (see Woelkerling, 1972) and are not considered per-
manent components of the Sargasso Sea flora.

ORDER PUNCTARIALES
Family Punctariaceae

Genus Colpomenia Derbes and Solier, 1856

Colpomenia sinuosa (Roth) Derbes and Solier. Taylor 1928:

110, pl. 7, Fig. 1, pl. 19, Figs. 3-4; 1960 :260, pl. 36, Fig. 1.

Womersley 1967 :244.

TYPE LOCALITY: Cadiz, Spain.

TYPE: probably lost.

DISTRIBUTION: widespread.

SPECIMEN EXAMINED: Sargasso Sea: 32^09'N-64*58'W, 16.v.1970,
Woelkerling (WJW 2655).

The plant collected almost certainly drifted out into Sar-
gasso Sea waters from Bermuda and does not represent à
permanent component of the flora.

14 Ећодога [Уо]. 77

Genus Petalonia Derbes апа Solier, 1850

Petalonia fascia (Mueller) Kuntze, Lund 1947 :81, Fig. 10.
Taylor 1957 :167, pl. 14, Fig. 5; pl.15, Fig. 8.
Ilea fascia (Mueller) Fries. Kylin 1947:77, Fig. 61A.
TYPE LOCALITY: Denmark.
TYPE: not located.
DISTRIBUTION: widespread.

SPECIMEN EXAMINED: Sargasso Sea: 39°11'N-69°24'W, 10.у.1970,
Woelkerling (WJW 2549).

This taxon is an invading element attached to Fucus
vesiculosus (see Woelkerling, 1972).

Genus Punctaria Greville, 1830
Punctaria latifolia Greville. Collins, Holden, and Setchell
1895:82; 1901:873; 1907:1388. Taylor 1957:166, pl. 15,
Fig. 5.
TYPE LOCALITY: Great Britain.
TYPE: not located.
DISTRIBUTION: widespread.

SPECIMEN EXAMINED: Sargasso Sea: 39°05'N-69°48’W, 10.v.1970,
Woelkerling (WJW 2573).

A single, rather small and battered plant was found at-
tached to an invading Fucus element (see Woelkerling,
1972).

Punctaria plantaginea (Roth) Greville. Rosenvinge et Lund
1947:11, Fig. 2; 1959:133, Fig. 28. Taylor 1957:166, pl.
15, Fig. 4; pl. 16, Fig. 4.

TYPE LOCALITY: Kattegat Channel between Denmark and

Sweden.

TYPE: probably destroyed.

DISTRIBUTION: widespread.

SPECIMENS EXAMINED: Sargasso Sea: 88°53'N-69°39'W, 10.v.1970,

Woelkerling (WIW 2560); 39°05'N-69°48’W, 10.v.1970, Woelkerling
(WJW 2575).

Two small plants occurred as epiphytes on Fucus; they
are not considered permanent components of the Sargasso
Sea flora (see Woelkerling, 1972).

1975] Sargasso Sea — Woelkerling 15

Genus Scytosiphon С. Agardh, 1811

Scytosiphon lomentaria (Lyngbye) C. Agardh. Rosenvinge
et Lund 1947:27, Fig. 9; 1959:103, Fig. 20. Taylor 1957:
168, pl. 15, Fig. 2; pl. 16, Fig. 3.

TYPE LOCALITY: Denmark.
TYPES C:
DISTRIBUTION: nearly cosmopolitan.

SPECIMENS EXAMINED: Sargasso Sea: 38753'N-69^39"W, 10.v.1970,
Woelkerling (WJW 2562); 39°05'N-69°48’W, 10.v.1970, Woelkerling
(WJW 2574) ; 39°11’N-69°24'W, 10.v.1970, Woelkerling (WJW 2548).

These collections occurred as epiphytes on Fucus and
probably do not represent permanent components of the
Sargasso Sea flora. The species does, however, occur in
Bermuda and along the southeastern U. S. Coast (Taylor,
1960), and may eventually be found to occur on Sargassum
fluitans or S. natans in the Sargasso Sea.

Family Striariaceae

Genus Isthmoplea Kjellman, 1877

Isthmoplea sphaerophora (Harvey in Hooker) Kjellman.
DeToni 1895:569. Kylin 1947:67, Figs. 56D-E. Taylor
1957 :156, pl. 9, Figs. 4-5. Woelkerling 1972:298.

Ectocarpus sphaerophorus Carmichael. Harvey 1846:

pl. CXXVI.

TYPE LOCALITY: Appin, Scotland.

TYPE: TCD.

DISTRIBUTION: cooler waters of Eastern North America

and Europe.

SPECIMEN EXAMINED: Sargasso Sea: 28?53'N-69739'W, 10.v.1970,
Woelkerling (WIW 2566).

The single collection contains a number of fertile plants
attached to Polysiphonia lanosa, in turn an epiphyte on a
plant of Ascophyllum, which had drifted out into the
Northwestern fringes of the Sargasso Sea (see Woelker-
ling, 1972).

16 Ећодога [Vol. 77

ORDER FUCALES
Family Fucaceae

Genus Ascophyllum Stackhouse 1809
Ascophyllum nodosum (L.) Le Jolis. Kylin 1947:84. Taylor

1957 :195, pl. 27, Figs. 1-2.

TYPE LOCALITY: Atlantic Ocean.

TYPE: LINN.

DISTRIBUTION: widespread in colder waters of the north-

ern hemisphere.

SPECIMENS EXAMINED: Sargasso Sea: 38°53’N-69°39'W, 10.v.1970,
Woelkerling (WJW 2558); 39°05'N-69°48’W, 10.v.1970, Woelkerling
(WJW 2568); 39°11'N-69°24’W, 10.v.1970, Woelkerling (WJW 2546).

The plants were found adrift along the northwest fringes
of the Sargasso Sea; there is some question as to whether
they should be regarded as permanent components of the
Sargasso Sea flora (see Woelkerling, 1972).

Genus Fucus Linneaus, 1753
Fucus vesiculosus L. Harvey 1852:71. Kylin 1947:83, Tab.
17, Figs. 53-54. Taylor 1957:192, pl. 25, Figs. 1-3.
TYPE LOCALITY: Atlantic Ocean.
TYPE: LINN.
DISTRIBUTION: widespread in colder waters of northern
hemisphere.

SPECIMENS EXAMINED: Sargasso Sea: 38^55'N-69^39'W, 10.v.1970,
Woelkerling (WJW 2563); 39°05’N-69°48’W, 10.v.1970. Woelkerling
(WJW 2570); 39°11’N-69°24’W, 10.v.1970, Woelkerling (WJW 2557).

The plants were found adrift along the northwest fringes
of the Sargasso Sea with Ascophyllum and Sargassum
natans and bore a number of epiphytes (see Woelkerling,
1972).

Family Sargassaceae
Genus Sargassum С. Agardh, 1820

The tropical American Atlantic species of Sargassum are
poorly known and specific limits and distinctions remain

1975] Sargasso Sea — Woelkerling 17

very unclear (see Taylor, 1960, p. 268). Moreover, keys to
species occurring in this region (e.g. Howe, 1920; Taylor,
1928, 1960) are based largely on vegetative features rather
than on reproductive structures, which appear to be of more
fundamental significance (Setchell, 1931; Womersley, 1954).
The situation is further complicated by the fact that the
two species of Sargassum most commonly found in the Sar-
gasso Sea apparently have never been found in a fertile
state (see, however, Parr, 1939).

During the course of this study, seven apparently distinct
species of Sargassum have been found adrift in the western
Sargasso Sea. Only three of these, however, have been
identified with any certainty. The remaining four have not
been definitely identified as yet and, following the approach
of Winge (1923), are referred to here as Sargassum A, B,
etc., to avoid further name confusion until such time as a
critical monographic study of the genus is undertaken for
this region.

Differences between the seven taxa are summarized as
follows:

Conspectus of Sargassum Taxa in
the Western Sargasso Sea

1. Plants sterile, pelagic, without evidence of a basal hold-

fog о nee a yo же Га в „о 2

2. Stems smooth; vesicles often apiculate; leaves linear,

up to 4 mm wide. .................... S. natans.

2. Stems muriculate; vesicles at most muticous; leaves
lanceolate, up to 8 mm wide. .......... S. fluitans.

1. Plants commonly fertile, normally attached, usually
showing evidence of a holdfast. ................ 8.

3. Fruiting branches often carpophyllaceous (i.e. of
mixed receptacles, leaves, and vesicles). ........ 4.

4. Receptacles pedicellate; eryptostomata tending to

be in a single row adjacent to the costa. ......
*Sargassum D."

18 Rhodora [Vol. 77

4. Receptacles not pedicellate; cryptostomata scat-

tered. ...................... “Sargassum В.”

3. Fruiting branches not carpophyllaceous (i.e. com-
posed solely of receptacles). .................. 5.

5. Receptacles pedicellate. ... .............. 6.

6. Receptacles commonly spiny; costa not spiny

or dentate. ..............................

. Sargassum hystrix var. buxifolium.

6. Receptacles not spiny; costa prominently den-
tate to spiny. ............ “Sargassum A.”

5. Receptacles sessile. .......... “Sargassum С.”

Sargassum fluitans (Boergesen) Boergesen 1914a:6, Taylor
1928 :127, pl. 18, Fig. 9; pl. 19, Fig. 5; 1960:281, pl. 39,
Fig. 2, pl. 40, Fig. 7.

Sargassum hystrix J. Ag. var. fluitans Boergesen 1914a:
11, Fig. 8. Winge 1923:23, Fig. 6 (as “Sargassum
III").

TYPE LOCALITY: Sargasso Sea.
TYPE: C.
DISTRIBUTION: known only from the Sargasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 21758'N-68*20'W, 24.iv.1970,
Moore (WJW 2639); 26*57'N-72?58'W, 26v. 1970, Moore (WJW
2649); 28°N-70°W, 4.vii.1970, Volkmann (WJW 2738); 31°N-69°
29'W, 81111970, Volkmann (WJW 2394); 31?N-70^W, 5.vii.1970,
Volkmann (ХЈУ 2731); 32°09'N-64°58’W, 16.v.1970, Woelkerling
(WJW 2654) ; 33*58'N-69*56'W, 15.v.1970, Woelkerling (WJW 2602);
24°N-70°W, 10.1.1970, Volkmann (WJW 2224), 7.111.1970, Volkmann
(WJW 2418), 6.vii.1970, Volkmann (WJW 2742), 14.viii.1970, Moore
(WJW 2933); 36*N-70*36'W, 9.xii.1970, Moore (WJW 2947); 37°М№-
70^ W, 12.v.1970, Woelkerling (ХЈУ 2616) ; 37°30'N-70°W, 8.vii.1970,
Volkmann (WJW 2712); 38°34'N-69°11'W, 19.v.1970, Woelkerling
(WJW 2606).

Sargassum fluitans is the less frequently encountered of
the two species of Sargassum endemic to the Sargasso Sea
but apparently enjoys almost as wide a distribution. The
specimens examined during this study were commonly cov-
ered with bryozoans and to a lesser extent with epibiotic
algae.

1975] Sargasso Sea — Woelkerling 19

Sargassum hystrix J. Agardh var. bwrifolium (Chauvin)
J. Agardh 1889:91, tab VII, Fig. 1. Boergesen 1914 :221.
Chapman 1963:45. Earle 1969:225, Fig. 118. Grunow
1915:399. Howe 1920:594. Taylor 1928:128, pl. 18,
Fig. 1, pl. 19, Fig. 9; 1960:279, pl. 38, Fig. 2, pl. 40,
Fig. 6.

TYPE LOCALITY: Caribbean Area (see DeToni 1895:53).

TYPES LD (7)

DISTRIBUTION: Florida to Brazil, Caribbean Islands, Sar-

gasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 36^N-70*36'W, 9.xii.1970,
Moore (WJW 2912); 39°30'N-71°W, 6.x.1970, Volkmann (WJW
2878).

The specimens examined agree well with the descriptions
and illustrations of Earle (1969) and Taylor (1960), and
probably are the same as “Sargassum УТ” of Winge (1923,
p. 25, Fig. 10). The material was collected in the immediate
vicinity of the Gulf Stream, the same region reported by
Winge (1923), and apparently is known mainly from drift
specimens (Earle, 1969; Taylor, 1928, 1960).

Both specimens bore non-carpophyllaceous receptacles
which were simple or once funcate, terete or slightly com-
pressed, verrucose or occasionally with odd spines, pedi-
cellate, and more or less racemose.

Sargassum natans L. Boergesen 1914a:7, Figs. 3-7. Taylor
1928 :128, pl. 18, Figs. 2-4, pl. 19, Fig. 13; 1960: pl. 37,
Fig. 2, pl. 40, Figs. 3, 8. Winge 1923:24, Figs. 3-5 (as
“Sargassum I & III").

TYPE LOCALITY: Sargasso Sea.

TYPE: LINN.

DISTRIBUTION: known only from the Sargasso Sea.
SPECIMENS EXAMINED: Sargasso Sea: 20°50'N-67°15’W, 24.iv.1970,

Moore (WJW 2645); 21?58'N-68'20'W, 24.iv.1970, Moore (WJW

2640); 26°50’N-71°48’W, 5.iii.1970, Volkmann (WJW 2422); 26°57'N-

72°58'W, 26.iv.1970, Moore (WJW 2653); 28°N-70°W, 411.1970,

Volkmann (WJW 2437). 4.vii.1970, Volkmann (WJW 2739), 31°N-

69°29'W, 3.111.1970, Volkmann (WJW 2393); 31°N-70°W, 5.vii.1970,

Volkmann (WJW 2732); 32°09'N-64°58’W, 16.v.1970, Woelkerling

(WJW 2662); 34°N-70°W, 10.1.1970, Volkmann (WJW 2223), 7.iii.

20 Ећодога [Vol. 77

1970, Volkmann (WJW 2417), 13.v.1970, Volkmann (WJW 2607),
30.vi.1970, Volkmann (WJW 2723), 6.vii.1970, Volkmann (WJW
2/43), 14.viii.1970, Volkmann (WJW 2931); 35°54'N-70°30'W, 13.viii.
1970, Moore (WJW 2904); 36°N-70°36’W, 9.xii.1970, Moore (WJW
2950); 86°28’N-70°29'W, 15.viii.1970, Moore (WJW 2905); 37°N-
70°W, 12.v.1970, Woelkerling (WJW 2617) ; 31*30'N-70^W, 8.vii.1970,
Woelkerling (ХЈУ 2711); 38°22'N-70°58’W, 12.x.1970, Volkmann
(WJW 2897) ; 38'34'N-69^11'W, 19.v.1970, Woelkerling (WJW 2605);
39°07'N-70°35’W, 16.viii.1970, Moore (WJW 2909) ; 39^11'N-69*24'W,
10.v.1970, Woelkerling (WJW 2644).

Sargassum natans plants comprise the vast bulk of the
Sargasso Sea macroscopic vegetation, estimated by Parr
(1939) to be up to 40 million metric tons. It also harbored
the greatest variety and quantity of epibiotic algae. The
considerable quantities of S. natans which wash up on the
shores of Bermuda, in contrast, are apparently devoid of
epiphytes (Woelkerling, personal observations).

“Sargassum A"

The single specimen [39°30’N-70°W, 6.x.1970, Volkmann
(WJW 2867)] referred to this “taxon” bears a small, dis-
coid holdfast and a sparsely branched, nearly terete main
axis with a few, scattered, long laterals whose stems are
muriculate. The leaves are lanceolate, up to 3 mm broad
and 30 mm long, finely serrate, with a prominent dentate to
spiny costa. Cryptostomata are lacking. Vesicles up to 5
mm in diameter are mostly single and pedicellate and scat-
tered among the leaves.

The receptacles are not carpophyllaceous, and are simple
or up to several times furcate, terete, verrucose, not spiny
or dentate, pedicellate, racemose, and up to 10 mm long.

While these features are most closely associated with
Sargassum filipendula var. montagnei as described in Taylor
(1960), definite specific affiliation of the specimen in ques-
tion remains uncertain.

“Sargassum B"

The single specimen [34°N-70°W, 10.1.1970, Volkmann
(WJW 2222)] lacks a holdfast and has a smooth stem with
a number of short lateral branches. The leaves are mostly

1975] Sargasso беа — Woelkerling 21

lanceolate ovate, пр to 5 mm broad and 30 mm long, serrate,
have costae without ridges or wings and bear scattered
cryptostomata. Vesicles up to 6 mm broad are short pedi-
cellate and are scattered among the leaves and receptacles.

The receptacles are generally carpophyllaceous and usu-
ally 2-4 times furcate, are terete, not dentate or spiny, are
not pedicellate and are borne in dense cymose clusters.

Although this taxon shows many of the features of Sar-
gassum vulgare C. Agardh as described in Taylor (1960),
final specific identification remains uncertain.

“Sargassum C”

The single specimen [33°58.5’N-69°56.5’W, 15.v.1970,
Woelkerling (WJW 2589)] lacks a holdfast and has mu-
riculate stems bearing lateral branches of variable length.
The leaves are linear lanceolate, up to 3 mm broad and 30
mm long, entire or finely serrate, and bear inconspicuous
costae and scattered cryptostomata. Shortly pedicellate
vesicles up to 5 mm in diameter are scattered along the
branches.

The receptacles generally are not carpophyllaceous, are
one to several times furcate, terete, verrucose, generally
not pedicellate, and are racemose.

This specimen could not be linked to any of the species
described by Taylor (1960). In some respects it is similar
to what Taylor (1960) calls Sargassum filipendula var.
montagnei, but differs in the nature of the costa and in
having non-pedicellate receptacles.

“Sargassum D"

The one specimen [349N-70?W, 10.1.1970, Volkmann
(WJW 2225)] lacks a holdfast and has a stem that is mu-
riculate in the younger portions and smooth in the older
portions. Lateral branches vary in length and bear lanceo-
late leaves up to 3 mm broad and 30 mm long which are
mostly entire, costate, and bear cryptostomata which tend
to lie in a single row on each side of the costa. Shortly
pedicellate vesicles up to 5 mm in diameter are scattered
along the lateral branches.

22 Rhodora [Vol. 77

The receptacles are carpophyllaceous, simple or once
furcate, terete, not dentate or spiny, pedicellate, and gen-
erally cymose.

Of the species described by Taylor (1960), this specimen
most closely approximates Sargassum acinarium (L.) C.
Agardh, but apparent differences in receptacle morphology
leaves some doubt as to the specimen’s true affinities.

A second specimen [37°N-70°W, 12.v.1970, Woelkerling
(WJW 2673)] shares many features with “Sargassum D"
including the linear distribution of cryptostomata, but the
racemose nature of the receptacles leaves some doubt as to
its exact relationships until further material becomes avail-
able for study.

CLASS RHODOPHYCEAE
SUBCLASS BANGIOPHYCIDAE
ORDER BANGIALES
Family Bangiaceae

Genus Asterocytis Gobi, 1879
Asterocytis ramosa (Twaites in Harvey) Gobi. Boergesen

1915:3, Fig. 1. Chapman 1963:49. Kylin 1944:6, Fig.

1d-f. Rosenvinge 1909:77, Fig. 17. Taylor 1928:132,

pl. 20, Figs. 1-2; 1960:287. Wille 1900:7, Tab. 1, Figs.

8-14.

Hormospora ramosa Twaites in Harvey 1846-51:Pl.

CCXII.

TYPE LOCALITY: Wareham, Dorsetshire, Great Britain.

TYPE: TCD.

DISTRIBUTION: widespread.

SPECIMENS EXAMINED: Sargasso Sea: 37^N-70^W, 12.v.1970, Woel-
kerling (ХЈУ 3957); 39*07'N-70*35'W, 16.у111.1970, Moore (WJW
2920).

In both cases the plants were epiphytic on Cladophora
which, in turn, was growing on Sargassum natans.

Hamel (1924), Boergesen (1927), and Pham Hoang-Ho
(1969), among others, have referred this species to the
synonomy of Asterocystis ornata (C. Agardh) Hamel.

1975] Sargasso беа — Woelkerling 28

Kylin (1944) and Taylor (1957, 1960), however, prefer to
separate the two taxa on grounds that A. ramosa 1з marine
and A. ornata freshwater. The validity of such a separation
requires further investigation, and until new evidence
comes to light, the two taxa will be kept distinct for pur-
poses of the present study.

Genus Erythrocladia Rosenvinge, 1909

Erythrocladia subintegra Rosenvinge 1909:73, Figs. 13-14.
Boergesen 1915:7, Figs. 3-4. Collins and Hervey 1917 :95.
Pham-Hoang-Ho-1969:80, Figs. 2-12. Taylor 1960:290.
TYPE LOCALITY: Hirshals, Skagerak, Denmark.

TYRE AG:

DISTRIBUTION: Europe, western tropical Atlantic, South-
east Asia.

SPECIMENS EXAMINED: Sargasso Sea: 31^N-69^29'W, 3.111.1970,

Volkmann (WJW 2387, 2391, 2392); 34°N-70°W, 10.1.1970, Volk-

mann (WJW 2212), 711.1970, Volkmann (WJW 2405, 2406) ; 36°N-
70*36'W, 9.xii.1970, Moore (WJW 2939, 2940).

All specimens occurred as epiphytes on hydroids which,
in turn, were growing on Sargassum fluitans, S. natans, or
Sargassum sp.

Erythrocladia recondita Howe et Hoyt 1916:112, pl. 12,
Figs. 1-5, pl. 13, Fig. 1. Hoyt 1920:467, pl. CXVI, Fig. 1,
pl. CXVII, Figs. 1-5.

TYPE LOCALITY: Beaufort, North Carolina,
TYPE: NY.
DISTRIBUTION : type locality, Sargasso Sea.

SPECIMEN EXAMINED: Sargasso Sea: 39°30'N-71°W, 6.x.1970, Volk-
mann (WJW 2866).

The specimens occurred epizoically on hydroids which in
turn grew on an unidentified Sargassum.

Genus Erythrotrichia Areschoug, 1850
Erythrotrichia carnea (Dillwyn) J. Agardh. Boergesen
1915:7. Collins and Hervey 1917:94. Hoyt 1920:466,
Fig. 24. Rosenvinge 1909:67, Fig. 8. Taylor 1957 :202,
pl. 28, Figs. 13-15; 1960:292. Woelkerling 1972 :298.

24 Ећодога [Vol. 77

TYPE LOCALITY: Great Britain.

TYPE: NMW.

DISTRIBUTION : widespread.

SPECIMENS EXAMINED: Sargasso Sea: 28°N-70°W, 4.iii.1970, Volk-
mann (WJW 2433); 31°N-69°29'W, 3.111.1970, Volkmann (WJW
2360; 2390) ; 34°N-70°W, 10.1.1970, Volkmann (WJW 2216), 7.11.1970,
Volkmann (WJW 2404); 39?*11'N-69*24'W, 10.v.1970, Woelkerling
(WJW 2551).

With the exception of the last cited specimen, all plants
occurred epizoically on hydroids which in turn were at-
tached to Sargassum fluitans, S. natans, or Sargassum sp.
In the other collection, the plants occurred epiphytically on
Fucus (Woelkerling, 1972).

SUBCLASS FLORIDEOPHYCIDAE
ORDER NEMALIALES
Family Acrochaetiaceae

Four species of Audouinella (A. daviesii, A. hallandica,
А. microscopica, A. saviana [== A. thuretii]) and two spe-
cies of Colaconema (C. infestans, C. secundata) occur in
the western Sargasso Sea and are the subject of a recent
detailed morphotaxonomic study (Woelkerling, 1973).

ORDER CRYPTONEMIALES
Family Corallinaceae
Subfamily Corallineae

Genus Jania Lamouroux, 1812

Jania adherens Lamouroux. Boergesen 1917:195, Figs. 184-
187. Chapman 1963:86, Fig. 85. Taylor 1928:205. Howe
1920 :589. 1960:413, pl. 49, Figs. 1-2.

TYPE LOCALITY: Mediterranean Sea.

TYPE: not located.

DISTRIBUTION: widespread in tropical and warm temper-
ate waters.

SPECIMEN EXAMINED: Sargasso Sea: 32^09'N-65^58'W, 16.v.1970,
Woelkerling (МЈУ 2656).

The single collection occurred as an epiphyte on a plant
of Colpomenia (q.v.) which probably drifted out from the

1975] Sargasso беа — Woelkerling 25

Bermuda Islands; consequently this species of Jania prob-
ably does not represent a permanent component of the Sar-
gasso Sea flora.

Jania capillacea Harvey 1853:85. Boergesen 1917:198, Fig.
188. Chapman 1963:86, Fig. 86. Collins, Holden, and
Setchell 1895:150. Howe 1920:589. Taylor 1928 :206,
pl. 29, Figs. 2, 10. 1960:413, pl. 49, Figs. 1-2.

TYPE LOCALITY: Bahia Honda, Florida.

TYPE: TCD.

DISTRIBUTION: tropical western Atlantic.

SPECIMENS EXAMINED: Sargasso Sea: 31°N-69°29'W, 3.iii.1970,

Volkmann (WJW 2217); 36°N-70°36’W, 9.х1.1970, Moore (WJW
2945); 39*07'N-70?35'W, 16.viii.1970, Moore (WJW 2926).

Jania capillacea has been found growing on Sargassum
fluitans, S. natans, and on several unidentified Sargassum
taxa. Prat (1935) previously reported this taxon from the
Sargasso Sea.

Subfamily Melobesieae

Genus Fosliella Howe, 1920

Fosliella farinosa (Lamouroux) Howe 1920:587. Chapman
1963:91, Fig. 92. Dawson 1960:30, pl. 21, Fig. 1, pl. 22,
Fig. 1l. Taylor 1960:388. Womersley and Bailey 1970:
309.

Melobesia farinosa Lamouroux. Lemoine in Boergesen
1917:170, Fig. 165. Hoyt 1920:523, Fig. 47. Taylor
1928 :211.

TYPE LOCALITY: Adriatic Sea.

TYPE: CN.

DISTRIBUTION: nearly cosmopolitan.

SPECIMENS EXAMINED: Sargasso Sea: 31?N-69*29'W, 3.111.1970,

Volkmann (WJW 2355); 34°N-70°W, 13.v.1970, Woelkerling (WJW

2608).

The plants occurred epiphytically on Sargassum fluitans
and S. natans.

Fosiella lejolisii (Rosanoff) Howe 1920:588. Masaki 1968:
23, pls. XII, XLIX, L. Taylor 1957 :253, pl. 36, Figs. 6-8.

26 Ећодога [Vol. 77

Melobesia lejolisii Rosanoff 1866:62, pl. 1, Figs. 1-13, pl.
7, Figs. 9-11. Rosenvinge 1917 :238, Figs. 156-159.

TYPE LOCALITY: Cherbourg, France.

TYPE: not located.

DISTRIBUTION: Europe, Atlantic North America, Japan,
Sargasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 28°N-70°W, 4.111.1970, Volk-
mann (WJW 2436); 31°N-69°29'W, 3.ii1.1970, Volkmann (WJW 2356,
2878); 32°09’N-64°58'W, 16.v.1970, Woelkerling (WJW 2659, 2672);
33°58.5'N-69°56.5'W, 15.v.1970, Woelkerling (WJW 2590, 2600);
34°N-70°W, 7.111.1970, Volkmann (WJW 2400, 2402); 36*N-70*36"W,
9.xii.1970, Moore (WJW 2946, 2948); 87°N-70°W, 12.v.1970, Woel-
kerling (WJW 2634); 39°07'N-70°35'W, 16.viii.1970, Moore (WJW
2928).

Fosliella lejolesii occurs as an epiphyte on Sargassum
fluitans, S. natans, Sargassum sp. as well as on other algae,
and in terms of numbers of individuals, is probably the
most common red alga in the Sargasso Sea. Only Ceramium
gracillimum (q.v.) approaches the same quantitative fre-
quency.

According to Taylor (1960, p. 387) Fosiella lejolisii is
distinguished from L. affinis and L. bermudense by having
thallus cells 6-7 wm broad rather than 9-18 um broad or
10-12 wm broad. However, cells up to 18 um broad were
found in Sargasso Sea collections, and this suggests that
the relationships of the three taxa require critical reinvesti-
gation.

ORDER CERAMIALES
Family Ceramiaceae
Genus Antithamnion Naegeli, 1847
Antithamnion antillarum Boergesen 1917 :226, Figs. 213-216.
Taylor 1960:499. Womersley and Bailey 1970 :322.
TYPE LOCALITY: St. Thomas, Virgin Islands.
TYPE: C.

DISTRIBUTION: Virgin Islands, Sargasso Sea, tropical
Pacific Ocean.

1975] Sargasso беа — Woelkerling 21

SPECIMENS EXAMINED: Sargasso Sea: 28°N-70°W, 411.1970, Volk-
mann (WJW 2426); 31?N-69?29'W, 3.11.1970, Volkmann (WJW
2376).

In both cases, plants occurred epiphytically on Sargassum
natans. One (WJW 2376) bore tetrasporangia; the other
was sterile. The genus Antithamnion in this area needs
critical reinvestigation in light of recent studies of Wollas-
ton (1968, 1971) on southern Australian and Pacific Coast
species.

Genus Ceramium Roth, 1797

Ceramium gracillimum (Kuetzing) Griffiths and Harvey.
Dawson 1962:57, pl. 20, Figs. 2-3. Feldmann-Mazoyer
in Boergesen 1952:42, Fig. 21. Nakamura 1965:136,
pl. 1, 5-6, Fig. 6.

Ceramium gracillimum var. byssoideum (Harvey) Ma-
тоуег 1938:323, Chapman 1963:178, Fig. 186. Feld-
mann-Mazoyer 1940:293, Fig. 109.

Ceramium byssoideum Harvey 1853:218. Taylor 1928:
190, pl. 27, Figs. 20, 21; 1960 :528, pl. 67, Figs. 1-3.
Ceramium transversale Collins and Hervey 1917:145,

pl. 5, Figs. 29-31. Boergesen 1918 :243.

TYPE LOCALITY: Trieste, Italy.

TYPES б,

DISTRIBUTION: widespread in tropical and warm temper-
ate seas; England.

SPECIMENS EXAMINED: Sargasso Sea: 21°58.5'N-68°20’W, 24.iv.1970,
Moore (WJW 2642); 26?50'N-71?48'W, 5.111.1970, Volkmann (WJW
2419) ; 26°57'N-72°58'W, 26.iv.1970, Woelkerling (WJW 2652) ; 28°N-
70°W, 4.111.1970, Volkmann (WJW 2428); 31°N-70°W, 5.vii.1970,
Volkmann (WJW 2733); 382°09'N-64°58’W, 16.v.1970, W oelkerling
(WJW 2661, 2669); 33°58.5’N-69°56.5'W, 16.v.1970, Woelkerling
(WJW 2591); 34°N-70°W, 13.v.1970, Woelkerling (WJW 2609), 6.vii.
1970, Volkmann (WJW 2746); 35°54’N-70°30'W, 13.viii.1970, Moore
(WJW 2903); 37°N-70°W, 12.v.1970, Woelkerling (WIW 2619);
37°30’N-70°W, 8.vii.1970, Volkmann (WJW 2705); 38°22'N-70°58'W,
12.x.1970, Volkmann (WJW 2890).

Ceramium gracillimum is a common epiphyte on Sargas-
sum fluitans, S. natans, and Sargassum sp. as well as on

28 Ећодога [Vol. 77

other algae attached to Sargassum and often occurs in
considerable numbers. One collection (WJW 2619) bore
tetrasporangial individuals; the remainder were sterile.

Taylor (1960, p. 528) maintains Ceramium byssoideum
and C. gracillimum as distinct taxa; however, following
Feldmann-Mazoyer (1940), Nakamura (1965) and others
the former is referred to the conspecificity of the latter.
Likewise, C. transversale is regarded as conspecific in ac-
cordance with Feldmann-Mazoyer (1940).

Ceramium fastigiatum Harvey in Hooker [non C. fastigi-
atum Roth = Polysiphonia fastigiata (Roth) Greville].
Boergesen 1918:241, Fig. 231. Chapman 1963:177, Fig.
184a-c. Nakamura 1965:129, pl. 1(3), Fig. 4. Taylor
1928 :191; 1957 :309, pl. 47, Figs. 3-5, 7, pl. 48, Figs. 2-4,
pl. 49, Figs. 3-4, pl. 50, Fig. 4, pl. 51, Figs. 6-7; 1960 :526,
pl. 67, Figs. 4-6.

TYPE LOCALITY: Great Britain.
TYPE: TCD.
DISTRIBUTION : widespread.

SPECIMENS EXAMINED: Sargasso Sea: 28°N-70°W, 4.111.1970, Volk-
mann (WJW 2427); 31°N-69°29’'W, 3.111.1970, Volkmann (WJW 2348,
2370, 2378) ; 34°N-70°W, 7.111.1970, Volkmann (WJW 2395); 39*07'N-
70°35'W, 16.viii.1970, Moore (WJW 2923).

This taxon occurs as an epiphyte on Sargassum fluitans,
S. natans, and Sargassum sp. All specimens examined were
sterile. The author citations of Taylor (1960) and Boerge-
sen (1918) (i.e. C. fastigiatium (Roth) Harvey) are in-
correct; Roth's taxon was described from Germany and
has been referred by Greville (1824) to Polysiphonia (see
DeToni, 1903, p. 945-6) whereas Harvey's taxon was de-
scribed from Great Britain (Harvey in Hooker, 1833).

Genus Crouania J. Agardh, 1842
Crouania attenuata (C. Agardh) J, Agardh, Boergesen
1917:230, Figs. 219-221. Chapman 1963:167, Fig. 173.
Collins and Hervey 1917:142. Harvey 1853:226, Tab.
XXXI, D. Taylor 1928:193, pl. 27, Figs. 7-9, pl. 32,
Fig. 9.

1975] Sargasso беа — Woelkerling 29

TYPE LOCALITY: Mediterranean Sea.

TYPE: LD.

DISTRIBUTION: Mediterranean, England, tropical western

Atlantic Ocean, Japan.

SPECIMENS EXAMINED: Sargasso Sea: 37°N-70°W, 12.v.1970, Woel-
kerling (WJW 2626); 39°30'N-71°W, 6.x.1970, Volkmann (WJW
2882).

Male (WJW 2626) and tetrasporangial (WJW 2882)
individuals occurred as epiphytes on Sargassum natans and
on Sargassum sp.

Genus Griffithsia С. Agardh, 1817
Griffithsia radicans Kuetzing 1862:11, tab. 33, Fig. A-C.

Taylor 1960 :515.

TYPE LOCALITY: Brazil.

TYPE: b.

DISTRIBUTION: Brazil, Sargasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 37°N-70°W, 12.v.1970, Woel-
kerling (WJW 2629); 39*30'N-71^W, 6.x.1970, Volkmann (WJW
2881).

The plants occurred as epiphytes on Sargassum natans
and Sargassum sp. and appeared to have some tetrasporan-
gial initials. They are referred to this species because of
their agreement with the description given by Taylor
(1960).

Genus Spermothamnion Areschoug, 1847
Spermothamnion investiens (Crouan in Maze et Schramm)

Vickers. Boergesen 1909:17, Fig. 10; 1917:200, Figs. 189-

190; 1920:461, Fig. 422. Collins and Hervey 1917:132.

Howe 1920:578. Taylor 1960 :520.

TYPE LOCALITY: Guadeloupe.

TYPE: РС.

DISTRIBUTION: North Carolina, Caribbean Islands, Sar-
gasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 39°30’N-71°W, 6.x.1970,

Volkmann (WJW 2863).

Female and tetrasporangial plants were found as epi-
phytes on “Sargassum A” and may not, therefore, be à

30 Ећодога [Vol. 77

permanent component of the Sargasso Sea flora. Tropical
and subtropical western Atlantic members of the Sper-
mothamneae are in need of thorough reinvestigation in
light of the recent study of Gordon (1972).

Genus Wrangelia C. Agardh, 1828
Wrangelia argus (Montagne) Montage. Boergesen 1916:
116, Figs. 125-6. Gordon 1972:40. Taylor 1928:144, pl.
20, Fig. 13, pl. 22, Fig. 6, pl. 32, Fig. 4; 1960 :502, pl. 66,
Figs. 7-8.
TYPE LOCALITY: unknown.
TYPE: probably c.
DISTRIBUTION: see Gordon 1972, p. 40.
SPECIMENS EXAMINED: Sargasso Sea: 31°N-69°29'W, 311.1970,

Volkmann (WJW 2371); 87°N-70°W, 12.v.1970, Woelkerling (WJW
2628); 39*30'N-71^W, 6.x.1970, Volkmann (WJW 2880).

Male and tetrasporangial plants occurred as epiphytes
on Sargassum natans and on Sargassum sp.

Family Dasyaceae
Genus Dasya C. Agardh, 1824

Dasya rigidula (Kuetzing) Ardissone. Howe 1920:576.
Taylor 1960 :558, pl. 72, Fig. 4.
TYPE LOCALITY: Spalato, Adriatic Sea.
TYPE: L.
DISTRIBUTION: Bermuda, Caribbean Islands, Mexico, Sar-
gasso Sea, Venezuela, Adriatic and Mediterranean Seas.

SPECIMENS EXAMINED: Sargasso Sea: 34^N-70^W, 7.11.1970, Volk-
mann (WJW 2398); 36°N-70°36’W, 9.xii.1970, Moore (WJW 2941);
39°30'N-71°W, 6.x.1970, Volkmann (WJW 2861, 2884).

Male, female, and tetrasporangial plants occurred epi-
phytically on Sargassum fluitans and on Sargassum Sp.

Genus Heterosiphonia Montagne, 1842
Heterosiphonia wurdemanni (Bailey in Harvey) Falkenberg
1901:638, pl. 16, Fig. 11. Boergesen 1919:324, Figs. 326-
328. Collins and Hervey 1917:131. Howe 1920:575.
Taylor 1928:178, pl. 25, Fig. 3; 1960:565, pl. 72, Fig. 9.

1975] Sargasso Sea — Woelkerling 31

Dasya wurdemanni Bailey in Harvey 1853:64, Tab. XV,
C.

TYPE LOCALITY: Key West, Florida.

TYPE: TCD.

DISTRIBUTION: western tropical Atlantic; Mediterranean.

SPECIMENS EXAMINED: Sargasso Sea: 34°N-70°W, 10.1.1970, Volk-
mann (WJW 2213).

A single tetrasporangial plant occurred epiphytically on
“Sargassum B"; its status in the Sargasso Sea flora must
remain in doubt until further collections come to hand.

Family Delesseriaceae
Genus Hypoglossum Kuetzing, 1843

Hypoglossum tenuifolium (Harvey) J. Agardh. Howe 1920:
564. Taylor 1960 :545, pl. 68, Fig. 2.
Delesseria tenuifolia Harvey 1853:97, Tab. XXII, Fig. B.
Boergesen 1919 :344, Figs. 340-343.
TYPE LOCALITY: Key West, Florida.
TYPE: TCD.
DISTRIBUTION: western tropical Atlantic Ocean.

SPECIMENS EXAMINED: Sargasso Sea: 34°N-70°W, 10.1.1970, Volk-
mann (WJW 2220).

A single plant was found epiphytically on “Sargaassum
B”, and until further collections come to hand, its status as
a permanent component of the Sargasso Sea flora must re-
main in doubt.

Family Rhodomelaceae
Genus Chondria C. Agardh, 1817

Two collections — one tetrasporangial [37°N-70°W,
12.v.1970, Woelkerling (WJW 2632)] and one sterile
[399077N-70935'W, 16.viii.1970, Moore (WJW 2910) ] —
contained very small (less than 1 ст long) plants of
Chondria attached to Sargassum natans. While both plants
appear to come close to the diagnosis of Chondria dasyphila
(Woodward) C. Agardh provided by Taylor (1960, p. 616),
specific affinity could not be determined with certainty on
the basis of the material at hand.

32 Ећодога [Vol. 77

Genus Herposiphonia Naegeli, 1846

Herposiphonia secunda (C. Agardh) Ambronn. Boergesen
1920 :469, Figs. 428-429. Chapman 1963:125, Figs. 130
a-b. Collins and Hervey 1917:126. Howe 1920:574.
Taylor 1928:176, pl. 25, Figs. 8-10; 1960:604, pl. 72,
Figs. 10-11.

TYPE LOCALITY: LD.

TYPE: LD.

DISTRIBUTION: western tropical Atlantic Ocean, Mediter-

ranean Sea, Adriatic Sea.

SPECIMENS EXAMINED: Sargasso Sea: 31^N-69?20'W, 8.11.1970,
Volkmann (WJW 2362); 32°09'N-64°58’W, 16.v.1970, Woelkerling
(WJW 2660); 36°N-70°36'W, 9.xii.1970, Moore (WJW 2935); 37°N-
70° W, 12.v.1970, Woelkerling (WJW 2630) ; 39*07'N-70*35'W, 16.viii.
1970, Moore (WJW 2911); 39*30'N-71^W, 6.x.1970, Volkmann (WJW
2864).

Male, female and tetrasporangial plants have been found
on Sargassum fluitans, S. natans, and on Sargassum sp.

Herposiphonia tenella (C. Agardh) Naegeli. Boergesen
1918:286, Figs. 287-289; 1920:472, Fig. 430. Chapman
1963:127, Fig. 133. Collins and Hervey 1917:126. Howe
1920:573. Taylor 1928:177, pl. 25, Fig. 11; 1960:604,
pl. 72, Fig. 12.

TYPE LOCALITY: Sicily.

TYPE: LD.

DISTRIBUTION: western tropical Atlantic Ocean; Mediter-
ranean and Adriatic Seas.

SPECIMENS EXAMINED: Sargasso Sea: 33^58.5'N-69^56.5'W, 15.v.
1970, Woelkerling (WJW 2596); 39°30'N-71°W, 6.x.1970, Volkmann
(WJW 2875).

The tetrasporangial specimens (WJW 2875) grew epi-
phytically on Sargassum hystrix; the sterile material was
found on “Sargassum C." Until specimens attached to S.
fluitans and/or S. natans are collected the status of this
taxon as a permanent component of the Sargasso Sea re-
mains in doubt.

1975] Sargasso беа — Woelkerling 33

Genus Laurencia Lamouroux, 1813

Tropical Atlantic American species of Laurencia are in
need of critical study in light of the recent investigations
of Saito (1967, 1969). Saito (1967, p. 72-73) recognizes
5 subgenera of Laurencia based on the presence or absence
of secondary pit connections in the cortex, shape of cortical
cells, presence or absence of lenticular thickenings in
medullary cell walls, and plant form (compressed or cy-
lindrical). Taylor (1960) does not provide full informa-
tion on these features, and his keys are based partly on
features of questionable taxonomic significance, thus mak-
ing specific identification of taxa difficult.

Seven collections of Laurencia have been made during
the present study, but in view of their small size (mostly
under 1 em tall) and with one exception their sterile condi-
tion, species affinities have not been determined. One of
these [34°N-70°W, 30.vi.1970, Volkmann (WJW 2729) ]
belongs to the subgenus Palisadae; the other six [319N-
69929'W, 3.11.1970, Volkmann (WJW 2361); 83958.5/N-
69956.5/W, 15.v.1970, Woelkerling (WJW 2593); 34°N-
TOOW, 6.vii.1970. Volkmann (WJW 2745) ; 37°N-70°W,
12.v.1970, Woelkerling (WJW 2636); 37°30’N-70°W,
8.vii.1970, Volkmann (WJW 2710); 39°30’N-71°W, 6.x.
1970, Volkmann (WJW 2874)] belong to the subgenus
Chondrophycus. The specimens occurred as epiphytes on
Sargassum natans and on Sargassum sp.

Genus Lophosiphonia Falkenberg, 1897

Lophosiphonia cristata Falkenberg 1901:499, Tab. 9, Figs.
7-10. Boergesen 1918:297, Figs. 295-298. Chapman
1964:125, Fig. 129. Hollenberg 1958:68. Taylor 1960:
606.

TYPE LOCALITY: Scogliera, Puntadel Posilipo, Gulf of
Napal.

TYPE: not located.

DISTRIBUTION: Mediterranean, Bermuda, Bahamas, Ja-
maica, Virgin Islands, Sargasso Sea.

84 Rhodora [Vol. 77

SPECIMEN EXAMINED: Sargasso Sea: 31°N, 69°29’W, 8.11.1970,
Volkmann (WJW 2868).

The single collection contained tetrasporangial plants
epiphytic on Sargassum fluitans.

Genus Polysiphonia Greville, 1824

As noted by Taylor (1960, p. 572-3), much taxonomic
uncertainty exists over tropical American Atlantic species
of Polysiphonia, and a critical revision of the genus for this
region is badly needed. Although several sources (includ-
ing Taylor, 1960) have been consulted during this study,
taxonomic identifications of Sargasso Sea collections have
been made mainly from the publications of Hollenberg
(1968, 1968a). The relationships of Hollenberg’s Pacific
taxa to the species names employed by Taylor (1960) for
tropical American Atlantic taxa remain uncertain.
Polysiphonia delicatula Hollenberg 1968:62, Fig. IF.

TYPE LOCALITY: Pokai Bay, Oahu, Hawaii.

TYPE: US (D1911662).

DISTRIBUTION: Hawaiian Islands, Tuamotu Archipelago,

Marshall Islands, Caroline Islands, Sargasso Sea.

SPECIMENS EXAMINED: Sargasso Sea: 28°N-70°W, 4.iii.1970, Volk-
mann (WJW 2428); 33°58'N-69°56.5'W, 15.v.1970, Woelkerling (WJW
2595); 37^N-70^W, 12.v.1970, Woelkerling (WJW 2631, 3229).

Female and tetrasporangial plants occurred as epiphytes
on Sargassum natans and Sargassum sp.

Polysiphonia lanosa (L.) Tandy, Taylor 1957:341, pl. 56,
Fig. 4, pl. 57, Figs. 14-15, pl. 59, Fig. 4. Woelkerling
1972 :298.

Polysiphonia fastigiata auct. поп. (Roth) Greville: Col-
lins, Holden, and Setchell 1895:145; 1907:1444. Far-
low 1881: 175. Harvey 1853:54.

TYPE LOCALITY: unknown.

TYPE: LINN.

DISTRIBUTION: North Atlantic Ocean.

SPECIMENS EXAMINED: Sargasso Sea: 38753'N-69?39'W, 10.v.1970,
Woelkerling (WJW 2565).

19751 Sargasso беа — Woelkerling 35

Polysiphonia lanosa occurred as an epiphyte on Asco-
phyllum nodosum (see Woelkerling, 1972) and is regarded
as an invader.

Polysiphonia poko Hollenberg 1968:70, Figs. 3A, 15, 22.
TYPE LOCALITY: North Island, Pacific Ocean.
TYPE: US (H65-113.1).
DISTRIBUTION: tropical Pacific Islands (see Hollenberg),
Sargasso Sea.
SPECIMENS EXAMINED: Sargasso Sea: 34°N-70°W, 10.1.1970, Volk-

mann (WJW 2207); 37?30'N-70^W, 8.vii.1970, Volkmann (WJW
2703); 39°30’N-71°W, 6.x.1970, Volkmann (WJW 2858).

Female and tetrasporangial plants occurred as epiphytes
on Sargassum natans and Sargassum sp.

SUMMARY

The epibiotic and pelagic algal flora of the western Sar-
gasso Sea has been extended to include ten Chlorophyceae,
25 Phaeophyceae, and 33 Rhodophyceae. Nearly 75% of
these taxa are newly reported for the Sargasso Sea. Per-
tinent references and taxonomic and ecological data are
provided for each taxon, and indication is provided as to
whether each species is likely to be a permanent component
of the Sargasso Sea flora or merely a temporary invading
element.

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FELDMANN-MAZOYER, G. 1940. Recherches sur les Ceramiacees de
la Meditierranee Occidentale. Algiers.

Соврох, E. 1972. Comparative morphology and taxonomy of the
Wrangelieae, Sphondylothamniae, and Spermothamniae (Cera-
miaceae, Rhodophyta). Aust. Jour. Bot. Suppl. Ser. 4: 1-180.

GREVILLE, R. K. 1824. Flora Edinensis. Edinburgh.

GRUNOW, А. 1915. Additamenta ad cognitionem Sagassorum. Verh.
Zool.-bot. Ges. Wien 65: 329-448.

HAMEL, G. 1924. Floridees de France II. Rev. Algol. 1: 427-457.

1931-1939. Pheophycees de France. Paris.

Harvey, W. H. 1833. Cryptogamia algae. Div. II. Confervoideae.
In: W. J. Hooker, The English Flora of Sir James Edward
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1846, 1849, 1851. Phycologia Britannica. Vols. 1-8.
London.

38 Rhodora [Vol. 77

1852. Nereis Boreali-Americana. Part I. Melano-
spermae. Smithson. Contr. Knowl. 3(4): 1-150. pls. 1-12.
1853. Nereis Boreali-Americana. Part II. Rhodosper-
mae. Smithson. Contr. Knowl. 5(5): 1-258. Pls. 13-36.
HENTSCHELL, E. 1921. Ueber den bewuchs auf den treibenden
Tangen der Sargassosee. Mitt. Zool. Staatsinst. Zool. von Ham-
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HEYDRICH, Е. 1892. Beitraege zur Kenntnis der Algenflora von
Kaiser Wilhelms Land (Deutsch Neu Guinea). Ber. dt. bot. Ges.
10: 458-485.
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1968a. An account of the species of Polysiphonia of
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1975] Sargasso беа — Woelkerling 39

Masaki, Т. 1968. Studies on the Melobesioidae of Japan. Mem.
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40 Ећодога [Vol. 77

1957. Marine algae of the Northeastern coast of North
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1906. Algologische untersuchungen an der biologischen
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1973. The Audouinella complex (Rhodophyta) in the
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WOLLASTON, E. 1968. Morphology and taxonomy of southern Aus-
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1971. Antithamnion and related genera occurring on
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WOoMERSLEY, Н. B. S. 1954. Australian species of Sargassum, sub-
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1967. А critical survey of the marine algae of south-
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& A. BarLEv. 1970. Marine algae of the Solomon
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DEPARTMENT OF BOTANY
UNIVERSITY OF WISCONSIN
MADISON, WISCONSIN 58706

SAXIFRAGES ON MOUNT WASHINGTON

In Rhodora 69: 483-486, three brief articles were pub-
lished regarding the rediscovery of Saxifraga Aizoon Jacq.
and Saxifraga cernua L. in Huntington Ravine on Mt.
Washington. These plants were first discovered by Dr. John
Churchill in 1939 but were apparently not seen again until
1967 when Steele, Hodgdon and James Teeri relocated them.
Although it was not possible to reach the plants because of
the steepness of the cliff, observations were made through
field glasses. Saxifraga Aizoon was growing in some abun-
dance on a nearly flat ledge about twenty feet above a van-
tage point which was reached without too much difficulty.
The plants in full bloom with conspicuous typical lime
encrusted leaves, were unmistakable even when viewed at
a distance of 100 feet from the bottom of the cliff. On a
steep shelf at a distance of forty feet from the vantage
point, Hodgdon detected a single flower which he believed
to be Saxifraga cernua, a plant he had become familiar with
on an Alaskan expedition. Steele, although not familiar
with the plant was skeptical and unwilling to accept the
identification. The plant was completely inaccessible so
there seemed to be no way to resolve the question without
the aid of rockclimbers.

Upon learning of the dilemma, Countryman undertook
to secure the aid of members of Norwich University’s
Mountain Rescue Team. Accordingly, another expedition
was organized in 1970 consisting of Countryman, Hodgdon,
Steele and three technical rock climbers from Norwich.
The vantage point below the Saxifraga Aizoon shelf was
easily reached but it was then discovered that nearly all
the soil and vegetation was gone from the ledge, presum-
ably having been removed by avalanche or flood, a matter
which emphasized the extreme ecological instability of the
whole area. The date of this expedition, Aug. 2, was past
the flowering time of the saxifrages and at first none were
in evidence. Soon Hodgdon, with the aid of field glasses,

41

42 Rhodora [Vol. 77

was able to locate some plants of 5. Aizoon оп a ledge above
the original shelf. Ray Quirk, a rock climbing instructor
at Norwich, undertook to investigate, assisted by John P.
Quirk and Jon W. Walsh, both undergraduates at that
institution. The ledge was very steep and the rocks un-
stable, but with the use of direct aid in the form of pitons
and a sling and tension from below, Quirk was eventually
able to reach the site. Guided by instructions from below
he was able to obtain small but unmistakable portions of
both saxifrages, these being the first collections from Mt.
Washington since 1939. It would have been desirable to
investigate another ledge but the lateness of the hour pre-
vented this.

After this trip, in discussing the 1967 expedition, a
friendly dispute broke out between Steele and Hodgdon as
to the exact location of the Saxifraga cernua observed at
the time. To settle this point, and because one promising
shelf still had not been investigated, another expedition was
organized by Steele in 1971, with a date selected for what
was believed to be the height of the flowering season of
both plants. Two rock climbers accompanied the expedition.
From the vantage point eighty feet up the cliff, S. cernua
was readily observed in flower on a vegetation covered
shelf below the original S. Aizoon ledge. With the aid of a
rope and a good belay from Paul Henle, Nathaniel Steele
was able to reach the slope. He made some careful observa-
tions and collected one fragment of a plant plus some
leaves. There were seven plants of S. cernua in bloom plus
some patches of leaves. The slope was steep and unstable
with many loose rocks. It would not be difficult for all the
vegetation to be dislodged in a slide.

Steele (senior) searched the whole area carefully with
field glasses and eventually located a small clump of S.
cernua leaves in a small pocket near the original S. Aizoon
ledge. On the higher ledge explored in 1970, a considerable
amount of S. Aizoon was observed in full bloom. The onset
of a thunderstorm prevented any further exploration, but
presumably S. cernua was there also.

1975] Saxifrages — Steele 43

The narrow gully continues very steeply above this site
nearly to the head of the ravine, but overhanging rock
makes it impossible to see into it from the bottom; thus it
is possible there are colonies of saxifrage farther up, seeds
from which might occasionally wash down апа colonize
suitable areas. It would appear this possibility could be
confirmed only by a team of strong rock climbers.

For any who might be inclined to investigate this or other
rare alpine plants, the authors suggest the following as a
result of their experiences. High powered field glasses are
very useful. Careful notes should be taken at the time as to
what is observed. Drawings and photographs of the locality
would be most helpful. A camera equipped with a tele-
photo lens would make an excellent record of the plant. All
information thus obtained could be put on an herbarium
sheet.

It may seem strange that Dr. Churchill was able to dis-
cover new plants in an area that had long been heavily
botanized. He was fortunate in that he was doing his
explorations during the short flowering season, His very
conscientious exploration and mountaineering ability no
doubt led him to the vantage point eighty feet from the
bottom from which he could observe the plants without field
glasses. How he was able to manage the last twenty feet
without a rope and a belay from a second remain a mystery
to the various technical rock climbers who have visited the
area. The area is ecologically unstable and there is a slight
possibility that he collected his plants from a station lower
down that is now extinct. Because of the instability the
authors feel that further exploration is inadvisable in that
it might lead to destruction of the very limited habitat.

Records for the plants now exist in the herbaria of the
University of Michigan, the New England Botanical Club,
the University of New Hampshire, Norwich University and
the Steele private herbarium in Tamworth, New Hampshire.

FREDERIC L. STEELE
THE WHITE MOUNTAIN SCHOOL
LITTLETON, NEW HAMPSHIRE 03561

RUDBECKIA AURICULATA (PERDUE) KRAL,
A SPECIES DISTINCT FROM R. FULGIDA AIT.

ROBERT KRAL

During field work toward a flora of Alabama I have come
across several additional populations of a very tall, smooth-
ish, rhizomatous Rudbeckia, which was originally described
by Perdue (1961) from a single population in Covington
County as var. auriculata of R. fulgida Ait. Admittedly,
R. fulgida is extremely variable, presenting a broad spec-
trum of radiate extremes (Perdue treats seven in his 1957
treatment of the complex) over a broad range of habitats
and geography. However, this particular Rudbeckia varies
from the other “fulgidas” so much, and so consistently, that
to treat it as another variety of R. fulgida would be incon-
sistent with species concepts in Rudbeckia. Perhaps I can
best demonstrate this situation by offering descriptions of
R. fulgida and Р. auriculata (Р. fulgida var. auriculata
Perdue), these based on 45 samples of the former and ten
of the latter. The description of R. fulgida is broadened to
include the three varieties now known to occur within Ala-
bama and Tennessee, which are possibly sympatric with
R. auriculata.

R. fulgida Ait. var. fulgida. Plate 1(2) ; Plate 2(3).
R. fulgida Ait., Hort. Kew. 3: 251. 1789.
R. chrysomela Michx. Fl. Bor. Am. 2: 143. 1803.
R. truncata Small, Bull. Torr. Bot. Club 25: 478. 1898.
R. acuminata Boynt. & Beadle, Fl. S.E.U.S. ed. 1. 1256.

1903.

R. foliosa Boynt. & Beadle, Fl. S.E.U.S. ed. 1. 1256. 1903.
R.tenax Boynt. & Beadle, Fl. S.E.U.S. ed. 1. 1257. 1903.

Stoloniferous-rhizomatous perennials to 1.3 m. tall, peren-
nating by one or more slender, stoloniferous offshoots, these
later thickening and becoming rhizomatous. Rosette leaves

44

1975]

PLATE 1.
1.

surface.

Rudbeckia — Kral

FLATE |

он =.

d
ak
S

SM

(сё

[ С

Pales, akenes and corollas of Rudbeckia.

Rudbeckia auriculata. la. akene; 1b. disc corolla; 1c. pale, outer
Drawn from Kral 48579.

Rudbeckia fulgida var. fulgida.

2a. pale, outer surface; 2b.
akene and disc corolla.

Drawn from Kral 33107.

Rudbeckia fulgida var. umbrosa.

3a. akene and dise corolla;

3b. pale, outer suface. Drawn from Kral 32945.

46 Ећодога [Vol. 77

with blades ovate to lanceolate or elliptic, altogether 3-11
(-25) cm. long, on petioles shorter than to as long as the
blades, the margins ranging from entire to undulate, cre-
nate, dentate, or serrate, the bases cuneate to attenuate,
the surfaces glabrous to (usually) appressed-strigose.
Stems erect, proximally hirsute or strigose or smooth, dis-
tally strigose, strigillose or smooth, and terete or sulcate-
costate. Heads few to many, long pedunculate usually from
the axils of upper stem leaves and forming an open “can-
delabra-like" cyme. Lower stem leaves similar in outline
and indumentum to rosette leaves, grading upward to
shorter, narrower, sessile or even clasping. Phyllaries ob-
long, slightly to much shorter than the rays, green, ciliolate-
scabrid, reflexed. Rays short-oblong to elliptic, 0.9-1.7 cm.
long, spreading or slightly recurved, often strigillose on the
backs, orange-yellow (frequently drying greenish). Disc
1.0-1.8 em. broad, the pales oblanceolate or cuneate, rarely
elliptical, 4-5 mm. long, short-acuminate to acute or obtuse,
the margins entire to sparsely ciliolate, the outer surfaces
distally a deep reddish-brown becoming proximally tan
with the 3 nerves, particularly the 2 submarginal laterals
a deep lustrous reddish-brown, smooth or sparsely strigil-
lose especially toward the tips. Disc corollas 3.0-3.5 mm.
long, the tube shorter than the narrowly campanulate or
broadly tubular throat, the lobes short-triangular and erect,
the whole corolla deep purple-brown, Akenes asymmetri-
cally short-oblong, 2.0-2.5 mm. long, curvate proximally,
usually 4-angled, dark gray, the pappus a low, slightly un-
even crown less than 0.5 mm. long or almost obsolete.

This entity inhabits prairies, swales, open woods, fields,
and roadsides, on a variety of soils and in dry to quite wet
situations from New Jersey to Illinois, south to Florida and
Louisiana. It is definitely the most widespread, certainly

PLATE 2. Habit sketches of Rudbeckia.
1. Rudbeckia auriculata.

Rudbeckia fulgida var. umbrosa.

Rudbeckia fulgida var. fulgida.

bo

МУ

FLATE 2

48 Ећодога [Vol. 77

the most variable taxon of the complex. It and the other
varieties of R. fulgida all display a fine orange-yellow ligule
color that distinguishes them well from other Rudbeckias
in the late summer and early autumn fields.

R. fulgia Ait. var. umbrosa (Boynt. & Beadle) Cronq.,
Rhodora 47: 400. 1945. Plate 1(3); Plate 2(2).
R. umbrosa Boynt. & Beadle, Biltmore Bot. Stud. 1: 16.
1901.
R. chapmanii Boynt. & Beadle, Biltmore Bot. Stud. 1: 14.
1901.

Differs from var. fulgida in being stouter, the stem often
angular, usually smoother, taller (frequently 1 m. tall),
with stouter rhizomes. Rosette and lower stem leaves pro-
portionately large, long-petiolate with ovate or lance-ovate,
cordate-based blades, these either glabrous or strigose or
scabrid, and with entire to crenate-dentate margins. Heads
larger, the rays longer, usually elliptic-oblong, and often
fully 3 em. long, the disc often to 2 ст. broad. Pales соп-
sistently ciliate distally save for most inner ones. Akenes
similar in shape but usually longer, to 3 mm.

This variety is perhaps the showiest of the Alabama-
Tennessee fulgidas, in my opinion an excellent horticultural
subject because of its late and long-blooming habit and its
full inflorescence. Its best habitats are sunny, moist to
quite wet swales and seeps from Virginia west to Ohio and
south to Alabama and Georgia.

R. fulgida Ait. var. spathulata (Michx.) Perdue, Rhodora
59: 297-298. 1957.
R. spathulata Michx. Fl. Вог. Ат. 2: 144. 1803.

Most similar to var. fulgida, with rosette and basal stem
leaves attenuated to the petiole, not cordate. Stems ranging
from glabrous to variously hirsute or strigose; mid- and
upper stem leaves ranging from broadly spatulate to ob-
lanceolate or pandurate, and from glabrous to strigose.
Heads as small as in var. fulgida with rays short (rarely
as long as 2 ст.).

1975] Rudbeckia — Kral 49

This variety grows in low, moist to wet, sunny places or
in low open woods from Virginia west to Tennessee and
south to Florida and Alabama. In habit, rosette and stem
leaves, inflorescence, and head it is so similar to extremes
of R. fulgida var. fulgida as to be indistinguishable, and its
status as a variety to me appears hardly tenable.

R. auriculata (Perdue) Kral, stat. nov. Plate 1(1); Plate
Zu
R. fulgida Ait. var. auriculata Perdue, Rhodora 63: 119-
120. 1961.

Robust, rhizomatous, glabrous or subglabrous perennial
to 3 m. tall, perennating by short, stoloniferous offshoots,
these becoming stout (to 1 em. or more thick) by the sec-
ond season. Rosette and lower stem leaves to 65 cm. long,
the blades oblong to oblanceolate or ovate-lanceolate, short-
acuminate to rounded or acute, entire to sinuate, crenate,
dentate, coarsely serrate or denticulate, on petioles from
1/3-1/2 the total leaf length, the surfaces smooth to scab-
rous and usually strigose on the veins, midrib and petiole.
Stems erect, stiffish, proximally fully 1 ст. thick, subterete,
multicostate to sulcate. Middle and upper cauline leaves
sessile, the blades progressively reduced up the stem, ellipti-
cal to ovate or pandurate, acute, the margins entire to
dentate, denticulate or coarsely serrate, the bases auricu-
late-clasping, the surfaces from smooth to sparsely scabrid
or strumose. Heads many in an open, broadly convex,
paniculate cyme. Phyllaries oblong-linear, spreading,
shorter than rays, smooth save for scaberulous margins.
Pales oblong-cuneate, ca. 6 mm. long, acute, ciliate, the
backs hirsutulous, proximally tan with maroon mid- and
sub-marginal nerves, distally a deep reddish-brown. Rays
bright yellow, oblong-linear, ca. 2 cm. long, spreading or
spreading-ascending, the backs frequently strigillose. Disc
1.0-1.7 ст. broad, the corollas purple-brown, 3.0-3.5 mm.
long, the short tube gradually expanding into the narrowly
funnelform throat, the lobes short-triangular, erect. Akenes
oblong-curvate, 4.0-4.5 mm. long, a rich, lustrous reddish-

50 Ећодога [Vol. 77

brown, 4 ribbed, the pappus prominently unevenly 4-6-
toothed, the teeth pale tan, chartaceous, narrowly triangu-
lar, the longer ones fully 2 mm. long.

TYPE: Alabama: COVINGTON CO.: moist soil along Ala-
bama Highway 55, 11 mi. south of McKenzie (2 mi. n. of
Red Level), 24 July 1958, R. E. Perdue 2177. Holotype at
GH, Isotype at US.

Rudbeckia auriculata inhabits bogs, swamps, seeps,
ditches and swales, mostly in full sun, sometimes in partial
shade at edges of swamp woodlands. So far, most of the
samples are from the lower Coastal Plain in Alabama, but
I have found one outlier in the Appalachians, northeast of
Birmingham in St. Clair County in the broad limestone
valley between Oak Mountain and Blount Mountain. Oddly
enough, one of its neighbors there is Leavenworthia exigua
var. lutea, which appears in patches in spring on low
outcrops of calcareous rock. Because Alabama is even now
rather poorly studied for its flora, other populations con-
necting the montane ones with the Coastal Plain ones will
probably be discovered. The plants are so tall (some reach-
ing 3 m. in height!) as to be conspicuous even at a distance,
the only other Rudbeckia of comparable height in the area
being R. laciniata.

Rudbeckia auriculata, to summarize, differs from R. ful-
gida in the following ways:

1. The plants are usually at least twice as tall, arising
from much stouter rhizomes, and are prevalently smooth-
stemmed.

2. The foliage is, overall, larger, with cauline leaves
prevalently panduriform, auriculate-clasping; this char-
acter is stressed by Perdue (1.с.) in his varietal distinction.
The phyllaries are hairy on the backs with spreading or
ascending hairs.

3. The rays are a bright yellow, not an orange-yellow as
in the fulgidas of the southeastern U.S. It is to be suspected
that their pigment chemistry is quite different.

1975] Rudbeckia — Kral 51

4. The disc corollas are shorter than the akenes. In Rud-
beckia fulgida the reverse is true, i.e., the akenes are
shorter than the disc corollas.

5. The akenes are narrower in outline and longer than
are those of Rudbeckia fulgida, and are a lustrous red-
brown rather than gray-tinted.

6. The pappus is prominent, produced upward into nar-
rowly triangular scales, these longer than is true for any
other southeastern Rudbeckia. In R. fulgida the pappus is
usually a low, uneven crown or is nearly absent.

Thus, when one considers the many striking character
differences, this is not only a species distinct from Rud-
beckia fulgida, but indeed might well comprise a part of
another section of the genus. Only a revisional attempt
would disclose this fact, but the taxa nearest to R. auricu-
lata in pappus character and in some respects of head char-
acteristics are, in southeastern U.S., R. nitida and R. тоћтп.

A cytological comparison of some southeastern Rud-
beckias is to be initiated during the summer of 1974, for
there is sufficient phenotypic difference to assume a con-
siderable chromosomal difference, at least from R. fulgida
and its varieties.

The following are some recently collected localities for
populations of Rudbeckia auriculata:

Alabama: BARBOUR CO.: swampy ditch beside low oak forest, Lugo,
12 Sept. 1968, R. Kral with K. E. Blum 33300. COVINGTON CO.: tall
plants of peaty swale by Ala. 55, just n. of Red Level, 30 Апг. 1970,
Kral 40841; sandy silt of creek bottoms by Ala. 54, 7.8 mi. п.е.
Flerala, plants to 8' tall, 24 Jul. 1968, Kral 81970. CRENSHAW CO.:
n. side Luvergne, sandy creek bottom by U.S. 231, plants to 3 m.,
16 Aug. 1968, Kral 32421. GENEVA CO.: sandy peat of longleaf pine-
land ditch due e. of Samson, just outside limits by Ala. 52, 3 Sept.
1969, Kral 36837; 5 mi. s. of Samson along Ala. 187, moist ditch,
grown up waste area, roadsides, 18 Aug. 1970, Н. D. Moore 640.
PIKE CO.: sandy shores of Pike Co. public lake, s. of Troy, 11 Sept.
1968, Kral 33174. ST. CLAIR on: ca. 4 mi. n. Leeds, abundant clone
in wet cleared avea of pasture, 27 Sept. 1972, Kral 48579.

52 Ећодога [Vol. 77

Field work toward this paper was supported largely by a
research grant to the author from the National Science
Foundation (NSF GB-6688X).

REFERENCES

PERDUE, К. W., ЈЕ. 1957. Synopsis of Rudbeckia subgenus Rud-
beckia. Rhodora 59: 293-299.
1961. A new variety of Rudbeckia fulgida. Rhodora
63: 119-120.
SMALL, J. K. 1903. Flora of the Southeastern United States. 1394
pp. New York, published by the author.
1933. Manual of the Southeastern Flora. 1554 pp. Re-
print, Univ. of North Carolina Press, Chapel Hill.

BIOLOGY DEPARTMENT
VANDERBILT UNIVERSITY
NASHVILLE, TENNESSEE 37235

APPARENT ECOTYPIC DIFFERENCES
IN THE WATER RELATIONS OF
SOME NORTHERN BOG ERICACEAE?

PETER J. MARCHAND?

The apparent paradox of xeromorphy in northern bog
plants has become a popular subject again, this time how-
ever, with a new interpretation. The development of
sclerophyllous leaf tissue (increased cutinization and lig-
nification, increased fiber/protein ratio) in plants of both
wet and dry habitats has recently been linked with phos-
phorous deficient soils (Beadle, 1966; Loveless, 1962), a
feature that both deserts and northern bogs have in com-
mon. In the light of this discovery, leaf sclerophylly and
the evergreen habit often associated with it are now being
interpreted as reflecting a specialized metabolism tolerant
of low nutrient levels, though the physiology of this adap-
tation is not yet fully understood (Small, 1972a, 1972b;
Beadle, 1968, 1966; Monk, 1966; Loveless, 1962).

From a morphological viewpoint much of the northern
bog flora seems particularly well adapted for the conserva-
tion of water. Narrow, leathery, and heavily cutinized
leaves with revolute margins and vertical orientation, char-
acters typical of many Ericaceae, are perfect water con-
serving adaptations. Yet these features cannot be explained
on the basis of plant-water relations in the northern sphag-
num bog. Early hypotheses relating these xeromorphic
characters to a condition of “physiological drought,” for
reasons mostly associated with the anaerobic bog substrate,
were dismissed long ago by Caughey (1945) for lack of ex-
perimental confirmation. More recent in sitw determinations
of the water status of bog plants by Small (1972a) and

1Contribution No. 2, The Center for Northern Studies.
2Present address: The Center for Northern Studies, Wolcott, Ver-
mont 05680.

58

54 Ећодога [Vol. 77

Marchand (1972) have likewise failed to turn up any evi-
dence of water stress in the bog environment. While normal
midday xylem water potentials in several species have been
found to be quite low (—15 to —20 bars), stomatal response
appears to be unaffected in this range (Small, 1972a; Mar-
chand, 1972). Assuming that stomatal functioning is an
adequate measure of plant sensitivity to its environment,
there seems enough justification indeed to abandon our
ideas of water stress in bogs.

This is not to suggest, however, that the subject of water
relations of bog plants is closed. To the contrary, the acid
and waterlogged substrate of the sphagnum bog poses some
unique ecological problems, and the data of Small (1972a),
as well as that of Marchand (1972), invite further investi-
gation into possible differences in the water metabolism of
bog plants and their counterparts on well drained sites.
Specifically, the following questions may be asked: Why
are xylem potentials of small bog shrubs so low, sometimes
exceeding —20 bars, when soil water supply in this habitat
is seemingly unlimited? In an environment where soil-
water supply is more restricted, how low a water potential
might these species tolerate, before stomatal closure is
induced? Do the observed water potentials represent an
optimum “operational” level for the species in question or
is the level induced by some edaphic characteristic of the
habitat? Presented here are some field observations which
suggest possible answers to these questions,

METHODS

The Lee Hill Bog in Lee, New Hampshire, is typical of
northern sphagnum peat bogs, supporting a flora dominated
by several members of the family Ericaceae along with
Picea mariana (Mill.) BSP. and Eriophorum virginicum L.
Because the bog is situated in a kettle amidst glacial de-
posits, however, the surrounding upland area contrasts
abruptly in both edaphic character and floristic composition.
Around the perimeter of the bog, the well-drained till sup-

1975] Bog Ericaceae — Marchand 55

ports a “mesic” pine-oak community. Both the bog and
adjacent upland sites, however, support well established
populations of Vaccinium corymbosum L. and Gaylussacia
baccata (Wang.) K. Koch, thus affording an opportunity
to study simultaneously the water relations of these species
under similar atmospheric conditions but under markedly
different edaphic conditions. In Wolcott, Vermont, a some-
what similar situation exists in which the water status of

Ledum groenlandicum Oeder and Kalmia angustifolia L.

growing in a wet peat substrate was compared with that
of the same species growing in a well-drained upland field.

At each site, xylem potentials were measured using a
portable pressure bomb (PMS Instruments Co.) as de-
scribed by Waring and Cleary (1967). The pressure bomb
is designed to measure the amount of tension in the xylem
water column resulting from transpirational “pull” against
flow resistances in the soil-plant continuum. This xylem
tension is related to the free energy of the water in the
plant, the water potential. When the stem of a transpiring
plant is cut, the cohesion of the water column is broken
and the tension is relaxed. The water column then quickly
recedes from the cut surface. By placing the branchlet into
a sealed chamber with only the cut end protruding, and
applying pressure, the water column can be forced back to
the cut surface, thus re-establishing the initial condition.
The amount of pressure required to do this is equal in
absolute value to the initial xylem tension or water potential
of the plant.

In this study, exposed branchlets about 10 cm. in length
and bearing several leaves were used for the water poten-
tial determinations. The sample number ranged from just
three stems per species when variation was low, as during
the early morning hours, to six when the variation within
species was higher. At the time of sampling, soil-water
potentials were determined using a tensiometer (Soilmois-
ture Equipment Corp. No. 2900) and wet- and dry-bulb
temperatures were recorded with a sling psychrometer.

56 Ећодога [Vol. 77

Since pressure bomb measurements by themselves are
difficult to interpret, stomatal response to the observed
water potentials was measured with a diffusion porometer.
This instrument is essentially an electric hygrometer whose
resistance varies inversely with humidity. In use, a small
plexiglass chamber containing a hygroscopic element is
clamped onto a leaf and, in effect, the leaf is allowed to
transpire under prescribed conditions. As water vapor
diffuses into the initially dry chamber, the time rate of
humidity increase over a narrow range is noted. From this,
the diffusion resistance of the leaf, mostly a function of
stomatal aperture, is calculated based on a calibration
curve for known resistances.

The porometer used in this study was modified from the
design of Kanemasu et al. (1969). In order to increase
instrument sensitivity for use on very small leaves the
chamber volume was reduced slightly, a higher sensitivity
element (Hygrodynamics No. 4-4816K) was used, and the
cup aperture was reduced to 6.35 mm. diameter. With
these modifications, the time required for attachment and
measurement on the leaf was usually of the order of 5 to 15
seconds when the stomates were fully open. In calibration,
storage, and field use, the recommendations of Morrow and
Slatyer (1971a, 1971b) were followed closely.

RESULTS

The data from the different sites are compared by using
the model of Elfving et al. (1972) where transpiration is
described by the ratio of vapor pressure deficit (VPD) to
stomatal diffusion resistance (rs). A plot of xylem water
potential against VPD/rs for the species studied here indi-
cates that for any given transpiration rate, under non-stress
conditions, xylem potentials are lower (more negative) in
the bog plants than in their upland counterparts (Fig. 1).

The rapid development of a very steep water potential
gradient in the bog plants was observed early in the day. At
the Lee Hill Bog, where soil water potential was zero bars,

1975]
=
kel
oo
=
e
o
а
N
од
=
eo
E
E
с
&.|,^
>
=<
e
Figure 1.

Bog Ericaceae — Marchand

57

S |
ur
Я pe
d |o
| 2 |
| E
| EL
| JE
E | p
| $247)
E Ki a
4 О |
| О
J // N
- d E
——rá rr

(s1eq) ч.

Xylem water potential (ух) as a function of estimated

transpiration rate (VPD/rs) for bog and upland ecotypes under non-
stress conditions. Symbols: О L. groenlandicum; O К. angustifolia;
ш (7. baccata; ху corymbosum.

58 Ећодога [Vol. 77

transpiration from Gaylussacia baccata was inhibited dur-
ing the first hours after sunrise by a heavy dew formation
on the leaves. As soon as the dew evaporated and trans-
piration began, a very steep gradient was established with
xylem water potential decreasing from —1 to —15 bars
within one hour and further decreasing to —22 bars in the
next four hours (Fig. 2). The leaf stomates remained open
(rs = 1.1 sec./em.) throughout the day however. In the
drier upland habitat where soil water potential was — 0.8
bars, transpiration from С. baccata began at sunrise with
xylem water potentials decreasing gradually, eventually to
be checked by an increase in stomatal resistance to an
average of 4.8 sec./em. by mid-morning and 8.3 sec./cm.
by mid-afternoon (Fig. 2).

While a soil-water potential of —0.3 bars is not usually
considered very dry, the difference in soil-water availability
between the above two sites is best indicated by the slow
rate of recovery or increase of xylem potential in the up-
land population as compared to that of the bog population,
beginning with the approach of sunset and continuing to
daybreak (Fig. 2). In spite of such differences in soil-
water availability, stomatal behavior, and transpiration
rate (an increase in rs from 1.1 sec./cm. to 8.3 sec./em. is
enough to cause a five-fold decrease in the transpiration
rate), the mid-day depression of water potential in both
populations was similar: —19.4 + 2.8 bars in the upland
population апа —22.0 = 2.7 bars in the bog population
(Fig. 2).

DISCUSSION

When the rate of movement of water through the plant
falls behind the rate of transpiration loss at the leaf sur-
faces, due either to internal resistances to flow or an in-
sufficient soil-water supply, the result is a decrease in plant
water potential. If there were no internal resistances in
the plant, then in any situation where soil-water is non-
limiting, the plant water potential would always be very
high (near zero). Conversely, where soil-water is freely

1975] Вог Ericaceae — Marchand 59

Time (hours)
8

\

Oi eee
© 10-0. 0 © 1D
oe a У

(SUE) "t

Figure 2. Diurnal trend of xylem water potential (vx) in Gay-
lussacia baccata, Lee Hill Bog and adjacent south-facing hillside.
Stomatal diffusion resistance (rg) is given in sec./em. Vertical bars
represent standard deviations.

60 Rhodora [Vol. 77

available and xylem water potentials become appreciably
low (more negative), as in the bog community, then in-
ternal resistances are indicated.

For steady-state flow through the plant, the transpiration
flux is considered to be directly proportional to the water
potential gradient and inversely proportional to internal
flow resistances. Therefore, relating the measured xylem
water potentials to the transpiration rate as estimated by
the ratio VPD/rs (for non-stress conditions) indicates that
flow resistances within the bog plants may be higher than
in the upland ecotypes. The rapid decrease of xylem poten-
tial with the commencement of transpiration in the bog
plants gives some measure of the magnitude of the driving
force (water potential gradient) needed to move water
through the plant in the presence of high internal resist-
ance.

Assuming that there is no resistance to water movement
in the saturated peat soil, the resistances indicated for these
bog plants may be considered to lie between the root-soil
interface and the stem xylem tissue just beneath the leaf
layer (the point at which the xylem water potential was
measured). It is believed that poor soil aeration, a condition
typical of bogs (see, for example, Armstrong and Boatman,
1967), inhibits root absorption, but is this alone enough to
account for the resistance indicated here? Studies of Mack-
lon and Weatherley (1965) suggest that root resistance
plays little part in establishing leaf water potentials lower
than —5 bars.

Flow resistances in the xylem, on the other hand, are
generally considered to be the lowest in the soil-plant-
atmosphere continuum. Increased resistance in the xylem
may possibly result from decreased diameters of conducting
elements or through the blocking of these by sclerids or
tyloses, creating in effect a more tortuous translocation
pathway in bog plants. Preliminary microscopic examina-
tion of the xylem tissue, however, showed vessel elements
to be on the order of 15 to 25 microns in diameter, essen-
tially the same as in the upland ecotypes and not small

1975] Bog Ericaceae — Marchand 61

enough to increase flow resistance, and revealed no obstruc-
tions in the vessels (Terrell Comstock, unpublished data).
The sovrce of the high internal flow resistances in bog
plants, if indeed they exist, thus escapes recognition for
the moment.

The values for VPD in this analysis were taken from
the wet-bulb depression only (i.e, atmospheric VPD)
which presupposes that the leaf and air were in thermal
equilibrium. If leaf temperatures were elevated above air
temperature, then the caleulated values for VPD/rs would
be low. Presumably however, with all plants fully exposed,
the direction of the error would be the same in all cases.
Moreover, when it is considered that the water potential of
the upland plants must also reflect flow resistances outside
the plant, since it can be assumed that the soil adjacent to
their roots is not saturated, then the relative differences in
water potential levels seen here take on even greater sig-
nificance with respect to indicated internal differences.

Of considerable interest in these data is the fact that the
midday xylem potentials in the two Gaylussacia populations
remained relatively close. This suggests the possibility of
a closely regulated optimum water potential levei for the
species, indicating also very sensitive stomatal control, with
the stomates of the upland plants closing before any de-
tectable decrease in xylem water potential. This same
phenomenon has been observed in yellow birch (C. A.
Federer, personal communication) and is somewhat sug-
gestive of the kind of direct response of stomates to evapo-
rative conditions of the atmosphere, independent of leaf
water potentials but conditioned by soil water supply, as
reported by Schulze et al. (1972).

As for the question of xeromorphy in bog plants, leaf
sclerophylly appears of advantage only insofar as it may
be related to the evergreen habit, possibly preventing win-
ter desiccation when water uptake is impaired. There is not
enough evidence yet, however, to conclude that the water
metabolism of bog plants is entirely similar to that of their
upland counterparts. While Small (1972a) demonstrated

62 Вћодога [Vol. 77

some similarity in the water status of woody plants in a
bog and nearby mesic habitat, his mesic site was described
as including a marsh and field adjacent to the bog and
many of the species studied at that site, such as Ilex verti-
cillata (L.) Gray, Nemopanthus mucronata (L.) Trel., and
Alnus rugosa (DuRoi) Spreng. are typically confined to
wetland areas. Lacking any data to the contrary, this sug-
gests that the two environments may not have differed
significantly in terms of soil-water availability. The data
presented here indicate that dissimilarities in water metab-
olism between bog and upland ecotypes may indeed exist in
relation to soil-water availability or other edaphic differ-
ences,

ACKNOWLEDGEMENTS

This study was supported in part by funds provided by
the Office of Water Resources Research, U.S. Department
of the Interior, by a Grant-in-Aid of Research from the
Society of the Sigma Xi, and by a grant from the Center
for Northern Studies, Wolcott, Vermont.

LITERATURE CITED

ARMSTRONG, W., & D. J. BoATMAN. 1967. Some field observations
relating the growth of bog plants to conditions of soil aeration.
Jour. Ecol. 55: 101-110.

BEADLE, N. C. W. 1968. Some aspects of the ecology and physiology
of Australian xeromorphie plants. Aust. Jour. Sci. 30: 348-355.

1966. Soil phosphate and its role in molding segments
of the Australian flora and vegetation, with special reference to
xeromorphy and sclerophylly. Ecology 47: 992-1007.

CAUGHEY, M. С. 1945. Water relations of pocosin or bog shrubs.
Pl. Physiol. 20: 671-689.

ELFVING, D. C, M. В. KAUFMANN, & A. E. HALL. 1972. Inter-
preting leaf water potential measurements with a model of the
soil-plant-atmosphere continuum. Physiol. Plant. 27: 161-168.

KANEMASU, Е., 6. THURTELL, & C. B. TANNER. 1969. Design, cali-
bration and field use of a stomatal diffusion porometer. РІ.
Physiol. 44: 881-885.

LovELESS, A. R. 1962. Further evidence to support a nutritional
interpretation of sclerophylly. Ann. Bot. 26: 551-561.

1975] Bog Ericaceae — Marchand 63

MACELON, A., & P. E. WEATHERLEY. 1965. Controlled environment
studies of the nature and origins of water deficits in plants.
New Phytol. 64: 414.

MARCHAND, P. J. 1972. Stomatal resistance and transpiration po-
tential of bog plants. Unpublished Master of Science Thesis,
Univ. N.H.

MoNK, C. D. 1966. An ecological significance of evergreenness.
Ecology 47: 504-505.

Morrow, P. A., & R. О. SLATYER. 1971a. Leaf temperature effects
on measurements of diffusive resistance to water vapor transfer.
Pl. Physiol. 47: 559-561.

& ——————— 19715. Leaf resistance measurements
with diffusion porometers: Precautions in calibration and use.
Agr. Meteorol. 8: 223-233.

SCHULZE, E. D., О. L. LANGE, U. BuscHBOM, L. KAPPEN, & M.
EvENARI. 1972. Stomatal responses to changes in humidity in
plants growing in the desert. Planta 108: 259-270.

SMALL, E. 1972a. Water relations of plants in raised sphagnum
peat bogs. Ecology 53: 726-728.

1972b. Ecclogical significance of four critical ele-
ments in plants of raised sphagnum peat bogs. Ecology 53: 498-
503.

WARING, В. H., & B. D. CLEARY. 1967. Plant moisture stress:
evaluation by pressure bomb. Science 155: 1248-1254.

DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM, NEW HAMPSHIRE 03824

THE DECIDUOUS MAGNOLIAS
OF WEST FLORIDA

RONALD F. MILLER

The deciduous magnolias are perhaps the most distinctive
relicts of the old Arcto-Tertiary forest. Pleistocene glacia-
tion apparently eliminated these once-widespread plants
everywhere from the temperate zone except in the eastern
United States and in an arc stretching from Sikkim and
Nepal to the Japanese Islands. Abundant rainfall, moderate
temperatures, and rich mixed hardwoods seem essential for
their greatest development in such regions as the southern
Appalachians and the Cumberlands in this country. There
Magnolia acuminata and M. fraseri contribute a minor
though noteworthy element to the mixed mesophytic forests,
growing alongside tuliptrees, hemlocks, lindens, buckeyes,
and other characteristic species. In the lower valleys,
especially along watercourses, M. tripetala and (in the
Cumberlands) M. macrophylla display their huge leaves.
Even the casual observer soon learns to associate the
deciduous magnolias with our mountain forests in their
cool, well-watered lushness. It may come then as a surprise
to discover that each of these four eastern deciduous mag-
nolias, either in itself or in a closely related species, can
be found amid the predominantly piney woodlands of the
Florida Panhandle.

This is not to suggest that these species can be found on
the Coastal Plain only in West Florida, but that their re-
markable collocation and the mode of their survival there
in a presumably alien environment are of special taxonomic
and ecological interest. This region differs from the rest of
the lower Coastal Plain stretching from Virginia to eastern
Texas mainly in its deeply incised terrain. The pine-covered
uplands and the evergreen bays characterizing this zone
are interrupted here and there by bluffs, steep-sided water-
courses, and sharply rolling country, all of which provide

64

1975] Magnolias — Miller 65

a foothold for a strongly deciduous and often surprisingly
Appalachian plant аззетђју: On these slopes, especially
where the soil is rich and water-retentive, more northern
genera such as Fagus, Halesia, Oxydendrum, and Tilia mix
with such subtropical species as Magnolia grandiflora and
Quercus laurifolia;? some of the riverbluffs supported
American chestnut until quite recently (Elias, 1971). The
understory will often prove to be a thick tangle of the
Appalachian Kalmia latifolia and the coastal bay-plant
Illicium floridanum. In the calcareous regions, where the
deciduous component is most prevalent, herbaceous plants
such as Trillium and Sanguinaria put in an appearance.
Throughout West Florida the indicator plant for such mesic
sites is the silky camellia, Stewartia malacodendron.'
These anomalous patches of forest suggest a time, un-
doubtedly during a glacial epoch, when a deciduous forest
covered much more of the area. Phytogeographers have
long noted the more northerly elements in the ravine flora

'Зее Thorne (1949) for a discussion of Appalachian species in the
ravines of Southwest Georgia. E. L. Braun (1950) mentions the
presence of a more deciduous element in the incised Tunica Hills
section of eastern Louisiana,

“There seems to be some confusion about the proper nomenclature
for the common “laurel сак” of the coastal zone. Kurz and Godfrey
(1962) argue that this plant should be called 0. hemisphaerica
Bartram. In this paper I have chosen to follow the conventional
ascription of Preston (1961).

*Old-timers in Okaloosa and northern Escambia Counties recall the
chestnut tree quite well. Unfortunately, the crowns do not seem to
sprout as well as they do further north, and the frequency of chink-
apins in the area keeps the blight pathogen in plentiful supply.

4Walter’s pine (Pinus glabra) is also always present in these
deciduous forests, but it is not a good indicator since it is also an
element of what may be called the hammock association, a mixture
dominated by this pine, Magnolia grandiflora, and the evergreen
oaks Quereus virginiana and Quercus laurifolia. This climax associa-
tion will eventually take over dry pine flats and ridges if the leaf
litter is allowed to build up and if the woods are protected from fire
and lumbering. The hammock association intergrades with the de-
ciduous forests on more gradual slopes, especially where the soil is
quite sandy.

66 Ећодога [Vol. 77

of the famous Apalachicola River bluffs in Gadsden and
Liberty counties, with their endemic Torreya and Taxus
and such Appalachian species as Cornus alternifolia. A
Florida endemic such as Rhododendron minus var. chap-
manii also points to a time when the flora of the area was
more like that of the southern highlands of the present day
(see James, 1961). Apparently only steep and (often)
north-facing terrain offered sufficient protection from the
hot winds and intense insolation that favored the develop-
ment of evergreen oaks and pines instead of deciduous
woodlands. Only in these protected enclaves do the decidu-
ous magnolias maintain their precarious Floridian existence.
In a few years clear-cut silviculture and residential
sprawl may make fieldnotes on the adaptations of these
plants to an alien environment unobtainable, The follow-
ing observations on various species are offered for whatever
ecological or taxonomic insights they may afford.

MAGNOLIA ASHEI Weatherby

The status of this, the rarest of American magnolias, has
been open to dispute since its description by C. A. Weath-
erby in 1926. Weatherby differentiates it from Magnolia
macrophylla by rather comparative traits: a little less hair
on the undersides of the leaves, smaller flowers and leaves,
the smaller size of the plants. Only its rather narrow in-
fructescence, as opposed to the strikingly globular one of
M. macrophylla, offers any certainty to the identification
of herbarium specimens (Kurz & Godfrey, 1962). I am
personally very familiar with both plants, yet am not at
all sure I would be able to distinguish a vigorous leaf or
large flower of М. ashei from one of ordinary M. macro-
phylla. Even the variations in the purple blotches at the
center of the flowers — something rather difficult to observe
in herbarium specimens — seem to run through identical
ranges in the two plants.

The two plants are quite separable in the field, however.
Magnolia macrophylla is essentially a tree; it competes for
space in the canopy in much the same way a hickory or a

1975] Magnolias — Miller 67

sweetgum will do, though these taller-growing trees may
eventually overtop it. The young plants are therefore re-
markably upright, and blossoming occurs only on mature
shoots high above the ground. The collector who wishes
to examine a M. macrophylla flower must locate either an
isolated horticultural specimen or one released by clearing
or roadbuilding. Magnolia ashei on the other hand behaves
аз a large, coarse shrub that seldom climbs out of the
understory. It competes for light in the manner of the
deciduous azaleas, by leaning toward holes in the canopy
and by taking on a loose-jointed, often horizontal growth
form. On rare occasions an individual specimen will become
what might be termed a tree, but that plant will hardly
ever exhibit the clean upward sweep of the average M.
macrophylla. Though the plant’s crowns sprout vigorously,
individual shoots of M. ashei, from the evidence of all the
dead wood, seem to be quite short-lived, and the species
apparently is incapable in the wild of that sustained devel-
opment on a single shoot necessary for competition in the
upper layers of the forest. Horticultural specimens of M.
macrophylla grow continuously from a single bole if undis-
turbed; M. ashei specimens can hardly be kept from cop-
picing.

Magnolia ashei seems to have adapted to its existence as
a shrub by assuming a tolerance for shade and by blooming
in the understory. The plant is remarkably free-flowering
at small sizes; in the wild I have observed small shoots (ca.
1 meter) capped by a blossom while standing in the dense
shade of Quercus laurifolia. In cultivation, the plant be-
comes strikingly floriferous. One in my care has set blooms
on coppice shoots less than 10 ст. tall, and another has
borne five blossoms on a single branched shoot not one
meter tall or half a meter across. Such behavior in M.
macrophylla is, to the best of my knowledge, unheard-of.

The present-day ranges of these two plants preclude
genetic exchange. In Alabama I have never observed
Magnolia macrophylla south of the rolling limestone hills
stretching from northern Mobile County to Monroeville

68 Ећодога [Vol. 77

and then eastward. In Mississippi and Louisiana the plant
goes further south into areas underlain by more recent
Tertiary formations, but along the Alabama-Florida border
only a few north-south river valleys and bluffs interrupt the
sterile Citronelle (late Pliocene or early Pleistocene) and
later Pleistocene deposits blanketing the terrain. Neither
taxon seems to have invaded the few upland hardwood areas
in this region, possibly because both prefer rather more
calcareous soils. I have never seen M. ashei north of the
line traced by U. 5. Highway 90.

The center of distribution of Magnolia ashei seems to be
the Knox Hill region in eastern Walton County. In this
strongly rolling area the sterile overburden is replaced by
a dark, rich, water-retentive clay or loam derived from
distinctly calcareous marl. The plant is actually common
on a few square miles surrounding Knox Hill. To the east
it can be found on hills and along watercourses near Ver-
non, Washington County (geologically a rather similar
area); on limestone hills along Econfina Creek in northern
Bay County; on high slopes along the east bank of the
Ochlockonee River near Smith Creek, Wakulla County ; and
on the Apalachicola bluffs between Bristol and Chattahoo-
chee. The stations a few miles south of Chattahoochee bring
M. ashei closest to M. macrophylla: patent M. macrophylla
may be found in the ravines south of Ft. Gaines, Georgia,
some 50 miles to the north. As best I can discover, the
southern border of Clay County, Georgia, marks the south-
ernmost extension of M. macrophylla, so intergradation
seems highly unlikely.^ It may be of interest that two other
separable but closely related taxa, Rhododendron minus
and its Florida form, R. minus var. chapmanii, reach their

"The region contains rich collecting spots for Miocene shells of the
Alum Bluff series. See Cook and Mosson (1929). Harper (1914)
describes the soil and vegetation of this area.

"At least in Georgia. І am much less familiar with the southeastern
corner of Alabama, so intermediates could conceivably be found in
Henry or Houston counties. If they are there, however, they are
elusive.

1975] Magnolias — Miller 69

southernmost and northernmost stations within less than
10 miles of the comparable stations for M. macrophylla and
М. ashei.

To the west of the Knox Hill region, M. ashei inhabits
occasional very precipitous ravines and steepheads within
Eglin Air Force Reservation. The westernmost station lies
just off State Highway 87, about 20 miles east of Pensacola.
It was from this region that W. W. Ashe obtained the
series upon which Weatherby based his description of the
species. Oddly enough, the soil on these slopes is sandy
and apparently sterile, quite unlike the heavy soi! of sta-
tions to the east. The deciduous element of the attendant
vegetation is also attenuated, with Fagus notably absent,
though Ostrya, Oxydendrum, and Stewartia are almost
always present. Gordonia lasianthus 18 quite frequent
along these streams.’

Magnolia ashei is thus probably a relict that through iso-
lation has adapted to life as a shrub rather than a tree.
The plant inhabits mixed evergreen and deciduous forests,
and indeed the dense canopy offered by the evergreen mag-
nolia and the evergreen oaks may have precluded competi-
tion in the manner open to M. macrophylla in regions where
these evergreen hardwoods are not so common. Except in a
few square miles the plant is remarkably rare; a compar-
ably vulnerable, unusual, and attractive bird or mammal
would surely have been the occasion for dozens of pilgrim-
ages and popular articles.

‘The presence of Magnolia ashei in East Texas has been reported.
Several years ago I observed a big-leafed magnolia in the yard of a
gentleman living at * Devil's Pocket," a rather flat, swampy region
in southern Newton County. He said this transplanted specimen was
ali that remained of an older plant that once stood in his pasture
on a hammock. Тће plant has since died, so I have been unable
to re-examine the specimen in the light of my field experience with
Florida M. ashei. As I recall, the plant bore reduced leaves, but the
form was definitely upright in a manner I later learned to associate
only with Magnolia macrophylla. I saw no infructescences. Several
years of searching and conferring have failed to turn up а single

other plant in that county or in Texas. Magnolia macrophylla, it
might be noted, is fairly common in adjacent western Louisiana.

70 Rhodora [Vol. 77

MAGNOLIA ACUMINATA L.

Magnolia acuminata is a more decidedly Appalachian
plant than M. macrophylla, yet it occurs in West Florida,
perhaps in the form of M. acuminata var. subcordata. The
distinction between the typical variety and var. subcordata
seems more plausible to the reader of taxonomic keys than
to the observer in the field, since great variations in hairi-
ness and flower color may occur within a given locality
(see Hardin, 1954). In general, the West Florida plants
bear greenish-yellow flowers intermediate between the
showy canary-yellow blooms of some plants in piedmont
Georgia and the greenish forms prevalent in the higher
mountains. Although the hairiest leaves I have ever seen
on any cucumber-tree grew on a Florida specimen, closeby
trees bore leaves indistinguishable from those on central
Mississippi and northern Alabama plants. As with many
other species, hairiness seems to increase gradually though
irregularly southward, as water loss from the leaves be-
comes more and more of a threat to survival. I wonder if
the cucumber-trees would be split at all if it were not for
the historical accident that a singularly yellow form existed
as a horticultural curiosity for almost a century until the
piedmont Georgia plants were relocated by Berckmans in
1913 (Sargent, 1933).

In Florida Magnolia acuminata is to be found only in the
Knox Hill region mentioned in the previous discussion. It
is much less frequent there than M. ashei, however, growing
only in a narrow band marking the transition between
beech and hickory woods along a few ravines and slopes in
the richest areas. There it becomes a fairly large tree.
Kurz and Godfrey (1962) report that a search of the area
located only six trees. Though I have located several
hundred, distribution is sporadic and the trees are easily
missed without a meticulous search in rather difficult
country.

The Citronelle deposits mentioned earlier separate these
Florida plants from the closest Alabama cucumber-trees,

1975] Magnolias — Miller ТІ

а large population inhabiting {һе Sepulga River valley to
its mouth in upper Escambia County, Alabama. The mouth
of the Sepulga marks the southernmost outcropping of
Oligocene limestone in that part of the state, so calcareous
soil again seems to be the key to distribution. The lime-
stone region in the Panhandle near Marianna and Tallahas-
see apparently does not support cucumber-trees, but then
cucumber-trees are either exceedingly rare or entirely ab-
sent in immediately adjacent Alabama and southwestern
Georgia. Southwestern Alabama and southern Mississippi,
on the other hand, are replete with cucumber-trees. Possi-
ble reasons for this anomaly elude me.

Genetic isolation seems to have resulted in no genetic
drift: trees from western Florida and southern Alabama
occupy virtually identical sites and, from every appearance,
are indistinguishable. Magnolia acuminata seems to be an
intermediate in the plant succession — like sweetgum, say
— whereas M. ashei is a more stable component of what
seems to be a climax association. Its extremely rapid
growth, handsome foliage, and tolerance for drought would
make the coastal strain a fine shade tree for use in the
lower South.

MAGNOLIA PYRAMIDATA Bartr.

This is the most common deciduous magnolia in West
Florida. Magnolia pyramidata can be found in beech woods
and on deciduous slopes throughout the area, both in the
rich Knox Hill region and on the less fertile streambanks
and bluffs along the Alabama line. Very steep north-facing
slopes are its most typical habitat, where it can often be
found amid mountain laurel thickets.

Magnolia pyramidata is the coastal equivalent of M. fra-
seri of the mountains. Whether these are separable either
as species or as varieties is a moot question. Certainly if
typical M. acuminata and M. acuminata var. subcordata
are to be separated, these two plants should also be. On
occasion, the mountain plant becomes a fair-sized tree with

72 Ећодога [Vol. 77

a thick, low-branched bole; the Florida plant is much
smaller, with an ascending form and (usually) a single
trunk. I do not remember ever seeing a bole bigger than
10 em. in diameter in West Florida. The leaves of the
mountain plants tend to be much larger, also, though oc-
casionally vigorous Florida specimens will belie the pub-
lished keys. In East Texas (Jasper and Newton counties)
putative M. pyramidata at times resembles М. fraseri in
the size of leaves and the form and size of the trees, and is
found, oddly enough, only on the top of a few sandy ridges;
no one familiar with M. pyramidata in Florida could guess
where to seek the Texas colonies. There is a gradual transi-
tion between M. pyramidata of Florida and M. fraseri of
the mountains, since the Alabama and Georgia plants form
a continuous sequence from north to south in these states.

In Florida this plant is occasional and hardly ever pro-
duces what might be termed colonies. The plant is by no
means uncommon, however, and a belief that it is reflects
not the state of nature but our limited knowledge of the
complexities of coastal vegetation. Like Stewartia mala-
codendron, which is often termed rare in handbooks, M.
pyramidata seems quite frequent once its habitat is under-
stood and sought out.

MAGNOLIA TRIPETALA L.

This is apparently the first record of the occurrence of
this magnolia in Florida. So far I have found it only along
about half a mile of high north-facing bluff and in a con-
tiguous ravine system on the Shoal River, approximately
two miles west of Dorcas, Okaloosa County. This particular
section is steep, but no more so than several other bluffs
and ravines along the Shoal and nearby Yellow rivers. The
bluffs are composed of Miocene marl, and the segment bear-
ing M. tripetala does seem to be covered by an unusual
quantity of redbud (Cercis canadensis), which in Florida
is a good indicator of limy soil. Тће ravine system itself,
where perhaps 95% of the M. tripetala plants grow, is

1975] Magnolias — Miller 73

a delightfully unspoiled area, with two small springfed
streamlets running over large blocks of marl, uncut hard-
woods such as beech and white oak and linden, an interest-
ing herbaceous layer, and here and there the big leaves of
the magnolias. This surprising and isolated stand of this
species is so small it could easily be destroyed entirely by
clearing or even lumbering.

The closest extensive colony of Magnolia tripetala on the
Coastal Plain seems to be located on the north-facing bluffs
along Hog Creek, Randolph County, Georgia. There M.
tripetala is a vigorous competitor in the subcanopy, resem-
bling in form and habit the specimens of M. macrophylla
with which it grows; in competition with M. macrophylla
it seems to favor the very steepest bluffs and the deepest
and darkest ravines. The Florida plants, on the other hand,
grow in a better-lighted area and mix with an understory
of Stewartia and small dogwood and redbud trees — the
last of which is certainly not a remarkably tolerant species.
I have observed no stem more than 6 or 8 cm. in diameter,
nor is any plant more than 10 meters tall. The great ma-
jority are between 3 and 5 meters.

Though an analogy to the reduced stature of Magnolia
ashei suggests itself, it seems likely that reduced vigor
rather than genetic divergence accounts for the difference
between ordinary M. tripetala and the Florida plants. For
one thing, there are dead stems leading from a high per-
centage of the crowns, so perhaps the leaders cannot sur-
vive long enough to grow into a well-developed tree. The
form of the plant is upright, unlike the horizontal tendency
of the truly shrubby M. ashei. Most significantly, they
show none of the ability of M. ashei to bloom vigorously at
a small size, and that would surely be a necessary con-
comitant to a genuine adaptation to existence as a shrub.
In fact, no M. tripetala seedlings could be discovered in a
rather meticulous search; only sprouts upon older root-
stocks were discovered. The plants do show one singular
habit, however. Approximately half the clumps are attended
(at a distance of a meter or so) by one or more small satel-

74 Rhodora [Vol. 77

lite plants which can be traced by what appear to be hori-
zontal runners back to the centra] crown. I have never
observed this tendency to spread by vegetative means in
other magnolias, and a check of the Hog Creek colony has
revealed only a few sprouts which might be thought com-
parable. At the present time I am cultivating several of
the Florida offsets in hopes of comparing their behavior
with a seedling M. tripetala taken from a vigorous colony
found in coastal North Carolina near New Bern.

Other stations close to the Shoal River colony seem to
be (1) on the Pascagoula River in southern Mississippi
and (2) in Butler County, Alabama (W. H. Duncan, per-
sonal communication). Since I have, unfortunately, never
located either station, I cannot comment on the vigor or
habitat at other locations on the southern coastal plain.

This deciduous magnolia seems to be the ultimate exam-
ple of an Arcto-Tertiary relict which has persisted in an
island of mesophytic forest on a protected north-facing
slope amid the pinewoods of Florida. How long this colony
has been there or whether it is the remnant of a larger
colony would be impossible to say. For some time now it
may even have been regenerating itself almost entirely by
vegetative means. Its chances of surviving the chain saw
and the log sledder and the bulldozer seem easier to esti-
mate. It does seem unfortunate, though, that all that will
remain of such a dogged adaptation to an alien environment
will be a few dried sheets in a herbarium and an aberrant
dot on a тар,5

*Specimens of the plants discussed in this article have been de-
posited in the herbarium of the University of Georgia. In particular,
I would like to express my appreciation to its director, Professor
Wilbur H. Duncan, for his unfailing generosity with his expertise
and for his remarkable tolerance for amateur enthusiasm.

1975] Magnolias — Miller 75

LITERATURE CITED

BRAUN, E. L. 1950. Deciduous forests of eastern North America.
Hafner Publishing Co., New York. 596 pp.

Cook, C. W., & S. Mosson. 1929. The geology of Florida. In:
12th Ann. Rep. Florida State Geol. Survey, pp. 31-227.

Еллав, T. S. 1971. The genera of Fagaceae in the southeastern
United States. Jour. Arnold Arb. 52: 159-195.

HARDIN, J. №. 1954. An analysis of variation within Magnolia
acuminata L. Jour. Elisha Mitchell Sei. Soc. 70: 298-312.
НАЕРЕЕ, R. M. 1914. Geography and vegetation of northern Flori-

da. Im: 6th Ann. Rep. Florida State Geol. Survey, pp. 163-416.
JAMES, C. W. 1961. Endemism in Florida. Brittonia 13: 225-244,
Kurz, H., & R. К. GODFREY. 1962. Trees of northern Florida.

University of Florida Press, Gainesville. 311 pp.

Preston, R. J., Је. 1961. North American trees. Iowa State Uni-
versity Press, Ames. 395 pp.
SARGENT, C. S. 1933. Manual of the trees of North America. Ed. 2.

Houghton Mifflin Co., Boston & New York. 934 pp.

THORNE, R. F. 1949. Inland plants of the Gulf Coastal Plain of

Georgia. Castanea 14: 88-97.

WEATHERBY, C. А. 1926. A new Magnolia from West Florida.

Rhodora 28: 35-36.

DEPARTMENT OF ENGLISH
UNIVERSITY OF WEST FLORIDA
PENSACOLA, FLA. 32503

INVESTIGATIONS OF
NEW ENGLAND MARINE ALGAE VII:
SEASONAL OCCURRENCE
AND REPRODUCTION OF MARINE ALGAE
NEAR CAPE COD, MASSACHUSETTS"?

DOUGLAS C. COLEMAN AND ARTHUR C. MATHIESON

In an earlier paper (Coleman and Mathieson, 1974) we
described the horizontal distribution of seaweeds at seven
sites from Scituate to Woods Hole, Massachusetts, including
the Cape Cod Canal. In the following account we summar-
ize the seasonal occurrence and reproduction of seaweeds at
the same locations. Most previous collections and observa-
tions of Cape Cod seaweeds have been restricted to the
summer. Conover (1958) and Sears (1971) have con-
ducted the only detailed seasonal investigations of Cape
Cod marine algae. Conover described the productivity and
seasonal composition of the algae in the Great Pond Estuary
of Falmouth in relation to a variety of environmental pa-
rameters. Sears described the subtidal benthic algae at
several sites in southern Cape Cod. Neither Sears nor
Conover gave any consideration to the seaweeds in the
Cape Cod Canal, which connects Cape Cod Bay and Buz-
zards Bay. The Canal is a transitional zone separating two
distinctive water masses and marine floras (Coleman and
Mathieson, 1974). The temperature discontinuity between
the two sides of the Cape may reach 10°C. during the sum-
mer; thus, Cape Cod is one of the major phytogeographical
boundaries on the Atlantic Coast of North America (Far-
low, 1870, 1882; Harvey, 1852-1858; Humm, 1969, Setchell,
1922).

‘Published with the approval of the Director of the University of
New Hampshire Agriculture Experiment Station as Scientific Con-
tribution Number 665.

2Јаскѕоп Estuarine Laboratory Contribution No. 14.

76

1975] Саре Cod Algae — Coleman & Mathieson TT

Monthly collections and observations of marine algae
were made at each of the seven sites (Fig. 1) during 1969.
Specific details of collections, identifications, and descrip-
tions of stations have been previously summarized (Cole-
man and Mathieson, 1974), and they will not be repeated
in the present paper.

|| Li
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САРЕ СОР ВАҮ 2 Уј
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41°47 pn CAPE COD CANAL И
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Figure 1. Map of Cape Cod, Massachusetts, showing the seven
stations.

78 Rhodora [Vol. 77

SEASONAL OCCURRENCE AND LONGEVITY

Figures 2 and 3 summarize the monthly occurrence of
seaweeds at each station. There was a conspicuous increase
in the number of species during the spring and summer,
except at stations 2, 3, and 4. The time of peak numbers

STATION 1

80

60:

40:

20:
WY
ud
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MONTHS
Figure 2. Monthly variation in number of species at stations 1-3.

Black == red algae; hatched area == brown algae; white == green

algae.

1975] Cape Cod Algae — Coleman & Mathieson 79

STATION 4

NUMBER OF SPECIES

Е МАМЈ JAS ON Рр

Figure 3. Monthly variation in number of species at stations 4-7.
Black — red algae; hatched area == brown algae; white == green

algae.

80 | Ећодога [Vol. 77

varied at different stations. Seasonal changes were usually
most obvious from January to June, except at stations 3
and 4. A gradual decline in the number of species was
observed from June to December at each station. The ma-
jority of species at each site belonged to the Rhodophyta
and Phaeophyta; the lowest numbers were members of the
Chlorophyta. The red algae showed a greater increase in
numbers during the spring and early summer than did
either the browns or greens. Seasonal changes in numbers
were greatest at Scituate, Wings Neck and Woods Hole.
See Tables I-III for specific details of occurrence at stations
1, 4 and 7 — i.e., a northern, a southern, and a Canal station.

Of the 106 taxa collected at the seven stations, 49 were
designated as annuals and 57 as perennials (Table IV).
Perennials accounted for the largest number of species at
each station, except at Wings Neck and Woods Hole. The
ratio of annuals : perennials at each station was as follows:
station 1 — 42 : 58, station 2 — 44 : 56, station 3 — 40 : 60,
station 4 — 42 : 58, station 5 — 43 : 57, station 6 — 54 : 46,
station 7 — 54 : 46. The conspicuous annuals and peren-
nials on each side of the Cape are summarized in Tables V
and VI. The dominance of annuals and reduced numbers of
perennials to the south is apparent.

Three groups of perennials can be distinguished as
follows, depending upon their abundance north and south
of the Cape: 1) species either restricted to or more abun-
dant north of the Cape; 2) species either restricted to or
more abundant south of the Cape; 3) species common on
both sides of the Cape. The first group included Choreo-
colax polysiphoniae, Gigartina stellata, Petrocelis midden-
dorfü, Plumaria elegans, Fucus distichus ssp. distichus and
Chaetomorpha atrovirens. The second group included Cal-
lithamnion baileyi, Sargassum filipendula, and Codium
fragile ssp. tomentosoides. The last group included Ceram-
ium rubrum, Chondrus crispus, Corallina officinalis, Spha-
celaria cirrosa and Chaetomorpha linum.

Distinct winter, spring and summer annuals were evi-
dent at the seven sites. The first group included Bangia

1975] Саре Cod Algae — Coleman & Mathieson 81

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Figure 4. Monthly variation of annuals at stations 1, 4 and 7.

82 Rhodora [Vol. 77

fuscopurpurea, Petalonia fascia and Codiolum gregarium.
The second group included Desmarestia viridis, Desmo-
trichum balticum, Punctaria plantaginea, Enteromorpha
linza, Monostroma grevillei and Monostroma pulchrum.
The third group included Dasya pedicellata, Champia par-
vula, Chondria sedifolia, Grinnellia americana, Hypnea
musciformis and Seirospora griffithsiana, Figure 4 illus-
trates the monthly number of annual species at stations 1,
4 and 7. The largest number of annuals occurred during
the spring and summer months, except at station 4, and a
decrease was evident during the winter months.

Differential, spatial, and seasonal successions of annuals
were recorded north and south of the Cape. Three patterns
of seasonal succession were evident (Table VII). Some
species occurred earlier south than north of the Cape,
others disappeared earlier south than north, while a third
group was found throughout the year on both sides of the
Cape. Some examples of differential spatial] succession may
also be noted. The winter annuals Bangia fuscopurpurea
and Petalonia fascia appeared simultaneously in the Canal
and on the south shore of the Cape, but they were not found
at Scituate until several weeks later. Other spring and
summer annuals such as Ceramium strictum, Agardhiella
tenera, Lomentaria baileyana, Chorda filum and Leathesia
difformis appeared successively at Woods Hole, the Canal
stations and finally at Scituate.

SEASONAL REPRODUCTION

Species common to both sides of the Cape showed dif-
ferences in reproductive periods (Tables I-III). The an-
nuals can be divided into three groups as follows according
to their reproductive patterns: 1) species reproducing ear-
lier south than north of the Cape; 2) species terminating
reproduction earlier south than north of the Cape; 3) spe-
cies reproducing throughout the year north of the Cape,
but with limited reproductive periods to the south. Exam-
ples of each group are as follows: 1) Agardhiella tenera,

1975] Саре Cod Algae — Coleman & Mathieson 83

Chorda filum, Leathesia difformis, Ceramium strictum,
Chordaria flagelliformis, and Asperococcus echinatus; 2)
Bangia fuscopurpurea, Dumontia incrassata, Porphyra um-
bilicalis, Leathesia difformis, Petalonia fascia, and Mono-
stroma grevillei; 3) Porphyra umbilicalis, Petalonia fascia,
and Scytosiphon lomentarius. Table VIII summarizes all of
the species found in each group.

The perennials can also be divided into three groups
according to their reproductive patterns: 1) species pri-
marily reproducing during the colder months; 2) species
primarily reproducing during the warmer months; 3) spe-
cies reproducing throughout the year. Examples of each of
the groups are as follows: 1) Chondrus crispus, Petrocelis
middendorfii, Laminaria spp., and Polyides rotundus; 2)
Callithamnion baileyi, and Cystoclonium purpureum var.
cirrhosum; 3) Ascophyllum nodosum, Fucus ssp., Pilayella
littoralis, Sphacelaria cirrosa, and Ahnfeltia plicata. The
generalized groupings of species according to reproductive
periods were not always consistent at all stations. For
example, Pilayella littoralis and Chondrus crispus had an
extended reproductive period north of the Cape, but ex-
hibited limited periods of reproduction to the south. See
Tables I-III for specific details of reproduction at stations
1, 4 and 7.

DISCUSSION

Davis (1913a, b) suggests that the wide range of tem-
peratures near Cape Cod results in distinct annual popula-
tions. He distinguishes two groups of annuals as follows:
(1) winter-spring, and (2) mid-summer or early autumn.
We observed four types of annuals: winter, spring, summer
and aseasonal. Sears (1971) also recorded aseasonal an-
nuals during his study of the subtidal marine algae in
southern Cape Cod. The latter group showed no seasonal
specificity; the plants reproduced throughout the year and
they were represented by successive generations of young
plants. Among others, Davis (1931a, b), Setchell (1920),

84 Ећодога [Vol. 77

Williams (1948) and Chapman (1964) state that species
may survive adverse temperatures in a resting stage.
Sears’ (1971) observations substantiate the latter sugges-
tion, for he observed germlings of several red algae over-
wintering in the deep subtidal zone off Martha’s Vineyard,
Massachusetts.

Cold water perennials might be expected to enter a com-
parable state during the warmer months. Setchell (1917)
states that the northern fucoid alga Ascophyllum nodosum
enters a heat labor state during the summer south of the
Cape. We have observed a similar response for A. nodosum
south of the Cape, for it becomes bleached and unhealthy
looking during the summer.

The ratios of annuals to perennials varied at different
stations. North of Cape Cod perennials were dominant,
while to the south annuals were more abundant. Inter-
mediate ratios were found in the Canal. The seasonal tem-
perature regimes at the seven sites are correlated with the
percentage of annuals and perennials at each site. Thus, the
temperature range south of the Cape is about 22°C., while
to the north it is about 17°C.; intermediate values are evi-
dent in the Canal (Coleman and Mathieson, 1974). The
high summer temperatures south of the Cape allow warm
temperature annuals to occur in abundance. In contrast the
winter flora at the same sites is dominated by northern
(boreal) annuals. Williams (1948, 1949) has recorded a
similar seasonal variation of annuals at Cape Lookout,
North Carolina. He emphasizes that areas with wide tem-
perature fluctuations, such as Cape Lookout, support a wide
range of annuals.

Geographical differences of seasonal succession were evi-
dent. For example, warm water annuals occurred earlier
south of the Cape than north, while many cold water an-
nuals remained longer north than south of the Cape. The
early appearance of warm water annuals south of the Cape
resulted from higher spring temperatures in Buzzards Bay
than Cape Cod Bay. Likewise, some cold water annuals
remained longer north of the Cape, because of the lower

1975] Cape Cod Algae — Coleman & Mathieson 85

spring temperatures in Cape Cod Bay. It is suggested that
differences in seasonal successions at the stations are pri-
marily dependent on temperature differentials.

Variable reproductive periodicities were also evident for
the same species at different stations. For example, several
warm water annuals initiated reproduction earlier south
than north of the Cape, while cold water annuals repro-
duced longer north than south. The reproductive patterns
of perennials also showed similar patterns as the annuals,
again indicating the importance of temperature differences.

Acknowledgements

We would like to thank Dr. A. Hodgdon for his critical
review of the manuscript. In addition we express our grati-
tude to Dr. R. Fralick, Dr. R. Turner and Mr. John Hans-
man for their assistance with diving and collection of speci-
mens, and to the Army Corps of Engineers at Cape Cod for
diving access to the Cape Cod Canal.

LEGEND: TABLES LIII— а = alpha spore, b == beta spore,

C == carpospore, б == gametangia, M = monospore, PS == pluriloc-
ular sporangium, R — receptacle, S — spermatium, T — tetraspore,
US — unilocular sporangium, Z — zoospore, X — present, — — ab-

sent.

[Vol. 77

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TABLE IV. LONGEVITY OF SPECIES
Chlorophyta
TAXON Longevity

Chaetomorpha atrovirens Taylor

Chaetomorpha linum (Muller) Kützing

Chaetomorpha melagonium (Weber et Mohr) Kützing

Cladophora flexuosu (Muller) Harvey

Cladophora gracilis (Griffiths ex Harvey) Kützing

Codiolum gregarium A, Braun

Codiolum petrocelidis Kuckuck

Codium fragile (Sur.) Hariot ssp. tomentosoides
(van Goor) Silva

Enteromorpha intestinalis (L.) Link

Enteromorpha linza (L.) J. Agardh

Monostroma grevillei (Thuret) Whittrock

Monostroma pulchrum Farlow

Rhizoclonium tortuosum Kützing

Spongomorpha arcta (Dillwyn) Kützing

Spongomorpha spinescens Kützing

Ulothrix flacca (Dillwyn) Thuret

Ulva lactuca L.

Urospora collabens (C. Agardh) Homes et Batters

Urospora penicilliformis (Roth) Areschoug

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Phaeophyta

TAXON Longevity

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Ascophyllum nodosum (L.) Le Jolis

Asperococcus echinatus (Mertens) Greville A
Chorda filum (L.) Stackhouse A
Chorda tomentosa Lyngbye A
Chordaria flagelliformis (Muller) C. Agardh A
Desmarestia aculeata (L.) Lamouroux Р
Desmarestia viridis (Muller) Lamouroux A
Desmotrichum balticum Kitzing A
Desmotrichum undulatum (J. Agardh) Reinke A
Dictyosiphon foeniculaceus (Hudson) Greville | Р
Ectocarpus siliculosus (Dillwyn) Lyngbye A
Elachista fucicola (Velley) Areschoug Р
Fucus distichus L. emend. ssp. distichus (C. Agardh) Powell Р
Fucus distichus L. степа. ssp. edentatus (С. Agardh) Powell Р
Fucus distichus L. emend. ssp. evanescens (С. Agardh) Powell Р

98 Rhodora

Phaeophyta
TAXON

Fucus spiralis L.

Fucus vesiculosus L.

Fucus vesiculosus var. sphaerocarpus J. Agardh
Fucus vesiculosus var. spiralis Farlow
Giffordia granulosa (Smith) Hamel

Giffordia secunda (Kiitzing) Batters
Laminaria digitata (Hudson) Lamouroux
Laminaria saccharina (L.) Lamouroux
Leathesia difformis (L.) Areschoug
Myrionema strangulans Greville

Petalonia fascia (Muller) Kuntze

Pilayella littoralis (L.) Kjellman

Punctaria latifolia Greville

Punctaria plantaginea (Roth) Greville

Ralfsia fungiformis (Gunner) Setchell et Gardner
Ralfsia verrucosa (Areschoug) J. Agardh
Sargassum filipendula C. Agardh

Scytosiphon lomentarius (Lyngbye) Link
Sphacelaria cirrosa (Roth) C. Agardh
Sphaerotrichia divaricata (С. Agardh) Kylin

Rhodophyceae
TAXON

Agardhiella tenera (J. Agardh) Schmitz

Ahnfeltia plicata (Hudson) Fries

Antithamnion americanum (Harvey) Farlow

Bangia fuscopurpurea (Dillwyn) Lyngbye

Bonnemaisonia hamifera Hariot

Callithamnion baileyi Harvey

Callithamnion roseum (Roth) Lyngbye

Ceramium rubrum (Hudson) C. Agardh

Ceramium strictum Harvey

Champia parvula (C. Agardh) Harvey

Chondria sedifolia Harvey

Chondrus erispus Stackhouse

Choreocolax polysiphoniae Reinsch.

Clathromorphum circumscriptum (Strømfelt) Foslie

Corallina officinalis L.

Cystoclonium purpureum (Hudson) Batters var.
cirrhosum Harvey

[Vol. 77

Longevity

> ia. Ma. ar)

>

PUP PUP ey PP

Longevity

A?
Р
А

"d
m

"d VU eene

1975] Саре Cod Algae — Coleman & Mathieson 99

Rhodophyceae (Continued)
TAXON Longevity

Dasya pedicellata (C. Agardh) C. Agardh

Dermatolithon pustulatum (Lamouroux) Foslie

Dumontia incrassata (Muller) Lamouroux

Gigartina stellata (Stackhouse) Batters

Gloiosiphonia capillaris (Hudson) Carmichael ex Berkeley

Goniotrichum alsidii (Zanardini) Howe

Griffithsia tenuis C. Agardh

Grinnellia americana (C. Agardh) Harvey

Hildenbrandia prototypus Nardo

Hypea musciformis (Wulfen) Lamouroux

Lithophyllum corallinae (Crouan) Heydrich

Lithothamnium glaciale Kjellman

Lomentaria baileyana (Harvey) Farlow

Lomentaria orcadensis (Harvey) Collins ex Taylor

Melobesia lejolisii Rosanoff

Petrocelis middendorfii (Ruprecht) Kjellman

Phyllophora brodiíaei (Turner) Endlick

Phyllophora membranifolia (Goodenough ex Woodward)
J. Agardh

Phymatolithon lenormandi (Areschoug) Adey

Plumaria elegans (Bonnemaison) Schmitz

Polyides rotundus (Hudson) Greville

Polysiphonia denudata (Dillwyn) Greville ex Harvey
in Hooker

Polysiphonia elongata (Hudson) Sprengel

Polysiphonia harveyi Bailey

Polysiphonia lanosa (L.) Tandy

Polysiphonia nigrescens (Hudson) Greville

Polysiphonia novae-angliae Taylor

Polysiphonia urceolata (Lightfoot ex Dillwyn) Greville

Porphyra leucosticta Thuret

Porphyra miniata. (C. Agardh) C. Agardh

Porphyra umbilicalis (L.) J. Agardh

Rhodochorton penicilliforme (Lightfoot) Rosenvinge

Rhodomela confervoides (Hudson) Silva

Rhodymenia palmata (L.) Greville

Seirospora griffithsiana (Harvey) Dixon

Trailliella intricata (J. Agardh) Batters

~

ну бе ну НИ Маре re ну Ру МУР кышы WUNNEN р р >

100 Rhodora [Vol. 77

TABLE V. DOMINANT ANNUALS NORTH AND SOUTH
OF THE CANAL

ANNUALS

МОЕТН

Bangia fuscopurpurea
Dumontia incrassata
Lomentaria baileyana
Polysiphonia harveyi
Porphyra umbilicalis

Asperococcus echinatus
Chorda filum

Chorda tomentosa
Chordaria flagelliformis
Ectocarpus confervoides
Leathesia difformis
Petalonia fascia
Scytosiphon lomentaria

Codiolum petrocelidis
Enteromorpha intestinalis
Monostroma grevillei
Monostroma pulehrum
Spongomorpha arcta
Spongomorpha spinescens
Ulothrix flacca

Ulva lactuca

Urospora collabens
Urospora penicilliformis

SOUTH

Agardhiella tenera
Antithamnion americanum
Bangia fuscopurpurea
Cailithamnion roseum
Ceramium strictum
Champia parvula
Chondria sedifolia
Dasya pedicellata
Dumontia incrassata
Grinnellia americana
Hypnea musciformis
Lomentaria baileyana
Polysiphonia harveyi
Porphyra umbilicalis
Seirospora griffithsiana
Trailliella intricata

Asperococcus echinatus
Chorda filum

Chorda tomentosa
Chordaria flagelliformis
Desmotrichum undulatum
Ectocarpus confervoides
Leathesia difformis
Petalonia fascia
Punctaria latifolia
Punctaria plantaginea
Scytosiphon lomentaria
Sphaerotrichia divaricata

Enteromorpha intestinalis
Monostroma grevillei
Monostroma pulchrum
Spongomorpha arcta
Ulothrix flacca

Urospora penicilliformis

1975]

TABLE VI.

Cape Cod Algae — Coleman & Mathieson

101

DOMINANT PERENNIALS NORTH AND SOUTH

OF THE CANAL
PERENNIALS

Моктн
Ahnfeltia plicata
Ceramium rubrum
Chondrus crispus
Choreocolax polysiphoniae
Clathromorphum circumscriptum
Corallina officinalis
Cystoclonium purpureum var.

cirrhosum

Gigartina stellata
Hildenbrandia prototypus
Lithophyllum corallinae
Lithophyllum macrocarpum
Petrocelis middendorfii
Phyllophora membranifolia
Phymatolithon lenormandi
Plumaria elegans
Polyides carpinus
Polysiphonia lanosa
Polysiphonia nigrescens
Polysiphonia urceolata
Rhodochorton penicilliforme
Rhodomela confervoides
Rhodymenia palmata
Ascophyllum nodosum
Desmarestia aculeata
Elachista fucicola
Fucus distichus ssp. distichus
Fucus distichus ssp. edentatus
Fucus spiralis
Fucus vesiculosus
Laminaria digitata
Laminaria saccharina
Pilaiella littoralis
Ralfsia fungiformis
Ralfsia verrucosa
Sphacelaria cirrosa
Chaetomorpha atrovirens
Chaetomorpha linum
Chaetomorpha melagonium
Rhizoclonium tortosum

SOUTH
Callithamnion baileyi
Ceramium rubrum
Chondrus crispus
Corallina officinalis
Cystoclonium purpureum

var. cirrhosum
Fosliella lejolisii
Lithophyllum macrocarpum
Phyllophora membranifolia
Polysiphonia nigrescens
Polysiphonia urceolata

Ascophyllum nodosum
Dictyosiphon foeniculaceus
Elachista fucicola
Fucus vesiculosus

var. sphaerocarpus
Pilaiella littoralis
Sargassum filipendula
Sphacelaria cirrosa

Chaetomorpha linum
Cladophora flexuosa
Cladophora gracilis
Codium fragile ssp.
tomentosoides

[Vol. 77

Rhodora

102

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104 | Ећодога [Vol. 77

REFERENCES

CHAPMAN, V. J. 1964. The Algae. Macmillan & Company, Ltd.
London. 472 pp.

COLEMAN, D. С. & A. C. MATHIESON. 1974. Investigations of New
England Marine Algae VI. The distribution of marine algae
near Cape Cod, Massachusetts. Rhodora 76: 587-568.

CONOVER, J. T. 1958. Seasonal growth of benthic marine plants as
related to environmental factors in an estuary. Publ. Inst. Mar.
Sci. 5: 97-147.

Davis, B. M. 1918a. General characteristics of the algal vegeta-
tion of Buzzards Bay and Vineyard Sound in the vicinity of
Woods Hole. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fisheries
31: 443-544.

————————. 1913b. A catalogue of the marine flora of Woods Hole
and vicinity. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fisheries,
31: 795-833.

FaARLOW, W. С. 1870. XVII. List of sea-weeds or marine algae of
the south coast of New England. U. 5. Comm. of Fish and
Fisheries. Commissioner's Report 1871-1872. Washington: Gov-
ernment Printing Office. Pp. 281-294.

1882. I. The Marine algae of New England. Ibid.,
report 1879. Pp. 1-210.

Harvey, W. Н. 1852-58. Nereis Boreali-Americana. I. Melano-
spermae. Smithsonian Contrib. Knowl. 3: 1-150, pl. 1-12. 1852;
II. Rhodospermae. Ibid. 5: 1-258, pl. 13-36. 1853; III. Chloro-
spermae. Ibid. 10: ii. + 1-140, pl. 37-50. 1858.

HuMM, H. J. 1969. Distribution of marine algae along the Atlantic
Coast of North America. Phycologia 7: 43-53.

SEARS, J. К. 1971. Morphology, systematics and descriptive ecology
of the sublittoral benthic marine algae of southern Cape Cod
and adjacent islands. Ph.D. Thesis. 295 pp. University of Mass.

SETCHELL, W. A. 1917. Geographic distribution of the marine algae.
Science 45: 197-204.

———————. 1920. Stenothermy and zone-invasion. Am. Nat. 54:
385-397.

1922. Cape Cod in its relation to the marine flora of
New England. Rhodora 24: 1-11.

WILLIAMS, L. G. 1948. Seasonal alternation of marine floras at

Cape Lookout, North Carolina. Am. Jour. Bot. 35: 682-695.
1949. Marine algal ecology at Cape Lookout, North
Carolina. Bull. Furman Univ. 31: 1-21.

JACKSON ESTUARINE LABORATORY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM, N.H. 03824

A LIST OF THE MONOCOTYLEDONEAE OF BELIZE
INCLUDING A HISTORICAL INTRODUCTION
TO PLANT COLLECTING IN BELIZE

DAVID L. SPELLMAN, JOHN D. DWYER,
AND GERRIT DAVIDSE

The country of Belize, formerly British Honduras, which
has had internal self-government since 1964, lies south of
the Yucatan Peninsula about 800 miles south of New
Orleans, Louisiana. The country is bounded on the north
by Mexico’s Territory of Quintana Roo, on the west and
south by Guatemala, and on the east by the Caribbean Sea.
In length, the country extends from the Rio Hondo in the
north to the Sarstoon River in the south, a distance of 174
miles. At its widest point, it is 70 miles from Belize City
on the sea to the western border. In area Belize is ap-
proximately the size of the State of Massachusetts. The
country encompasses 8,750 square miles exclusive of 116
square miles of offshore islands known locally as cays. It is
divided into the following political districts: Belize, Cayo,
Corozal, Orange Walk, Stann Creek, and Toledo (Fig. 1).

Belize 1з almost bisected by the Belize River. The north-
ern half of the country is similar to the southern part of
the Yucatan Peninsula, being a fairly level plain with a
maximum elevation of about 120 m. above sea level. The
southern part of the country is mountainous for the most
part. The dominant physiographic features in the south
are the Maya and Cockscomb Mountains. The reader is
referred to a recent paper (Kessler et al., 1974) dealing
with the geological history of this area. Average elevations
in both these mountain masses lie between 850 т. and
910 m. above sea level, with the country’s highest point
reaching over 1130 m.

Climatically the country is described as sub-tropical even
though under the Koeppen classification it is classified as
“Ат” (tropical monsoon) north of the Maya Mountains

105

106 Ећодога [Vol. 77

-—
|

Toledo

Guatemala

Honduras

Figure 1. Map of Belize showing administrative districts, and
relationship to neighboring countries (inset).

1975] Belize Monocotyledoneae — Spellman et al. 107

and “Af” (tropical rainforest) in the mountainous south.
There is a marked dry season from February to May. Pre-
cipitation is greatest in the south and in the central moun-
tain mass, and decreases to the north. Average annual
precipitation ranges from 432 cm. in the Toledo District in
the south to 132 cm. in the Corozal District in the north.

According to Lundell (1945), in terms of area Belize has
a floral diversity which is perhaps unequaled among the
countries of Middle America. Standley & Record (1936)
state that the most significant element of the flora is West
Indian. There are a large number of plants native to Cuba,
Jamaica, and other parts of the West Indies, which on the
continent are known only from Belize. This provides stu-
dents of plant geography with a unique opportunity for
investigation. Pine stands, swamps, and marshes occur in
the northern plain. In the south the vegetation approaches
rainforest in composition, and resembles in some respects
forests in adjacent Guatemala with an admixture of Pan-
amanian and South American elements.

Sprague & Riley (1924) predicted a flora of about 4000
species for the country, an estimate with which Lundell
(1945) concurred. Standley & Record (1936) regarded
endemism as low, citing a total of 138 species which could
be so designated. If this be the rate of endemism, it will be
about four percent. This is strikingly low, considering that
Lundell estimated about fifteen percent for the entire Yuca-
tan Peninsula.

Sprague’s statement in 1924 that the flora of Belize was
perhaps less known than that of any other British Colony
still holds true today. Prior to 1883, there were probably
not more than twenty sheets of dried plants in the Kew
Herbarium (Sprague & Riley, 1924). During the period
1883 to 1894, this number was increased by about 200
through the efforts of Daniel Morris, E. D. M. Hooper,
J. Robertson, and Sir Alfred Maloney, the then Governor
of the Colony.

The first collection of any significant size was that of
Morton E. Peck of the Yale University School of Forestry.

108 Rhodora [Vol. 77

Professor Peck visited the country during the years 1905
to 1907, collecting about 1000 numbers. No complete ac-
count of the Peck collections has ever been published, but
partial accounts have been given of the grasses by Hubbard
(1913) and of the Dicotyledoneae by Blake (1917) and
Sprague & Riley (1924). The Peck collections include more
than 50 type specimens.

Following Peck’s visit to the country, virtually no bota-
nizing occurred for two decades. A resurgence of activity
took place in 1926, however, when a cooperative study was
initiated by the Forestry Department of the country and
the Yale School of Forestry, chiefly through the efforts of
Samuel J. Record and his colleagues at Yale University.
This led to collections by several of the University personnel
as well as by various members of the local Forestry Depart-
ment and employees of private timber companies.

In 1928 and 1929 C. L. Lundell collected approximately
700 numbers at Honey Camp in the Orange Walk District,
under the auspices of the Tropical Research Foundation.
In 1929, the significant collections of W. A. Schipp were
started in Stann Creek and Toledo Districts. Schipp’s bo-
tanical activities in Belize were discussed at length by
R. M. Lowden (1970). It appears that Schipp’s sole means
of support was obtained through the sale of herbarium and
seed collections. His Flora of British Honduras for 1933-
1934 is a collector’s item; it is a mimeographed catalogue
listing some 1129 flowering plants and offering for sale
seeds and preserved plant parts. His collections from 1929
to 1941 contain at least 3500 numbers.

The period 1931 to 1958 is noteworthy for the quantity
of plants collected in Belize as well as their quality. During
the Maya Expeditions of the Carnegie Institution and the
University of Michigan, H. H. Bartlett and C. L. Lundell
botanized extensively in Cayo District and made smaller
collections in Belize District (Lundell, 1940). Lundell’s
collections of 1928 through 1936 amount to about 3000
numbers. у

1975] Belize Monocotyledoneae — Spellman et al. 109

During the Maya Expedition of 1931, Bartlett trained
Percy Gentle of Belize City as a botanical collector (Bart-
lett, 1935). Lundell later assumed the responsibility of
directing Gentle’s field activities. Gentle remained an active
collector until shortly before his death in 1958. His col-
lection represents the largest by a single individual, just
under 10,000 numbers being gathered between 1951 and
1958 (Lundell, 1960).

The Cambridge University Expedition of 1959 and 1960
accounts for two significant sets of collections. The first of
these was a series of more than 700 numbers collected by
the Expedition botanist D. R. Hunt. Hunt’s collections are
from Toledo, Stann Creek, and Cayo Districts. Most of his
numbers seem to be from the Mountain Pine Ridge area of
Cayo District. The second collection was made by the
geographer of the Expedition, D. R. Stoddart. Stoddart’s
collecting activities were in conjunction with his mapping
of certain of the cays. While his collections were not as
extensive as those of Hunt, they are important, since they
represent the only extensive collections from the offshore
islands. Subsequent visits by Stoddart, with F. R. Fosberg,
and M.-H. Sachet and D. Spellman, have yielded rather
complete collections from 65 of the cays.

From 1961 to 1969 several botanists visited the country ;
these include George Proctor of the Institute of Jamaica,
John Dwyer of St. Louis University and the Missouri Bo-
tanical Garden, Thomas Elias, then a graduate student at
Southern Illinois University (Carbondale), Martin Kell-
man of Simon Fraser University, and Edward Tyson and
his students from Florida State University.

The year 1970 yielded two sets of collections. The senior
author collected about 800 numbers in January and in July
while working with the University of Edinburgh Expedi-
tion to British Honduras (Furley, 1972). Sidney McDaniel
of Mississippi State University made extensive collections
in March.

From 1972 to the present time the bulk of plant collecting
in Belize has been done by botanists from the Missouri

110 Rhodora [Vol. 77

Botanical Garden. Thomas Croat, Curator of Phanerogams,
collected 1794 numbers in 21 field days in the period from
the 23га of May to June 19, 1973. Alwyn Gentry, Assistant
Curator at the Garden, collected about 1000 numbers during
the same time. John Dwyer, Research Associate at the Gar-
den, made several trips to Belize since 1972, collecting about
2750 numbers, almost 1000 numbers being gathered at the
same time and in the same localities visited by Croat and
Gentry. In eleven days in the field in January, 1974, Dwyer
returned to Belize to collect about 450 numbers in the cen-
tral and northern sections of the country. In ten days of
field work in January, 1974, Liesner collected 293 numbers
in the Districts of Belize, Cayo, and Orange Walk. By
arrangement with the Belize Government, representative
collections from the Croat, Dwyer, Gentry, and Liesner
lots are being selected and forwarded from the Garden for
permanent deposit in the recently established herbarium at
Belmopan, the new capital of Belize.

The following are other botanists who over the past four
years have done some collecting in Belize:

Dr. Derek Burch, University of South Florida

Dr. William Crankshaw, Ball State University

Father Leonard Dieckman, S.J., St. Johns College, Belize
City

Dr. Richard Keating, Southern Illinois University (Ed-
wardsville)

Dr. Yon Linhart, University of Colorado

Dr. Robert Long, University of South Florida

Dr. Richard Pippen, Western Michigan University

Dr. Paul Sorenson, Northern Illinois University

Dr. Mario Sousa, The National University of Mexico

Mr. Bruce Vanderveen, graduate student, St. Louis Uni-
versity

Dr. Richard Wunderlin, University of South Florida.

The following list includes most of the botanists who col-
lected in Belize during the past century or more and the
year or years in which they collected :

1975] Belize Monocotyledoneae — Spellman et al. 111

Aguilar, Mercedes

Aitken, J. B. — 1927
Anderson, Roger — 1970
Bartlett, H. H. — 1931
Brown, C. S. — 1927-1929?
Brunt, Martin

Burns, J. A.

Campbell, A. J. Е. — 1922
Carrick, J.

Castillo, A. — 1932
Chanek, M.

Contreras, Elias — 1968
Crankshaw, William — 1970
Croat, Thomas — 1973

Day, David Fisher — 1864
Dieckman, Rev. L. — 1970-1972
Donald, G. H.

Dwyer, John — 1967; 1972-1974
Egler, F. E. — 1942

Elias, Thomas — 1967

Fralish, James — 1970

Gentle, Percy H. — 1931-1958
Heyder, Н. M. — 1927

Hooper, E. D. M.

Hope, M. O.

Hummel, C.

Hunt, D. R. — 1959-1960
Karling, J. S. — 1927-1928
Keating, Richard — 1971-1972
Kellerman, W. A. — 1906-1907
Kellman, M. C. — 1969
Kinloch, J. B. — 1932-1955
Kluge, H. C. — 1924?

Lamb, A. F. A. — 1947?
Lancaster, probably Thomas Leonard, a New Zealander
Lazor, Robert — 1970

Liesner, Ron — 1974

Long, Robert — 1970

112. Ећодога [Уо]. 77

Lundell, C. L. — 1928-1929

McDaniel, Sidney — 1972

Maxwell, Richard — 1967

Meyer, William C. — 1930

Molino, Antonio — 1954

Maloney, Sir Alfred — 1893-1894

Morris, Sir Daniel — 1893-1894

Newey, Walter W. — 1970

O'Neil, Rev. Hugh — 1936

Peck, M. E. — 1905-1907

Pelly, R. S. — 1933

Pippen, R. — 1972-1973

Proctor, G. R. — 1969

Record, S. J. — 1926; 1930

Robertson, Rev. J. — 1883-1884

Rose-Innes, R. — 1970

Sachet, M.-H. — 1972.

Sampson, H. C. — 1928

Schipp, William A. — 1929-1938

Schmidt, K. P. — 1923

Smart, H. P. — 1920; 1930?

Sorenson, Paul — 1972

Sousa, M. — 1973

Spellman, D. L. — 1970-1972

Stanton, Rev. W. A., S.J. — 1896; ca. 1906

Stevenson, Duncan — 1927-1928

Stevenson, Neil S. — 1927-1928

Stocker, C. L. — 1924

Stoddart, D. R. — 1959-1972

Thompson, J. E. — 1927-1929; 1931

Tyson, Edward — 1970

Vaughn, Joseph — 1971

Vanderveen, B. — 1973

Winzerling, H. W. — 1926-1927

Wunderlin, Richard — 1971

We have been unable to determine the given names of
the following collectors: Mrs. Armstrong; Disney; Einth;
Espat; and Saunders.

1975] | Belize Monocotyledoneae — Spellman её al. 113

LIST OF THE MONOCOTYLEDONEAE

The flora of Belize has been treated in part or in a com-
prehensive fashion in a number of publications. Schipp's
(1933-1934) Flora of British Honduras, Price List of Seeds
& Herbarium Material was the first attempt to list all of
the vascular plants from the country. This was followed
by the checklist of Standley & Record entitled The Forests
and Flora of British Honduras, which is considerably more
complete. As a result of the Maya Expeditions in the 1930's,
Lundell and other contributors published detailed taxonomic
papers on several plant families or parts thereof. These
appeared in two volumes entitled Botany of the Maya Area
(Publ. Carnegie Inst. Wash., 1936, 1940). "These treat-
ments, although wider in geographical coverage than Belize,
are a valuable source of information for the flora of Belize.
Since 1946 the Flora of Guatemala, now nearing completion,
has attempted to include the plants of Belize as well as those
of Guatemala and currently represents the most authorita-
tive source of information for the flora of Belize.

The following list of Monocotyledoneae is presented as a
way of updating current knowledge of this part of the
Belize flora. The list was prepared by searching the litera-
ture, by examining collections in herbaria, by identifying
recently collected materials, and by checking field notes and
unpublished lists. Any novelties encountered in studying
very recent collections will be described and published sep-
arately by specialists. Virtually all collections made recently
by botanists from the Missouri Botanical Garden are de-
posited in the herbarium of the Garden (Mo).

We have attempted to cite individually most of the col-
lections made in Belize since 1959, especially the extensive
ones made by the staff of the Missouri Botanical Garden in
the last few years. Since many of the collections made be-
fore 1959 have been treated in the Botany of the Maya Area
or formed a basis for the treatments in the Flora of Guate-
mala, as well as in monographs and other taxonomic papers,
we have not cited these specimens specifically. We have

114 Ећодога [Vol. 77

examined all collections cited in this paper with the excp-
tion of those of D. R. Hunt.

The names of the collectors encountered in the list are
abbreviated as follows:

C = Thomas Croat

D = John Dwyer

Dieck = Leonard Dieckman

G = Alwyn Gentry

н — D. К. Hunt

K — Martin Kellman

Lies — Ron Liesner

Lin — Yon Linhart

Lon — Robert Long

P — George Proctor

Sor — Paul Sorenson

Sp — Spellman

V — Bruce Vanderveen

D&L — Dwyer and Liesner

D & Р = Dwyer and Pippen

DEM = Dwyer, Elias and Maxwell

L&D — Liesner and Dwyer

Sp & St — Spellman and D. R. Stoddart*

*All collections by Spellman and Stoddart, cited as Sp & St, are
from the cays.

The authors of this paper would like to express their
thanks to the Belize Government, especially to the Premier,
the Hon. George Price, for his sustained interest in the
project, as well as to Mr. E. O. Bradley, Acting Chief
Forest Officer, for his assistance. The extensive collecting
done by the botanists of the Missouri Botanical Garden in
1973-1974 has been facilitated greatly by the physical facili-
ties provided at Ridge Lagoon Plantation, located at Mile
1115 on the Northern Highway, Belize by Robert Baum-
gartner, an American residing in Belize. The assistance
rendered by the faculty at St. Johns College, Belize City is
gratefully acknowledged. Тће senior author derived valu-
able field experience from his association with the Univer-

1975] Belize Monocotyledoneae — Spellman et al. 115

sity of Edinburgh Expedition to Belize in July-August 1970
under the direction of Dr. Peter Furley. He was also fortu-
nate in having botanized the Cays of Belize with Dr. Ray-
mond Fosberg and Dr. David Stoddart during June-July,
1972. Father Leonard Dieckman, S.J., and Mrs. Lydia
Waight of Belize City were kind enough to check the his-
torical introduction. Special thanks are due to Dr. Thomas
Croat for his assistance in identifying the Araceae and
Bromeliaceae. Special thanks are due the National Geo-
graphie Society for financial support given to Dr. Dwyer
during the years 1973 and 1974 as well as the support re-
ceived from the Associated Universities for International
Education.

In the list, those species which we judge have not been
previously recorded from Belize in published accounts, are
marked by an asterisk before the binomial.

Alismataceae
Echinodorus andrieuxii (Н. & A.) Small — P 29929
E. grandiflorus (Cham. & Schlecht.) Micheli — D 10978
E.nymphaeafolius (Griseb.) Buch.
Sagittaria lancifolia L.

Amaryllidaceae
Bomarea edulis (Tuss.) Herb. — P 29659
Crinum erubescens Ait.
Curculigo scorzonerifolia (Lam.) Baker — Н 22
*“Furcraea guatemalensis Trel. — Sor 7052
Hippeastrum puniceum. (Lam.) Urban — Lin 205
Hymenocallis cf. caribaea Herb. — Sp 1787; Sp 1843
H. littoralis (Jacq.) Salisb. — P 29938; Sp & St 2268; Sp
& St 2288; Sp & St 2360; Sp & St 2449; Sp & St 2478
Zephyranthes lindleyana Herb.

Araceae
Anthurium sp. — G 1833
А. aemulum Schott — D 9859; D 11560; D & L 12343;
G 8025; Sp 1604
A. bakeri Hook. f. — C 24225; C 24453

116 Ећодога [Vol. 77

A. concinnatum Schott

A. crassinervium (Jacq.) Schott — L & D 1472

A. denudatum Engler — C 23533

A.gracile (Rudhe) Lindley — G 7954; V 616; V 618;
V 621

A.scandens (Aubl) Engler — C 23592; C 24309; D
12690; G 7912; D 9946; P 30356

A. scolopendrinum (Hamilt.) Kunth

A. tetragonum Hook. ex Schott. — Sp 1948; V 658

A. trinerve Miq. — G 7413

A. verapazense Engl. — P 29825

Dieffenbachia aurantiaca Engl. — D 11184

D. sequina, (L.) Schott

Monstera acuminata Koch

M. adansonii Schott — G 7434

M. belizensis Lundell

M. magnispatha Matuda

M. tuberculata Lundell

Montrichardia arborescens (L.) Schott — L & D 1448;
L&D 1644

Philodendron belizense Standl.

P. brevispathum Schott — G 8423

P. fragrantissimum (Hook.) Kunth — C 24510

P. guttiferum Kunth — D 11306; G 8018; G 8241; P
29700

Р. hederaceum (Jacq.) Schott — Sp 1967

P. inequilaterum Liebm. — С 24305; D 11103

P. panamense Krause — D 9891; G 8033; G 8132

P. radiatum Schott — C 23534; C 24335; C 24512

P.scandens C. Koch & Cello — C 23697; C 24511; D
12718

P. tripartitum (Jacq.) Schott — C 23670; C 24509

Pistia stratiotes І. — D & L 12158

Rhodospatha acuminata C. Koch

R. nervosa Lundell

Spathiphyllum blandum Schott — G 8017; G 8147; G 8255

S. friederichsthali Schott — C 24332; C 24451A

S. phryniifolium Schott

1975] Belize Monocotyledoneae — Spellman et al. 117

Syngonium podophyllum Schott —D & L 12043; G 8133;
1, & Р 1468; Р 29596; Р 29916; Sp 1584
Xanthosoma violaceum Schott — D & L 12078; P 29941

Bromeliaceae

Aechmea bracteata (Sw.) Мег — С 23922; C 23950; C
24686; D 10116; D & L 12035; D & L 12184; P 80288

A. bromeliaefolia (Rudge) Baker

A. dactylina Baker — G 6791

A. mexicana Baker — D & L 12129

A. nudicaulis L. Griseb.

A. tillandsioides (Hart.) Baker — C 24678; P 30178; V
607

Ananas comosus (L.) Merr.

Androlepis donnell-smithii (Baker) Mez — C 23322; C
24314

Bilibergia viridiflora Wendl.

Catopsis aloides (Cham. & Schlecht.) Baker

C. berteroniana (Schult.) Mez — P 29818

C. morreniana Mez — Sp 1988; Sp 2002

C. sessiliflora (К. & Р.) Мег var. dioica L. B. Sm.

Guzmania lingulata (L.) Мег — C 23768; D 9934

Pitcairnia imbricata (Brong.) Regel

P. hemsleyana Mez

P. aff. integrifolia Gawl — C 23848

P. petiolata (Koch & Bouche) Baker — P 29439

Tillandsia balbisiana Schult. — D & L 12199; P 29529;
P 29598

T. bulbosa Hook. — C 24041; С 24803; D & L 12187A;
D&L 12362; G 7974; P 29645

T. dasyliriifolia Baker — C 23274; DEM 623; H 612

T. digitata Mez

T. excelsa Griseb.

T. fasciculata Sw. — D & L 12167 ; Sp 1981

T. festucoides Brong. ex Mez — С 23607; G 7788; P 30223

T. filicifolia Cham. & Schlecht.

T. juncea (R. & P.) Poir. — G 7961

T. leiboldiana Schlecht.

118 Rhodora [Vol. 77

T. monadelpha (E. Morren) Baker — С 7974

T. polystachia L.

T. schiedeana, Steud. — C 23760; C 24827; D 9847

T. streptophylla Scheidw. — С 23881 A; D & L 12186;
D & L 12365; L & D 1600; P 29528

T. sublaxa Baker

T. utriculata L.

T. valenzuelana A. Rich. — C 23766; C 23758

T. vestita Schlecht. & Cham.

Vriesa disticha (L.) Small

V. paniculata (L.) Mez

V. schippii L. B. Sm.

Burmanniaceae
Apteria aphylla (Nutt.) Barnh. ex Small
Burmannia capitata (Walt.) Mart.—D & L 12369; P
29684
B. flava Mart.
Dictyostegia campanulata Karst.
D. orobanchioides (Hook.) Miers
Gymnosiphon tenellus (Benth.) Urban

Cannaceae
Canna edulis Ker — K 451; P 29465
С. indica L. — DEM 141; Sp 1411

Commelinaceae
Callisia repens L.
C. ciliata H.B.K.
Campelia zanonia (L.) H.B.K. — D 9854; D 9883
Commelina diffusa Burm.f, — K 411
C. elegans H.B.K.
С. erecta L. — С 23801; С 24100; G 8008
C. longicaulis Jacq.
Dichorisandra hexandra (Aubl.) Standl.
Phaeosphaerion persicariaefolium (DC.) C. B. Clarke
Rhoeo spathacea (Sw.) Stearn — P 29646; Sp 1977; Sp
1980 ; Sp 1987 ; Sp 2007
Tradescantia belizensis Standl.

1975] Belize Monocotyledoneae — Spellman et al. 119

T. geniculatum Jacq. — P 30078

T. lundellii Standl.

Tripogandra cumanensis (Kunth) Woodson — P 30153

T. grandiflora (D. Sm.) Woodson — H 399; К 592; P
29692; Sp 2008

T.serrulata (Vahl) Handlos — C 23717; С 7981

Zebrina pendula Schnizl.

Cyclanthaceae
Asplundia chiapensis (Mart.) Harl.
Carludovica palmata R. & P.
C. utilis (Oerst.) Benth. & Hook.

Cyperaceae
* Abildgaardia ovata (Burm.f.) Kral — D 11050
Bulbostylis arenaria (Nees) Lindm. — Н 189; P 29431
B. capillaris (L.) C.B. Clarke
B. junciformis (H.B.K.) Lindm. — H sn; H 13; H 244
B. paradoxa (Spreng. Lindm. — D & P 10124; Н 81;
P 29432
B. tenuifolia (Rudge) Macbr. — С 28962
B. vestita, (Kunth) C.B. Clarke — D 11608; Н 18
Calyptrocarya glomerulata (Brong.) Urban — С 8595
Carex polystachya Sw. ex Vahl var. bartletti (O'Neill)
Standl. & Steyermark — H 248; P 29647
Cladium jamaicense Crantz — С 23296; C 24995; D
12642; DEM 629; Dieck 294; б 8527; бр & St 2239
Cyperus articulatus L.
. brevifolius (Rottb.) Endl. ex Hassk.
C. compressus L.
C. cyperoides (L.) Kuntze — H 283
C. diffusus Vahl — C 24865; G 7997; G 8032; G 8443
C. digitatus Roxb. — P 29926
C. eggersii Boeck.
C
C
C

Q

. elegans L.
‚ esculentus L.
. flavus (Vahl) Nees
C. haspan L. — C 28915; C 23998; С 24865; D 10756;
H 100; P 29675; P 29716

120 Rhodora [Vol. 77

С. hermaphroditus (Jacq.) Standl. — C 24853
C. humilis Kunth — P 29580
С. imbricatus Retz.
. ischnos Schlecht.
. lanceolatus Poir.
. lentiginosus Millsp. & Chase
. ligularis L. — С 24066; C 24106; C 24533; DEM 516;
Lin 30; Sp & St 2211; Sp 2450
. luzulae (L.) Retz. — С 23522; С 23800; D 12705;
DEM 489; G 7996; G 8000; G 8431; P 29934; H 176;
H 254; Sp 1591
.macrocephalus Liebm. — C 23370
. mutisii (H.B.K.) Griseb. — Н 477
. ochraceus Vahl — С 24007; С 24660; P 29928; Sp 1814
. odoratus І. — C 24009; C 24579; G 7999; DEM 274;
Dieck 161; P 29927
. pallens (Liebm.) Standl. & Steyermark
. peruvianus (Бат. F.N. Williams — C 24111; DEM
515; Sp & St 2271; Sp & St 2343; Sp 2400; Sp 2469
C. planifolius L. Rich. — Lin 1; Lin 2; Lin 31; Lin 71;
Sp & St 2202; Sp & St 2210; Sp & St 2216; Sp & St
2237; Sp & St 2257; Sp & St 2270; Sp & St 2272; Sp
& St 2289; Sp & St 2324; Sp & St 2372; Sp & St 2385;
Sp 2400; Sp 2408; Sp 2422; Sp 2503; Sp 2537; Sp 2560
C. polystachyos Rottb. — C 24105; C 24865A; D 11465
C. prolixus H.B.K. — D 10855
C. rotundus L. — C 24085
C. semiochraceus Boeck.
C. surinamensis Rottb. — G 7996
C. tenuis Sw. — K 419; K 426
C. unioloides R. Br. — Н 98
Eleocharis caribaea (Rottb.) Blake — D & L 1255; С
1868; DEM 607
E. cellulosa Torr.
E.elegans (H.B.K.) R. & S. — C 24868; D 10760; DEM
259; H. 175
*E. elongata Chapm. — H 379
E. filiculmis Kunth — DEM 238; Н 422; D & L 1670

о сода

сусу сода

1975] Belize Monocotyledoneae — Spellman et al. 121

E. flavescens (Poir.) Urban
E. interstincta (Vahl) Е. & S.
. maculosa (Vahl) R. & S. — E 170; Н 415
. minima Kunth — H 445
. minutissima Britt.
. montana (H.B.K.) R. & 5. — D & L 12328
. mutata (L.) К. & S. — Vaughn et al. 2768
. nigrescens (Nees) Steud.
. pachystyla (C. Wright) C.B. Clarke — Н 377
E. retroflexa (Poir.) Urban
Fimbristylis annua (АП.) В. & S.
F. autumnalis (L.) R. & S.
Е. castanea (Michx.) Vahl
F. complanata (Retz.) Vahl — D 10855
Е. dichotoma (L.) Vahl — D 9525А; Н 618
F. milacea (Т...) Vahl
К. spadicea (L.) Vahl — C 24095; D 11288; D& P 10037;
Vaughn eft al. 275
Ё. spathacea Roth — Sp & St 2445; Sp & St 2277; Sp &
St 2877; Sp 2415; Sp 2511; Sp & St 2565
F. vahlii (Lam.) Link.
Fuirena bulbipes Blake — C 24043; C 24125; D 11377;
G 7877; G 8529
*F. campotricha C. Wright — H 97; H 329
Е. incompleta Nees
Р. simplex Vahl
F. umbellata Rottb. — P 29579
Hypolytrum longifolium (L. Rich.) Nees ssp. тсата-
guense (Liebm.) Koyama — D & L 12067
Lagenocarpus guianensis Lindl. & Nees
Lipocarpa maculata (Michx.) Torr.
Rhynchospora barbata (Vahl) Kunth — С 23273; C
24082; D 10912A ; G 8530; H 342; H 621; L & D 1683
R. cephalotes (L.) Vahl — C 23516; С 24786; G 8411;
H 250
* R. chapmanii M. А. Curt. — DEM 442
К. colorata (L.) Н. Pfeiff. — C 23256; D 10931; С 7865;
H 393

ч ш ES ESSE

122 Rhodora [Vol. 77

R. corymbosa (L.) Britt. — C 24067
R. curvula Griseb. — D & P 10050; H 337
R. cyperoides (Sw.) Mart. — C 23254; C 23341; C 23511;
C 23820; С 23952; С 23968; С 24768; D 11070; D
11482A; D 12767; D & P 10017; G 7871; G 8528;
Н 169; Н 237; Sp 1636
R. divergens M. A. Curt. — C 23260; C 24080A; D 10726;
D 12420; D 12615
R. exaltata Kunth — D 11687
R. exima (Nees) Boeck. — D 9050
R. fascicularis (Michx.) Vahl — D 10406; D & P 10039
R. globosa (H.B.K.) В. & S. — D 11670; D 12709; D & P
10015; H 84; H 841
*R. globularis (Chapm.) Small — С 23966; D & P 10076;
G 7872; H 11; H 309; H 366
R. hirsuta Vahl — L & D 1666
*R. intermixta C. Wright — D 10924; D 10925; D 11601;
D 11609; D 10473A; D & P 10038; D & P 10132A;
G 7867
R. marisculus Nees — Н 291
R. micrantha Vahl
R. nervosa (Vahl) Boeck. — C 23532; С 24608; D 9829;
DEM 238; DEM 355; G 8496; H 293; K 524; Sp 1531;
Sp 1711
* R. oligantha Gray — Н 426
*R. plumosa Ell. — C 24036; DEM 651; D & P 10132A
* R. podosperma C. Wright — Н 196
R. pusilla (Sw.) Griseb.
R.radicans (Cham. & Schlecht.) H. Pfeiff. — C 23366;
C 23399; D 11131; G 7995
* R. rariflora. (Michx.) ЕП. — С 24036A
R. robusta (Kunth) Boeck. — DEM 617; H 99
R. rugosa (Vahl) Gale — D 11612; DEM 651
R. setacea (Berg.) Boeck. — C 23277; DEM 651; L&D
1681
R. tenuis Link — DEM 442
* R. torresiana Britt. & Standl. — Н 230; P 29676
R. watsoni (Britt.) Davidse

1975] Belize Monocotyledoneae — Spellman et al. 123

Scirpus chilensis Nees & Meyen
*Schoenus nigricans L. — Н 386; L & D 1675

Scleria areolata Lundell

8. bracteata Cav. — С 23294; С 23933; С 23964; С 24001;
D 12403; Р & L 12233; Н 156; L & D 1424

S. ciliata Michx. — C 23964; H 192

S. eggersiana Boeck. — С 24034 A

S. georgiana Core—C 23414; С 24080; D 10404; D
10717; Н 321; L & D 1660; L & D 1664; L & D 1676;
L & D 1684

S. hirtella Sw. — L & D 1679

S. interrupta L. Rich.

S. latifolia Sw. — С 24508; С 24836; DEM 529; D & L
12081

S. lithosperma (L.) Sw. — C 24921; D 11013; D 12753

S. macrophylla Pres]

S. microcarpa Nees — D 11467; H 494

S. micrococca (Liebm.) Steud. — H 191

S. mitis Berg. — DEM 470

S. muehlenbergii Steud. — DEM 231; H 455

S. pauciflora Muhl.

S. pinetorum Britt.

S. pterota Presl — C 23672; C 23936; C 23971; C 24034;
C 24536; С 24943; D 11012; D 11216; D 11380; DEM
545; G 8527; Spellman et al. 276

S. pterota Presl var. melaleuca (Reichenb.) Uittien — Н
259; K 503

S.secans (L.) Urban — C 24143; C 24475; G 8527; H
493 ; Spellman et al. 276

S. setacea Poir.

S. verticillata Muhl. ex Willd.

Dioscoreaceae

N.B. A goodly number of very recent collections have not
been identified (and hence not cited) due to the current
state of confusion in the genus.

Dioscorea bartlettii Morton — Sp 1895; Sp 1904
D. ђеттош ата Prain & Burkhill — Н 267

124 Ећодога [Vol. 77

D convolvulacea Cham. & Schlecht. — Sp 1903; Sp 1918
D. esurientium Uline

D. macrostachya Benth.

D. polygonoides H. & B. ex Willd.

D. spiculiflora Hemsl.

Eriocaulaceae
Eriocaulon fulginosum C. Wright — Н 888
E. schiedeanum Koern.
E. schippii Standl.
Paeplanthus gentlei Moldenke
P. lamarckii Kunth
Syngonanthus gracilis (Koern.) Ruhl
S. hondurensis Moldenke
S. lundellianus Moldenke
S. o’neillii Moldenke
Tonina fluviatilis Aubl.

Gramineae

Acroceras zizanioides (H.B.K.) Dandy

Andropogon bicornis L. — H 89

A. elliotti Chapm.

A. gerardi Vitman — Sp 1574; Sp 1644

A. glomeratus (Walt.) B.S.P.—C 24044; Lin 34; Lin
115; Lin 122; Sp & St 2210; Sp & St 2342; Sp & St
2376; Sp 2389

A. lateralis Nees

A. leucostachyus H.B.K. — С 24006; D 11506; D 12769;
G 7861; G 7866

A. selloanus (Hack.) Hack. — Н 144; Н 430

A. virginicus L. — Sp 1642

Anthephora hermaphrodita (L.) Kuntze

Aristida arizonica Vasey

A. capillacea Lam. — H 339

A. divaricata H. & B.

A. implexa Trin. — H 79

A. laxa Cav. —L & D 1446; L & D 1667

A. liebmannii Fourn.

A. longifolia Trin.

1975] Belize Monocotyledoneae — Spellman et al. 125

A. orizabensis Fourn. — D 9107; H 419
A. purpurascens Poir. — G 7862
A. recurvata H.B.K. — H 303; H 417
* A. setifolia H.B.K. — D 11604
A. tenuispica Hitche. — C 23965; C 24081; D 11518
A. ternipes Cav.
* A. vilfifolia Henr. — D & P 10051
Arundinella berteroniana (Schult.) Hitche. & Chase —
H 418
A. deppeana Nees — D & L 12229; Н 385
Axonopus aureus Beauv. — H 141
. ciliatifolius Swallen — H 83
. compressus (Sw.) Beauv. — H 441
. elongatus Swallen — H 490
. poiophyllus Chase — H 391
.purpusii (Mez) Chase
. rhizomatosus Swallen
Bambusa vulgaris Schrad. ex Wendl.
B. swalleniana McClure — С 23424
*Bothriochloa bladhii (Retz.) S. T. Blake — D 9921A
Brachiaria fasciculata (Sw.) Blake — С 23453; С 23456;
C 24045; C 24918; D 11551; DEM 401; К 421; Sp 1816
B. mutica (Forsk.) Stapf — К 430
Cenchrus brownii К. & S. — С 24980; К 435; Sp 1795
C. echinatus L. — C 24096
*C. incertus M.A. Curt. — Sp & St 2249; Sp & St 2364
Chloris barbata Sw. — C 24069
C. ciliata Sw.
C. radiata (L.) Sw. — C 23501; H 47
Cynodon dactylon (L.) Pers. — Dieck 184; К 697; Sp 244
*Dichanthium annulatum (Forsk.) Stapf — D 11181
Digitaria cayonensis Swallen
D. ciliaris (Retz.) Koel.
D. fuscescens (Presl) Henr. — D 11602
D. horizontalis Willd. — C 23306; C 24108; Sp 1721
D. insularis (L.) Mez ex Ekman — К 583
D. multiflora Swallen
Distichlis spicata (L.) Greene

> e e deu

126 Rhodora [Vol. 77

Echinochloa colonum (L.) Link — C 23311; D 11407
E. polystachya (H.B.K.) Hitche. — D 10745
*Echinolaena gracilis Swallen — D 10424B ; H 334

Eleusine indica (L.) Gaertn. — C 23305; C 23307; С
23311; C 23502; G 8088; К 422; D 9840A; D 98534;
D 10403; Sp 1815

Eragrostis acutiflora (H.B.K.) Nees — С 8092

E. bahiensis Schrad. — D 9105

E.ciliaris (L.) R. Br. — Н 453; К 576; Sp & St 2971;
Sp 2446

E.domingensis (Pers.) Steud. — D 11457; D & P 10034;
D & P 10036; D 11457; D 12149; Dieck 117; H 378;
Lin 73; Lin 116; Lin 117; Lin 119; Sp & St 2252; Sp
& St 2378; Sp 2405; Sp 2454

E. elliotii Wats.

E. hirsuta (Michx.) Nees — H 468

E. hypnoides (Lam.) B.S.P.

E.lugens Nees

E.maypurensis (H.B.K.) Steud.— D 10403; DEL
12147; D & L 12212; H 194

E. mexicana (Hornem.) Link

E. prolifera (Sw.) Steud.

Е. tenella (L.) R. & S.

Eriochloa punctata (L.) Desv. ex Hamilt.

Eriochrysis cayennensis Beauv.

Eustachys petraea (Sw.) Desv.— Lin 35; Lin 75; Lin
121; Lin 163; Sp & St 2362; Sp & St 2236; Sp & St
2250; Sp & St 2410; Sp 2437

Gymnopogon spicatus (Spreng.) Kuntze

Gynerium sagittatum (Aubl.) Beauv.

Hackelochloa granularis (L.) Kuntze

*Hemarthria cf. altissima (Poir.) Stapf. & С.Е. Hubb. —
Sp & St 2368

Homolepis aturensis (H.B.K.) Chase—C 23634; C
24060; H 101

Hymenchne amplexicaulis (Rudge) Nees — C 23432

Hypogynium virgatum (Desv.) Dandy —C 24192; D
11067; D & L 12364; DEM 613; H 271; L & D 1672

1975] Belize Monocotyledoneae — Spellman et al. 127

Ichnanthus axillaris (Nees) Hitchc. & Chase

1. brevivaginatus Swallen

I. lagotis (Trin.) Swallen — H 434

I. lanceolatus Scribn. & Smith — D 11379

I. mexicanus Fourn.

I. nemoralis (Schrad.) Hitchc. & Chase

I. pallens (Sw.) Munro ex Benth.

I. scaberrimus Swallen

I. standleyi Hitche.

I. tenuis (Presl) Hitche. & Chase

I. villosus Swallen

Imperata brasiliensis Trin. — К 563

I. contracta (H.B.K.) Hitchc.

Isachne sp. — C 24019

I. polygonoides (Lam.) Doell.

Ischaemum latifolium (Spreng.) Kunth — C 24184; C
24878; D 10466; D 11379; D & L 12192; L & D 1432

Lasiacis divaricata (L.) Hitche. var. divaricata — C
23649; C 23707; C 3924; C 24420; D 9957; D 10971;
D 11155; D & L 12030; D & L 12209; Sp 1713

L. grisebachii (Nash) Hitche. var. grisebachii — С
28884; C 23398; DEM 544 B; L & D 1483; Sp 1598;
Sp 1661

L. oaxacensis (Steud.) Hitch. var. oaxacensis — DEM
544 A; K 651; L & D 1588

L. procerrima (Hack Hitchc. — C 24416; D 11194; D
11416; D & L 12075

L. rugelii (Griseb.) Hitche. var. rugelii— C 24770; C
24990; D 12757; D & L 12284; H 266; P 29950

L. ruscifolia (H.B.K.) Hitche. var. ruscifolia — D 11005

L.sloanei (Griseb.) Hitche. — C 23912; К 588; Sp 1888

L.sorghoidea (Desv.) Hitche. & Chase var. sorghoidea
— К 697

Leersia hexandra Sw.

L. ligularis Trin. var. breviligularis (Prod.) Pyrah — C
23496

Leptochloa domingensis (Jacq.) Trin

L. filiformis (Lam.) Beauv.

128 Ећодога [Vol. 77

L. uninerva (Presl) Hitche. & Chase

L. virgata (L.) Beauv. — С 23454; К 405; Sp 1731

Leptocoryphium lanatum (H.B.K.) Nees — G 7869

Merostachys pauciflora Swallen

Mesosetum blakei Swallen — Н 346

M. filifolium F.T. Hubb. — D 10041; D & P 10020; G
7870; H 85

Olyra latifolia L. — С 23943; C 24671; DEM 373; G 7847;
Sp 1586; Sp 1659; Sp 1924; Sp 1965

О. yucatana Chase — С 23395; С 23478; С 23604; С
23894; L & D 1511

Oplismenus burmannii (Retz.) Beauv. — K 532

O. hirtellus (L.) Beauv.

O. setarius (Lam.) R. & S. — P 29842

Oryza alta Swallen

O. latifolia Desv.

О. sativa L.

Panicum agrostoides Spreng.

. albomarginatum Nash — Sp 1631

P. altum Hitche. & Chase

P. aquaticum Poir.

P. bartlettii Swallen — C 23332; C 23887; К 551

P. cayennense Lam.

P.

P.

P.

Y

cayoense Swallen
chamaelonche Trin.
cyanescens Nees ex Trin. — C 24185A ; G 8051; H 211;
H 344
P. erectifolium Nash — C 23259; C 24077; D 10716
P. frondescens Mey. — D 11469
P. furtivum Swallen — H 451
P. fusiforme Hitche. — H 308; DEM 449
P. ghiesbreghtii Fourn.
P. hirsutum Sw. — C 23483; C 23664; C 24285; K 461
P. hirtum Lam.
P. ichnanthoides Fourn.
P. lancearium Trin.
P. lanuginosum Ell. — H 10

1975] | Belize Monocotyledoneae — Spellman её al. 129

P. laxum Sw.—C 23537; С 23967; С 24025; С 24121;
C 24185; C 24860; C 24930; D 10755; DEM 273; K
698; Lin 74; Lin 123; H 442; H 458
. longiligulatum Nash
. lundellii Swallen
. maximum Jacq. — D 12717; K 662
. milleflorum Hitche. & Chase
. neuranthum Griseb.
. olivaceum Hitche. & Chase — D 9110
parvifolium Lam. — H 495
parviglume Hack.
. patulum (Scrib. & Merr.) Hitchc.
. pilosum Sw. — C 24880; Sp 1600; Sp 1693
. polycaulon Nash — H 9
polygonatum Schrad. — C 24516; C 24517; G 7956
. pulchellum Raddi — DEM 178
. rudgei R. & S. — DEM 454; H 332
. sellowii Nees — H 463
. sphaerocarpon ЕП.
. stagnatile Hitche. & Chase
. stenodoides F.T. Hubb.
. tenerum Beyr. ex Trin. — H 497
. trichanthum Nees
. trichoides Sw. — C 24867; G 7989; G 8450; K 455
. tuerkheimii Hack.
. virgatum L. — K 687
. viscidellum Scribn. — С 28891; C 23398A; С 240774;
H 258
P. wrightianum Scribn. — Sp 1514
Paspalum blodgettii Chapm. — C 23939; C 24046A; Н
448

P. botterii (Fourn.) Chase — С 23626; С 23668; С 23700;
C 24769

P. caespitosum Flügge — C 23626; C 23668; C 23771; D
10779

P. clavuliferum C. Wright
P. conjugatum Berg. var. conjugatum — С 23302; C

24098; C 24262; C 24698; DEM 176; H 285; Sp 1666

Му Му му ду Му Му Му му Му МУ МУ МУ МУ МУ МУ МУ МУ МУ Ба» У Ma M У

180 Rhodora [Vol. 77

P. conjugatum var. pubescens Doell — Sp 1599
P. conspersum Schrad. ex Schult.
P. convexum Н. & B. — Н 195
Р. corcovadense Raddi — G 8607; Н 440
P. coryphaeum Trin.
P. decumbens Sw. — C 24421; C 24520
Р. distichum L. — Sp 2465; Sp 2487; Sp 2509; Sp 2531;
Sp & St 2464
P. fasciculatum Willd. ex Flügge — C 24130; C 24135;
C 24183
*P. fimbriatum H.B.K.—C 23455; С 24612; DEM 242;
S 1722
P. humboldtianum Flügge — H 462
P. langei (Fourn.) Nash — C 23828
*P. larum Lam. — Н 442; Н 458; Sp & St 2276
* P. lineare Trin. — Sp 1518
P. millegrana Schrad. — C 24769; D 11291; D 12469
P. minus Fourn. — H 165; H 439
P. multicaule Poir. — Н 336
P. notatum Flügge
P. nutans Lam.
P. orbiculatum Poir.
P. paniculatum L. — C 23789; H 154
P. peckii F.T. Hubb.
P. pectinatum Nees — Н 80
P. pilosum Lam.
P. plicatulum Michx. — C 24033A ; C 24130
P. pulchellum Kunth — C 23997; C 24033; D 11092; G
1859; G 7864; Н 88; Sp 1641
P. repens Berg.
P. serpentinum Hochst. ex Steud. — C 24033; G 7863
P. vaginatum Sw. — C 24047; C 24099
P. virgatum L. — C 23309; C 23457; C 24107; D 98594;
D 11339; DEM 175; DEM 276; С 7957; Lin 36; Lin
72; Lin 120; Lin 204A; Sp 1732
Pennisetum complanatum (Nees) Hemsl. — DEM 114
P. nervosum (Nees) Trin.
P. setosum (Sw.) L. Rich.

1975] Belize Monocotyledoneae — Spellman et al. 131

Pharus latifolius L. — C 24542; Sp 1615; Sp 1602

P. parvifolius Nash — G 8175

Phragmites australis (Cav.) Trin. ex Steud. — D 105385;
Н 297 ; L & D 1440

Rhipidocladum bartlettii (McClure) McClure

Rhynchelytrum repens (Willd.) С.Е. Hubb. — L & D 1445

Rottboellia exaltata L.f.

Saccharum officinarum L.

Sacciolepis myuros (Lam.) Chase

Schizachyrium brevifolium Nees ex Kunth

S. hirtiflorum Nees

S. microstachyum (Desv.) Roseng., Arr. & Izag.—D
11206, D 12632; DEM 540

S. semiberbe Nees — L & D 1671

S. tenerum Nees

Setaria scandens Schrad. ex Schult. — Н 488

S. geniculata (Lam.) Beauv. — C 24051; С 24984; Sp
1692; Sp 1768

S. grisebachii (Steud.) Fourn.

S. tenacissima Schrad. ex Schult.

S. tenax (L. Rich.) Desv. — H 421

S. vulpiseta (Lam.) К. & S.

Sorgastrum setosum (Griseb.) Hitche. — Н 20

Sorghum halepense (L.) Pers. — DEM 650

Spartina spartinae (Trin. Merr. — D 11397; D & P
10035; G 8372; Lin 168; Sp & St 2206; Sp 2291; Sp
2553; Sp 2562

Sporobolus buckleyi Vasey

S. cubensis Hitehc. — Н 82

S. indicus (L.) К. Br. — Н 108; Sp 1730

S. jacquemontii Kunth — C 23308; C 23875; C 24163;
C 24992; D 9838A; D 9995A; D 11167; D 11408; D
12431

S.virginicus (L.) Kunth — C 24093; D 11475; Lin 4;
Lin 168; Lin 204; Sp 2182; Sp 2526; Sp & St 2234;
Sp & St 2275; Sp & St 2286; Sp & St 2382; Sp 2414;
Sp 2462; Sp 2508; Sp 2526; Sp 2573

Stenotaphrum secundatum (Walt.) Kuntze

132 Ећодога [Vol. 77

Streptochaeta sodiroana Hack.

Thrasya camplostachya (Hack.) Chase — Н 437; Н 614

T. thrasyoides (Trin.) Chase

Trachypogon angustifolius (H.B.K.) Nees ex Hack. — H
86; H 488

Т. montufari (H.B.K.) Nees

Triplasis purpurea (Walt.) Chapm.

Tripsacum dactyloides L.

T. latifolium Hitchc. — D & L 12211; Н 315

T. laxum Nash

Zea mays L.

Haemodoraceae
Xiphidium caeruleum. Aubl.

Hydrocharitaceae
Thalassia testudinum Konig

Hypoxidaceae
Hypoxis decumbens L. — H 153; Sp 1516; Sp 1629

Iridaceae
Cipura paludosa Aubl. — D 12801
Marica gracilis Herb.
Nemastylis bequaertii Standl.
N. sylvestris (Loes) Loes (= Eustylis sylvestris Loes)

—D&P 10064
Sisyrinchium tinctorium H.B.K. — DEM 216; P 29857
Juncaceae
*Juncus marginatus Rotsk. — D 10754
Liliaceae

Allium cepa L.

A. sativum L.

Dracaena americana J. D. Sm.—D 9967; D 12602; P
30028; Sp 1709A; Sp 1713A; Sp 1891

Echeandia sp. — Н 109

Yucca elephantipes Reg.

Marantaceae
Calathea albicans Brong.
C. altissima (P. & E.) Koern.

1975] | Belize Monocotyledoneae — Spellman et al. 133

C. barbillana Cut — С 7889; С 8179; G 8479

C. insignis Peters — C 24374; G 7943; G 8594

С. lutea (Aubl.) Mey. — С 24268; D 9877; DEM 538; С
8034; P 30229

С. macrosepala Schum.

С. microcephala (P. & E.) Koern. — C 24440; С 8181;
G 8588; V 635

Ischnosiphon morlaei (Eggers) Schum. — C 24400

Maranta arundinacea L. — C 23874; C 24444; D 9975;
DEM 131; G 1885; С 8148; К 678; Sp 1568

Myrosma guapilesense J. D. Sm.

Pleiostachya pruinosa (Reg.) Schum.

Stromanthe lutea (Jacq.) Eichl. — D 11311

Thalia geniculata L. — C 23991

Mayacaceae

Mayaca aubletii Michx.
M. fluviatilis Aubl. — P 29427

Musaceae

Heliconia acuminata L. Rich.

H. aurantiaca Ghiesbr.

H. bourgaeana O.G. Peters.

H. champneiana Griggs

H. collinsiana Griggs

H. hirsuta L.f. var. rubriflora R.R. Sm.

H. latispatha Benth. — С 23305; С 24903; DEM 581; Sp
1681; P 29923

H. mariae Hook.f.

H. pendula Wawra

H. psittacorum L.f. — Sp 1963

H. schiedeana Kl. — D 10815; DEM 235

H. spissa Griggs — P 30154

H. cf. subulata R. & P. — Sp 1603

H. vaginalis Benth. — D 9870; G 8144; G 8248; V 599

H. wagneriana O.G. Peters. — C 24273

Naiadaceae
Nais guadalupensis (Spreng.) Morong

184 Rhodora

Orchidaceae

[Vol. 77

Bletia purpurea (Lam.) DC — D 10464; С 7606; H 498

Brassavola nodosa (L.) Lindl. — H 411
Campylocentrum micranthum (Lindl.) Rolfe
C. sullivanii Fawc. & Rendle

Catasetum integerrimum Hook. — H 126
Cattleya skinneri Batem. — H 50

Chysis bractescens Lindl.

Coryanthes picturata Reichenb. f. — P 30276
C. speciosa Hook.

Corymborchis flava Kuntze

Cranichis sylvatica A. Rich. & Gal. — H 289
Cycnoches chlorochilon КЛ.

С. warscewiczii Reichenb. f. — Sp 1952
Diacrium bidentatum (Lindl.) Hemsl.

Dicrium bilamellatum (Reichenb. f.) Hemsl. (= Caular-

thron bilamellatum (Reichenb. f.) Schult.)
Dichaea tuerckheimii Schltr.
Elleanthus linifolius Presl
Encyclia bractescens (Lindl.) Hoehne
Е. cochleata (L.) Dressler — С 7818; Sp 1581
Epidendrum alatum Batem.
.anceps Jacq. — P 30091
. belizense Reichenb. f.
. boothianum Lindl.
. boothii (L.) L.O. Wms.
. ciliare L.
. clowesii Batem. ex Lindl.
cochleatum L. — G 7818; Н 127; P 29905
. condylochilum Lehm. & Kranzl
difforme Jacq.
imatophyllum Lindl. — P 30275
isomerum Schltr. — P 29774
. nocturnum Jacq.
paniculatum R. & P.
. polyanthum Lindl.
. polybulbon Sw.
. pygmaeum Hook.

Ej by by ty р рр Ыр?

1975] Belize Monocotyledoneae — Spellman et al. 135

E. radiatum Lindl.

E. rigidum Jacq. — Р 29597

E. stamfordianum Batem.

E. stenopetalum Hook.

E. verrucosum Sw. — P 29488

Erythrodes purpurea Ames

Eulophia alata (L.) Fawe. & Rendle

E. longifolia (Kunth) Schltr.

Galeandra batemanii Rolfe

G. baueri Lindl.

Gongora maculata Lindl.

G. cf. quinquenervis R. & P. — p 9939; С 8039; С 8184

Habenaria lankesteri Ames

Н. mesodactyla Griseb.

H. odontopetala Reichenb. f.

H. cf. pauciflora (Lindl.) Reichenb. f.

H. repens Nutt.

H. setifera Lindl.

Ionopsis utricularioides (Sw.) Lindl.

Isochilus crassiflorus A. Rich. & Gal.

I. linearis (Јаса.) К. Br. var. carnosiflorus (Lindl.) Correll

Laelia digbyana Benth.

L. tibicinis (Batem. ex Lindl.) L.O. Wms. — H 327

Lycaste cochleatam Lindl. ex Paxt.

Macradenia brassavolae Reichenb. f.

Masdevallia tubuliflora Ames

Mavillaria crassifolia (Lindl.) Reichenb. f.

M. densa Lindl.

M. friedrichsthalii Reichenb. f. — Р 29774; P 29827; P
29687

М. macleei Batem

M. ringens Reichenb. f.

M. tenuifolia Lindl. — P 50188

M. uncata Lindl. — P 29686

Mormodes buccinator Lindl.

M. ringens (Lindl.) Schlecht.

Notylia barkeri Lindl.

N. trisepala Lindl & Paxt.

186 Rhodora [Vol. 77

Oncidium ascendens Lindl.

O. carthagenense (Jacq.) Sw. — H 132

O. cebolleta (Jacq.) Sw. — H 137

O. ensatum Lindl. — DEM 461

O. luridum Lindl.

О. pusillum (L.) Reichenb. f. — D 9938; H 410

О. sphacelatum Lindl. — H 396?

Ornithocephalus inflexus Lindl.

O. pottsiae Wats.

Pleurothallis blaisdellii Wats.

P. brighamii Wats. — C 24156

P. marginata Lindl. — H 136

P. racemiflora (Sw.) Lindl.

P. yucatanensis Ames & Schweinf.

Polystachya cerea Lindl.

P. clavata Lindl. — H 128?

P. luteola (Sw.) Hook.

P. masayensis Reichenb. f.

P. minor Кале. & Rendle

Ponera striata — Lindl.

Rhyncholaelia digbyana (Lindl.) Schltr. — Н 133

Scaphyglottis behrii (Reichenb. f.) Benth & Hook. ex
Hemsl.

S. cuneata Schltr.

S. wereklei Schultz var. major Schweinf.

Sobralia decora Batem.

S. fragrans Lindl.

S. macrantha Lindl. — DEM 460; Н 51

Spiranthes acaulis (J. E. Sm.) Cogn.

S. graminea Lindl. — Sp 1623?

S. lanceolatus Fawc. & Rendle

S. orchidioides (Sw.) A. Rich. — DEM 537

S. squamulosus Fawc. & Rendle

S. tortilis (Sw.) L. Rich. — H 220

Stelis ciliaris Lindl.

Stenorrhynchos lanceolatus (Aubl. Griseb.— Н 404

Trigonidium egertonianum Batem. ex Lindl.— P 30222

Vanilla fragrans (Salisb.) Ames

1975] | Belize Monocotyledoneae — Spellman et al. 137

Palmae
Acrocomia belizensis Bailey
A. mexicana Karw. ex Mart.
A. panamensis Bailey — C 23916; Sp 1832
Asterogyne martiana Wendl. ex Hook. f. — С 24524
Asterocaryum mexicanum Liebm. — С 23735; C 24165;
C 24455; D 10906; G 8143

Bactris major Jacq. (including B. balanoides (Oerst.)
Wendl.) — C 28487

B. trichophylla Burret — С 23972; C 24340; C 24788; D
11101; Sp 1686

Calyptrogyne | donnell-smithii (Dammer) Burret — C
24381

Chamaedorea adscendens (Dammer) Burret

C. arenbergiana Wendl.

C. elegans Mart.

C. ernesti-angusti Wendl. — С 23335; С 24306; D 10148;
D & L 10907; G 7735; G 7831; G 7980; G 8129; Sp
2005
. erumpens Moore
. geonomaeformis Wendl.

. lindeniana Wendl. sp. vel aff. — С 23318; C 24296;
C 24382; С 7734

. neurochlamys Burret

. oblongata Mart. — G 7828

. schippii Burret — G 7831A

C. wendlandiana Hemsl. — C 28816; C 24564; D 11107;

D 11441; G 7854; G 7976; G 8309; G 8599
Cocos nucifera L.

Crysophila argentea Bartlett — C 23789A; D & L 12251;

L & D 1621; Sp 1892
C. warscewiczii (Wendl.) Bartlett, sp. vel aff. — С 23605
Desmoncus ferox Bartlett — C 23338; С 24746; D 11352
D. leiorhachis Burret
D. quasillarius Bartlett
D. schippii Burret — D 10958; G 8035; Sp 1685; Sp 1882
Euterpe macrospadix Oerst.

AAQ AQQ

188 Ећодога [Vol. 77

Geonoma binerva Oerst.

G. longipetiolata Oerst.

С. охусатра Mart. (including G. mexicana Liebm.) —
C 24468; D 10921; D 11312; D & L 12096; G 7987

Opisandra maya Cook.

Orbignya cohune (Mart.) Dahlg. ex Standl. — С 23444;
Sp 1687

Paurotis schippii Burret

P. wrightii (Griseb. & Wendl.) Britt. ex Britt. & Schafer
— С 23267; С 23953; Р 10478; Р 10977; Dwyer et al.
365; G 8055; Sp 1617; Sp & St 2356

Reinhartia gracilis (Wendl.) Burret var. gracilis

R. gracilis var. gracilior (Burret) H.E. Moore

R. latisecta (Wendl.) Burret

Roystonea oleracea (Jacq.) Cook

R. regia (H.B.K.) Cook

Sabal mayarum Bartlett

S. morrisiana Bartlett — Sp 1893

S. nematoclada Burret

Scheelea sp. — C 23436; G 8596

Schippia concolor Burret — C 24814; G 8422

Synnechanthus fibrosus Wendl. — С 24527; С 24556; Р
11309; Sp 2015A

Thrinax radiata Lodd. — Sp & St 2145; Sp & St 2222;
Sp & St 2298; Sp & St 2357; Sp & St 2417; Sp & St
2476; Sp & St 2557. We follow Reed (Principes 18:
39. 1974) in the usage of this name in preference to
T. parviflora Sw. and T. floridana Sarg.

Pontederiaceae
Eichornia crassipes (Mart.) Solms
Heteranthera reniformis R. & P. — P 29942
Pontederia lanceolata Nutt. — P 30311
P. rotundifolia L.

Potamogetonaceae
Potamogeton lucens L.
P. pectinatus L.
Ruppia maritima L.

1975] Belize Monocotyledoneae — Spellman et al. 139

Smilacaceae
Smilax domingensis Willd. — K 488

. gentlei Lundell

. lanceolata L.

. lundellii Killip & Morton

. mexicana Griseb. ex Kunth

. mollis Н. & B. — G 8492; Sp 1663; Sp 1878

. munda Killip & Morton

. ornata Lem.

. spinosa Mill. — DEM 1913; G 8043

S. velutina Killip & Morton — D & L 12210; D & L 12275;
H 436; P 29626; P 29765

Triuridaceae
Sciaphila picta Miers

о о и л л о о 0

Турћасеае
Typha angustifolia L.
Xvridaceae
Xyris ambigua Bey. ex Kunth — Sp 1627
X. caroliniana Walt. — Н 172
X. jupicai L. Rich. — С 24021; DEM 32; D & P 10065;
Sor 7091; Sor 715
X. navicularis Griseb. — D 10407
X. smalliana Nash — C 24015; D 10410; D 11610; D 11068
X. subnavicularis Malme — Н 199?
X. subulata R. & P. — Dieck 101

Zingiberaceae

Alpinia speciosa (Wendl.) Schum.

Costus congestus Rowlee — Sp 1966; Sp 2001

C. pulverulentus Presl — D & L 12098; G 7731; С 7850;
G 7910; G 7919; G 8002; G 8115

C. cf. ruber Griseb. — Sp 1683; Sp 1911; Sp 1917

C. spicatus (Jacq.) Sw.

C. villosissimus Jacq. — DEM 185; Sp 1684

Hedychium coronarium Koenig — D 11195; С 8233

Renealmia aromatica ( Aubl.) Griseb.

R. exaltata L.f. — D 9931; L & D 1458

R. mexicana Kl.

140 Rhodora [Vol. 77

LITERATURE CITED

BARTLETT, H. H. 1935. A method of procedure of field work in
tropical American phytogeography based on a botanical recon-
naissance in parts of British Honduras and the Peten forest
of Guatemala. Misc. Paper I. Publ. Carnegie Inst. Wash. 461:
1-25.

BLAKE, S. F. 1917. Descriptions of new spermatophytes chiefly
from the collections of Professor M. E. Peck in British Hon-
duras. Contr. Gray Herb. 52: 59-106.

FURLEY, Р. A. (editor). 1972. University of Edinburgh Expe-
dition to Central America 1970. General Report, vol. 1.

HaArLcRow, M., & M. L. HarcRow. 1967. Orchids of Belize. 151 pp.
The Government Printer, Belize.

HUBBARD, Е. T. 1913. On the Gramineae collected by Professor
M. E. Peck in British Honduras. 1905-1907. Proc. Am. Acad.
Arts 49: 493-502.

KESSLER, S. E. C. Е. KIENLE, & J. Н. BATESON. 1974. Tectonic
significance of intrusive rocks in the Maya Mountains, British
Honduras. Bull. Geogr. Soc. Am. 85: 549-552.

LowDpEN, R. M. 1970. William A. Schipp’s botanical explorations in
the Stann Creek and Toledo districts, British Honduras (1929-
1935). Taxon 19: 831-861.

LUNDELL, C. L. 1940. The 1936 Michigan-Carnegie Botanical Expe-
dition. Misc. Paper XIV. Publ. Сатпеоле Inst. Wash. 522: 1-57.

1945. The vegetation and natural resources of British
Honduras, pp. 270-273 in F. Verdoorn (editor), Plants and Plant
Science in Latin America. Waltham, Mass.

1960. Plantae Mayanae — I: Notes on collections from
the lowlands of Guatemala. Wrightia 2: 49-63.

Publ. Carnegie Inst. Wash. 1936. Botany of the Maya Area. Misc.
Papers I-XIII. Publ. 461. Washington, D. C.

. 1940. Botany of the Maya Area. Mise. Papers XIV-
XXI. Publ. 522. Washington, D. C.

ScurPP, W. A. 1933-1934. Flora of British Honduras. Price List
of Seeds & Herbarium Material from William А. Schipp, Stann
Creek, British Honduras.

SPRAGUF, T. A, & І. А. M. RILEY. 1924. Materials for a Flora
of British Honduras: I. Bull. Misc. Inform. 1924: 1-20.

STANDLEY, P. C, & S. J. Recorp. 1936. The Forests and Flora of
British Honduras. Field Mus. Nat. Hist. Bot. Ser. 12: 1-432.

DAVID L. SPELLMAN GERRIT DAVIDSE
JOHN D. DWYER MISSOURI BOTANICAL GARDEN
ST. LOUIS UNIVERSITY ST. LOUIS, MISSOURI 63110.

ST. LOUIS, MISSOURI 63110.

NOTES ON THE LEGUMINOSAE II.
FACULTATIVE DWARFISM IN
CROTALARIA SAGITTALIS L.'

DONALD К. WINDLER*

In his study of the North American species of Crotalaria,
based on herbarium specimens, Senn (1939) described a
dwarf Crotalaria sagittalis L. from rocky, xeric habitats in
the American Southwest and Mexico to which he gave the
name, variety Blumeriana. During the fall of 1967, while
studying unifoliolate species of Crotalaria in North Amer-
ica (1970), I was able to observe a number of Mexican
populations of this dwarf form in the field. In most loca-
tions visited, larger plants of C. sagittalis were often found
growing in more favorable areas, near populations of vari-
ety Blumeriana. The consistency of this association seemed
to place the status of the variety in question.

In the summer of 1968 plants reared from seeds of two
“Blumeriana” populations (Windler & Windler 2950 — State
of Jalisco; Windler & Windler 2981 — State of Oaxaca)
were planted along with representatives of other species in
a common garden plot at the North Carolina Botanical
Gardens. A total of 47 plants, progeny of eleven seed par-
ents from the two field populations, were grown. All plants
grown in the garden plots were substantially larger than
their field grown parents (see Figures 1 & 2). Plants from
the field populations rarely grew to one decimeter in height
and were mostly less than 0.7 dm. tall. Garden grown off-
spring of these plants grew to heights of three to seven
decimeters. In addition to stem length, substantial increases
were also observed in fruit size and number of fruits per
plant. Garden plants from the dwarf seed parents com-

1A portion of a thesis submitted in partial fulfillment for the Ph.D.
at the University of North Carolina at Chapel Hill.

2Contribution number 4 from the Towson State College Herbarium,
Baltimore, Maryland 21204. Cost of publication of illustrations
assisted by 'the Towson State College Faculty Research Fund.

141

142 Ећодога [Vol. 77

BOTANICAL COLLECTIONS OF
DONALD R WINDLER

ETET P

Figure 1. A. Herbarium specimen of Crotalaria sagittalis “var.
Blümeriana”. B. A branch of a specimen grown from seed produced

by the plant shown in Figure 1А.

1975] Crotalaria — Windler 143

Figure 2. A. Herbarium specimen of Crotalaria sagittalis “var.
blumeriana". B. А branch of a specimen grown from seed produced
by the plant shown in Figure 2A,

144 Ећодога [Vol. 77

pared favorably in general size with those from non-dwarf
seed parents.

During preparation for the garden plantings, seeds were
scratched and soaked overnight before being placed in the
soil. At planting time, the 47 seeds from the dwarf popula-
tions were swollen and the radical was strongly exserted
from the seed coat. In contrast, among the more than 1,000
seeds from 62 other North American Crotalaria populations,
all were swollen at planting time but none showed an ex-
serted radicle.

DISCUSSION

Based upon the above information, the plants that Senn
recognized as var. Blumeriana appear to be ecophenes of
Mexican Crotalaria sagittalis ecotypes that are adapted to
xeric, rocky conditions where moisture becomes available
but is soon depleted. Under these conditions organisms
must produce seeds which germinate quickly, and the re-
sulting plants must quickly develop to a physiological stage
conducive to flowering and fruit set if the population is to
continue to exist. Plants of most other Crotalaria popula-
tions in North America have not evolved under similar
pressures and have not developed the quick germination
response shown by the Mexican dwarf plants.

The ability to germinate quickly and produce seed at a
small plant size appear to be genetically controlled, but the
latter characteristic is not expressed unless the plant is
developing in a xeric situation. Consequently no formal
recognition should be accorded to the epithet Blwmeriana.

LITERATURE CITED

SENN, Н. 1939. The North American Species of Crotalaria. Rho-
dora 41: 317-366.

WINDLER, D. R. 1967. Notes on the Leguminosae I (Neptunia and
Cassia). The Southwestern Naturalist 12: 336-337.

1970. Systematic studies in Crotalaria sagittalis and

related species in North America (Leguminosae). Unpublished
dissertation. University of North Carolina at Chapel Hill.

BIOLOGY DEPARTMENT
TOWSON STATE COLLEGE
BALTIMORE, MD. 21204

PECTIS HUMIFUSA NEW TO THE FLORA
OF THE UNITED STATES

During research preparatory to a revision of section
Pectis of the genus Pectis L. (Compositae), a new record
for the flora of the United States was discovered. A speci-
men of Pectis humifusa Swartz, a taxon distributed among
several islands of the Caribbean region, was found to have
been collected in southern Florida. This first mainland
record for Р. humifusa is a significant range extension,
since the closest previous collection site is over 900 miles
away in the Dominican Republic. The previously known
range for this species extends from Tobago north through
the Lesser Antilles to Puerto Rico and the Dominican Re-
public. The following are the collection data for this new
record: UNITED STATES: Florida, Collier Co., Naples, in
white sand along route 41, the Tamiami Trail, 5 mi. SE of
town, 24 Mar 1956, J. A. Churchill s.n. (MSC).

Three other species of Pectis are known to occur in Flor-
ida: P. leptocephala (Cass.) Urban, P. linearifolia Urban
and P. prostrata Cav. All are low herbs with yellow, five-
rayed heads, as is Р. humifusa. However, of the three pre-
viously known taxa, only P. prostrata bears a close resem-
blance to Р. humifusa. The two taxa are similar in their
prostrate habit, relatively broad leaves (usually more than
2 mm. wide) and very short-peduncled or sessile heads.
The two species differ in several respects, however, and are
easy to distinguish. Both species tend to root at the nodes,
but the tendency is much more developed in P. humifusa.
The foliage of Р. humifusa is blue-green and the leaves are
mostly 4-12 mm. long and 1.5-4 mm. wide, ranging from
oblanceolate to obovate. The leaves of P. prostrata are more
olive-green, commonly exceed 20 mm. in length and are at
most oblanceolate. The heads of P. humifusa are more open
at anthesis than those of P. prostrata and are commonly

145

146 Ећодога [Vol. 77

campanulate rather than cylindric or urceolate. The in-
volucral bracts of P. humifusa are obovate, obtuse, and
keeled only part of their length whereas those of P. pro-
strata are usually lanceolate or oblanceolate, acute, and are
keeled nearly to the tips. Pectis humifusa is a plant of
sandy areas, and P. prostrata occurs in a variety of habi-
tats and is a common roadside weed in much of Latin
America.

DAVID J. KEIL
DEPARTMENT OF BOTANY
THE OHIO STATE UNIVERSITY
COLUMBUS, OHIO 43210

MARITIME AND MARINE LICHENS
FROM NAHANT' °

E. E. WEBBER

The Marine Science Institute of Northeastern University,
Nahant, Massachusetts, became operational in 1966; its first
classes were offered during the summer of 1967. At that
time, Dr. Ivan Mackenzie Lamb of the Farlow Herbarium
made an initial collection (with determinations) of the
lichens common to the environs of the laboratory. The
majority of lichens collected encrust the extensive outcrops
of rock which essentially encircle this research facility.

LICHEN DETERMINATIONS

A. Below are listed those maritime lichens occurring on
the metamorphic rocks above the sea:

Acarospora fuscata (Schrad.) Arn.
Caloplaca scopularis (Nyl.) Lett.
(Sect. Gasparrinia)
Candelariella aurella (Hoffm.) Zahlbr.
Lecanora dispersa (Pers.) Somm.
Lecanora (Aspicilia) gibbosula Mogn.
Lecanora rubina (Vill.) Ach.
(Sect. Placodium)
Parmelia conspersa (Ach.) Ach.
(Sect. Xanthoparmelia)
Rhizocarpon grande (Flörke ex Flot.) Arn.
Rhizocarpon distinctum Th. Fr.
Dimelaena oreina (Ach.) Norm.

1This list is published with the permission of Dr. I. M. Lamb,
whose extreme generosity in this regard is sincerely appreciated.

?Contribution No. 25 from the Marine Science Institute, Nahant,
Mass. 01908.

147

148 Ећодога [Vol. 77

B. The following marine lichens appear as black, tar-like
coatings on rock; the first is found in the spray zone,
while the second is abundant in the intertidal:

Verrucaria erichsenii Zach.
Verrucaria mucosa Wahlenb. ex Ach.

The above twelve lichens were collected during the course
of one afternoon (Lamb, personal communication). Un-
doubtedly, additional species remain to be discovered.

DEPT. OF BIOLOGY
KEUKA COLLEGE
KEUKA PARK, N.Y. 14478

PHYCOLOGICAL STUDIES FROM THE
MARINE SCIENCE INSTITUTE
NAHANT, MASSACHUSETTS:

1. INTRODUCTION AND PRELIMINARY
TABULATION OF SPECIES AT NAHANT

E. E. WEBBER

This is the initial paper in a series dealing with the at-
tached marine algae at Nahant. These observations and
studies began in the summer of 1968, and are continuing.

The present paper summarizes the history of marine
phycology in New England; this summary is followed by
citations emphasizing recent research dealing with the local
marine algal vegetation. A tabulation of the species found
in the vicinity of Nahant is then presented. No attempt is
made at this time to discuss the plants listed since discus-
sions of specific taxa will constitute the topics of future
papers in this series,

INTRODUCTION

Published accounts of the marine algal vegetation of New
England began essentially in the late 1800’s. The Nahant
area figured prominently in these studies. The efforts of
such local investigators as Farlow, Collins, Davis, and
others (see Taylor, 1957 for a thorough bibliography),
extending from about 1880 to the early part of this century,
resulted in the cataloging of the seaweeds common along
the open coast. Early field work emphasized the macroscopic
algae of the littoral and upper littoral zones of the shore-
line. Study of New England marine algae progressed
through the 1920’s and 1930’s, culminating in the appear-

!Contribution No. 24 from the Marine Science Institute. I wish
to thank Dr. A. C. Mathieson for his valuable suggestions during the
writing of this paper.

149

150 Rhodora [Vol. 77

ance of Taylor’s (1937) phycological survey of the north-
eastern coast of North America. Twenty years later, recog-
nizing the need for both an updating of information and the
inclusion of new data, Taylor published a revision (1957)
of his volume. This major contribution is still the basic
reference for phycologists interested in the benthic marine
algae of our northeastern coast.

From 1957 to the present, marine phycological studies in
New England have progressed rapidly along several fronts.
The utilization of SCUBA diving equipment has enabled
the marine phycologist to study more accurately the hereto-
fore inaccessible sublittoral zone; in addition, this technique
has permitted field research on a year-round basis. The
following may be cited as recent examples of SCUBA-aided
research in New England waters: Lamb & Zimmermann,
1964; Hehre & Mathieson, 1970; Sears & Wilce, 1970;
Hehre et al., 1970; Adey, 1970; Wilce, 1970; and Mathieson,
et al., Nova Hedwigia, in press.

Severa’ important phycological discoveries have resulted
from recent intensive field studies. For example, the ap-
pearance of Lomentaria clavellosa in North America was
reported from Massachusetts by Wilce & Lee, 1964; more
recently, this species has been found in New Hampshire
(Hehre, 1972).

Also from New Hampshire has come the first record of
the occurrence of Halicystis ovalis along the northeastern
coast of North America (Mathieson & Burns, 1970). This
raises the larger question: is there a Halicystis-Derbesia
alternation occurring in nature among the New England
populations of these algae, for Derbesia is common to the
sublittoral of our area (Sears & Wilce, 1970) ?

In addition, year-round field studies have established the
presence of two rare and monotypic brown algal crusts
(Petroderma maculiforme and Porterinema fluviatile)
known previously only from the Arctic, and now recorded
for New England from Ipswich, Massachusetts (Wilce,
Webber, & Sears, 1970). More specifically, this publication

1975] Nahant Algae — Webber 151

presents the first report of Petroderma in Massachusetts,
as well as the first record of the occurrence of Porterinema
in North America. This latter taxon had been described
previously only from the Baltic. For detailed discussions
of the morphology, cytology, reproductive periodicities, and
geographical distributions of these two “fleshy crusts”, the
reader is referred to the above paper. A comprehensive
updating and summation of both brown and red algal
“fleshy crusts” in New England has been presented by
Wilce (1971).

Complementing the renewed vigor in research on the
algae of the sublittoral zone, recent attention also has been
focused on estuaries and their adjacent salt marshes. In
New England, particularly, these ecosystems represent
promising areas of study for the marine phycologist. Pub-
lished accounts of the benthic algal vegetation of our salt
marshes are few; the majority of species common to such
interesting habitats are microscopic, and many, undoubt-
edly, have been overlooked by previous investigators. Pub-
lications relative to New England salt marsh algae are,
therefore, recent in their appearance. While such publica-
tions are essentially ecologically oriented, they also deal
with problems in systematics, life history studies, and
polymorphisms of certain of the species encountered. The
following papers may be mentioned as examples of recent
studies centered about salt marsh ecosystems: Blum, 1960,
1968; Blum & Conover, 1953; Blum & Wilce, 1958; Webber,
1967, 1968, 1971; Webber & Wilce, 1971, 1972; and Mathie-
son & Fralick, 1972. The works of Drouet (1968) and
Blum (1972), although not dealing entirely with New Eng-
land salt marsh algae, do contain some phycological data
applicable to this area.

An even greater void in our understanding of the attached
marine algae relates to the autecology of particular species.
To date, comparatively few autecological studies have been
undertaken. Examples of New England seaweeds in both
field and laboratory settings include the following: Kan-
wisher, 1957, 1966; Conover, 1958; Mathieson & Burns,

152 Rhodora [Vol. 77

1971: Burns & Mathieson, 1972a, 1972b; Fralick & Mathie-
son, 1972, 1973; and Jordan & Vadas, 1972.

PRELIMINARY SPECIES TABULATION

The species listed in the following tabulation have been
collected and examined by me. I acknowledge the advice of
Dr. F. Drouet concerning several of the bluegreen algal
determinations; Dr. C. van den Hoek for assistance with
species of Cladophora; Dr. H. W. Johansen for advice on
the Coralline algae; and Dr. R. T. Wilce for confirmation
of the Peyssonnelia determination.

The systematic treatment of the Cyanophycophyta fol-
lows that of Drouet and Daily (1956) ; Drouet (1962, 1963,
1964, 1968) ; Fan (1956); and Tilden (1910). The treat-
ments of the Chlorophycophyta, Phaeophycophyta, and Rho-
dophycophyta all essentially follow Taylor (1957), with the
following exceptions: van den Hoek (1963) was followed
for Cladophora, while the recent studies of Bliding (1963,
1968) were consulted for the Ulvales; species designations
in Ectocarpus are after Rosenvinge & Lund (1941) and
Cardinal (1964), Petroderma folows Waern (1952), and
Powel's study (1957) was used for Fucus, and that of
Wilce (1965) for Laminaria. The Chrysophycophyta (ex-
cluding diatoms) are treated in this paper following Taylor
(1957) for the Vaucheriales, and Parke and Dixon (1968)
for the Ochromonadales.

Cyanophycophyta

Anabaena torulosa (Carm.) Lag.
Calothrix crustacea (Thur.) Fan
Coccochloris stagnina Spreng.
Entophysalis deusta (Menegh.) Dr. et D.
Lyngbya lutea (Ag.) Gom.

Microcoleus chthonoplastes Thur.

M. lyngbyaceus (Kütz.) Crouan

М. vaginatus (Vaucher) Gom.
Nodularia harveyana (Thwaites) Thur.
N. spumigena Mert.

1975] Nahant Algae — Webber 153

Chlorophycophyta

Blidingia minima (Nag. ex Kütz.) Kylin
Brachiomonas sp.

Bryopsis plumosa (Huds.) C. Ag.
Capsosiphon fulvescens (C. Ag.) S. et G.
Chaetomorpha linum (Muell.) Kitz.

C. melagonium (Weber & Mohr) Kiitz.
Chlorochytrium moorei Gardner
Cladophora albida (Huds.) Kutz.

Codiolum gregarium A. Br. f. intermedium (Fos.) Collins
Enteromorpha flexuosa (Wulfen ex Roth) J. Ag.
E. intestinalis (L.) Link

E.intestinalis (L.) Link f. clavata J. Ag.

Е. linza (L.) J. Ag.

E. linza (L.) J. Ag. var. oblanceolata Doty
Percursaria percursa (C. Ag.) Bory
Prasinocladus lubricus Kuck.

Prasiola stipitata Suhr

Rhizoclonium tortuosum Kitz.
Spongomorpha arcta (Dillw.) Kütz.

S. lanosa (Roth) Kitz.

S. spinescens Kütz.

Ulothrix flacca (Dillw.) Thur.

Ulva gigantea (Kütz.) Bliding

Phaeophycophyta

Agarum cribosum (Mert.) Bory
Alaria esculenta (L.) Grev.
Ascophyllum nodosum (L.) Le Jolis
Asperococcus echinatus (Mert.) Grev.
Chorda filum (L.) Stack.

Chorda tomentosa Lyngbye

Chordaria flagelliformis (Müll.) C. Ag.
Corynophlaea sp.

Desmarestia aculeata (L.) Lam.

D. aculeata (L.) Lam. var. attenuata Taylor
D. viridis (Müll.) Lam.

154 Rhodora [Vol. 77

Dictyosiphon chordaria Aresch.

D. foeniculaceus (Huds.) Grev.

Ectocarpus confervoides var. confervoides (Roth) Kjellm.
E. confervoides var. siliculosus (Dillw.) Kjellm.
E. fasciculatus (Griff.) Harv.

Elachistea fucicola (Vell.) Aresch.

Fucus distichus L. ssp. edentatus (de la Pyl.) Powell
F. distichus L. ssp. evanescens (C. Ag.) Powell
Е. vesiculosus L.

Laminaria saccharina (L.) Lam. sensu Wilce

L. digitata (Huds.) Lam.

Leathesia difformis (L.) Aresch.

Petalonia fascia (Muell.) Kuntze

Petroderma maculiforme (Woll.) Kuck.
Pylaiella littoralis (L.) Kjellm.

Ralfsia clavata (Harv. in Hook) Crouan

R. fungiformis (Gunn.) S. et. С.

R. verrucosa (Aresch.) J. Ag.

Scytosiphon lomentaria (Lyngbye) Link
Sphacelaria cirrhosa (Roth) C. Ag.
Streblonema aecidioides (Rosenv.) Fos.

Rhodophycophyta

Acrochaetium sp.

Agardhiella tenera (J. Ag.) Schmitz

Ahnfeltia plicata (Huds.) Fries

Antithamnion floccosum (Müll.) Kleen

Asparagopsis hamifera (Hariot) Okamura

Bangia fuscopurpurea (Dillw.) Lyngbye

Ceramium deslongschampii Chauvin var. Hooperi (Harv.)
Taylor

C. rubriforme Kylin

C. rubrum (Huds.) J. Ag.

Chondrus crispus Stack.

Corallina officianalis L.

Clathromorphum circumscriptum (Strom.) Fos.

Cystoclonium purpureum (Huds.) Batters

1975] Nahant Algae — Webber 155

C. purpureum (Huds.) Batters var. cirrhosum Harv.
C. purpureum (Huds.) Batters f. stellatum Collins
Dumontia incrassata (Müll.) Lam.

Euthora cristata (C. Ag.) J. Ag.

Gigartina stellata (Stack.) Batters
Gloiosiphonia capillaris (Huds.) Carm.
Gracilaria verrucosa (Huds.) Papenfuss
Hildenbrandia prototypus Nardo
Lithothamnium sp.

Membranoptera alata (Huds.) Stack.
Petrocelis middendorfii (Rup.) Kjellm.
Peyssonnelia sp.

Phycodrys rubens (L.) Batters

Phyllophora brodiaet (Turner) J. Ag.
Phymatolithon laevigatum (Fos.) Fos.
Phymatolithon sp.

Polyides caprinus (Gunn.) Papenfuss
Polysiphonia lanosa (L.) Tandy

P. nigrescens (Huds.) Grev.

P. novae-angliae Taylor

P. urceolata (Lightfoot) Grev.

Porphyra umbilicalis (L.) J. Ag.

Ptilota serrata Kütz.

Rhodocorton purpureum (Lightfoot) Rosenv.
Rhodymenia palmata (L.) Grev.

Chrysophycophyta

Apistonema — like plants

Ки тета sp. (?maritima (Anand) Parke)
Urococcus foslieanus Hanser.

Vaucheria compacta (Collins) Collins

LITERATURE CITED

ADEY, W. H. 1970. The crustose corallines of the northwestern
North Atlantic, including Lithothamnium lemoinae n. sp. Jour.
Phycol. 6: 225-229.

BLIDING, С. 1963. А critical survey of European taxa in Ulvales.
I. Opera Bot. (Suppl. Bot. Not.) 8: 1-160.

156 Ећодога [Vol. 77

1968. A critical survey of European taxa in Ulvales.
II. Ulva, Ulvaria, Monostroma, Kornmannia. Bot. Not. 121:
535-629.

BLUM, J. L. 1960. A new Vaucheria from New England. Trans.
Amer. Micros. Soc. 79: 298-301.

1968. Salt marsh Spartinas and associated algae. Ecol.
Monogr. 38: 199-221.
1972. Vaucheriaceae. No. Am. Flora. II. 8: 1-64.

,& J. T. CONOVER. 1953. New or noteworthy Vaucheriae
from New England salt marshes. Biol. Bull. 105: 395-401.

, & R. T. Witce. 1958. Description, distribution, and
ecology of three species of Vaucheria previously unknown from
North America. Rhodora 60: 283-288.

Burns, К. L., & А. C, MATHIESON. 1972a. Ecological studies of
economie red algae. II. Culture studies of Chondrus crispus
Stackhouse and Gigartina stellata (Stackhouse) Batters. Jour.
Exp. Mar. Biol. Ecol. 8: 1-6.

& . 1972b. Ecological studies of economic red
algae. III. Growth and reproduction of natural and harvested
populations of Gigartina stellata (Stackhouse) Batters in New
Hampshire, Ibid. 9: 77-95.

CARDINAL, A. 1964. Etude sur les ectocarpacées de la Manche. Nov.
Hedw. 15: 1-86. 41 figs.

Conover, J. T. 1958. Seasonal growth of benthic marine plants as
related to environmental factors in an estuary. Inst. Mar. Sci.,
Univ. Texas. Port Aransas 5: 97-197.

DmRovET, К. 1962. Gomont’s ecophenes of the bluegreen alga Micro-
coleus vaginatus (Oscillatoriaceae). Proc. Acad. Nat. Sci. 114:
191-205.

——————. 1968. Ecophenes of Schizothriz calcicola (Oscillatori-
aceae). Ibid. 115: 261-281.

1964. Ecophenes of Microcoleus chthonoplastes. Rev.
Algol. 4: 815-324.

1968. Revision of the classification of the Oscillatoria-
ceae. Monogr. 15. Acad. Nat. Sci., Philadelphia. 370 pp.

, & W. А. Папу. 1956. Revision of the coccoid Myxo-
phyceae. Butler Univ. Bot. Studies 12: 1-218.

FAN, К. C. 1956. Revision of the genus Calothrix Ag. Rev. Algol.
N.S. 2: 154-178.

FRALICK, К. A., & A. С. MATHIESON. 1972. Winter fragmentation
of Codium fragile (Suringar) Hariot ssp. tomentosoides (van
Goor) Silva (Chlorophyceae, Siphonales) in New England. Phy-
cologia. 11: 67-70.

& . 1973. Ecological studies of Codium fra-
gile in New England. Mar. Biol. 19: 127-132.

1975] Nahant Algae — Webber 157

НЕНЕЕ, E. J. 1972. Lomentaria clavellosa (Turner) Gaillons: an
addition to the marine algal flora of New Hampshire. Rhodora
74: 158.

, J. R. Conway, & R. A. STONE. 1970. Flora of the Wolf
Islands, Part III: The marine algae. Ibid. 72: 115-118.

,& A. C. MATHIESON. 1970. Investigations of New Eng-
land marine algae. III: Composition, seasonal occurrence, and
reproductive periodicity of the marine Rhodophyceae in New
Hampshire. Ibid. 72: 194-239.

JORDAN, A. J, & В. L. Vapas. 1972. Influence of environmental
parameters on intraspecific variation in Fucus vesiculosus. Mar.
Biol. 14: 248-252.

KANWISHER, J. L. 1957. Freezing and drying in intertidal algae.
Biol. Bull. 113: 275-285.

1966. Photosynthesis and respiration in some seaweeds.
Pp. 407-420 In: Н. BARNES, (Ер.), Some Contemporary Studies
in Marine Science. George Allen and Unwin Ltd., London.

LAMB, I. M. & М. Н. ZIMMERMANN. 1964. Marine vegetation of
Cape Ann, Essex County, Massachusetts. Rhodora 66: 217-254.

MATHIESON, A. C., & R. L. BURNS. 1970. The discovery of Halicystis
ovalis (Lyngbye) Areschoug in New England. Jour. Phycol. 6:
404-405.

& . 1971. Ecological studies of economic red
algae. I. Photosynthesis and respiration of Chondrus crispus
Stackhouse and Gigartina stellata (Stackhouse) Batters. Jour.
Mar. Biol. Eeol. 7: 197-206.

————————, & В. A. Евашск. 1972. Investigations of New England
marine algae. V. The algal vegetation of the Hampton-Seabrook
Estuary and the орсп coast near Hampton, N. H. Rhodora 74:
406-435,

————— E. J. HEHRE, & №. B. REYNOLDS. Investigations of New
England Marine Algae. I. A floristic and descriptive ecological
study of the marine algae at Jaffney Point, New Hampshire.
Nov. Hedw. (in press).

— —, N. В. REYNOLDS, & E. J. HEHRE. Investigations of New
England marine algae. II. The species composition, distribution
and zonation of seaweeds in the Great Bay Estuary System and
the adjacent open coast of New Hampshire. Ibid. (in press).

PARKE, M., & P. S. Dixon. 1968. Check-list of British marine algae
— second revision. Jour. Mar. Bio. Assoc. U. К. 48: 785-832.

PowELL, Н. T. 1957. Studies in the genus Fucus L. I. Fucus
distichus L. emend, Powell. Ibid. 36: 407-432.

RosENvINGE, L. K., & S. LUND. 1941. The marine algae of Denmark.
Contributions to their natural histery. II. Phaeophyceae. Part
I D. Kel. Danske Vidensk., Biol. Skrifter., Bd. 1. København.

158 Rhodora [Vol. 77

Sears, J. R, & К. T. WILCE. 1970. Reproduction and systematics
of the marine alga Derbesia (Chlorophyceae) in New England.
Jour. Phycol. 6: 381-392.

TAYLOR, W. R. 1937. Marine Algae of the Northeast Coast of
North America. ix + 427 pp. Univ. Mich. Press. Ann Arbor.

1957. Marine Algae of the Northeast Coast of North
America. (Revised ed.). viii + 509 pp. Univ Mich. Press. Ann
Arbor.

TILDEN, J. 1910. Minnesota Algae. I. Myxophyceae of North
America. Minneapolis.

VAN DEN Hork, C. 1963. Revision of the European species of
Cladophora. E. J. Brill. Leiden. 248 pp. + 55 pls.

WAERN, M. 1952. Rocky-shore algae in the Gregrund Archipelago.
Acta Phytogeogr. Suecica 30: 1-298. Uppsala.

WEBBER, E. E. 1967. Bluegreen algae from a Massachusetts salt
marsh. Bull. Torrey Bot. Club 94: 99-106.

1968. New England salt marsh Vaucheriae. Rhodora
70: 274-277.

1971. Observations on Microcoleus lyngbyaceus (Kütz.)
Crowan from marine habitats in New England. Ibid. 73: 238-
243.

, & К. T. WILCE. 1971. Benthic salt marsh algae at
Ipswich, Massachusetts. Ibid. 73: 262-291.

& . 1972. The ecology of benthic salt marsh
algae at Ipswich, Massachusetts. I. Zonation and distribution
of algal species. Ibid. 74: 475-488.

WILCE, К. T. 1965. Studies in the genus Laminaria. Pp. 247-256
In: Bot. Gothoburgensia. III. Proc. Fifth Mar. Biol. Symposium.
Göteborg.

—————. 1970. Cladophora pygmaea Reinke in North America.
Jour. Phycol. 6: 260-263.

1971. Some remarks on the benthic chrysophytes and
the fleshy red and brown crusts. Pp. 17-25 In: Symp. Cold
Water Inshore Marine Biology — Some Regional Aspects. Mar.
Sci. Inst., Nahant, Mass.

‚© R. W. LEE. 1964. Lomentaria clavellosa in North
America. Bot. Mar. 6: 251-258.

— — — ——, E. E. Wesrer, & J. R. SEARS. 1970. Petroderma and

Porterinema in the New World. Mar. Biol. 5: 119-135.

DEPT. OF BIOLOGY
KEUKA COLLEGE
KEUKA PARK
NEW YORK 14478

PHRAGMITES COMMUNIS IN SOUTH CAROLINA

Phragmites communis Trinius, the tall reed, is a coarse
rhizomatous perennial reed forming extensive colonies in
open marshy habitats in almost every state of the eastern
United States. It occurs sporadically in the southeastern
states but has not been previously recorded in South Caro-
lina or Georgia (Radford et al., 1968). It does occur
sporadically in North Carolina, favoring a few of the
northeastern and southeastern counties.

A large colony of Phragmites was discovered by the
author in Georgetown County, South Carolina, covering
an area of several hundred square meters. The colony is
located approximately 100 meters west of the Waccamaw
River (at high tide) and 125 meters north of Route 17
South. Vouchers of immature specimens were collected and
are on deposit at the herbarium of the University of South
Carolina, Columbia, South Carolina.

LITERATURE CITED

RADFORD, A. E., Н. E. AuLES, & С. R. BELL. 1968. Manual of the
vascular flora of the Carolinas. 1183 pp. Univ. North Carolina
Press, Chapel Hill.

RICHARD STALTER
DEPARTMENT OF BIOLOGY
ST. JOHN'S UNIVERSITY
JAMAICA, NEW YORK 11432

159

BUTOMACEAE: А NEW FAMILY RECORD
FOR NORTH DAKOTA

Butomus umbellatus L., flowering rush, was collected on
July 25, 1972, four miles south of Valley City, North Da-
kota (SE!4 Sec. 9, R58W, T139N). Scattered colonies in
flower were found growing in shallow water along the
shoreline and on silted gravel bars of the Sheyenne River
for a distance of about four miles (Godfread 4640, 4882,
and 4926, NDA). These stations were revisited in 1973 and
1974 and the colonies were increasing in size.

Butomus umbellatus is a native of Eurasia but has be-
come established along the St. Lawrence River Valley.
Herbarium records indicate that the species existed in Que-
bec in 1916. Rousseau (1966) indicates that B. umbellatus
and another Eurasian introduction, Lythrum salicaria L.,
purple loosestrife, are common along the shoreline of the
St. Lawrence River, appearing to be as much in equilibrium
as if in their native habitat. It is interesting to note that
L. salicaria is also well established along the Sheyenne
River near Valley City.

Since its introduction into North America, Butomus
wmbellatus has become established at widely scattered lo-
calities. In 1956 it was found in Idaho. It was collected in
Wisconsin for the first time in 1957, and it is known from
Flathead County, Montana (Hahn, 1973). In 1972 it was
reported from Rice County, Minnesota, while Van Bruggen
(1972) reports B. wmbellatus in South Dakota from Faulk
County, 150 miles south of the North Dakota plants along
the Sheyenne River. The establishment of В. wmbellatus in
North Dakota is apparently rather recent. Mr. and Mrs.
Russell Pederson, who own adjacent farmland, first noticed
the plants five or six years ago.

LITERATURE CITED

HAHN, B. E. 1973. Flora of Montana Conifers and Monocots, De-
partment of Botany, Montana State University, Bozeman, Mon-
tana. 143 pp.

160

1975] Butomaceae — Godfread & Barker 161

RovssEAU, J. 1966. Movement of plants under the influence of man.
Pp. 81-89. In: R. L. Taylor and R. A. Ludwig [ed.], The Evo-
lution of Canada's Flora. University of Toronto Press, Toronto.

VAN BRUGGEN, T. 1972. A Manual for the Identification of the
Vascular Plants of South Dakota. University of South Dakota
Educational Media, Vermillion, South Dakota. 465 pp.

CAROLYN GODFREAD
WILLIAM T. BARKER

DEPARTMENT OF BOTANY

NORTH DAKOTA STATE UNIVERSITY
FARGO, N.D. 58102

GERANIUM SIBIRICUM L. (GERANIACEAE)
IN WISCONSIN

This species, adventive from Eurasia, has heretofore been
reported to extend from New York and Pennsylvania to
Illinois (Fernald, Gray’s Manual, ed. 8, 1950.). I report
here its discovery in Wisconsin, from records in the her-
baria at WIS and OSH. The curators of the herbaria at MIL
and UWM have kindly informed me that they have no rec-
ords of the plant from Wisconsin. The four collections
listed below from two counties warrant the inclusion of
the plant as an element in the Wisconsin flora.

Wisconsin: DANE CO.: large colony along town road close
to village of Dane, Н. C. Greene, 17 Aug. 1954 (wIs); on
disturbed sandy peat in grazed area of south Lake Wabesa
marsh, Don Samuelson, 5 Sept. 1972 (wis) ; in shady, damp
soil along Elm Drive, U. of Wisconsin-Madison Campus,
Thomas 002, 3 Sept. 1972 (у/15). COLUMBIA СО.: along a
sandy roadside on Chryslaw Road, to the NW. of the junc-
tion of State Route 113 and county trunk J, Sect. 19, R7E,
TION, Zaudtke 053, 9 Sept. 1972 (osH).

NEIL A. HARRIMAN
DEPARTMENT OF BIOLOGY

U. OF WISCONSIN — OSHKOSH
54901

162

EDITOR’S NOTE ТО CONTRIBUTORS

During the past two years the number of manuscripts
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163

164 Ећодога [Vol. 77

Address manuscripts, proofs, and related correspondence
to:
Dr. A. Linn Bogle
Editor-in-Chief, Rhodora
Botany & Plant Pathology Dept.
University of New Hampshire
Durham, N. Н. 03824

Volume 77, No. 809, including pages 1-164, was issued May 30, 1975.

Cover illustration

Rhododendron canadense (L.) Torr.
from
Curtis’ Botanical Magazine t.474

RHODORA March, 1975 Vol. 77, No. 809

CONTENTS

On the Epibiotic and Pelagic Chlorophyceae, Phaeophyceae, and
Rhodophyceae of the Western Sargasso Sea

TTT éen TTT TF EE E Ae ЫА Сыл 1
Saxifrages on Mount Washington
hi. EEN E Ae EN

Rudbeckia auriculata (Perdue) Kral, a Species Distinct from
R. fulgida Ait.
КОР с, ыл EE wee, CX HANDLE 44

Apparent Ecotypic Differences in the Water Relations of Some
Northern Bog Ericaceae

FM V. то се на ТОО ыс а `
The Deciduous Magnolias of West Florida
ZOOM M INE ОРООН ЗАТИМ bis пау КО DR e uh И 64

Investigations of New England Marine Algae VII: Seasonal
Occurrence and Reproduction of Marine Algae Near Cape Cod,
Massachusetts
Douglas C. Coleman and Arthur C. Mathieson ............ E UR 76

А List of the Monocotyledoneae of Belize Including a Historical
Introduction to Plant Collecting in Belize
David L. Spellman, John D. Dwyer, and Gerrit Davidse ........ 105

Notes on the Leguminosae II: Facultative Dwarfism in Crotalaria
sagittalis L.

ТИИИП AE Co О p Ul, Б. cranii EE Жы 141
Pectis humifusa New to the Flora of the United States

ТАМ D. JOE ЕНИН ЧАИ ОИНИ nebenan €".
Maritime and Marine Lichens from Nahant

Se КАЛАН уилн EE" "CC с 147

Phycological Studies from the Marine Science Institute, Nahant,
Massachusetts I: Introduction and Preliminary Tabulation of
Species at Nahant

BW WEE лове ШИ нерви ован T
Phragmites communis in South Carolina

ПИО ЯШИ. sed iiec А РОНЕ. EH 159
Butomaceae: A New Family Record for North Dakota

Carolyn Godfread and William Т. Barker .................................... 160
Geranium sibiricum L. (Geraniaceae) in Wisconsin

ПИК PU INI. Sah ats ois tia hades sn rrt ssh E coca cscs 162

ol, 77

[000га

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

June, 1975 No. 810

The Nem Lugland Botanical Club, uc.

Botanical Museum, Oxford Street, Cambridge, Mass. 02138

Conducted and published for the Club, by

ALFRED LINN BOGLE, Editor-in-Chief
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Vol. 77 June, 1975 No. 810

DE PLANTIS TOXICARIIS E MUNDO NOVO
TROPICALE COMMENTATIONES XII
NOTES ON BIODYNAMIC PIPERACEOUS PLANTS

RICHARD EVANS SCHULTES

Recent field work has served to corroborate the suspicion
long held by botanists that the Piperaceae represent a fam-
ily of plants of very basic importance in the ethnopharma-
cology of primitive societies of the New World tropies.
The following notes are offered as a contribution to our
growing knowledge of the use of piperaceous species for
their variety of biodynamic activity.

Peperomia emarginella (Sw.) С. De Candolle, in DC. Prodr.
16, pt. 1 (1869) 437.

COLOMBIA: Comisaría del Putumayo, Mocoa and
vicinity. December 8, 1942. R. E. Schultes et C. E. Smith
3028.

Called in the Ingano language cungamanda-ambe, this
herb “pounded and mixed with tobacco and urine" is used
“to poultice bites of the cungamanda ant”.

Peperomia glabella (Sw.) A. Dietrich var. melanostigma
Dahlstedt, in Kgl. Sv. Vet, Akad. Handl. 33;
pt. 2 (1900) 122.
COLOMBIA: Comisaria del Putumayo, Mocoa and
vicinity. Alt. 820 m. “For mal de оўо”. December 6, 1942.
R. E. Schultes et C. E. Smith 2059.

165

166 Ећодога [Vol. 77

This herb is a supposed remedy for conjunctivitis. In
the Ingano language of Mocoa, it is known as tre-gwen or
gwinan.

Peperomia serpens (Sw.) Loudon, Hort. Brit. (1830) 13.

COLOMBIA: Comisaría del Putumayo, Río Sucum-
bios, Santa Rosa. “Remedy for bite of conga ant. Kofán
пате: u-nu-sč'-hč-pa”. April 7-8, 1942. R. E. Schultes 3589.

The aromatic leaves and stems of Peperomia serpens are
employed locally to relieve the irritant sting of the conga
ant. In Kofán, the term 36'-ће-ра refers to a medicinal or
poisonous plant.

Piper Allenii Trelease, in Ann, Mo. Bot. Gard. 25 (1938) 826.

PANAMA: Provincia del Darién, trail between Pin-
ogana and Yavisa. Altitude about 15 m. March 17, 1937.
P. A. Allen 270.

Allen reports that the “roots are used by Indians to
deaden pain" and the leaves “аз a snake bite remedy".

Piper auritum Humboldt, Bonpland et Kunth, Nov. Gen, et
Sp. 1 (1815) 54.
EL SALVADOR: Vicinity of San Salvador. Alti-
tude 650-850 m. February 2-7, 1922. P. C. Standley 20550.
According to Standley, the juice of the crushed leaves of
this highly aromatic species is employed to remove ticks.
The shrub is locally called Santa María.

Piper Bartlingianum C. De Candolle, in De Candolle Prodr.
16, pt. 1 (1869) 257.
DUTCH GUIANA: Fetikruk. On sandy hills. Au-
gust 10, 1939. Geykes sine тит.
The collector reports Piper Bartlingianum as an ingredi-
ent of “оега]і poison" amongst the Wayana Indians.
Geykes sine num. appears to represent the same species
as A. C. Smith 2826 and 2827 from British Guiana. It is
said to be employed as one of the elements of Waiwai In-
dian arrow poison.

1975] Piperaceae — Schultes 167

Piper dactylostigmum Yuncker, in Inst. Bot. S&o Paulo Bol.
No. 3, (1966) 35, fig. 29.

BRAZIL: Estado do Amazonas, Manáos and vicin-
ity, Reserva Ducke, “Climber, adpressed to trunk of tree.
Leaves and stem rapidly numb the tongue when chewed".
April 13, 1972. R. E. Schultes et W. Rodrígues 26150.

The rapid, strong and long lasting numbness of the
tongue produced when the leaves and stems of this species
are chewed is well known to the inhabitants of the forested
areas around Manáos. So far as could be ascertained, how-
ever, this property does not constitute the basis of any
medicinal or other folk use of Piper dactylostigmum.
Numbing of the tongue and mucous membranes of the
mouth, which, though not common, is known for other
species of the genus, sometimes leads to local medicinal
use. The leaves and twigs of Piper corcovadensis (Mio)
DC. and P. Jaborandi Vell., for example, are chewed in the
region of Rio de Janeiro to relieve toothache because of
their strong numbing effect (Mors, W. B. and C. T. Rizini:
Useful plants of Brazil (1966) 89).

Piper erythroxyloides R. E. Schultes et García-Barriga sp.
nov.

Frutex glaber, usque ad 11% ped. altus, erectus. Rami
simplices vel pauce ramificati, internodiis superioribus sat
gracilibus elongatisque, subgranulosis, glabris, 4.5-6 cm.
longis. Folia membranacea, elliptica, apice acuminata, basi
aequilater subrotundata, 14-18 cm. longa, 6-8.5 em. lata,
omnino pinnatim venosa, venis primariis glabris, utrinque
8-10, supra glabra, subtus minute punctulosa et irregu-
lariter albo-squamulosa; petiolo subearnosulo, usque ad 1
em. longo, longitudinaliter striato, scobiculato. Pendunculus
subcarnosulus, usque ad 1 ст. longus, glabrus vel minutis-
sime albido-papilloso-pilosus, bracteis crassis, suborbicu-
laribus, cupulatisque, plus minusve 1 mm. longis. Drupa
carnosa, extus papillosa, conica, 2 mm. in diametro, 1 mm.
longa, in stylum crassum, 0.75 mm. longum desinens.

168 Ећодога [Vol. 77

COLOMBIA: Departamento de Santander del Norte,
La Motilonia, Río Catatumbo у Río Brandy. Altitude 80 m.
“0.5 m. alto, erecto. Amentos verde-oscuros. A] masticar
el tallo se le siente un piquante fuerte y luego se duerme la
lengua y los labios. Los indios bari (motilones) lo mastican
frecuentemente como la coca. Por lo tanto es narcótico".
May 24-26, 1965. Hernando García-Barriga et Gustavo
Lozano-C 18414. TYPUS in Herb. Nac. Colomb.; TYPUS
DUPLICATUS in Econ. Herb. Oakes Ames.

Piper erythroxyloides appears to be most closely allied to
P. dariense С. DC. of northern Colombia and Panama. It
differs in various important respects. The leaves of Piper
erythroxyloides are basally rounded and apically short
acuminate, with a blunt tip (not cuneate and long acumi-
nate with a sharp tip); the internodes are not so slender
and are somewhat longer; the inflorescence is usually
longer, and thicker; the flowers are more congested, not
loosely arranged; the style is thicker; and the stigmas are
relatively longer. Perhaps the most noticeable difference
lies in the shape of the drupe: in Piper erythroxyloides, the
fruit is conic and rough-papillose, whereas in P. dariense
it is globose-tetragonous and either smooth or obscurely
papillose.

According to García-Barriga and Lozano, the stems of
Piper erythroxyloides are chewed by the Мо опе (Bari)
Indians as а kind of narcotic which has local effects in the
mouth that resemble those of coca: whence the specific
name erythroxyloides. Upon mastication, the stems induce
a strong burning sensation which is followed by numbness
of the tongue and lips.

In addition to this use, the plant is chewed in the belief
that it prevents dental caries.

Local Indian names of Piper erythroxyloides are achi-
kaira and chanquira.

It is of interest to note that in February, 1971, Mr. Alfred
M. Ajami of Harvard University submitted to the Botanical
Museum several internodes of what appear, so far as such

1975] Piperaceae — Schultes 169

incomplete material can indicate, to represent the same
species: Piper erythroxyloides. According to Mr. Ajami,
the Bari Indians of the central Venezuelan region west of
Lake Maracaibo, who chew the internodes as an aphrodisiac,
experience a strong analgesic effect in the mouth and con-
spicuous dilation of the pupils. Inasmuch as the tribe is
apparently the same group as the Colombian ‘Motilones”
and the two localities are not distant, the information com-
municated by Mr. Ajami assumes greater significance. It
is hoped that complete botanical specimens from the Vene-
zuelan locality may be forthcoming.

Piper hispidum Swartz, Prodr. Veg. Ind. Occ. (1788) 15.

ECUADOR: Parroquia Concepión, Playa Rica. Alti-
tude 91 m. “Forest near stream; undergrowth. Perennial
herb 4 m. high; aments erect. Common”. December 6,
1936. Y. Mexia 8407.

The collector reports that the leaves of this plant are
“crushed in water to kill head lice". It is locally known as
pipilongo. This collection is the type of Piper pediculicidum
Trelease.

Piper cf. interitum Trelease ex Macbride, in Field Mus. Publ.
Bot. 13, no. 357 (1936) 176.

PERU: Departamento de Loreto, alto Rio Purts,
Zapote. “Culina Indian name: tetsi. Substitute of tobacco
snuff. The leaves are dried and reduced into powders. Tree
about 10 m. high. Lowland forest". October 15, 1968. Lau-
rent Rivier 21.

This interesting report of the preparation of a snuff from
the leaves of a species of Piper — а snuff used as a “substi-
tute" for tobacco — provides a significant addition to our
growing understanding of plants employed in South Amer-
ica in the preparation of powders for inhalation. The num-
ber of species utilized is much larger than has been sus-
pected, and, in this case, the indication of psychoactive
constituents — probably essential oils — adds to the ethno-
pharmacological interest of the report.

170 Ећодога [Vol. 77

Piper sp.

BRAZIL: Estado do Amazonas, Rio Livramento,
Humayta, near Livramento. October 12-November 6, 1934.
В. А. Krukoff 6972.

The stem of this vine-like species of Piper, according to
the collector’s notes, contains “а substance producing local
anaesthesis”. The plant is employed locally by the Indians
“to cure toothache” and is called cipó de dor dente (‘‘tooth-
ache vine’’).

Pothomorphe umbellata (L.) Miquel, Comm. Phyt. (1840)
36.

COLOMBIA: Comisaria del Putumayo, Rio Guam-
ués, San Antonio. Secondary growth near path. “Коѓап
Indian name: a-nama-he sé’-hé-pa. Curare, used alone or
mixed; for monkeys and wild pigs (saquita). Bark of lower
stem and root is scraped, then boiled. Herb 5-6 feet”. Sep-
tember 5, 1966, H. V. Pinkley 421. Same locality. Febru-
ary 18, 1966, Pinkley 119.

The preparation of arrow poisons amongst the Kofan
Indians of the border region of Colombia and Ecuador is
ethnobotanically extremely complex. Although piperaceous
plants are known to be employed as ingredients of arrow-
poisons over a wide area in tropical America (Hegnauer,
R.: Chemotaxonomie der Pflanzen 5:321. 1969), Pinkley’s
report of the utilization of Pothomorphe umbellata “alone
or mixed" assumes special significance, Most, if not all, of
the piperaceous species employed in these preparations are
not known to have curare constituents, and their use as
additives may be based on superstition or symbolism. If a
species be utilized “alone” — that is, as the only ingredient
in a curare — it must have a biodynamically active con-
stituent.

CHROMOSOME COUNTS OF COMPOSITAE
FROM THE UNITED STATES, MEXICO,
AND GUATEMALA!

DAVID J. КЕП, AND Тор Е. STUESSY

Chromosome numbers can be extremely useful in system-
atic studies, particularly for helping to reveal evolutionary
relationships. For the past fifteen years numerous chromo-
some reports from plants have been published, especially
in the Compositae, and these counts have been compiled
in several major sources (Darlington & Wylie, 1955; Cave,
1958-65 ; Ornduff, 1967-69; Fedorov, 1969; Moore, 1970-72).
However, a rapid glance through these references indicates
not only that many species never have been counted, but
also that many taxa are known only from a single plant in
one population. In view of the common occurrence of
euploid and aneuploid races in plants as illustrated by
several detailed investigations (e.g., Lewis, 1962, 1970;
Stuessy, 1971а), it is desirable to have several to many
counts from each species before accurate judgments can be
made regarding evolutionary relationships (Stuessy, 1971b;
Kovanda, 1972; Strother, 1972). The present paper helps
to remedy these deficiencies in the Compositae by: (1) re-
porting first chromosome counts for several genera, species,
and varieties; and (2) reporting additional populational
chromosome counts for taxa documented previously.

MATERIALS AND PROCEDURES

The meiotic chromosomal material for this study was
collected during the past several years by the senior and
junior authors on various field excursions. Immature capit-
ula were killed and fixed in modified Carnoy’s fluid (4
chloroform: 3 absolute alcohol: 1 glacial acetic acid) and
refrigerated in the laboratory until later prepared by con-

‘Publication No. 843 from the Department of Botany, The Ohio
State University, Columbus.

171

172 Ећодога [Vol. 77

ventional acetocarmine squash techniques. Voucher speci-
mens collected by Keil and assistants are on deposit in the
herbarium of The Ohio State University (05); vouchers
collected by Stuessy are in the herbarium of the University
of Texas at Austin (TEX).

RESULTS

The chromosome counts obtained in the present study
are listed in Table 1. First counts are reported for two
genera, 16 additional species, and one variety; 112 addi-
tional counts are for taxa counted previously, seven of
which are new numbers. The first counts for genera are
from Epaltes Cass. (п = 10) and Tricarpha Longpre
(n — 8), and first counts for species are in Bidens L.,
Calea L., Guardiola Cerv. ex H. & B., Machaeranthera
Nees, Melampodium L., Otopappus Benth., Sclerocarpus
Jacq., Senecio L., Sigesbeckia L., Simsia Pers., Spilanthes
Jacq., Tridax L., and Zaluzania Pers.

DISCUSSION

Because many of the counts presented here corroborate
previous chromosomal reports, the discussions are restricted
either to first counts or to new reports for genera, species,
or varieties. The order of commentary will follow the se-
quence of tribes in the classification of Hoffmann (1890-94),
which is the same as that used in Table 1. References for
statements regarding the range of chromosomal variation
within genera will not be given; documentation for these
counts comes from the several major sources cited in the
introduction to this paper.

EUPATORIEAE. Counts for three herbaceous species of
Stevia Cav. represent new reports. Stevia elatior H.B.K.
is cited here as n = 12r & 121 (Fig. 1), whereas the two
previously recorded numbers have been n = 84ү (Powell &
Turner, 1963) and n = 33; (Grashoff, Bierner, & North-
ington, 1972). Our count for Stevia origanoides H.B.K.,

ә

1975] Chromosome Counts — Keil & Stuessy 177

reported here as п = 11 (Fig. 2), is the first for this taxon
at what appears to be the diploid level; the previous counts
were n = 341 and n = са. 43 + 11 (Grashoff et al., 1972).
Stevia plummerae A. Gray var. durangensis Robins, has
been reported before only once by Grashoff et al. (1972) as
п = са. 17, but our count is n = 44; (Fig. 3). As pointed
out by Grashoff et al. (1972), it is common to find varying
meiotic chromosomal associations and numbers in species
that have apomictic races, as are present in these three
taxa. It is not surprising, therefore, that our reported
counts add to this chromosomal diversity.

ASTEREAE. Several previous counts have been reported
for Erigeron karwinskianus DC.: 2n — 32 (Carano, 1924;
Battaglia, 1950) ; 2% = 36 (Fagerlind, 1947; Larsen, 1953,
1954; Kliphuis & Wieffering, 1972); п = 9 and n= 27
(Turner, Ellison, & King, 1961) ; and п = са. 27 (Turner,
Powell, & King, 1962). Considering the variation in chro-
mosome number that has been documented previously in this
species, as well as our new report of n = Бп & 171 (Fig. 4),
it is likely that E. karwinskianus is apomictic through at
least part of its range from Mexico to northern South
America (Solbrig, 1962). It is interesting that our count
comes from a population very near the locality cited by
Turner et al. (1961) for their counts of n = 9 and n = 271.

Machaeranthera coulteri (А. Gray) Turner & Horne (as
Psilactis coulteri A. Gray) was reported as n — 5 by Sol-
brig, Anderson, Kyhos, Raven, and Rüdenberg (1964).
However, based on the recent revision of sect. Psilactis of
Machaeranthera by Turner and Horne (1964), the geo-
graphie location of the voucher for the count seems more
appropriate for M. arida Turner & Horne than for M.
coulteri. The latter species, as recently interpreted, is
known only from the vicinity of Guaymas, Sonora,
where our voucher was collected. Our first count of n — b
(Fig. 5) for Machaeranthera coulteri is particularly inter-
esting because in the previously mentioned revision of sect.
Psilactis of the genus (Turner & Horne, 1964; cf. their

174 Ећодога [Vol. 77

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Figs. 1-22. Camera lucida drawings of meiotic chromosomes of
species of Compositae. Diplotene, Fig. 18; diakinesis, Figs. 5, 7-10,
12, 15, 19-21; metaphase I, Figs. 1, 3, 4, 6, 11, 14, 16, 17; metaphase
II, Figs. 2 (опе half of cell shown), 13. All figures same scale.
Bivalents black, univalents white. KC = Keil & Canne, KM = Keil
& McGill, К = Keil. Fig. 1, Stevia elatior, К 9396, п = 121 & 121;
Fig. 2, Stevia origanoides, KC 8884, n = 11; Fig. 3, Stevia plummerae
var. durangensis, KC 8927-1, n = 441; Fig. 4, Erigeron karwinskianus,
KC 9178, n = 5u & 171; Fig. 5, Machaeranthera coulteri, KC 8637,
n = 5; Fig. 6, Epaltes mexicana, KC 9211, п = 10; Fig. 7, Bidens

1976] Chromosome Counts — Keil & Stuessy 175

riparia var. refracta, KC 8710, п == 12; Fig. 8, Guardiola platyphylla,
KM 8558, n = 12; Fig. 9, Melampodium appendiculatum, KC 8706A,
n — 10; Fig. 10, Otopappus imbricatus, KC 9112, n — 16; Fig. 11,
Parthenium incanum, KM 7765A, n == 181 & 187; Fig. 12, Sclero-
carpus spatulatus, KC 8671А, n — 11; Fig. 13, Sigesbeckia jorullensis,
KC 8902, n = 30; Fig. 14, Simsia eurylepis, KC OASIS — bet
Fig. 15, Simsia grayi, KC 9081, n — 17; Fig. 16, Spilanthes phane-
ractis, KC 9035, n — 41; Fig. 17, Tricarpha durangensis, KC 8860A,
п = 8; Fig. 18, Tridax tenuifolia var. microcephala, KC 8808, n = 9;
Fig. 19, Zaluzania grayana, KM 8379A, n = 17; Fig. 20, Schkuhria
pinnata var. guatemalensis, К 9402А, п — 10; Fig. 21, Senecio
runcinatus, KC 9192, п = 22; Fig. 22, Pinaropappus roseus, KC 9177,
nec 20 mp ве e

176 Ећодога [Vol. 77

Fig. 3), M. coulteri on morphological grounds was placed
in the z — 5 cytophyletic group along with M. arida and
M. ствра (Brandg.) Turner & Horne, both known chromo-
somally as n — 5. More recently M. arizonica Jackson &
В. В. Johnson and M. parviflora A. Gray have been added
to this group and both species have been counted as n = 5
(Jackson & Johnson, 1967). This first chromosomal report
for M. coulteri substantiates its phyletic association with
these other species. All other taxa in sect. Psilactis are
known chromosomally as either n = 4 or n= 9 (Turner
& Horne, 1964).

INULEAE. The first report for Epaltes (E. mexicana),
n == 10 (Fig. 6), is in keeping with its present subtribal
disposition in the Plucheinae. Of the related genera of the
same subtribe (Hoffmann, 1890-94) that are known chromo-
somally (Blumea DC., Pluchea Cass., Pterigeron (DC.)
Benth., Pterocaulon EIl., Sphaeranthus L., and Tessaria
Ruiz & Pav.), all are based on 2 = 10 except Blumea which
appears multibasic with x = 9, 10, and 11. On morphologi-
cal and geographical grounds, in our opinion, Epaltes mexi-
cana Less. is quite similar to some species of Pluchea, the
former differing mainly in its smaller heads and flowers
and in its epappose achenes. As emphasized by Bentham
(1873) and Godfrey (1952), the generic boundaries in the
Plucheinae are not well defined and perhaps should be re-
evaluated.

HELIANTHEAE. Bidens riparia is reported for the first
time as n — 12 (Fig. 7) in a genus that has most frequently
counted numbers of n = 12, 24, and 36 (clearly based on
х — 12).

The first count for Calea zacatechichi Schlecht., » — ca.
19, is in keeping with previous reports for other species of
the genus (n = 9, 16, 18, 19, 24, 32). According to the
most recent revision of the Mexican and Central American
taxa (Robinson & Greenman, 1896), C. zacatechichi is most
closely related to C. nelsonii Robins. & Greenm. which has
been counted as п = ca. 18 (Turner et al., 1962). The

1975] Chromosome Counts — Keil & Stuessy 177

morphological and chromosomal heterogeneity within Calea
and the absence of a recent revision of the entire genus
suggest that a thorough modern study is much needed.

Guardiola, a genus of about ten species, has been placed
traditionally in the subtribe Melampodiinae (Hoffmann,
1890-94). However, recent studies by the junior author
suggest that on morphological and cytological evidence it
belongs more properly in the Coreopsidinae (Stuessy,
1973). Our first count of n = 12 (Fig. 8) for С. platyphylla
A. Gray is consistent with the recent reports of n = 12
for both С. tulocarpus A. Gray (Grashoff et al., 1972) and
G. mexicana H. & B. (Solbrig, Kyhos, Powell, & Raven,
1972) ,> and with the base number of x = 12 for several
other members of this subtribe.’

The count of п = 10 (Fig. 9) is a first report for
Melampodium appendiculatum Robins. In a recent revision
of the genus (Stuessy, 1972) this species is placed in series
Cupulata of sect. Melampodium; three other related species
(M. cupulatum А. Gray, M. rosei Robins., and M. tenellum
Hook. & Arn.) also are known chromosomally as n = 10
(Stuessy, 1971b). The addition of this new count increases
the number of species surveyed within the genus to 27 out
of 37,

The first generic report for Otopappus (О. scaber 8. F.
Blake) has been published recently by Solbrig et al. (1972)
as п — 16. Our first report for О. imbricatus (Sch.-Bip.)
5. F. Blake of » = 16 (Fig. 10) confirms this chromosomal
level for the genus. The related genera Salmea DC. and
Notoptera Urb. (Blake, 1915) are known respectively as
n = 18 + 2 frag. (Turner et al., 1962) and » — ca. 15 & 16
(Turner et al., 1962; Turner & King, 1964), although very
few taxa have been examined from each..

"The count published by Solbrig et al. was listed for С. atriplici-
folia A. Gray, but in the most recent published revision of the genus
(Robinson, 1899) this epithet is regarded as synonymous with 6.
mexicana.

3 (e.g., Bidens L., Coreopsis L., Cosmos Cav., Glossocardia Cass.,
Thelesperma Less.)

178 Rhodora [Vol. 77

Parthenium L., and particularly P. argentatum A. Gray,
has been studied extensively for many years (cf. Hammond
& Polhamus, 1965), including a comprehensive revision by
Rollins (1950). Parthenium incanum H.B.K. has been re-
ported previously as having a polyploid series of n = 18,
27, 36, and 45, but our new count is n = 18п & 18r (Fig.
11). This interploid number could represent the product of
hybridization between P. incanum and other species of the
genus that grow in the vicinity, such as P. argentatum,
but our voucher specimens show no morphological indica-
tion of such intergradation. Alternatively, the meiotic con-
figuration could indicate a hybrid between n = 18 and
n = 86 cytotypes of the same species. The plants under
consideration also could be apomictie, a condition that is
known to occur in populations in the northern range of P.
incanum (Rollins, 1950) where our material was collected.

The count of n = 11 (Fig. 12) for Sclerocarpus spatu-
latus Rose is consistent with previously reported numbers
of n — 11, 12, 14, and 18 in the genus as recently defined
by Feddema (1971). The closely related genus, Aldama
LaLlave & Lex., is known chromosomally as n = 17 (Turner
et al., 1962; Powell & Cuatrecasas, 1970; Feddema, 1971).

Sigesbeckia L. of the subtribe Helianthinae is a small
genus of less than ten species. It is worthwhile to mention
that a close morphological resemblance exists with T'ri-
gonospermum Less. (McVaugh & Anderson, 1972; Stuessy,
1973) and perhaps also with Rumfordia DC., the former
of the subtribe Melampodiinae and the latter of the Helian-
thinae. Our count of n = 15 is a first report for S. agrestis
Poepp. & Endl All but two other reports in the genus
[n == 12 (Subramanyam & Kamble, 1967) and 2n = 20
(Hsu, 1967) for S. orientalis L.] have been either n = 15
or n == 30. Infraspecifie euploidy is known to occur in S.
orientalis (Mehra, Gill, Mehta, & Sidhu, 1965) and it is
now documented for S. jorullensis H.B.K. by our counts of
n = 15 and 30 (Fig. 13). Only one count of n = 15 (Sol-
brig et al, 1972) has been recorded previously for this
species.

1975] Chromosome Counts — Keil & Stuessy 179

Of the approximately 35 species of Simsia recognized by
various authors (Blake, 1913, 1917, 1928; Cuatrecasas,
1954; Robinson & Brettell, 1972), ten have been counted
from morphologically diverse parts of the genus, and all
counts have been n = 17. Our first counts of n = 17 (Figs.
14 & 15) for S. eurylepis б. F. Blake and S. grayi Sch.-Bip.
ex S. F. Blake emphasize the chromosomal uniformity
within the genus.

Spilanthes with approximately 60 species (Moore, 1907)
is a taxonomically complex genus much in need of revision-
ary attention. It appears to belong in the subtribe Galin-
soginae rather than in the Helianthinae as traditionally
placed (Hoffmann, 1890-94). Chromosomally the situation
also is complex. Even though only six species have been
counted, four base numbers, r — 7, 12, 13, and 16, are
present. Our first report of n == ca, 45 for S. ocymifolia
(Lam.) A. H. Moore adds another chromosomal] level to the
already chromosomally diverse sect. *Salivaria" (= sect.
Spilanthes) known with n = 7, 12, 16, and 26. All previous
reports for sect. Acmella (Rich) DC. have been clearly
based on x = 13 (only n = 13 and n = 26 counts reported).
Our new report of » — 41 (Fig. 16) for S. phaneractis
(Greenm.) A. H. Moore increases the chromosomal diversity
of this section as well.

Tricarpha is a genus of two species recently described by
Longpre (1970). Our first count for the genus (from T.
durangensis Longpre) of » — 8 (Fig. 17) substantiates its
presumptive close relationship to Sabazia Cass. (п = 4, 8,
and 16) and Selloa Kunth ( = 8) as mentioned by Long-
pre (1970). The problem of generic delimitation in the
subtribe Galinsoginae, involving Tricarpha, Sabazia and
Selloa as well as Galinsoga Ruiz & Pavon, Stenocarpha
5. F. Blake, Tridax and Jaegeria Kunth, is much in need
of further study, despite the appearance in recent years of
several excellent revisions (Powell, 1965; Turner, 1965;
Torres, 1968; Longpre, 1970). Part of the difficulty in sort-
ing out the proper affinities of all the taxa in the Galinsogi-
nae is that previous workers have been working from the

180 Ећодога [Vol. 77

perspective primarily of a single genus and not from a
perspicacious overview of many of the genera within the
subtribe. An added difficulty is the absence of a recent
revision of Galinsoga (most recent treatment that of Robin-
son, 1894), the understanding of which clearly is central to
sorting out these generic relationships.

Our first report for Tridax tenuifolia Rose, n — 9 (Fig.
18), fits well with the established base number of x — 9
for sect. Tridax to which T. tenuifolia belongs (Powell,
1965).

Of the 14 species of Zaluzania recognized by Sharp
(1935), six have been counted with definite numbers of
п = 16 and 18. Our first report, n = 17 (Fig. 19), for
Z. grayana Robins. E Greenm, firmly establishes this as а
new chromosomal level for the genus (a previous count of
n = 17 = 1 for Z. montagnaefolia Sch.-Bip. was reported
by Powell and Turner, 1963).

HELENIEAE. Schkuhria pinnata (Lam.) Cabrera has been
counted before as 2n — 20 (Covas and Schnack, 1946), and
var. virgata (LaLlave) Heiser of the same species has been
reported as n = са. 20 (Turner et al., 1962). Recently
McVaugh (1972) transferred S. anthemoidea (DC.) Coult.
var. guatemalensis (Rydb.) Heiser to S. pinnata, and our
counts of n = 10 (Fig. 20) for this taxon are the first
reports. A count of n = 11 (Table 1) was obtained from
material tentatively identified as S. anthemoidea. However,
our voucher specimen differs from the characters of this
species as delimited by Heiser (1945) in having more
numerous disc florets and large ray florets, as in S. schkuh-
rioides (Link & Otto) Thellung in Fedde. In pappus struc-
ture, though, our plants are much more similar to S, anthe-
moidea than to S. schkuhrioides. 'This collection may repre-
sent a previously undescribed taxon.

SENECIONEAE. Our first reports of n = ca. 30 for Senecio
hartwegii Benth. and » — 22 (Fig. 21) for S. runcinatus
Less. are consistent with counts reported previously for
other taxa of the genus. Thirty-three species of Senecio

1975] Chromosome Counts — Keil & Stuessy 181

have been reported as n = 30 and six are known with n =
22. Although the genus is based either on xr = 5 (Barkley,
1962) or х= 10 (Ornduff, Raven, Kyhos, & Kruckeberg,
1965; Ornduff, Mosquin, Kyhos, & Raven, 1967), the diver-
sity of haploid numbers is great, representing 30 different
chromosomal levels from n = 5 to n = ca. 92.

CICHORIEAE. The small genus Pinaropappus Less. has
been counted from only one species, P. roseus Less., and
the reported counts are n — 9 and 18 (Darlington & Wylie,
1955; Turner et al., 1961; Powell & Turner, 1963; Powell
& Sikes, 1970). Our present count of n = 201 & 11 (Fig.
22) is a new report for this taxon. The meiotic configura-
tions of cells in our preparation were irregular with bridges,
lagging chromosomes, and varying numbers of univalents.

ACKNOWLEDGMENTS

Field work for this investigation was completed largely
with support from NSF grant GB-30240; publication costs
were defrayed partial by funds from NSF grant GB-
37678. Thanks are extended to Judith M. Canne and Lyle
A. McGill for assistance on collecting trips.

[Vol. 77

Rhodora

182

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192 Еһодога [Vol. 77

LITERATURE CITED

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CARANO, Е. 1924. Osservazioni sul meccanismo di divisione della
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CavE, M. [Ер.]. 1958-65. Index to Plant Chromosome Numbers,
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Covas, G., & B. ScHNACK. 1946. Nümero de cromosomas en Antó-
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CUATRECASAS, J. 1954. Notas a la flora de Colombia, XIII. Revista
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194 Ећодога [Уо]. 77

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1975] Chromosome Counts — Keil & Stuessy 195

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‚ & К. M. KING. 1964. Chromosome numbers in the
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DEPARTMENT OF BOTANY
THE OHIO STATE UNIVERSITY
COLUMBUS, OHIO 43210

А VEGETATIONAL SURVEY OF THE
VASCULAR PLANTS OF THE KENT ISLAND GROUP,
GRAND MANAN, NEW BRUNSWICK’

JOHN.W. McCAIN

In 1930, Mr. J. Sterling Rockefeller, interested in pre-
serving the eider ducks that breed along the coast of the
northeastern United States, purchased Kent Island, on
which the eiders nest. In 1934, he gave the island to
Bowdoin College as a breeding bird preserve and field
study station. Extensive ornithological work has been done
at this station over the years, but there have been only
limited botanical investigations. Dr. David Potter (1936,
1937) and John McCain (1973) compiled vascular species
lists, and Henry A. Gleason, Jr. (1937) sampled herbaceous
vegetation in the forested northern end of Kent Island. The
present study was designed to survey and map the vegeta-
tion of the entire island and to define the plant community
types in the non-forested areas. It is also intended to serve
as a reference for future ornithological research on Kent
Island.

Kent Island lies at 66°46’ E longitude and 44°35’ N lati-
tude, about fifteen miles east of the coast of Maine in the
Bay of Fundy. It is the outermost island in the Grand
Manan Archipelago and is politically part of Charlotte
County, New Brunswick, Canada. Its greatest length is
1.75 miles and its width varies from one-ninth to one-half
mile. The title deed lists the area at 150 acres.

(Thi study was supported by National Science Foundation Under-
graduate Research Participation Grants GY-4343 in 1968 and GY-
6095 in 1969 through Bowdoin College. The author wishes to
acknowledge the advice of Dr. Е. A. Stowell of Albion College and
Dr. Е. С. Voss and Dr. W. Н. Wagner of the University of Michigan,
and the support and encouragement of Dr. Clara L. Dixon of Albion
College and Dr. C. E. Huntington, director of the Bowdoin Science
Station on Kent Island. This paper is published with the approval
of the Director of the Bowdoin Science Station as Contribution
No. 48.

196

1975] Kent Island — McCain 197

All of Kent Island is now covered with water-worked
glacial till (Meserve, 1937). The only major relief is an
upland area with twenty-foot sand cliffs on the northern
end of the island and two fifty-foot hills at the far southern
end. The northern and western ends are heavily wooded,
and the west end is low and marshy. A 7.8 acre tidal basin
on Kent Island is flooded twice daily by tides averaging
14.4 feet. Kent is the largest of what local fishermen call
the Three Islands. Fifty acre Hay Island is forested except
for one large field on the south shore where a few now
uninhabited houses and a weathered barn still stand. It
can be reached on foot from Kent Island at low tide. Thirty
acre Sheep Island is a low flat island with vegetation resem-
bling the south end of Kent Island. It has no woody plants.

MATERIALS AND METHODS

I was able to spend six weeks during the summer of 1968
and ten weeks during the summer of 1969 on Kent Island
at the Bowdoin Science Station. In 1968, a general survey
of the plant cover of the entire Three Islands group was
made, and quantitative sampling of the vascular flora of
the forests was completed. Aerial photographs taken in
1965 were used to supplement the field studies in mapping
the vegetation of the islands (Fig. 1). To determine the
relative frequency of the forest herbs, a 100-foot grid was
established in the wooded areas of Kent Island using a
Brunton compass, and meter quadrats were set up at the
intersections and at the 50-foot marks.

The non-forested areas, identified in the 1968 survey,
were sampled in 1969 by the line-intercept method. А non-
forested area, as defined for the purposes of this study, is
one in which herbs or shrubs are the dominant species and
where trees, if present, do not form a closed canopy. Eleven
such areas were defined. In each area, either a two- or a five-
hundred foot transect was established along a compass line.
Absolute and relative dominance, and relative density were
calculated for plant species intercepted, according to the

198 Ећодога [Vol. 77

HAY ISLAND

BASIN ——

POND

SHEEP ISLAND

KENT ISLAND
[ | FiELDS N
МУ SHRUBS \
MARSHES 1000 FT. \|
: GULL HILLS d
WOODED FIG.I- VEGETATION MAP, KENT IS.

1975] Kent Island — McCain 199

method of Cottam and Curtis (1962). Absolute dominance
is expressed in this paper as "ie cover", a measure of the
number of herbs or shrubs of a given species intercepting
each segment of the transect line. Since several different
species may overlap one another, the total “% cover" will
often exceed 100%. Relative frequency was determined
through the use of nested quadrats offset by а random
numbers table at twenty-five foot intervals along the tran-
sect line. The sum of relative frequency, relative density,
and relative dominance (of woody species) is the Impor-
tance Value (IV) of a species on a given transect. The
sum of all species’ Importance Values on each transect
should equal 300 for shrubs, 200 for herbs.

Soil samples were taken from the A, layer at each
sixty-five foot mark, and used to determine soil moisture
and water retaining capacity (WRC) by the Hilgard-cup
method, and pH by the use of pHydrion paper. In soils
with high organie content, the WRC values often exceed
the dry weight of the soil itself.

Seymour (1969) was used for the identification of the
plant specimens with the exceptions of the two groups
indicated in the previous paper on the Kent Island flora
(McCain, Pike & Hodgdon, 1973).

RESULTS

Collections totaling 240 species and 10 varieties of vascu-
lar plants were made on the Three Islands in 1935 by
Dr. David Potter of Clark University, in 1968-69 by the
present author, and in 1970 by Dr. A. R. Hodgdon and
R. B. Pike of the University of New Hampshire. These
collections were listed in McCain et al. (1973). Ninety-four
of the genera in the total list (66.2%) are represented by
one species only, as are twenty-one of the families (39°).
Of the fifty-four families, six (Gramineae, Cyperaceae,
Polygonaceae, Caryophyllaceae, Rosaceae, and Compositae)
account for 48% of the species present on the island, and
for 100% of the species intercepted in the nesting areas of

200 Rhodora [Vol. 77

the islands’ sizable herring gull population. The presence
of these birds may be the main cause of the lack of floristic
diversity.

Eighty-five one-meter quadrats sampled in the forested
areas showed a ground layer homogeneous in species com-
position but widely varying in density of individuals. In
the more open areas, or along the forest edge, Dryopteris
spinulosa var. americana was clearly dominant and the
total higher plant cover averaged 88.13% under spruce-fir
and 70.56% under mountain ash and canoe birch (Betula
papyrifera var. cordifolia). Where the spruce are 20-30
feet tall, they form a very dense thicket. Here the higher
plant cover averaged only 32.86%, with Oxalis montana
and Maianthemum canadense dominant. In several of the
denser spruce thickets, higher plant cover was entirely
absent in the herb layer. The understory there suffered
not only from a lack of light but also from a shortage of
soil moisture. Most rain is intercepted by the dense canopy
and most of the throughfall is caught by the deep litter
layers before it can penetrate to the soil itself where it
would be available for the understory plants. Mean higher
plant cover in all eighty-five quadrats was 55.17%. Lower
plant cover averaged 31.43% in the dense areas, 10.89%
under the open stands.

The most important herb species in the wooded areas was
Aster acuminatus (%F = 60.0). Rubus idaeus var. cana-
densis was the most common woody species, occurring in
28.2% of the quadrats. In openings or blowdowns, it often
accounted for nearly 100% of the cover. Table 1 shows
the relative frequency values for the forest floor species.
Gleason (1937) sampled one hundred similar plots on Kent
and Hay Islands, finding corresponding values for the spe-
cies present. The major exception was that Oxalis montana
was the most abundant species in his study, occurring in
44% of his quadrats. In 1968, O. montana was present in
24.7% of the quadrats in the total sample, but was present
in 71% of the quadrats in denser areas where the spruce
was roughly 25-40 years old. This change in the presence

1975] Kent Island — McCain АС

of О. montana suggests a change in the age structure of
the tree population, with the sorrel giving way to larger
species, often Dryopteris or the mountain aster, in the older,
less dense woods.

Table 1. Kent Island species occurring in more than 5%
of forest area quadrats sampled, 1968.

Species % frequency
Shrubs
Rubus idaeus L. var. canadensis Richardson 28.2
Herbs

Aster acuminatus Michx. 60.0

Dryopteris spinulosa (O. F. Mueller) Watt. 38.8
var. americana (Fischer) Fern.

Rumex acetosella L. 38.8

Oxalis montana Raf. Se

Trientalis borealis Raf. 21.1

Maianthemum canadense Desf. 16.4

Cornus canadensis L. 10.2

Poa pratensis L. 9.4

Circaea alpina L. 9.4

Impatiens capensis Meerb. forma immaculata 8.3
(Weath.) Fern. & Schub.

Stellaria media (L.) Cyrillo 8.2

Anthoxanthum odoratum L. 7.1

Carex brunnescens (Pers.) Poiret 5.9
var. sphaerostachya (Tuck.) Kukenthal

Streptopus amplexifolius (L.) DC. 5.9

var. americanus Schultes

Four groups of non-forested stands can be clearly defined.
The first includes the old fields near the middle of Kent
Island and the main field on Hay Island. All are composed
of turf grasses like Agrostis stolonifera and Festuca rubra
and perennial composites such as Solidago rugosa var.
villosa. These fields are relatively sheltered both from the
salt spray and the weather but show the greatest influence

202 Ећодога [Vol. 77

of man both in terms of cutting and of introduced species.
Most of these areas were cleared for planting of forage
crops, notably Phleum pratense, in the nineteenth and early
twentieth centuries. Shrub cover is uniformly low: 1.0,
14.40, 16.60, and 16.71% on the transects studied. Herb
cover is the highest of the four groups, with an average
in these fields of 386.37%. The old field transects all in-
cluded a greater number of plant species than any of the
other transects studied.

The second group consists of raspberry and shrub com-
munities. Epilobium angustifoliwm is found in association
with Rubus idaeus var. canadensis in blowdowns and open
areas along the north and east shores of Kent Island. Shrub
cover values are uniformly high with a mean of 103.96%,
but herb cover dropped to 180.3796. Each of these areas
is the site of active gull nesting, and, as a result, has a very
rich soil with a high organic content.

Fresh-water and salt marsh communities along the west-
ern shore of Kent Island and on the north shore of Hay
Island are in the third group. Iris versicolor (Importance
Value — 19.46), Carex canescens var. disjuncta (IV —
23.18) and Epilobium glandulosum var. adenocaulon (IV
— 20.65) are the most important species of these areas.
Shrubs, but for Ribes lacustre, are seriously limited, except
where the marsh is the last remnant of an old raised bog.
In those places, Kalmia angustifolia and Ledum groenlandi-
cum are the dominants, growing over Sphagnum moss.
Only one small part of a formerly very extensive bog is
left on Kent Island because of the maturing of the forests
and man-made changes in the drainage patterns. There is
one heath remnant, perhaps twenty feet in diameter, at the
north edge of the bog area, the sole location found for five
species of the flora: Calopogon tuberosus, Arethusa bulbosa,
Sarracenia purpurea, Rubus chamaemorus and Empetrum
nigrum.

In the fourth group, the South Hills of Kent Island and
all of Sheep Island, are the main nesting areas of the
islands’ estimated 23,000 breeding pairs of herring gulls.

1975] Kent Island — McCain 203

This very disturbed area supports dense populations of
several plant species found only infrequently elsewhere on
the island, and then only as weeds. For example, common
ragweed (Ambrosia artemistifolia) normally loses out in
competition to the grasses and other composites. On the
South Hill, however, Ambrosia has an importance value of
58.02, second only to Aster foliaceus var. arcuans (ТУ =
64.62). Rumex acetosella, another weed, is similarly fav-
ored by these disturbed conditions (ТУ = 14.28). The
number of species found on the South Hill is also un-
doubtedly held down by the gulls. Only eight herb species
and one shrub (Rubus idaeus var. canadensis) appeared
on the 500 feet of South Hill transects. By comparison,
the four old field transects averaged 26 herb species and
3 shrub species.

Clearly the most frequent herbaceous species of the non-
forest areas is the ubiquitous blue aster A. foliaceus var.
arcuans. Оп all four field transects, on the north shore of
Hay Island, and on the gull hills at the south end of Kent
Island, it is the species with the highest importance value.
Aster foliaceus was present in some degree in all eleven of
the transect samples.

The most frequent shrub species, in both forested and
non-forest areas, is Rubus idaeus var. canadensis. Rasp-
berry plants covered over half of the line on the four shrub
community transects: the south field (%cover — 92), North
Clearing (86.5%) Downer’s Field (51%), and the Crock-
ett’s Point Field (69%). No other shrub was so clearly
dominant in any of the areas sampled. Ribes lacustre was
a dominant shrub (18% cover) in the Hay Island north
shore marshes, but, where Ribes and Rubus are both pres-
ent, the red raspberry is clearly more dominant than the
currant.

Soil moisture values vary from 19.5% in the North Field
to 69.0% on the South Hill (Table 2). The South Hill also
varies significantly in mean pH values from the rest of the
island. The other groups of stands have mean pH values
of 5.67, 5.77 and 5.60, while that of the South Hill is only

204 Ећодога [Vol. 77

4.93. WRC values are consistent within each of the four
groups, except for a high of 629.90% on Crockett’s Point
where much poorly decomposed Sphagnum moss appeared
in the soil profile. Soil data is summarized in Table 2. The
extreme values for soil moisture and pH in the South Hill
are certainly factors in the presence of a vegetation typical
of disturbed areas. However, careful study may show that
the mechanical destruction of plants due to the perching,
pulling and pecking of the gulls may be more harmful to
plant survival than the chemical alteration of the soil
caused by their droppings.

Table 2. Soil analysis data.

Tran- % Soil WRC
Stand sect Moisture pH (%)
North Field I. 19.5 6.05 121.18
Hay Field II. 26.2 5.58 148.50
West Field III. 29.8 5.40 82.96
East Field IV. 34.4 5.38 61.95
North Clearing V. 28.0 5.80 105.55
South Field VI. 34.7 5.70 22.64
Downer's Field VII. 39.5 5.80 29.42
Crockett's Point VIII. 41.0 5.80 629.90
West shore IX. 40.0 5.70 425.44
Hay north shore marsh X. 40.2 5.50 300.00
South Hill XI. 69.0 4.93 196.53

Mean Values

Transect Group
I-IV (Old fields) 27.48 5.67 103.50
V-VIII (Shrubs) 35.80 5.77 *52.54
IX-X. (Marshes) 40.10 5.60 362.72
XI. (South Hill) 69.00 4.93 196.53

*discounting high value for transect VIII.

1975] Kent Island — McCain 205

The remaining botanical area of the islands is a zone just
above the highest tide lines, favored mostly by halophytes,
the most important among them apparently being Cakile
edentula, Atriplex patula var. hastata and A. glabriuscula,
Lathyrus japonicus var. pellitus, Mertensia maritima, and
Arenaria peploides var. robusta. There are also a few
perennials mixed in at the upper edge of this zone, notably
Aster foliaceus and Urtica procera. A few plants have
ventured out onto the floor of the tidal basin where they
are covered twice daily by the waters of high tide: Sali-
cornia europaea, Glaux maritima, Limonium nashit, Suaeda
maritima and S. americana. Prominent growths of Spar-
tina alterniflora and Hordeum jubatum line the edge of the
basin. No statistical sampling has yet been done in these
zones.

DISCUSSION

Three characteristics of the flora are especially worth
noting. The first is the paucity of species. About one-half
of the genera (56.44%) and one-fourth of the families
(27.37%) in mainland New Brunswick are represented by
a single species (Fowler, 1885). On Kent Island, two-thirds
of the genera and two-fifths of the families are monotypic.
The recentness of colonization could be a factor in the lack
of diversity of flora, for all of the Grand Manan Island
group was covered during the last glaciation. Thus there
has been little time for proliferation of immigrants. The
second possible factor is the presence of the herring gull
colony. In work on three similar islands nearby in the Bay
of Fundy, Hodgdon and Pike (1969) concluded that “the
screening effect of bird populations is more apparent than
their transport of propagules". This seemed to be true on
Kent Island where the gulls probably prohibit by their
activities the establishment of many potential immigrants
and of many woody species. The continual disturbance
caused by the birds maintains the area in early stages of
succession, The abundance of a group of fast growing, sun

206 Ећодога [Vol. 77

loving species typical of pioneer habitats, including Achil-
lea millefolium, Rumex domesticus and R. acetosella, Urtica
procera, Coelopleurum lucidum and Ambrosia artimisiifolia,
indicated that these plants apparently benefited from the
gulls’ presence.

The second notable characteristic of the flora is the pres-
ence of a high number of boreal species for this latitude.
Nearly three-fourths of the Kent Island species have ranges
that extend north and west from this area. Such species as
Rubus chamaemorus, Empetrum nigrum, Vaccinium vitis-
idaea, Stellaria humifusa, and Euphrasia americana reach
their extreme southeastern limits in the Kent Island region.
The reason for their presence here is the cool, damp climate
of the Bay of Fundy. The warm Gulf Stream and the cold
Labrador Current meet near the mouth of the Bay. During
the summer the prevailing wind is from the south or south-
west over 50% of the time. Thus the air is considerably
warmer than the waters of the Labrador Current and of
the Bay of Fundy (which seldom exceeds 50°F. even in
midsummer) and the result is very frequent dense fog.

In the ten-day period beginning 26 July 1969, for exam-
ple, the greatest visibility was just under one mile and the
mean daily low visibility was about one hundred feet. The
fog-drip caused by the condensation of this heavy fog on
tree limbs increases the humidity at the herb level.

Table 4 shows pertinent weather data for the years
1965-69. Year-round weather records for the five-year
period 1937-1941 may be found in Cunningham’s report
(1942).

The third important characteristic of the flora is that
many of the new species reported in 1968-69 are weeds
evidently introduced by man, probably after the previous
studies were done in 1936. Thirty-one of these sixty-one
new species were found in the field where the houses are
on Hay Island, and on Kent Island near the wharf, along
the main path, or in the area of the dormitory in the cen-
tral part of the island. Conspicuous among these species
are Trifolium pratense, Convolvulus sepium, Chenopodium

1975] Kent Island — McCain 207

album, Galeopsis tetrahit, Chrysanthemum leucanthemum,
and Hieracium aurantiacum. These species are often spread
by man, and are too conspicuous to have been missed by
Potter in his work.

Nearly all of Kent Island was once wooded and has been
cut over at one time or another (Gleason, 1937). The
northern and western ends are now wooded, with white
spruce (Picea glauca) dominant in successional areas
where the forest is encroaching on the fields and in blow-
down areas. Balsam fir (Abies balsamea) is reproducing
better in the more established woods.

Table 3. Herbaceous and woody species cover.

Shrub cover Herb cover

Stand Transect (%) %

North Field 1. 16.71 421.56
Hay Field П. 1.00 489.50
West Field ПІ. 16.60 317.40
East Field IV. 14.40 317.00
North Clearing V. 86.50 212.50
South Field VI. 184.00 93.00
Downer’s Field VII. 69.00 203.00
Crockett’s Point Field VIII. 16.33 212.97
West shore marsh LS 10.00 389.00
Hay north shore marsh X. 7.50 156.00
South Hill XI. 16.80 175.40

Mean Values

Transect group

I-IV (Old fields) 12.18 386.87
V-VIII (Shrubs) 103.96 180.37
IX-X (Marshes) | 8.75 272.50

XI. (South Hill) 16.80 175.40

208 Ећодога [о]. 77

Table 4. Meteorological summary of Kent Island, New
Brunswick, Canada, 1965-1969.
(Bowdoin Science Station Weather Records)

Avg. Mean Avg. Mean Avg. Total Avg. Days

Month Max. Т.°Е. Min. T.?F. Prec., in. Fog/Month
June 57.2 44.6 2.49 13.0
July 61.8 48.8 2.44 17.8
August 62.3 50.5 3.08 15.6

Avg. total days with fog for the three month period: 46.4.
Mean annual precipitation (Cunningham, 1942): 44.86”.

The four classes of non-forested areas vary in terms of
soil water and plant cover. Table 2 lists soil moisture con-
tent, water retaining capacity, and pH by transects and
Table 3 lists woody and herbaceous species cover by tran-
sect. In the old fields, the driest areas studied, the herba-
ceous species seemed to be encouraged. As the moisture
content of the soil increased, the shrubs increased, but the
total water the site could hold decreased as did the number
of herbs. In the marshes the water content became high
enough to inhibit the number of shrubs, and another group
of herbaceous species took over.

Finally, the highly organic soil of the South Hill had a
high moisture content, but a lowered WRC. This is sur-
prising since water retaining capacity usually increases
with increased soil organic content. The increased acidity
of these areas may have limited both the herbs and shrubs
in variety. Where the gulls nest in areas with trees, they
are destroying the woody plants. The south hills of Kent
Island were formerly forested, as is evidenced by many
fragments of decaying wood found in the soil, but the con-
stant influence of the gulls has left only an area of twisted
and broken snags where Rubus idaeus has now taken over.
Many trees have been killed by the severe mechanical dam-
age caused by the gulls. Some others have become stunted

1975] Kent Island — McCain 209

and have formed an unnaturally dense and tangled crown,
thick enough and strong enough to support a man.

In some of the most heavily disturbed areas, the grass
pulling of the gulls keeps large areas of ground completely
barren of all vegetation. The largest such barren area in
1969 measured nearly fifty feet by thirty feet. The vegeta-
tion around these barren areas is almost purely Ambrosia
artemisiifolia. This weedy species emphatically underlines
the extreme disturbance by the gulls of this part of the
island. Nowhere else in the Three Islands group are con-
ditions nearly so favorable for this species.

LITERATURE CITED

CoTTAM, G., & J. T. Curtis. 1962. Plant ecology workbook. Burgess
Publishing Co., Minneapolis. 193 pp.

CUNNINGHAM, R. 1942. Meteorology. Bull. Bowdoin Sci. Sta. 6:
10-13.

FOWLER, J. 1885. Preliminary list of the plants of New Brunswick.
Bull. Nat. Hist. Soe. New Brunswick 4: 8-84,

GLEASON, Н. А. JR. 1937. А botanical report on Kent's Island.
Bull. Bowdoin Sci. Sta. 3: 27-29.
Норсрох, А. R., & R. B. Dr 1969. Floristic comparison of three
bird islands in the Gulf of Maine. Rhodora 71: 510-523.
McCAIN, J.. В. B. PIKE, & A. Е. Норсрох. 1973. The vascular
flora of Kent Island, Grand Manan, New Brunswick. Rhodora
15: 311-322.

MESERVE, F. W. 1937. Preliminary study of geology and geography
of Kent’s Island, 1935-1956. Bull. Bowdoin Sci. Sta. 3: 10-13.

Роттев, D. 1937. Plants found growing on Kent’s Island, 1935-
1936. Bull. Bowdoin Sci. Sta. 3: 19-26.

Seymour, F. C. 1969. The flora of New England. Chas. E. Tuttle
Co., Rutland. 596 pp.

WEATHERBY, C. A. & J. ADAMS. 1945. A list of the vascular plants
of Grand Manan, Charlotte County, New Brunswick. Contrib.
Gray Herb. Harvard Univ. Мо. 158. 96 pp.

DEPARTMENT OF BIOLOGY
ALBION COLLEGE
ALBION, MICH. 49224

А RED-PETIOLED FORM OF THALIA GENICULATA 1.
FROM CENTRAL FLORIDA

ALLEN G. SHUEY

In 1967 several unusual plants of Thalia geniculata were
noticed growing in a wet disturbed area caused by the
widening of State Road 192, located ten miles west of the
town of Melbourne Village, Brevard County, Florida. These
Thalias had become established along with other herbaceous
aquatics to the north of the road after construction on the
road had ceased. The disturbed site was of very sandy soil
which often became flooded by heavy rains. These plants
were noticeably different from the usual all-green form of
Thalia by the presence of bright red petioles and petiole
sheaths.

For unknown reasons the plants disappeared and it was
not until 1972 that the red form was again seen in the same
vicinity, this time in the ditch to the south of State Road
192 and west of where it was originally sited. Additional
searching later uncovered several small populations to the
east and west of the population found in 1972. The same
year a stand of red-petioled Thalias was seen in a cattle
pasture to the east of the Military Trail, northwest of the
city of Palm Beach, Palm Beach County, Florida. This
population has since been destroyed.

The most striking aspect of the red form of Thalia
geniculata is the bright rhubarb red coloration of the
lower petiole and petiole sheaths. This coloration is carried
farther by the presence of a red pulviniform area (usually
greenish or orangish in the green form) at the base of the
leaf blade, and by varying degrees of red on the joints and
axes of the inflorescence. Red may also occur at the margin
of the upper side of the leaf blade.

In the green form of Thalia geniculata, no red coloration
occurs except for a red spot which may be found on the
upper surface of the leaf blade where the blade meets the
pulviniform area at the top of the petiole.

210

1975] Thalia — Shuey AW

Thalia geniculata L. f. rheumoides Shuey, forma nova

А forma geniculata differt basibus vaginisque petiolorum
rubris et inflorescentiae furcis, axibus, bracteisque princi-
palibus rubris vel viridi-rubris, necnon area pulviniformi
ad apicem petioli cremea usque rubra.

Holotype: Florida: Brevard Co.: on south slope of water
filled ditch, base of plant at water line, growing in full sun
with grasses, other herbaceous plants, and the green form
of Thalia geniculata, south side of State Road 192, 10 mi.
west of the town of Melbourne Village, Shwey 105571
(USF).

The form name rhewmoides is derived from Rheum, the
generic name for rhubarb, and oides, Greek suffix meaning
similar to; this was selected because the red stem sheaths
are rhubarb red, looking like the petioles of the red vari-
eties of rhubarb. This form of Thalia is capable of per-
petuating itself — numerous small seedlings exhibiting red
coloration being noted at several sites.

The coloration of the petiole stalks and sheaths does not
vary to any great degree from one plant to another, but
the color of the axes of the inflorescence varies considerably.
Some plants exhibit slight coloration only around the joints,
bracts and bases of the inflorescence, while others tend
toward conspicuous coloration over much of the axis.

The coloration of the plants is very much influenced by
light intensity. Plants growing in full sun have deeply
colored petiole stalks and sheaths. Plants growing in the
shade, on the other hand, have sheaths which are almost
green, but retain the distinct red color near the margin of
the sheath. This is lacking altogether in the green form.

It is interesting that several plants dried for herbarium
material retained a strong red tinge around the joints of
the inflorescence and especially on the stem sheaths.

I wish to extend a note of thanks to Dr. Haven C. Sweet,
Assistant Professor of Biological Studies, Florida Techno-
logical University, for valuable help in preparing the text

212 Ећодога [Vol. 77

of this paper. Also, to Dr. Henry O. Whittier, Assistant
Professor of Biological Studies, Florida Technological Uni-
versity, and to Dr. С. S. Smith (Department of Ornamental
Horticulture, University of Florida) and Dr. D. Ward
(Director of the Herbarium, University of Florida) for
supplying valuable information.

DEPARTMENT OF BIOLOGICAL SCIENCES
FLORIDA TECHNOLOGICAL UNIVERSITY
ORLANDO, FLORIDA 32816

THE TAXONOMY OF TRIPOGANDRA
(COMMELINACEAE) '

WAYNE L. HANDLOS

INTRODUCTION

Although some species of commelinaceous plants —
Tradescantia and Zebrina — are very well-known in intro-
ductory courses in biology and botany for their use in
demonstrating cyclosis, plasmolysis, and anther squashes,
and while cytologists consider species of Tradescantia good
teaching material and popular research subjects because of
their large chromosomes, taxonomists have not agreed as
to generic limits or relationships within the family. The
problems of generic delimitation within the Commelinaceae
have existed almost since the time of Linnaeus. Concepts
have changed through time and even a careful and ob-
servant worker such as C. Kunth modified his concepts of
the genera during his working years as is evidenced by
his transferral of species from one genus to another. In
recent years the publications of Aristeguieta (1965),
Hutchinson (1959), Matuda (1956), Moore (1960, 1963),
Pichon (1946), Rohweder (1956), and Woodson (1942)
illustrate the different concepts each investigator has of
various genera, especially those American genera related
to Tradescantia.

Tripogandra has been included in what has been called
the Tradescantia alliance. The United States species of
this alliance were studied by Anderson and Woodson
(1935) and were found to form a uniform group of species.
Tradescantia outside of the United States contains diverse
elements, and a study of Tripogandra was considered in
the nature of an introduction to the whole alliance. As

1Based on a thesis submitted to the Graduate School, Cornell Uni-
versity, in partial fulfillment of the requirements of the degree of
Doctor of Philosophy. This study was supported in part by National
Science Foundation Grant GB-6277.

213

214 Ећодога [Vol. 77

delimited by Woodson (1942) on the basis of inflorescence
structure, Tripogandra was not well understood and con-
tained a number of species of questionable affinity. Moore’s
(1960) investigations showed that some species had a
modified androecium which suggested an important bio-
logical function probably related to insect attraction, as-
suring pollination and out-crossing. A detailed study of
the species included in Tripogandra was thought to be
one way of approaching the problem of generic delimita-
tion and of determining the relationship of Tripogandra
to its closest relatives and to the remainder of the family.

Here I have delimited Tripogandra to include only those
20 species which have dimorphic stamens and double cin-
cinni not subtended by foliaceous bracts. These species are
described in detail while the other species which have been
included in Tripogandra at some time in the past are not
considered congeneric and are not described in detail.
These species and reasons for exclusion are to be found
at the end of this paper.

HISTORICAL REVIEW

Tripogandra was first proposed by Rafinesque in 1837
to contain one species, Tradescantia multiflora. Rafines-
que’s publications were not widely available апа his
taxonomic decisions were not always accepted by other
botanists. As a result, the name Tripogandra wes not
adopted by the botanical community until much later.
When Rafinesque published Tripogandra he also proposed
the following as segregants from Tradescantia: Sarcoperis,
Siphonstima, Gibasis, Etheosanthes, Tripogandra (based
on Tradescantia multiflora Jacq.), Phyodina, Leiandra,
Heminema (based on Tradescantia multiflora Swartz), and
Aploleia. Rafinesque conciuded his proposals with the fol-
lowing comments:

“These 3 last Genera [Leiandra, Heminema, Aploleia]
lack the very essential characters of bearded Stam. that
once was the only distinction of Tradescantia from Com-

1975] Tripogandra — Handlos 215

melina, but these 2 Genera are in utter confusion, as the
above proves. Compare also my genera of Commelina, It
is deplorable to see Botanists forcing sp. into genera, in
spite of characters. There is not a single generic character
common to all the above G. 22 to 32! my reform and
revision were indispensable, and begun in 1815. ... this
whole Genus is a mass of linnean errors.”

Rafinesque was correct by present criteria in believing
that several generically different elements were present in
Tradescantia, but he did not solve all the problems. Both
Tripogandra and Heminema are based on the same type,
though Rafinesque credits two different authors. Rafines-
que may have been misled by the apparent though not real
differences between Swartz’s original description and
Jacquin’s description and illustration. Woodson (1942)
united Heminema and Tripogandra and his choice of
Tripogandra must be followed according to Article 57 of
the International Code of Botanical Nomenclature (1966).

Kunth in 1843 considered the entire family and described
sixty species of Tradescantia which he divided into several
groups, one being “Species anomalae." This group was
characterized by anthers of two shapes, with three longer
and three shorter filaments. All of the species in this
group are presently considered in the genus Tripogandra.

Schlechtendal (1853) proposed the name Descantaria
for the species which Kunth included in his “Species
anomalae.” Schlechtendal wrote that those species seen
by him were distinguished by three bearded perfect sta-
mens and three beardless imperfect stamens. No com-
binations were made by Schlechtendal however.

In 1866, Hasskarl used Disgrega as a generic name in
a key with Tradescantia disgrega in parentheses probably
indicating that this species belonged to the genus Disgrega,
but no formal transfer of the species was made.

Clarke (1881) dealt with the Commelinaceae as a family.
He divided the genus Tradescantia into three sections, one
of which was Descantaria, characterized by three longer

216 Ећодога [Vol. 77

and three shorter stamens more or less dissimilar. Clarke
(in Donnell-Smith, 1902) described the genus Donnellia
characterized by a three-valved capsule, locules bearing
one seed, and based on Callisia grandiflora which was
originally described as having two groups of dissimilar
stamens. He was prompted to write that in his previous
work on the family

“по attempt is made to deal with the genera ‘logically,’ the
same characters, which in the American genera (Tra-
descantia and its allies) constitute genera, only constitute
subgenera in Commelina and Aneilema. . . . convenience
has been preferred to any logical system."

Donnellia was found to be a later homonym and Rose
(1906) proposed the name Neodonnellia for the genus.

Clarke's work has caused concern among taxonomists
and Anderson and Woodson (1935) note that Clarke's

“revision of the genus Tradescantia reflects little credit
upon the author, when compared with the critical, if ec-
centric observation of Rafinesque nearly a half century
previously. Not only was Clarke satisfied to ignore the
complexity of the genus by unwarranted reducing to strict
synonymy or ambiguous varieties most of the species pro-
posed by his predecessors, but the literature was compli-
cated by the publication in synonymy of unpublished names
of other botanists, and the misinterpretation of numerous
others. The revision of the genus in the ‘Monographiae’ is
clearly a piece of chorework reflecting rather a none too
laborious attempt at compilation than a lively interest and
acuity."

Briickner (1927) chose Schlechtendal’s name, Descan-
taria, as the generic name for several of the species now
included in Tripogandra and formally transferred those
species. Brückner had suggested these transfers in 1926
when he had published a complete description of Descan-
taria and gave Schlechtendal credit for proposing the name.
No synonyms were given by Brückner until 1930, when

1975] Tripogandra — Handlos 217

he listed Descantaria Schlechtend., Heminema, Tripagandra
[sic] Rafin., and Disgrega Hassk. It is not clear why
Briickner chose a name which lacked priority. A possible
reason is that he had no first-hand knowledge of Rafines-
que’s publication. Briickner’s misspelling of Tripogandra
is the same as that in Index Kewensis, fase. 4, which may
have been Briickner’s only acquaintance with the name.

The American members of the Commelinaceae were
treated by Woodson (1942) in an attempt to better delimit
the genera. A number of transfers were made in the newly
delimited genera, and Tripogandra was discussed at length.
Woodson suggested new approaches to the family, stating
that

“the Commelinaceae always have been difficult subjects
for herbarium study because of their deliquescent flowers.
It is not easy to understand, therefore, why previous sys-
tematists of the family have focused almost their whole
attention upon floral structure in the delimitations of sub-
families, tribes, and genera.”

Woodson felt that stamens were too variable a character
to use for a major subdivision of the family and pointed
out Brückner's inconsistency in dividing the family into
two subfamilies and then placing Descantaria in the Hex-
andrae and Neodonnellia in the Triandrae. І believe
Woodson was correct in considering Descantaria and Neo-
donnellia as congeneric. Woodson suggested that a major
subdivision of the family could be made using inflorescence
structure. In his concept of the tribe Commelineae, the
ultimate units of the inflorescence are individual scorpioid
cymes while in the Tradescantieae the basic structures “аге
paired sessile scorpioid cymes which appear as a 2-sided
unit superficially, . . ." Woodson’s concept of Tripogandra,
which was placed in the Tradescantieae, was based pri-
marily on inflorescence structure so it included more species
than I have included. Woodson included species of Lep-
torhoeo and Cuthbertia which have six similar stamens
because these species lacked foliaceous bracts subtending

218 Rhodora [Vol. 77

the paired cymes. Previous authors and I have considered
the two dissimilar whorls of stamens and their position
as characters which delimit the most natural grouping
of species.

Briickner (1930), Hutchinson (1934, 1959), Woodson
(1942), Pichon (1946), Rohweder (1956), and Brenan
(1966) have all dealt with the problems of generic defini-
tion and the characters used within the family to delimit
them. Different emphases were used by each author but
in general a consideration of several characters was found
necessary for more satisfactory treatments. Tripogandra
and some of its species have been considered in recent
years in studies of restricted scope by Macbride (1936,
1944), Standley and Steyermark (1944, 1952), Matuda
(1956), Moore (1960), Bacigalupo (1964, 1967), and
Aristeguieta (1965), all of whom have been influenced
to some degree by Woodson’s work.

The genera of the Commelinaceae have been divided
among various groups by the different authors listed above.
No general consensus has been reached and all schemes
are unsatisfactory to some extent. Brenan (1966) sum-
marized the earlier schemes of classification and has di-
vided the genera into fifteen groups. Tripogandra is con-
sidered in Group XI, which includes Rhoeo, Campelia,
Callisia, Aploleia, Tradescantia, Phyodina, and Cymbis-
patha. Using Brenan’s criteria, Tripogandra seems more
closely allied to the species of this group than any other.

MORPHOLOGY

The morphology of Tripogandra has been investigated
to varying degrees by taxonomists who have used certain
characters in classifying the species, but the genus has
never been systematically investigated by a morphologist.
Tomlinson (1966) has been a primary investigator of
morphology and anatomy having looked at the epidermis,
hairs and stomatal patterns of three species. Brückner
(1926) described in some detail the species 7. glandulosa

1975] Tripogandra — Handlos 219

(as Tradescantia pflanzii). More recently Rohweder
(1963а) investigated shoot development and the course of
vascular bundles near the apex of Tripogandra pflanzii
(—T. glandulosa).

The following observations have been made on herbarium
specimens and on living plants collected in México or grown
in the greenhouse.

Habit. The piants of Tripogandra, whether annual or
perennial, are succulent, weak-stemmed herbs. The plants
may have an erect stem which is either branched or not.
In erect, annual species — T. amplexicaulis, T. angustifolia,
T. guerrerensis, and T. palmeri — branches may or may
not develop from axillary buds. Plants in sunny, moist
locations branch frequently. Crowded plants or those in
shaded locations are commonly unbranched. Because the
base of the plant is small and there is no secondary growth
it is unable mechanically to support the weight of later
growth. The base of the plant, therefore, becomes decum-
bent and additional support and anchorage is gained by
the production of adventitious roots at the nodes of the
decumbent stem. The presence of an intercalary meri-
stem at the base of each internode allows the main stem
to remain upright through differential growth and bending
in these areas.

Some species, e.g. Tripogandra disgrega, T. saxicola,
and T. purpurascens, may be either erect or trailing. This
habit seems to be partly under genetic control; some plants
have sturdier stems and grow upright while others produce
weak, flexible stems which trail over the ground.

Perennial species such as Tripogandra montana and T.
serrulata, commonly trail over the ground to a length of as
much as two meters, producing adventitious roots at nearly
every node which touches the soil The flowering stems
usually are erect. Branching occurs at irregular intervals.
A vegetative branch is often produced from a node below
the inflorescence. After growth of this axillary branch, the
inflorescence appears to be borne in a lateral position, but

220 Ећодога [Vol. 77

close examination shows the vegetative shoot to be in an
axillary position and the inflorescence to be terminal.

Tripogandra grandiflora produces long upright shoots
which are often supported by surrounding woody vegeta-
tion. Sheathing bracts or cataphylls are borne on the lower
portions of the stem. Structures transitional between the
cataphylls and a typical vegetative leaf can be found on
most stems.

Root. All species of Tripogandra have fibrous roots.
Thickened storage roots such as are found in species of
other genera, ie. Tradescantia, Gibasis, Setereasea, Sepa-
rotheca, Dichorisandra, and Commelina, have not been
observed in any species of Tripogandra. The roots may
be produced only at the base of the plant or they may be
produced adventitiously from nodes along the length of
the stem. Root initials may be differentiated early and
remain dormant as has been reported by Tomlinson (1969)
for species of Tradescantia.

Stem. The stems of all species of Tripogandra are more
or less succulent structures. The stems as well as most
other organs contain a mucilaginous juice which is exuded
when the structures are crushed or broken. А common
feature of most species is the presence of a line or band
of hairs extending down one side of the internode (Fig. 4).
The hairs within this line are always uniseriate, i.e. com-
posed of several cells joined end to end. The terminal cell
is not enlarged or modified in shape from the cells below
it. This line of hairs is continuous with the line of hairs
present on the sheath of the leaf above and is always
found on the side of the stem opposite the leaf blade of
that sheath. The line of hairs on the internode may extend
the full length of the internode as in T. saxicola, may
extend for part of the length of the internode as in T. ser-
rulata, or occasionally may be lacking as in T. grandiflora
or T. amplexicaulis.

Elsewhere hairs may be more or less uniformly scattered
over the surface of the stem, as in Tripogandra purpur-
ascens subsp. australis, or the stem may be glabrous as in

1975] Tripogandra — Handlos 221

T. grandiflora, When present, the hairs may be uniseriate
and resemble those in the line on the internode or they
may have an enlarged distal cell and be called capitate as
in T. encolea.

Leaf. The leaves of Tripogandra species are simple. The
base is sheathing, forming a complete tube which is closely
appressed to the internode. The leaf surface may be
glaucous as seen in T. amplexicaulis апа T. amplexans,
but is more usually bright green and shiny. The dorsi-
ventral blade is broad and flattened in all species except
T. angustifolia, where the lamina is C-shaped or terete in
cross-section and in 7. purpurascens where the lamina is
complicate. The base of the blade may be variously modi-
fied. It may be narrowed and subpetiolate as seen in some
collections of Т. amplexicaulis and Т. disgrega. In some
species, particularly on the stem just below the inflores-
cence, the base may be amplexicaul and surround the stem
producing a superficially perfoliate appearance as in T.
amplexicaulis, Т. amplerans, and T. encolea. In other
species — Т. diuretica, T. montana, and T. serrulata —
the leaf base is oblique, often rounded on one side and
cuneate on the other.

Arrangement may be spiral as in Tripogandra guerre-
rensis and T. disgrega or two-ranked (distichous) as most
obvious in T. grandiflora.

The leaves may be glabrous or variously vestite and the
vesture may be constant or variable within a species. The
uniseriate hairs may be distributed on the leaves in the
following fashions:

1. Both surfaces (abaxial and adaxial) as in some plants
of T. disgrega and T. diuretica;

2. Abaxially (dorsally) as in T. brasiliensis;

3. Adaxially (ventrally) as in T. saxicola;

4. On the ventral surface over the midvein as in T.
serrulata and T. montana;

5. Adaxially in a line near the margin as in some plants
of T. multiflora.

222, Ећодога [Vol. 77

Distribution of hairs is most variable in Tripogandra
multiflora and is described further under that species.

The margins of blades are usually ciliate with uniseriate
hairs except for some plants of Tripogandra grandiflora
and T. angustifolia. Marginal hairs may vary in shape
from species to species and have been illustrated for
several species by Bacigalupo (1967). The distribution
of marginal hairs may be regular or irregular, the latter
condition being most apparent in 7. angustifolia. The
length of the hairs varies within species.

The orifice of the leaf sheath is villous in most species,
although it may be glabrous in Tripogandra amplexicaulis
and T. grandiflora. The hairs are multicellular and uni-
seriate and intergrade with those of the leaf margin.

Tomlinson (1969) has reported that a strand of collen-
chyma extends along the leaf margin in Tripogandra. In
living material this strand can be seen as a light-colored
line and is especially obvious in T. grandiflora.

The upper epidermis is colorless and the cells may be
larger than any within the leaf. This phenomenon has
been reported by Brückner (1926) for Tripogandra pflanzii
in his description of leaf anatomy; I have seen these en-
larged cells in Т. angustifolia and Т. montana. Tomlinson
(1966, 1969) has found this specialized epidermis in several
genera and assumes the function to be that of water
storage.

The presence of silica bodies in the epidermis has been
noted by Briickner (1926) and Tomlinson (1966, 1969).

The stomata have been described by Briickner (1926)
and Tomlinson (1966, 1969) as having two accessory cells
adjacent to the guard cells.

A weakly developed palisade layer may be seen in Tri-
pogandra montana but is lacking in T. angustifolia.

Inflorescence. The basic inflorescence unit which ap-
pears throughout the family is the cincinnus according
to Brenan (1966). The cincinnus as defined by Rickett
(1955) is “а monochasium in which flowers appear alter-
nately to right and left along one side of a sympodial axis.”

1975] Tripogandra — Handlos 223

The cincinni of a plant may be arranged in various
patterns to form more complex, compound inflorescences.
Brenan (1966) has illustrated some of the inflorescences
in other genera.

The basic unit of the inflorescence in Tripogandra is
also the cincinnus, but throughout the genus two cincinni
occur fused together as in several other genera, e.g. Tra-
descantia, Setcreasea, Rhoeo, and Zebrina. The type of
inflorescence has been called the “einfache Wickel zu
zweien" by Brückner (1926), paired cymes or 2-sided cin-
cinni by Woodson (1942), '*Wickelpaare" by Rohweder
(1956, 1963b), paired cincinni by Moore (1963) and
dichotomous сутез by Mericle and Mericle (1969),

The constant, paired nature of the cincinni in several
genera has been described by Brenan (1966) as “а peculiar
fusion of each pair into a characteristic bifacial unit con-
crescent with its peduncle.” Brenan has used the term
"paired cincinni" to include structures which are bifacial
as in Setereasea and Zebrina, as well as geminate as seen
in Gibasis genicvlata where the inflorescence is character-
istically composed of two separate and discrete cincinni
each on an elongate peduncle.

I have used the term “double cincinnus" to refer to the
bifacial, two-sided structure of the inflorescence which
occurs in Tripogandra, Тће difference may be subtle but
"double" seems to imply more in the nature of fusion than
does “paired.” Brückner (1926) used the term “Doppel-
wickel" which could be directly translated as double cin-
cinnus. According to Rohweder’s (1963b) translation,
Briickner’s definition of Doppelwickel was а structure
“composed of two cincinni arising from the same node
and being opposite in а strict morphological sense.”
Rohweder and Brenan consider this concept erroneous.
My use of the term double cincinnus, if it arises through
the fusion of two separate cincinni, may conflict with
Briickner’s concept of the origin of this structure. The
origin of the bifacial structure has not yet been demon-
strated.

224 Ећодога [Vol. 77

The problem of terminology is further confused by
Rickett’s (1955) illustration of paired cincinni in Myosotis
scorpioides. 'The structure shown corresponds to Brück-
ner's Doppelwickel which I believe can be seen in the
atypical inflorescences sometimes produced in Gibasis kar-
winskyana. I have chosen the term double cincinnus in
lieu of a better term.

According to the definition of a cincinnus, each flower
is terminal and the continuing axis is always an axillary
one. This seems to be true for Tripogandra. The pedicel
of each flower apparently bears a bract. In the axil of
this bract a bud develops which is terminated by a flower
which also bears a bract on its pedicel. The small bracts
found on the top of the peduncle of all Tripogandra species
represent these bracts.

The double cincinnus always seems to terminate a stem.
Other shoots terminated by double cincinni may or may
not be produced at lower nodes. A short shoot bearing
several double cincinni may develop in a leaf axil, as in
Tripogandra multiflora and Т. montana, giving the im-
pression that several double cincinni arise at a node. Close
inspection shows the true nature of the situation, The
double cincinni of T. guerrerensis are produced in a larger,
much branched inflorescence so the appearance of the whole
is that of a large panicle.

The number of flowers produced per double cincinnus
is variable. The cincinni of some species — Tripogandra
montana, T. serrulata — are long-lived, producing many
flowers over a long period of time, and the sympodial cin-
cinnus axis may become one or two centimeters long. On
the other hand, T. guerrerensis and T. amplexans, produce
only a few flowers on each ultimate unit over a short period
of time.

Тће peduncle of the inflorescence may be either glabrous
or variously vestite. The hairs may be in lines or bands
and/or scattered. Hairs borne in lines or bands are always
uniserate (T. serrulata), but the scattered hairs may be

1975] Tripogandra — Handlos 225

either uniseriate (T. multiflora) or capitate (Т. amplexi-
caulis). The peduncle length is variable, probably being
determined both genetically and environmentally.

Flower. The flowers of all Tripogandra species follow
the typical monocotyledonous pattern — three sepals, three
petals, six stamens in two whorls or three stamens and
three staminodes, and three carpels.

The sepals, which are usually green, contain one median
vascular bundle and are more or less boat-shaped or hooded
near the apex. The margin is hyaline and may or may not
be pigmented. The sepals may be glabrous or nearly so
(Fig. 3) as in T. guerrerensis and T. grandiflora, densely
pilose with capitate hairs as in T. glandulosa and T. pur-
purascens (Fig. 1), pilose with uniseriate hairs as in T.
saxicola (Fig. 2), or they may have only a few hairs at
the apex as in T. diuretica. The hairs, when present, are
often of diagnostic value. At anthesis the sepals may be
reflexed but they close when the petals deliquesce.

The petals in all species are larger than the sepals and,
depending on the taxon, vary in color from white to dark
pink or magenta. Both white and pink flowers have been
observed in Tripogandra angustifolia, T. purpurascens
and T. serrulata. The petals remain open for only a few
hours, commonly opening in the morning except for T.
saxicola which flowers in the afternoon, All petals deli-
quesce a few hours after opening. The cell membranes
apparently become permeable or break down during deli-
quescence allowing the cytoplasm to seep out of the cells
as drops of liquid. The cell walls remain but the petal as
a whole shrivels to a crumpled mass. The veins of the
petals are difficult to distinguish in living material.

The androecium is dimorphic in all species; in bud it is
actinomorphic but at anthesis it becomes zygomorphic as
described below. 'The outer whorl of the androecium is
always opposite the sepals and is composed of fertile sta-
mens with short filaments (Fig. 25). The filaments are
more or less subulate ог awl-shaped and may be glabrous

226 Rhodora [Vol. 77

as in Tripogandra grandiflora (Fig. 8) or bear one to
many multicellular hairs on the dorsal surface (Figs. 10,
5). The number of hairs is variable within a species and
has not been used as a taxonomic character. In dried ma-
terial these hairs are almost impossible to detect. In the
living state the cells of the hairs are variable in shape so
the hairs may appear as long, uniseriate structures as in
T. palmeri (Fig. 13) or as the classical moniliform hairs
as in T. purpurascens (Fig. 5). The cells of the hair may
be colorless, white, or pink.

The distal end of the filament forms the connective
of the anther and the connective is usually somewhat
expanded and thickened as is easily seen in Tripogandra
grandiflora (Figs. 8, 9). Anther sacs are borne on the
ends of the connective (Fig. 14). The anthers are usually
extrorse in bud, but during anthesis bending of the distal
end of the filament causes the anthers to become introrse
or, with less bending, pollen is exposed upward. Dehiscence
is longitudinal in all species (Fig. 8). The pollen is spheri-
cal to oblongoid and monosulcate.

The structures of the inner whorl of the androecium are
borne opposite the petals (Fig. 25) and may be either
staminodes which produce no functional pollen or stamens
with functional pollen. In all species the filaments of these
stamens or staminodes are longer than the filaments of
the outer whorl of stamens. The filaments may be glabrous
as in Tripogandra guerrerensis (Fig. 17) and T. palmeri
(Figs. 21, 22), or variously bearded. The hairs may be
borne on the dorsal surface as in T. saxicola (Fig. 20),
around the filament as in T. serrulata (Fig. 19) and T.
montana (Fig. 31) or in two more or less discrete patches
as in T. grandiflora (Fig. 28). In all species the hairs are
borne more abundantly on the middle or distal end of the
filament. The cells of the hairs are usually spherical, pro-
ducing the typical moniliform hairs, or cylindrical, pro-
ducing uniseriate hairs as in T. grandiflora (Fig. 28). The
cells of one hair are usually variable in shape and size,
those at the base commonly being cylindrical while the

1975] Tripogandra — Handlos 227

distal ones may be either cylindrical or spheroidal depend-
ing on the species. The cells may appear white, pigmented,
or colorless.

The filaments of the stamens or staminodes of the inner
whorl are bent to varying degrees. In all species two of
the filaments bend at the base around the filaments of two
outer stamens so the inner stamens or staminodes become
aligned near the third inner stamen or staminode in front
of the upper petal (Fig. 16). In addition, all filaments
are bent in the middle. In Tripogandra palmeri the bend
is C-shaped (Fig. 21); in Т. serrulata (Fig. 19), Т. grandi-
flora (Fig. 28), and T. montana (Fig. 31) the bend is an
open S-shape; while in T. guerrerensis (Fig. 17 dt
amplexicaulis, T. amplexans (Fig. 26), апа Т. angusti-
folia (Fig. 30) the bend is a more pronounced S-shape.
The filament may be more or less expanded and inflated
distally as in T. amplexicaulis (Fig. 23) or cylindrical as
in T. serrulata (Fig. 19).

The connective and filament form a continuous structure
in Tripogandra palmeri (Fig. 21) but the filament is very
thin distally in species such as T. guerrerensis, T. amplexi-
caulis, and T. angustifolia (Fig. 27), and the anther then
is more or less versatile. The connective may be elongate
and straight, bent in the middle to form a V- or C-shape,
or discoid. The anther sacs are borne on the ends of the
connective and form a small part of the anther. Dehis-
cence is longitudinal.

The pollen from anthers of the outer whorl of the
androecium may be fertile as in Tripogandra serrulata
or modified and sterile as in T. grandiflora or T. guerre-
rensis. Lee (1961) first described the pollen within an
anther of tetraploid T. grandiflora. This pollen varies in
shape from nearly spherical to oblongoid to sub-fusiform
and absorbs stain differentially from aniline blue-lacto-
phenol. The pollen of diploid Т. grandiflora stains uni-
formly, is more or less spherical, but is larger and has a
more sculptured surface as compared to the pollen of the
fertile stamens. The variation in pollen shape is greatest

228 Ећодога [Vol. 77

in T. amplexans but within any one anther the pollen
grains are uniform, Some collections produce spherical
pollen which is externally indistinguishable from that of
the outer whorl of stamens. In other collections the pollen
grains are oblongoid or fusiform.

The gynoecium of the Tripogandra flower is composed
of three fused carpels, Each carpel is supplied by three
vascular bundles, two ventrals and one dorsal. Placenta-
tion is axile and two orthotropous ovules are borne in each
of the three locules (Fig. 49). A short filiform style is
found in all species. The shape of the stigma may be
constant or variable within a species. The stigma may
be simple and represent the top of the style, may be slightly
enlarged (capitellate), greatly enlarged (capitate), or
slightly penicilliform. In some plants the stigma may be
somewhat three-lobed.

The pistil matures to form a loculicidal capsule which
splits down the dorsal surface of each carpel. One to six
seeds may be produced in each capsule. The lower ovule
never matures in Tripogandra palmeri, consequently a
maximum of three seeds is produced in each capsule of
this species.

The orthotropous ovules develop into seeds with a dorsal
embryotega. The embryotega represents that part of the
integuments which lies over the embryo. In all species
the position of the embryo is readily observed and is seen
to be situated on the side of the seed opposite the hilum.
The micropyle is also located on the side opposite the
hilum — the embryo, in fact, develops just below the
micropyle. As has been pointed out by Brückner (1926)
for the family and as shown by Chikkannaiah (1962, 1963,
1964, 1965a, 1965b) from embryological studies of Com-
melina, Murdannia, Floscopa, and Tinantia, the relation-
ship between the micropyle, the embryo and the embryo-
tega is a constant one. In Tripogandra palmeri (Fig. 72)
and T. grandiflora (Figs. 76, 77) the embryo and embryo-
tega protrude from the surface of the seed but the margins
of the seed are revolute and the embryo is more or less

1975] Tripogandra — Handlos 229

surrounded and protected from mechanical damage. In
other species of Tripogandra, the embryo is impressed
and completely surrounded by the remainder of the seed.
While the position of the embryotega is uniform within
the genus it varies within the family. No detailed studies
have been made of embryology and development of the
seeds of Tripogandra.

The seeds of most species of Tripogandra are trigonal
in outline. Tripoyandra amplexans has seeds which are
variable and they may be either trigonal (Fig. 59) or
rectangular (Fig. 58) in outline. The seeds of T. palmeri
and T. grandiflova are unique in having revolute margins.
Tripogandra palmeri has the margin revolute in three
places (Fig. 72), while T. grandiflora has the two opposite
margins revolute (Figs. 76, 77). Seed outline is also
dependent on the number of seeds which develop in a
locule. The seeds become trigonal if both ovules develop
(Fig. 76) but are more elliptical if only one develops (Fig.
77). The single ovules usually develop into larger seeds.

The appearance of the surface of the seed coat is of
taxonomic value. Following the terminology of Murley
(1951) the following categories of seed surface texture
may be recognized among the species of Tripogandra:

1. Reticulate: 7. serrulata (Figs. 38, 39), Т. montana
(Figs. 44-46) ;

2. Reticulate-foveate: Т. multiflora (Figs. 32-35), Т.
warmingiana (Fig. 61);

3. Ribbed reticulate-foveate: Т. glandulosa (Figs. 42,
43) ;

4. Areolate: T. palmeri (Figs. 72, 78), T. saxicola
(Figs. 56, 57);

5. Ribbed areolate: 7. amplexans (Figs. 58-60), T.
brasiliensis (Figs. 47, 48), T. disgrega (Figs. 52, 53),
T. guerrerensis (Figs. 68-71), T. purpurascens (Figs. 50,
51, 54, 55);

6. Farinose: T. grandiflora (Figs. 76-78) ;

7. Ribbed farinose: T. angustifolia (Figs. 74, 75).

280 Кћодога [Vol. 77

The seeds of Tripogandra amplexicaulis are somewhat
variable and are generally areolate but some collections
have seeds which are also distinctly alveolate (Figs. 64,
65).

Tripogandra diuretica (Figs. 40, 41) produces seeds
which are more or less intermediate between the ribbed
areolate and the reticulate-foveate conditions and which
could be called ribbed areolate-foveate.

The outline of the hilum may be punctiform (nearly
circular) as in Tripogandra angustifolia (Fig. 75), T.
disgrega (Fig. 53), T. diuretica (Fig. 41), T. multiflora
(Figs. 33, 35), and Т. purpurascens (Figs. 51, 55), linear
as in T. grandiflora (Fig. 78) and T. palmeri (Fig. 73),
or elliptical as in T, brasiliensis (Fig. 48) and T. guerre-
rensis (Figs. 69, 71). The outline of the hilum is constant
within a species and is useful taxonomically.

The germination of Tradescantia virginiana (as T. vir-
ginica) has been described and illustrated by Gravis
(1898) and that of Commelina virginica has been de-
scribed by Bates (1939) but no previous studies have been
made of Tripogandra. I have observed germination in
detail in two species, T. amplexans and T. purpurascens.
There are no significant visible differences in germination
among the three genera.

When the seeds are wet they swell slightly and within
à few days the seedling emerges. Studies by Chikkannaiah
(1962, 1963, 1964, 1965a) of Commelina, Murdannia and
Floscopa, as well as those by Gravis (1898) of Tradescan-
tia show the radicle of the embryo situated directly below
and adjacent to the micropyle and embryotega. I have
observed, as did Bates and Gravis, that the radicle emerges
from the seed first. The embryotega may be pushed to
one side or lifted free from the seed and cap the root tip
(Fig. 24). Through further growth the remainder of the
embryo emerges from the seed. The tip of the cotyledon
remains in the seed, presumably as an absorbing structure,
while the cotyledonary neck or petiole elongates further.
The first true leaf emerges from the sheathing base of the

1975] Tripogandra — Handlos 231

cotyledon. By elongation of the radicle and first node, the
seed and cotyledonary petiole may be lifted above the
ground level.

CYTOLOGY

The cytological relationships of various Mexican Com-
melinaceae as well as a review and sumary of previous
investigations are discussed in Handlos (1970).

Tripogandra is cytologically distinct from other genera.
Karyotypes show both telocentric and metacentric chromo-
somes as well as larger and smaller chromosomes. The
basie chromosome number is probably eight, though Jones
and Jopling (1972) indicate the additional possibility of
13. Diploid chromosome numbers range from 16 to 64.
Two species, T. montana and T. saxicola, with haploid
numbers of 21 may be allopolyploids based on the numbers
8 and 13.

BREEDING SYSTEMS AND POLLINATORS

The genus Tripogandra is characterized by a distinc-
tively modified androecium as was pointed out by Moore
(1960). He suggested that the arrangement of stamens
and staminodes, in addition to the self-sterility of the clone
studied, indicated dependence on insect pollinators and the
necessity of out-crossing in the species. Several other com-
melinaceous genera, including Aneilema, Cochliostema,
Commelina, Tinantia, also exhibit various androecial modi-
fications. Few studies have been made on the relationship
between insects and the androecium in the Commelinaceae.
Pollination has been studied in Commelina (Breitenbach,
1885), Tinantia (Knuth, 1906) and Tradescantia (Kerner
von Marilaun, 1894; Sinclair, 1968).

From a series of 166 interspecific crosses involving
Tripogandra amplexans, T. angustifolia, T. palmeri, and
T. purpurascens subsp. purpurascens made in 1966, no
interspecifie hybrids were obtained. Flowers were bagged
to exclude insects. All species studied proved to be self-

232 Ећодога [Vol. 77

fertile, Further observations of T. amplexicaulis, T. dis-
grega, T. diuretica, T. glandulosa, T. gwerrerensis, T.
montana, T. multiflora, T. saxicola, and T. serrulata grow-
ing in insect-free greenhouses show that these species are
also self-fertile. Of the 14 species observed, only T. grandi-
flora is self-sterile.

In 1967, I observed natural populations of Tripogandra
amplexans, T. amplexicaulis, Т. angustifolia, T. disgrega,
T. grandiflora, T. guerrerensis, T. montana, T. purpur-
ascens, and. T. saxicola in México for a total of 611% hours.
Two hundred thirty-seven insects were observed of which
158 were captured. These insects are now in the collec-
tions of the Department of Entomology, Cornel] Univer-
sity, as "Lot no. 994."

Hymenopterous insects formed the largest category of
visitors. There were 60 individuals in the Apidae, 48 in
the Halictidae, 20 in the Andrenidae, 4 in the Anthophori-
dae, 1 in the Coletidae. The bees formed the largest and
most active group of insects and should be considered the
most important pollinators. Bees only engaged in gather-
ing pollen since the plants do not produce nectar. Some
individual bees visited only the longer stamens (stami-
nodes), others visited only the shorter, while still other
bees crawled over all stamens and gathered pollen indis-
criminately.

In only one species did I observe that the position of the
staminodes caused bees to move in a restricted fashion in
the flower. In Tripogandra guerrerensis the petals are not
arranged symmetrically but instead at anthesis the two
lower petals bend away from each other and are located
nearer the upper petal; in this way an angle of approxi-
mately 909 is formed between either of the lateral petals
and the upright petal, while the two lower petals are sepa-
rated by 180°. The three staminodes arch out over the
stamens and ovary. A bee can approach the stamens only
by clinging upside down to the filaments of the staminodes.
The significant position seemed to be that assumed when
а bee was clinging to the staminodes and gathering pollen

1975] Tripogandra — Handlos 288

from the anthers. The insect’s abdomen then touched the
anthers and pollen adhered to its body. It seems likely
that pollen could be carried from flower to flower in this
fashion and cross-pollination would thus occur. The sig-
nificance of cross-pollination remains to be demonstrated
in this species because these plants are also self-fertile.
In other plants, Allard (1965) has shown that a large
amount of variability is maintained though only a small
percentage of outcrossing occurs. He believed that most
individuals of a self-pollinating population would be highly
homozygous but the recombination of genes introduced by
а low level of outcrossing would provide sufficient new
genotypes adapted to the microhabitats which occur in
an area and would allow for increasing fitness of the
species to a specific niche. Such reasoning may be applied
to the annual and perennial self-fertile species of Tripo-
gandra but further investigations should be conducted to
determine the basic facts.

Some bee species, even when collected in small numbers,
were found to visit two or more species of Tripogandra.
One species of Dialictus was collected from four different
species of Tripogandra; one species of Pseudopanurgus
and another species of Dialictus were collected from three
Tripogandra species, while three other Dialictus species,
one species of Evylaeus, Augochlora near smaragdina,
Trigona fulviventris, T. mosquito frontalis, and T. testacea
orizabaensis were collected on two species of Tripogandra.

Many of the bees collected in this study are solitary bees
which have been reported (Linsley, 1958) to be most
abundant in warm semi-desert regions of the world. Many
habitats in México can be described as warm and arid
or border semi-desert or desert regions which may account
for the many solitary bee species collected. The majority
of solitary bees are oligolectic (Linsley, 1958) (oligo-
tropic fide Faegri and van der Pijl, 1966), i.e. utilize only
a few related species of plants as food sources. The social
bees, Trigona and Apis, are probably polylectic (polytropic
fide Faegri and van der Pijl), i.e. utilize food from un-

234 Rhodora [Vol. 77

related plant sources. Trigona mexicana and T. acapulconis
were both collected at one location in México. Some indi-
viduals were found on Tripogandra and others were caught
on an abundant composite.

Tripogandra grandiflora, the only demonstrated self-
sterile species produces strongly fragrant flowers. Tripo-
gandra amplexicaulis, T. saxicola, and T. purpurascens
subsp. purpurascens are very faintly scented. The scents
produced may allow bees to identify and return to a specific
source of pollen or, in the case of T. grandiflora, may
allow bees to locate the plant because of its odor.

Of 23 dipterans collected, 18 were in the family Syrphi-
dae. Faegri and van der Pijl consider these flies to be
irregular and not very active pollinators. In Tripogandra
I would confirm this observation. The three dipterans in
the family Bombyliidae may effect some pollination but
the remaining two flies and three beetles were probably
not visiting the flowers for pollen and should not be con-
sidered to be important pollinators.

MEASUREMENTS

To prevent misunderstandings and to provide for con-
sistent results I am giving below the methods by which
I obtained my measurements. The length of the leaf
blade is the distance from the tip of the leaf to the top of
the sheathing leaf base. The length of the sheath is mea-
sured from the line of attachment at the node to the lowest
point on the orifice of the sheath; this point is always on
the side opposite the lamina. The diameter of the leaf
sheath reflects varying amounts of inaccuracy because
these measurements are from dried, pressed specimens.
If the specimen is flattened and well-pressed, the mea-
surement will be greater than the diameter in the living
plant; if the stem is not well-flattened, the measurement
may be close to that in the living state or even somewhat
amaller. In either case, the sheath seems to shrink less
than the stem during drying and is a more accurate esti-

1975] Tripogandra — Handlos 235

mate of stem diameter than a direct measurement of the
dried stem. The length of the peduncle is the distance
between the subtending leaf and the base of the bracts at
the distal end of the peduncle. The length of the pedicel
is measured on flowers at anthesis. Anthesis is a definable
period of a few hours duration and is used because it pro-
vides a uniform standard which is not subject to personal
bias. Pedicels may elongate in fruit but a measurement
standard would be more difficult to define. Measurements
of flower parts are made on flowers at anthesis for the
same reasons given above. The length of the filaments
is the linear distance from the base to the connective dis-
regarding curves and bends. The length of the anthers is
the distance represented by the length of the anther sacs,
while the width is taken as the distance between the anther
sacs on the same anther. Pollen viability is judged by pol-
len stainability in aniline blue-lactophenol. The length of
the ovary is the distance between the base of the ovary
and the base of the style. The length of the style includes
the stigma. The length of the capsule does not include the
persistent style.

SYSTEMATIC ACCOUNT

Tripogandra Rafinesque, Flora Telluriana 2:16. 1837
(‘1836’). TYPE: Tradescantia multiflora Swartz.

Heminema Rafinesque, Flora Telluriana 2:17. 1837
(‘1836’). TYPE: Tradescantia multiflora Swartz.

Descantaria Schlechtendal, Linnaea 26:140. 1853; Briick-
ner, Bot. Jahr. Syst. Beiblatt 137, 61:60, 61. 1926.
LECTOTYPE: Tradescantia multiflora Swartz.

Disgrega Hasskarl. Flora 49:215. 1866. TYPE: Trades-
cantia disgrega Kunth.

Donnellia Clarke in Donnell-Smith, Bot. Gaz. 33:261.
1902; non Austin (1880). TYPE: Callisia grandiflora
Donnell-Smith.

Neodonnellia Rose, Proc. Biol. Soc. Wash. 19:96. 1906.
TYPE: Callisia grandiflora Donnell-Smith.

236 Rhodora [Vol. 77

Fibrous-rooted annuals and perennials, stems usually
branching monopodially, erect or trailing, internodes cylin-
drical, glabrous or variously vestite, often with a line of
uniseriate hairs extending down one side.

Leaves with a tubular sheathing base; blade narrowly
ovate to ovate (linear in Tripogandra angustifolia) , usually
flat but complicate in T. purpurascens and C-shaped or
terete in Т. angustifolia, usually fleshy, glabrous or vari-
ously vestite with uniseriate hairs, base cuneate, rounded,
oblique, or amplexicaul, sometimes narrowed and sub-
petiolate, apex acute, sometimes acuminate ; sheath glabrous
to pilose, orifice glabrous or more often villous with long
uniseriate hairs, usually with a line of uniseriate hairs
extending down the side opposite the blade and continuing
to the internode below.

Inflorescences composed of a single terminal or а termi-
nal and a few axillary double cincinni, or a terminal panicle
of double cincinni; peduncles with or without 1-2 lines of
uniseriate hairs, otherwise glabrous to densely pilose, hairs
uniseriate or capitate; pedicels erect or reflexed in fruit,
glabrous to pilose, hairs uniseriate or capitate; bracts at
the base of the pedicels small, a thin band of tissue which
is glabrous to pilose with capitate or uniseriate hairs,
margin entire, denticulate, with a few uniseriate hairs,
or ciliate.

Flowers white to bright pink or magenta; sepals three,
cymbiform, ovate, elliptic, or obovate, apex acute or obtuse,
glabrous to pilose, margin hyaline, entire; petals three,
ovate, elliptic, or obovate, with acute, rounded, obtuse, or
irregular apex, deliquescing a few hours after anthesis
to a shapeless mass; androecium actinomorphic in bud
becoming zygomorphic at anthesis, composed of six stamens
or three stamens and three staminodes in two whorls, the
outer whorl always of stamens and opposite the sepals,
shorter, with glabrous or variously bearded filaments about
equalling the pistil; anthers extrorse in bud, becoming
either introrse at anthesis or horizontal and then shedding
pollen upward toward the stigma, dehiscing longitudinally,

1975] Tripogandra — Handlos 237

connective usually short and inconspicuous, pollen usually
fertile; inner whorl of stamens or staminodes opposite the
petals, longer, filaments glabrous or variously bearded
with uniseriate or moniliform hairs, variously curved and
bent, two filaments bending at the base toward the third
inner stamen-staminode around the filaments of the two
outer intervening stamens so the inner stamens-staminodes
are all in an erect position in front of the upper petal at
anthesis, anthers dehiscing longitudinally, connective short
and inconspicuous or elongate, straight or bent in a C-
or U-shape, pollen fertile or sterile.

Ovary globose or cylindrical, somewhat flattened on three
sides, glabrous, trilocular, placentation axile with two
orthotropous ovules per locule, style shorter than the ovary,
filiform, stigma simple, capitellate, or capitate.

Fruit a loculicidal capsule with persistent style; seeds
usually 2 per locule (1 in Tripogandra palmeri, 1-2 in T.
grandiflora), usually triangular, the surface variously
reticulate or roughened, hilum punctiform, elliptic, or
linear, embryotega dorsal, protuberant or impressed.

KEY TO THE SPECIES

a. Leaves terete or C-shaped in cross section, linear in
outline, less than 2.7 mm wide, red or green; testa
ribbed farinose. .....0........... 3. T. angustifolia.

a. Leaves flat or complicate, never terete, mostly broader
than semm, green, ЗЕЕ ЕТЕТ а b.
b. Nodes below the inflorescences bearing amplexicaul

leaves; leaves flat, never complicate. .......... с.
с. Plants perennial: upper internodes pilose; hilum
punctiform to elliptical. ........ 7. Т. encolea.

с. Plants annual; upper internodes glabrous or with
a line of uniseriate hairs; hilum linear. .... d.

d. Petals bright pink; plants tall, to 92 ст high;
leaves of stem below amplexicaul leaves to

14.8 cm long, occasionally subpetiolate; pe-
duncles pilose; seeds usually with an alveolate
surface, convex dorsally. . 2. T. amplexicaulis.

238

Ећодога [Vol. 77

d. Petals white or pale pink; plants shorter,

to 58.5 em high; leaves below amplexicaul
leaves to 7.8 ст long, rounded or cuneate at
the base; peduncles often glabrous; seeds
never with alveolate surface, flattened dor-

sally. .................. 1. T. amplexans.

b. Nodes below the inflorescences without amplexicaul
leaves ог if amplexicaul then also complicate. .. e.

e. Filaments of inner (longer) staminal whorl
glabrous. ........ ees f.

f. Sepals glabrous or nearly so; seeds two per

f.

locule, the upper larger, hilum linear, testa
ribbed, areolate; inflorescence an open pani-
cle. ................. 10. Т. guerrerensis.
Sepals with scattered capitate hairs or pilose;
seeds in each locule equal in size or only one
per locule and then the margins revolute;
inflorescence of double cincinni variously
arranged but never in an open panicle. .. g.
g. Hilum linear, seeds one per locule with
margin revolute; pedicel erect in fruit.
Lee eee ee eee eee eee 15. Т. palmeri.
Hilum punctiform, seeds usually 2 per
locule, lacking revolute margins; pedicel
reflexed in fruit. .................. h.
h. Leaf base oblique; testa reticulate but
not ribbed; capsule obovoid, slightly
stipitate. .......... 19. T. silvatica.

h. Leaf base not oblique, but instead
cuneate to rounded; testa ribbed;
capsule globose or obovoid, not stipi-
tate. ........................... i.

i. Testa prominently ribbed reticu-
late-foveate; longer filaments con-

cave in upper third, to 3.3 mm long;

seeds 0.8-1.4 mm long. ..........
TUM 8. T. glandulosa.

0%

1975]

Tripogandra — Handlos 239

i. Testa ribbed areolate; longer fila-
ments inflated in upper portion, to
8.0 mm long; seeds 1.2-21 mm
long; се... неваљан ELT j:
j. Calycine hairs long, some 1.5-4.5

mm long; leaves usually flat;
peduncle usually glabrous; dor-
sal seed surface convex. ......
ак канаат реч 5. Т. disgrega.
j. Calycine hairs short, the longest
less than 1.0 mm long; leaves
complicate; peduncle variously
glabrous, pilose, with or with-
out lines of hairs; dorsal sur-
face of seed flat or concave. . k.
k. Internodes glabrous except
for a line of uniseriate hairs
down one side, rarely the
upper internodes with scat-
тегей заты л d шеш.
нкү, 16а. Т. purpurascens
subsp. purpurascens.

k. Internodes all with scattered
capitate hairs and a line of
uniseriate hairs down one
side. . 16b. T. purpurascens
subsp. australis.

e. Filaments of inner (longer) staminal whorl
hénrded, never glabrous. hese e n l.

1.

Hairs of the filaments uniseriate, never moni-
liform; hilum linear, testa farinose, ......
а lena ee ee 9. T. grandiflora.
Hairs of the filaments moniliform; hilum
punctiform or elliptical e шылу; m.
m. Filaments of longer stamens inflated; leaf

base subpetiolate, oblique; testa ribbed

areclate, hilum elliptical. .............
4. Т. brasiliensis.

«+ + 9 9 9 9 9 9 а c9 à» а СИ

240

Ећодога [Vol. 77

m. Filaments of longer stamens terete, never
inflated; leaf base various; testa not
ribbed except T. diuretica, hilum puncti-
form (occasionally elliptical in T. saxi-

cola and T. diuretica). .............. n.
n. Lamina of leaf cuneate or truncate at
the base; petals white. .......... 0.
о. Sepals pilose; pedicels pilose. .. р.

p. Pedicels erect in fruit; pe-
duncles pilose with capitate
hairs. ...... 11. Т. kruseana.

p. Pedicels reflexed in fruit; pe-
duncles glabrous except for 1-2
lines of uniseriate hairs.
err 17. T. saxicola.

o. Sepals nearly glabrous, with a few
hairs at the point of union of adja-
cent sepals; pedicels glabrous, re-
flexed in fruit; peduncles glabrous
except for 1-2 lines of uniseriate
hairs. .......... 14. T. neglecta.

n. Lamina of leaf oblique at the base,
rarely cuneate or rounded and then
the petals bright pink; seeds trigonal,

never lobed. .................... q.

q. Leaves subpetiolate at the base of
the plant, to 4.4 ст long, to 1.85
em wide; sepal midvein pilose;
plants annual; seeds reticulate-
foveate; bracts at the base of each
pedicel with erose margin. .....
ZEIT 20. T. warmingiana.

q. Leaves never subpetiolate, to 14.5
ст long, to 3.5 ст wide; plants
perennial; sepals glabrous to pi-
lose, the midvein not distinctively
vestite; seeds ribbed, reticulate-

19751

Tripogandra — Handlos

241

foveate, or reticulate; bracts vari-

ous.
r.

KENNT KEEN sagen eee р.
Style nearly as long as the
ovary, 0.6-1.1 mm long; testa
of seeds ribbed areolate-fove-
ate; sepals 4.5-7.0 mm long.
DEER 6. Т. diuretica.
Style much shorter than the
ovary, 0.15-0.6 mm long; testa
of seeds reticulate or reticulate-
foveate, never ribbed; sepals
1.8-6.0 mm long, .......... 8.
з. Testa reticulate-foveate; pe-
duncles pilose (rarely gla-
brous or with 2 lines of
hairs); ovary 0.5-1.0 mm
long; sepals 1.8-4.0 mm
long; outer anthers 0.3-0.6
mm long. . 13. T. multiflora.
s. Testa reticulate; peduncles
glabrous or with 1-2 lines of
hairs, rarely pilose, if pilose
then ovary 1.2-1.7 mm long;
sepals 4.0-6.0 mm long and
outer anthers 0.8-1.8 mm
loin IS ee ee ан а -
t. Seed surface uniformly
brown; sepals narrowly
ovate, elliptical, or obo-
vate, 4.0-6.0 mm long,
1.6-2.8 тт wide; ре-
duncles 0.7-12.7 em long;
inner filaments 4.5-7.5
mm long, densely beard-
ed; petals bright pink or
magenta, 5.5-11.0 mm
long; ovary 1.2-17 mm
long; style 0.2-0.6 mm
long. .. 12. T. montana.

242 Ећодога [Vol. 77

t. Seed surface gray or
brownish with lighter
reticulations ; sepals
ovate, 2.5-4.6 mm long,
1.3-2.7 mm wide; ре-
duncles 0.4-5.1 cm long;
inner filaments 2.7-4.5
mm long, variously
bearded; petals white or
pink, 3.5-6.2 mm long;
ovary 0.6-1.3 mm long;
style 0.1-0.3 mm long. ..
DENEN 18. T. serrulata.

1. Tripogandra amplexans Handlos, sp. nov. HOLOTYPE:
México. MICHOACAN: Km. 201.2 of Hwy. 15, 8.1 km.
north of Tuxpan, 1920 m., 18 Sept. 1967, Handlos 428
(us!).

Herba annua; caulis erectus, usque ad 58.5 cm altus,
internodiis infra folia amplexicaulia usque ad 10.5 cm
longis. Folia ovata, laminis usque ad 7.8 cm longis, usque
ad 3.4 ст latis, inferis basi cuneatis, superis amplexicaulis,
glabris, margine ciliatis, apice acutis, vaginis usque ad
10.5 mm longis, usque ad 7.7 mm diam., glabris vel linea
unica pilorum instructis. Imflorescentiae simplices vel pani-
culatae ex 1-8 cincinnis duplicibus constantes, 1-5 foliis
amplexicaulibus et 0-4 bracteis vaginantibus subtentae;
pedunculi usque ad 6.8 cm longi, glabri vel pilis paucis
dispersis, distalibus, capitatis instructi; cincinni duplices
alabastra, flores, vel fructus usque ad 12 gerentes ` pedicelli
usque ad 7.0 mm longi, pilis capitatis pilosi, erecti vel
maturitate effusi, bracteis basi pedicellorum margine in-
tegris, glabris vel pilis paucis, dispersis, capitatis pubes-
centibus. Flores albi vel subrosei; sepala ovato- vel ellip-
tico-cymbiformia, 2.5-5.0 mm longa, 1.2-2.2 mm lata, pilis
paucis, dispersis, capitatis, incoloribus instructa, margine
integra et hyalina, apice acuta, rotundata vel obtusa;

1975] Tripogandra — Handlos 243

petala late ovata vel ovato-elliptica, 4.2-8.0 mm longa, 3.9-
6.5 mm lata, apice acuminata, acuta, vel obtusa; stamina
3, Sepalis opposita (Fig. 25), filamentibus 1.0-2.2 mm
longis, albis, dorsaliter medio 0-7 pilis albis, incoloribus
vel subroseis, moniliformibus pubescentibus (Fig. 6), an-
theris 0.3-1.0 mm longis, 0.3-1.0 mm latis, albis, polline
albido; staminodia 3, petalis opposita (Fig. 25), filamenti-
bus epipetalis, 2.0-6.0 mm longis, albis, glabris, sigmoideis
et distaliter dilatis (Fig. 26), antheris luteis, 0.2-1.0 mm
longis, 0.3-1.0 mm latis, connectivo C- vel V-formi, polline
luteo; orariwm 0.8-1.5 mm longum, 0.7-1.2 mm diam.,
glabrum, stylo 0.3-0.8 mm longo, stigmate simplici vel
capitellato. Capsula globosa, 2.7-3.5 mm longa, glabra,
seminibus triangularibus raro trapezoideis (Figs. 58, 59),
1.2-1.8 mm longis, costatis, areolatis, hilo lineari (Fig. 60).

Chromosome number: 73-16.

Distribution and habitat: western México in the states
of Jalisco, Michoacán, México, Morelos, and Guerrero; in
damp, rocky places in nearly neutral soil at elevations
from 800-2200 m.

Flowering: This species begins flowering in the rainy
season from early August and continues until October.
Flowers open from 8:00 AM to 9:00 AM and close between
10:30 AM and 12:50 PM. In the field these annuals exhibit
a great deal of variation with regard to size when flower-
ing. This is probably due to the amount of moisture avail-
able after germination, the effects of grazing, and the
fertility of the soil.

REPRESENTATIVE SPECIMENS

México. JALISCO: barranca SE cf Ciudad Guzman, 22 Oct. 1940,
Moore, Jr. 158 (BH, GH). MICHOACÁN: Zitacuaro, Dist. Zitácuaro,
1950 m., 6 Sept. 1938, Hinton et al. 13198 (ARIZ, GH, MICH, MO, NY,
US). MEXICO: just N of bridge, Puente Calderón, ca. km. 135 of
Hwy. 55, just north of Ixtapan de la Sal, 6 Aug. 1967, Handlos 312
(BH). MORELOS: railroad from Mexico City to Cuernavaca at km.
96; region of El Parque, ca. 2200 m., 5 Oct. 1958, Hawkes, Hjerting
& Lester 1623 (C, F). GUERRERO: Manchon, Dist. Mina, 1290 m.,
13 Aug. 1936, Hinton et al. 9206 (ARIZ, GH, NY, US).

~

244 Кћодога [Vol. 77

The range of Trivogandra amplexans is of special in-
terest because it is wholly within the range of the more
widespread and similar T. amplexicaulis. In many loca-
tions the two species occur together. I am unable to dis-
cern any differences in habitat preferences for these two
species.

The main reasons for considering these plants as two
distinct species, aside from morphological differences, are
their occurrence together with little evidence of hybridiza-
tion and the presence of a partial isolating mechanism in
the slightly different flowering seasons,

A list of distinguishing characteristics is presented here
to summarize the differences between the species. Tripo-
gandra amplexans is generally smaller in stature (to 58.5
ст tall), has smaller leaves (to 7.8 cm long, to 3.4 cm
wide) which are rounded or cuneate at the base, a smaller
inflorescence composed of fewer (to 8) double cincinni, a
glabrous peduncle, flowering period from August to Octo-
ber, fewer flowers (to 12 per double cincinnus), which
have smaller white or pale pink petals (to 8.0 mm long),
seeds with larger, smoother ribs, a deeply impressed
embryotega and a flatter dorsal surface.

Tripogandra amplexicaulis can be recognized by its
taller stature (to 92 em tall), its larger leaves (to 14.8
em jong, 5.0 em wide), a narrowed, subpetiolate laminar
base in some populations, a large paniculate inflorescence
composed of up to 14 double cincinni, a pilose peduncle,
flowering period from mid-August to December, more
flowers per inflorescence (to 18), larger petals (to 13.0 mm
long), flowers bright pink, seeds with an alveolate surface,
finer ribs, a slightly impressed embryotega and convex
dorsal surface.

There may be evidence of hybridization between these
two species because at Puente Calderón in the state of
México I found one plant that was vegetatively like Tripo-
gandra amplexans with small flowers but showing the
bright pink color of T. amplexicaulis. Other genetic

1975] Tripogandra — Handlos 245

mechanisms could explain the flower color difference but
for the moment the problem has not been resolved.

A few plants have been collected which exhibit charac-
teristics of both species and can only arbitrarily be placed
in either species (Figs. 29, 62, 63). Of these intermediate
collections, five are from the known margins of the range,
while one collected from a road cut on a steep hillside and
another from a lava field represent populations existing
in disturbed habitats. Further investigations should be
conducted to determine whether these plants represent
stabilized hybrids, populations with introgression into one
or the other of the parental species, specialized peripheral
populations, or whether some other explanation of their
intermediacy is plausible. A list of the collections which
appear intermediate follows.

SPECIMENS EXAMINED

México. JALISCO: ca. km. 781.2 of Hwy. 15, Guadalajara to Tepic,
са. 25 km. NW of Magdalena, 1000 m., 6 Oct. 1967, Handlos 451
(ви); km. 1050 of Hwy. 80, 14 km. SW of Autlán, 1120 m., 4 Oct.
1967, Handlos 442 (рн). MEXICO: Rincón del Carmen, Dist. Temas-
caltepec, 1340 m., 16 Sept. 1932, Hinton 1745 (BM). MORELOS: lava
fields near Yautepec (‘Yantepec’), 4500 ft., 22 Oct. 1902, Pringle
8697 (BM, C, F, GH, GOET, M, MEXU, MO, MSC, NY, PH, POM, UC, US).
GUERRERo: Rincón Viejo, 800 m., 17 Oct. 1963, Kruse 898 (MEXU);
Hwy. 95 between Ocotito and Tierra Colorado, 680 m., 21 Sept. 1967,
Handlos 484 (вн); Parotas, Dist. Mina, 800 m., 12 Sept. 1936, Hinton
et al. 9504 (ARIZ).

2. Tripogandra amplexicaulis (Klotzsch ex Clarke) Wood-

son, Ann. Missouri Bot. Gard. 29:152. 1942.

Tradescantia amplexicaulis Klotzsch ex Clarke in DC.,
Monographiae Phanerogamarum 8:804. 1881. TYPE:
México. Chiapas, etc. Sept. 1864-70. Ghiesbreght
887 (Lectotype, К; isolectotype, GH!).

Tradescantia dilatata Clarke in DC., Monographiae
Phanerogamarum 3:304. 1881, nom. nud. pro syn.

Tradescantia umbellata Pavon ex Clarke in DC., Mono-
graphiae Phanerogamarum 3:304. 1881, nom. nud.
pro syn.

246 Ећодога [Vol. 77

Descantaria amplexicaulis (Klotzsch ex Clarke) Brück-
ner, Notizbl. Bot. Gart. Berlin-Dahlem 10:56. 1927.

Plants annual; stem erect, to 92 cm tall, unbranched or
branching profusely at the base, occasionally decumbent
basally and rooting at the nodes; internodes below the
inflorescence to 11.2 ст long, glabrous in the upper part
of the plant or with a complete or partial line of uniseriate
hairs extending down the stem from the sheath above on
the side opposite the blade. Leaves narrowly ovate to
ovate; blades to 14.8 ст long, to 5.0 ст wide, glabrous on
both sides, base of variable shape, always amplexicaul
below the inflorescence, at the base of the plant either
cuneate, rounded, or narrowed and then appearing petio-
late, margin ciliate, apex acuminate, or less often, acute;
sheaths to 16.5 mm long, to 11.2 mm in diam., upper ones
completely glabrous, lower ones with a few long uniseriate
hairs at the orifice, and a complete or partial line of uni-
seriate hairs extending down the side opposite the blade.
Inflorescences terminating each stem, composed of a single
double cincinnus or paniculate with up to 14 double cin-
cinni, 1-6 amplexicaul leaves and 0-4 sheathing bracts,
сіпсіппі borne terminally and in the axils of the upper
bracts and amplexicaul leaves; peduncles to 9.1 cm long,
green and/or red, pilose distally, or rarely with only a few
seattered capitate hairs with the lowest cell enlarged;
double cincinni with up to 13 (-18) buds, flowers and/or
fruits; pedicels 3.0-7.3 mm long, to 1.0 mm in diam., erect
or spreading in fruit, green or red with a green base, pilose,
hairs capitate: bracts at the base of each pedicel pilose or
with scattered capitate hairs, margin entire. Flowers
bright pink; sepals ovate-cymbiform, 4.0-6.1 mm long, 1.5-
3.0 mm wide, green with a darker tip and midvein, or
green with a red base and margin, or red, with scattered
capitate hairs, margin entire and hyaline, apex more or
less acute; petals ovate to broadly ovate, 6.0-13.0 mm long,
ca. 3.4-11.5 mm wide, base cuneate, apex acuminate or
rounded; stamens 3, opposite the sepals, filaments 1.1-2.0

1975] Tripogandra — Handlos 247

mm long, pink, bearing a few (5-10) pink, moniliform
hairs in two rows in the middle of the dorsal side (Fig. 7),
anthers 0.8-1.8 mm long, 0.5-1.8 mm wide, whitish with a
pink or magenta line around the more or less parallel
anther sacs, dorsifixed, versatile, pollen whitish; stami-
nodes 3, opposite the petals, filaments 3.5-8.0 mm long,
slightly epipetalous, pink proximally, white distally, gla-
brous, inflated in the distal third just below the anthers
(Fig. 23), bent in an S-shape; anthers 0.7-2.2 mm long,
0.7-2.2 mm wide, dorsifixed, connective yellow, V-shaped,
with yellow, elongate more or less parallel anther sacs,
pollen sterile, yellow; ovary 0.6-1.4 mm long, 0.7-1.3 mm
in diam., white, glabrous, style 0.3-0.7 mm long, stigma
simple or capitellate. Capsule 3.1-4.5 mm long, 2.5-4.0 mm
in diam., green or brown, glabrous; seeds 2 per locule,
triangular, 1.4-2.2 mm long, dark brown or black, testa
areolate, sometimes alveolate (Figs. 64, 65), with ribs
radiating from the embryotega (Fig. 66), hilum linear
(Figs. 65, 67).

Chromosome number: n=16.

Distribution and habitat: Sonora to Chiapas, México
and Guatemala; at elevations from (300-) 600 to 2150 m
in thin layers of slightly acid soils which range from clay
to sandy loam, or humus.

Flowering: Flowering in the native habitat occurs from
mid-August through December; most flowering collections
seem to have been made in September and October. In
México and Ithaca, N.Y., flowers open before 8:00 AM
and close between 11:30 AM and 12:50 PM.

REPRESENTATIVE SPECIMENS

México. SONORA: Sierra Charuco, Río Mayo, 10 Sept. 1935, Gentry
1706 (ARIZ, F. GH, MO, UC, US). CHIHUAHUA: Guayanopa Canyon,
Sierra Madre Mts., 23 Sept. 1903, Jones (POM). SINALOA: 16.3 miles
S.W. of El Paraíso, on road between Villa Unión and El Salto, 27
Sept. 1953, Ownbey & Ownbey 1903 (Е, MICH, NY, UC, US). DURANGO:
La Bajada. Tamazula, 300-600 m., Nov. 1921, Ortega 4349 (08).
NAYARIT: hills back of Jalisco, 11 Nov. 1925, Ferris 5981 (ps).
JALISCO: km. 69 of Hwy. 41, N of Guadalajara, 5 Oct. 1967,
Handlos 448 (BH). MICHOACAN: 11 miles below Uruapan, kms. 95-

248 Ећодога [Vol. 77

96, on road to Apatzingán, 1160 m., 11 Sept. 1961, Moore, Jr. &
Bunting 8750 (BH, MEXU, UC). MEXICO: Rincón del Carmen, Dist.
Temascaltepec, 2 Dec. 1935, Hinton et al. 8762 (ARIZ, GH, MICH, NY,
US). MORELOS: km. 125.3 of Hwy. 95D, 6.2 km. west of bridge over
Rio Amacuzac, ca. 1200 m.. 21 Sept. 1967, Handlos 431 (BH).
PUEBLA: Barranca de Chochonotla, Municipio de la Unión, 4 km.
al E. de Xicotepec de Juárez, 27 Sept. 1964, González Quintero 1674
(MICH, MSC). GUERRERO: Cañon del Mano, railroad tracks north
of Iguala, ca. 3 km. N of El Naranjo, ca. 840 m., 13 Sept. 1967,
Handlos 417 (BH). CHIAPAS: along Hwy. 190 in the Zinacantán
paraje of Muctajoc, Municipio of Ixtapa, 3500 ft., 26 Oct. 1965,
Breedlove 13797 (DS, Е, MEXU). Guatemala. HUEHUETENANGO: along
road between San Sebastián H. and San Rafael Pétzal, 1900-2000 m.,
14 Aug. 1942, Steyermark 50537 (F). GUATEMALA: 1939, Aguilar
374 (F). JALAPA: mountains about Chahuite, northwest of Jalapa,
about 1650 m., 16 Nov. 1940, Standley 77472 (F). SANTA ROSA:
Cenaguilla, 4000 ft., Dec. 1892, Heyde & Lux 4284 (GH, M, NY, US).
JUTIAPA: hills between Jutiapa and Plan de Urrutia, north of
Jutiapa, 900-1200 m., 28 Oct. 1940, Standley 75525 (F).

The history of the name Tripogandra amplexicaulis
and the specimens associated with it must be considered
to understand its present application, In the original de-
scription, Clarke (1881) credits Klotzsch for the name
Tradescantia amplexicaulis in the following fashion,
“(Klotzsch ms. in herb. Berol.)". As the Berlin herbarium
was largely destroyed during World War II it is not
possible to examine the specimens that Klotzsch may have
seen. However, type specimens of the Commelinaceae in
the basement of the Berlin herbarium did not burn (Pil-
ger, 1957) and among them is a specimen labelled “уриѕ!””
collected by C. Ehrenberg in “Mejico” and annotated as
Tradescantia amplexicaulis by C. B. Clarke in his own
hand. This may be the specimen Klotzsch would have
designated as the type for his name. Klotzsch never pub-
lished his description so this specimen has no standing
under the current rules. Clarke does not mention the speci-
men in his description of the species. The Ehrenberg
specimen represents Tripogandra amplexans. Clarke listed
eight collections of Tradescantia amplexicaulis (Schaffner
108, 138; Botteri 531, 892; Ghiesbreght 887; Salvin;
Savage; Hoffman) which are syntypes because no holo-

1975] Tripogandra — Handlos 249

type was designated. Additionally he cited Tradescantia
umbellata Pavon as a synonym and I have seen one speci-
men so labelled from the British Museum and annotated
by Clarke as T. dilatata, his manuscript пете for T.
amplexicaulis.

The Pavon specimen (T. umbellata). Schaffner 108, and
Botteri 581 represent Tripogandra purpurascens. Botteri
892 and Ghiesbreght 887 represent the taxon presently
under consideration and the remaining specimens have
not been seen. The original description given by Clarke
does not overwhelmingly refer to any one species. То
preserve current usage of the name T. amplexicaulis I
therefore designate Ghiesbreght 887 (K) as the lectotype
for T. amplexicaulis. The following phrases from Clarke's
description are taken as referring to T. amplexicaulis
sensu strieto and distinguishing it from T. purpurascens.

“Pedunculis quasi paniculam efformantibus, . . . Folia
. acuminata, . . . vaginae ore glabratae; folia summa
fere ad vaginas reducta. . . . fructus tempore patenti-

erecta." The remainder of Clarke's original description
could apply to both species. I do not understand his state-
ment, “Ovarium apice vix aut minute pubescens." I have
never seen hairs оп the ovary of any specimens of Tripo-
датата.

In the past Tripogandra amplexicaulis and Т. amplexans
have been considered the same species. I prefer to separate
them on the basis of a number of morphological characters,
a difference in flowering season and the fact that I have
found both species growing side by side with little or no
evidence of hybridization as discussed under T. amplexans.

3. Tripogandra angustifolia (Robinson) Woodson, Ann.
Missouri Bot, Gard. 29:152. 1942.
Tradescantia angustifolia Robinson, Proc. Amer. Acad.
Arts 27:185. 1893. TYPE: México. SAN LUIS POTOSI:
Las Canoas, 14 Aug. 1891, Pringle 3902. (Holo-
type, GH!; isotypes. B!, BM !, ВЕ!, EL GOET!, м!, Mol,
MSC!, NY!, PH!, UC!).

250 Ећодога [Vol. 77

Descantaria angustifolia (Robinson) Briickner, Notizbl.
Bot. Gart. Berlin-Dahlem 10:56. 1927.

Ilustrations: Matuda, Anales Inst. Biol. Nac. México
26:368. 1956 (‘1955’).

Plants annual; stem erect, to 24 cm tall, unbranched
or branched; internodes to 5.0 cm long, green or red,
glabrous except for a line of uniseriate hairs extending
straight down the side opposite the leaf blade from the
sheath above. Leaves linear; blades to 7.1 ст long, 2.7 mm
wide, terete or C-shaped in cross section, glabrous dorsally
and ventrally, margin usually ciliate in the proximal por-
tion or occasionally to the tip, acute at the apex; sheaths
to 4.1 mm long, to 4.0 mm in diam., with a few long uni-
seriate hairs at the orifice, otherwise glabrous except for
a line of uniseriate hairs extending down the side opposite
the lamina. Inflorescences composed of 1-4 double cincinni
borne terminally and in the axils of the upper leaves;
peduncles to 7.5 cm long, green or red, glabrous or some-
times with 1-2 lines of uniseriate hairs extending down one
side; buds, flowers and/or fruits 2-10 per double cincinnus;
pedicels to 8.0 mm long, to 0.6 mm in diam. at anthesis,
green or red, glabrous, erect in fruit; bracts at the base
of each pedicel glabrous, margin entire, erose, or some-
times ciliate, lowest 1-2 bracts sometimes leaf-like. Flowers
pink, occasionally white; sepals elliptic- or ovate-cymbi-
form, to 4.5 mm long, to 2.0 mm wide, green, green with
a darker tip, or red, glabrous, margin entire, hyaline, apex
acute; petals broadly ovate, to 5.0 mm long, to 4.0 mm
wide, cuneate basally, apex acute; stamens 3, opposite the
sepals, filaments to 1.5 mm long, white or pink, nearly
glabrous except for a few minute hairs, anthers 0.3-0.8 mm
long, 0.4-0.7 mm wide, pink, connective bent in the middle
and U-shaped with the anther sacs parallel (Fig. 14),
pollen white; staminodes 3, opposite the petals, filaments
epipetalous, to 4.0 mm long, pink and/or white, glabrous,
bent in an S-shape distally (Fig. 30), anthers 0.7-1.1 mm
long, 0.7-1.1 mm wide, versatile, connective yellow, promi-

1975] Tripogandra — Handlos 251

nent, C-, U- or V-shaped, with the yellow anther sacs di-
vergent (Fig. 27), pollen white, cream, or yellow, sterile;
ovary 0.5-1.0 mm long, 0.4-1.0 mm in diam., green, gla-
brous, style 0.3-0.7 mm long, stigma simple or capitellate.
Capsule globose, 2.0-3.0 mm long, 1.5-2.4 mm in diam.,
brown, glabrous; seeds 2 per locule, triangular, 0.9-1.3 mm
long, brown or gray, testa ribbed farinose (Fig. 74), hilum
punctiform (Fig. 75), embryotega impressed,

Chromcsome number: n=8.

Distribution and habitat: central and southern México
and western Guatemala from the states of San Luis Potosí,
west to Guerrero and south to Chiapas in México and the
department of Huehuetenango in Guatemala; in thin alka-
line soil on limestone cliffs and ledges.

Flowering: In the native habitat, flowering occurs in
August through October. In a uniform environment, plants
collected at different locations flower at different times,
probably indicating physiological variation in response to
differing environmental factors in the various localities.
Flowers open between 7:00 AM and 8:15 AM — earlier
in sunny locations and later in shady locations. Closing
occurs between 11:00 and 11:30 AM.

REPRESENTATIVE SPECIMENS

México. SAN LUIS POTOSI: Pozo de Azuna, 15 km. al E. de Gua-
dalacázar, 1450 m., 26 Sept. 1955, Rzedowski 6695 (MEXU, MICH).
HIDALGO: between Jacala and Barranca Seca via Hilo Juanico, Dist.
Jacala, 1400-1600 m., 30 Oct. 1946, Moore, Jr. 1800 (BH, GH). VERA-
CRUZ: Banos del Carrizal, Aug. 1912, Purpus 6174 (F, GH, MO, NY,
UC, US). MEXICO: rounded hill with crater in the center about 0.5
km. М. of village of Tonatico, 23 Aug. 1965, Handlos 169 (BH).
MORELOS: near Yautepec, Aug. 1903, Rose & Painter 6575 (сн).
PUEBLA: km. 298.3 of Hwy. 190, ca. 12.8 km. south of Acatlán, 1340
m., 11 Sept. 1967, Handlos 406 (BH). GUERRERO: km. 231.6-7 of Hwy.
95, 37.7 km. north of Chilpancingo, 30 Aug. 1967, Handlos 388 (BH).
OAXACA: Cerro de San Antonio de la Cal, 1700 m., 18 Aug. 1907,
Conzatti 1995 (Е, MICH). CHIAPAS: km. 1062.5 of Hwy. 190, 4 km.
south of Berriozabal, 930 m., 18 Aug. 1967, Handlos 364 (вн).
Guatemala. HUEHUETENANGO: dry slopes between San Ildefonso
Ixtahuacán and Cuileo, 1350-1600 m., 16 Aug. 1942, Steyermark
500694 (F, MO, US).

252 Ећодога [Vol. 77

Tripogandra angustifolia is unique for its narrow leaves
which are nearly terete or C-shaped in cross-section. These
leaves are probably a special adaptation which allows the
plants to store water between rains and grow in very dry
areas. Small plants can survive and produce seeds with
as few as three or four leaves and two to four flowers in
a single terminal inflorescense. To illustrate how densely
plants of T. angustifolia grow, in 1967 I arbitrarily se-
lected an area of 100 square cm, ten cm to a side. I care-
fully removed each plant and discovered there were exactly
240 plants in that area! Each plant was surviving on an
average of 0.416 square em of space.

This species seems to be the only one possessing very
short hairs on the staminal filaments. These obscure hairs
may indicate that the ancestor of Tripogandra angustifolia
had a bearded filament and that as an adaptation for water
conservation these structures were reduced but not com-
pletely lost.

The seeds of Tripogandra angustifolia are distinctive for
their triangular outline and the conspicuously ribbed sides.
Maintenance of this species in cultivation has proved a
problem because seed germination has been very low.
Field-collected seeds have produced one or two or no seed-
lings at all where seeds nearly covered the surface of a
pot of soil. This contrasts with conditions in México where
plants grow in very dense stands, implying a high per-
centage of seed germination.

Tripogandra angustifolia is morphologically very uni-
form throughout its range which extends for some 1270
km from north to south. The variations which occur —
white or pink flowers, red or green stems or leaves — occur
within a population and do not distinguish one population
from another. One or two leaf-like bracts occur rarely
at the base of an inflorescence and may indicate an ancestral
inflorescence similar to that of present day Tradescantia
which has large leaf-like bracts. There are slight varia-
tions in intensity of flower color and petal shape in plants

1975] Tripogandra — Handlos 253

from different populations, but the significance of these
variations is unknown. The general lack of morphological
variation may be due to the specialized and uniform habitat
which this species inhabits — the thin layer of alkaline
soil found in cracks or on ledges of limestone rock through-
out southern México and parts of Guatemala — where it
seems to be specially adapted to survive. As soil depth
increases, other species of plants seem to be better adapted
to compete and T. angustifolia is crowded out.

4. Tripogandra brasiliensis Handlos, sp. nov. HOLOTYPE:
Brazil. MARANHAO: “Ilha de Balsas" region, between
the Balsas and Parnaiba Rivers. Ca. 6 km. north of
main house of Fazenda “Morros”, ca. 30 km. south of
Loréto, са. 300 m., 30 April 1962, Eiten & Hiten 4458
(NY!).

Herba аппџа (7) ; caulis erectus, usque ad 52.5 em altus,
internodiis usque ad 13.4 ст longis, linea unica pilorum
instructis, aliter glabris ve! pilis dispersis pubescentibus.
Folia angusto-ovata, petiolata, laminis usque ad 8.7 cm
longis, usque ad 2.1 em latis, apice acuminatis, basi obliquis,
margine ciliatis, dorsaliter pilosis vel pilis dispersis in-
structis, ventraliter glabris praeter lineam unicam pilorum
secus costam, vaginis usque ad 6.0 mm longis, usque ad
3.6 mm diam., orificio villosis, linea unica pilorum in-
structis, aliter glabris vel pilis dispersis praeditis. Inflo-
rescentiae terminales et in axillis foliorum summorum,
ex 1-2 cincinnis duplicibus constantantes; pedunculi usque
ad 3.9 ст longi, pilis paucis dispersis pubescentes; pedicelli
usque ad 3.0 mm longi, glabri, bracteis basi pedicellorum
glabris, margine integris. Flores albi; sepala ovato-cymbi-
formia, usque ad 4.2 mm longa, usque ad 2.7 mm lata,
glabra vel 1-2 pilis instructa, margine hyalina, apice
obtusa; petala non visa; stamina 6 in verticillis duobus,
3 sepalis opposita filamentibus brevis, usque ad 1.5 mm
longis, glabris, antheris usque ad 0.7 mm longis, usque
ad 0.6 mm latis, 3 petalis opposita filamentibus longioribus,
usque ad 3.9 mm longis, dilatatis et barbatis distaliter,

254 Ећодога [Vol. 77

antheris usque ad 0.4 mm longis, usque ad 0.9 mm latis,
connectivo elongato; ovarium usque ad 1.5 mm longum,
usque ad 1.0 mm diam., glabrum, stylo usque ad 0.3 mm
longo, stigmate simplici. Capsula globosa, usque ad 3.2 mm
longa, usque ad 3.3 mm diam., glabra, seminibus triangu-
laribus, usque ad 2.0 mm longis, costatis, areolatis (Fig.
4T), hilo elliptico (Fig. 48).

Vernacular name: baixáo do cipó, fide Eiten & Eiten.

Distribution: known from only three locations in
Maranhão and Minas Gerais, Brazil. The habitat of this
species, according to Eiten and Eiten, was tall forest along
a gully in a disturbed area with a pronounced dry season
and intermittent streams.

SPECIMENS EXAMINED

Brazil. MARANHAO: 30 April 1962, Етеп & Епеп 4458 (NY).
MINAS GERAIS: Lagoa Santa, Warming (Е p.p. US p.p.); Lagoa
Grande, Belo Horizonte, Mar. 1935, Cocheaw (R).

This species is recognizable from its probable annual
habit, petiolate leaves, peduncle with scattered uniseriate
hairs, an inflated filament on the longer stamens, and the
large ribbed areolate seeds with elliptical hilum. While
the collections known to me are few, the species is un-
doubtedly distinct from other South American species of
Ттродатата.

5. Ттїро апага disgrega (Kunth) Woodson, Ann. Missouri
Bot. Gard. 29:152. 1942.

Tradescantia disgrega Kunth, Enumeratio Plantarum
4:97. 1843. TYPE: México. Serro Colorado, Aug.
1828, Schiede 974 (816) (Lectotype, B!; isolectotype,
HAL!).

Tradescantia ehrenbergiana Klotzsch ex Clarke in DC.,
Monographiae Phanerogamarum 3:305, 1881, nom.
nud. pro syn.

Disgrega mexicana Hasskarl ex Clarke in DC., Mono-
graphiae Phanerogamarum 3:305. 1881, nom. nud.
pro sym.

1975] Tripogandra — Handlos 255

Descantaria disgrega (Kunth) Briickner, Notizbl. Bot.
Gart. Berlin-Dahlem 10:57. 1927.

Tradescantia disgrega forma glandulosa Standley &
Steyermark, Field Mus. Nat. Hist., Bot. ‘Ser. 23:36.
1944. HOLOTYPE: Guatemala. ZACAPA: Sierra de las
Minas, along trail between Rio Hondo and summit
of mountain at Finca Alejandria, 1000-1500 m., 11
Oct. 1939, Steyermark 29751 (Е!) ; photograph (F!).

Tradescantia disgrega forma pubescens Standley &
Steyermark, Field Mus. Nat. Hist., Bot. Ser. 23:37.
1944. HOLOTYPE: Guatemala. GUATEMALA: Borraja,
1085 m., Oct. 1928, Morales R. 1106 (F!); photo-
graph (F^.

Tripogandra disgrega forma glandulosa (Standley &
Steyermark) Standley & Steyermark, Fieldiana:
Bot. 24(3) :37. 1952.

Tripogandra disgrega forma pubescens (Standley &
Steyermark) Standley & Steyermark, Fieldiana:
Bot. 24(3) :37. 1952.

Plants annual, rooting at lower nodes; stem erect, to
53 em tall, branched or unbranched, some plants with the
lowermost portion decumbent; internodes to 12.5 ст long,
pilose or glabrous except for a line of uniseriate hairs ex-
tending down the side of the stem, continuous with the
line on the sheath above. Leaves narrowly ovate to ovate;
blades to 9.8 em long, 3.8 em wide, cuneate, rounded, or
subpetiolate at the base, glabrous to pilose on both sides,
the uppermost leaf usually with fewer or no hairs, margin
ciliate, apex acute or acuminate; sheaths to 15.0 mm long,
4.3 mm in diam., villous at the orifice, a line of uniseriate
hairs extending down the side opposite the blade, other-
wise glabrous to pilose. Inflorescences composed of 1-5
double cincinni borne terminally and in the axils of the
upper leaves; peduncles to 9.5 (-15) em long, glabrous or
with a few capitate hairs near the distal end; double cin-
cinni with up to 25 buds, flowers and/or fruits; pedicels
to 7.0 mm long, glabrous proximally but pilose distally,

256 Ећодога [Vol. 77

reflexed in fruit; bracts at the base of each pedicel pilose
or with scattered capitate hairs, margin entire. Flowers
pink; sepals ovate- or elliptic-cymbiform, to 6.0 mm long,
3.0 mm wide, green or with red at the tip and/or base,
pilose or with scattered capitate hairs 1.5-4.5 mm long,
margin hyaline, apex more or less acute; petals broadly
ovate, to 8.0 mm long, to 7.0 mm wide, base cuneate, apex
acuminate; stamens 6, in two whorls, the outer shorter,
filaments to 2.0 mm long with a few (ca. 10) moniliform
hairs borne on the middle of the dorsal side, anthers 0.7-1.6
mm long, 0.5-1.4 mm wide, dorsifixed, versatile, with
anther sacs slightly spreading (not parallel); stamens of
the inner whorl longer, filaments to 8.0 mm long, glabrous,
with a U- or open S-shaped bend and inflated in the upper
half, anthers to 1.5 mm long, to 1.6 mm wide, basifixed,
connective conspicuous, anther sacs either divergent or
becoming parallel and adjacent through bending of the
connective in a U-shape; ovary to 1.1 mm long, to 1.8 mm
in diam., glabrous, style 0.4-0.7 mm long, stigma simple
or capitellate (minutely penicilliform). Capsule globose,
to 3.5 mm long, to 3.5 mm in diam., green or brown at
maturity, glabrous; seeds 2 per locule, triangular or rarely
elliptical, 1.2-1.7 mm long, brown to black, testa areolate,
with ribs (sometimes obscure) radiating from the em-
bryotega (Fig. 52), hilum punctiform (Fig. 53).

Vernacular name: yerba del pollo fide Hinton.

Distribution and habitat: from Jalisco to Chiapas,
México, Guatemala, Honduras and Е1 Salvador; in moist
areas along streams or in pine forests, occasionally a weed
in cornfields.

Flowering: This species flowers from August to Novem-
ber in different parts of México, and from October to
February in Guatemala. The flowers open before 8:45 AM
and begin closing about 11:25 AM.

REPRESENTATIVE SPECIMENS

México. JALISCO: Sierra del Tigre, 3 miles south of Mazamitla,
2100-2200 m., 22 Sept. 1952, MeVaugh 13167 (MICH). HIDALGO:
vicinity of Molango on Lolotla road, Municipality Molango, District

1975] Tripogandra — Handlos 257

Molango, 1600 m., 9 Nov. 1946, Moore, Jr. 1984 (BH). VERACRUZ:
Orizaba, Botteri 326 (GH). MICHOACAN: roadsides in pine zone about
48 kms. from Pátzcuaro on road to Tacámbaro, ca. 6-7000 ft., 2 Sept.
1948, Moore, Jr. & Wood, Jr. 4847 (BH). MEXICO: Ypericones, Dist.
Temascaltepec, 23 Nov. 1935, Hinton et al. 8718 (GH, MICH, MO, NY).
MORELOS: in pine forest, km. 61, off Cuernavaca-Mexico City High-
way, 2 Oct. 1943, Lundell & Lundell 12496 (MICH, NY, UC). OAXACA:
15 km. al 5. de Sola de Vega, sobre la carretera a Puerto Escondido,
1800 m., 30 Sept. 1965, Rzedowski 21296 (MICH). CHIAPAS: sitio in
San Cristóbal las Casas, Municipio of San Cristóbal las Casas, 7100
ft, 20 Sept. 1965, Breedlove 12310 (ps). Guatemala. SAN MARCOS:
barrancos 6 miles south and west of town of Tajumulco, north-
western slopes of Volcán Tajumulco, 2300-2800 m., 26 Feb. 1940,
Steyermark 36621 (F). CHIMALTENANGO: above Las Calderas, 1800-
2100 m., 15 Dec. 1938, Standley 60014 (F). SACATEPEQUEZ: Ciudad
Vieja, Nov. 1914, Tejada 301 (US). GUATEMALA: near San Juan
Sacatepéquez, about 1800 m., 8 Dec. 1938, Standley 59258 (F). SANTA
ROSA: Santa Rosa, 3000 ft., Nov. 1892, Heyde & Lux 4285 (GH, NY,
US). Honduras. MORAZAN: colinas cultivadas de maíz de la Monta-
fita, 1500 m., 18 Nov. 1948, Molina R. 1610 (GH, US). EL PARAISO:
slopes above Yuscarán, Montserrat, 1500 m., 25 Nov. 1958, Hawkes,
Hjerting & Lester 2061 (c, к). El Salvador. SANTA ANA: Hda. Los
Planes nórdl. Metapán, 1800 m., 29 Oct. 1950, Rohweder 707 (F).
MORAZAN: eastern edge, finca of General J. T. Calderón, Montes de
Cacaguatique, са. 1340 m., 28 Dec. 1941, Tucker 646 (BH p.p., Е р.р.,
MICH p.p., MO, NY р.р., PH p.p. UC p.p., US р.р.).

Schlechtendal (1831) described but did not name a plant
of Tripogandra disgrega from a specimen (Schiede 974
(815)) housed at Halle. Kunth later named the species
and his description was drawn from a duplicate specimen
of Schiede's housed in Berlin *(Descr. juxta specimen
Schiede.)". In all respects the description and specimen
are in accord. I have seen two sheets of this collection
and because there is no evidence that Kunth saw the sheet
at Halle, I designate the Berlin specimen as the lectotype,
therefore making Schlechtendal's material at Halle an iso-
lectotype.

Kunth's original description cites another specimen,
Berlandier 948. There is such a specimen in the Berlin
herbarium but it is not annotated by Kunth so perhaps
he did not see this particular sheet. This collection is a

258 Rhodora [Vol. 77

specimen of Tripogandra purpurascens. These two species
are similar апа, as the Berlandier specimens are widely
distributed, the name T. disgrega has been applied con-
sistently to two different species. As Kunth drew his
description from the Schiede specimen, no name other than
T. disgrega can be applied to this species. There seems
to be no other course if one believes there are two species
but to call this one T. disgrega and to apply another name
to the species represented by Berlandier 948.

Tripogandra disgrega has some variable morphological
characteristics but I am unable to see any trends or pat-
tern in the variation. The species is rare and has not been
collected often. The vesture of the leaves varies consider-
ably but not in any consistent way. The leaves may be
glabrous on both sides or may have a varying amount of
hair ranging from scattered to pilose. When hairs are
found on the leaves they are also present on the inter-
nodes and on the leaf sheaths. Tripogandra disgrega forma
pubescens represents the extreme condition in pilosity.

There does seem to be some correlation between the
presence of hairs on the leaves and a narrowing at the
base of the blade producing a subpetiolate appearance.
More of the pilose-leaved plants have a narrowed blade
than do the glabrous-leaved plants, but the significance, if
any, of this correlation is not evident.

Tripogandra disgrega and T. purpurascens are very
similar ir most respects and on this basis I would con-
sider them very closely related. One difference separates
all specimens and a second difference separates most.
Tripogandra disgrega has long hairs on the sepals, the
longest from 1.5-4.5 mm long, while the hairs on the sepals
of T. purpurascens are shorter and vary from 0.2-1.0 mm
long on dried specimens. Most plants of T. disgrega have
a glabrous peduncle, but in three of 31 collections there
were scattered capitate hairs on the distal end. T'ripo-
gandra purpurascens is more variable, plants may have
glabrous peduncles or any single plant may have both

1975] Tripogandra — Handlos 259

glabrous and variously vestite peduncles, ranging from
scattered hairs to pilose and/or with one or two lines of
hairs.

The following differences between these species generally
hold but there are exceptions which may be due partly to
environmental influences, may be due also to genetic effects,
or could be the result of undetected hybridization. Tripo-
gandra disgrega usually has flat open leaves which may be
subpetiolate, the plants grow in shady habitats, the petals
are acuminate distally, the dorsal surface of the seeds is
rounded or convex, the seeds are often obscurely ribbed,
and in any given location the peak of flowering seems to
occur later in each season than the peak of flowering for
T. purpurascens. In contrast, T. purpurascens has compli-
cate (folded or canaliculate) leaves which are usually
rounded at the base and never narrowed and petiolate,
the plants grow in open sunny locations, the petals are
irregularly indented or acuminate at the apex, the seeds
are prominently ribbed and the dorsal surface is flat or
even concave with a deeply impressed embryotega, and
flowering seems to occur earlier in each season in a given
location than the peak of flowering for T. disgrega.

6. Tripogandra diuretica (Martius) Handlos, comb. nov.

Tradescantia diuretica Martius in Spix & Martius,
Reise in Brasilien 281. 1823. HOLOTYPE: Brazil.
Min. Ge. et S. Pauli, Martius (M!).

Tradescantia commelina Vellozo, Florae Fluminensis
140; 3:154. 1829 (1825). HOLOTYPE: apparently
lost,

Tradescantia diuretica B foliis vaginisque magis glabris
Schultes in Schultes & Schultes, Systema Vegeta-
bilium 7:1163. 1830. HOLOTYPE: Brazil. Martius
(м!).

Tradescantia gaudichaudiana Kunth, Enumeratio Р1ап-
tarum 4:98. 1843. HOLOTYPE: Brazil. Rio Janeiro,
1832, Gaudichaud 125 (B!).

260 Ећодога [Vol. 77

Tradescantia mollis Kunth, Enumeratio Plantarum

4:95. 1843. TYPE: Brazil. 1836, Sello 565 (Holo-
type, B!; isotype, B!).

Tradescantia sellowiana Kunth, Enumeratio Plantarum
4:93. 1843. TYPE: Brazil Vittoria-Bahia, 1836,
Sello 1006 (Holotype, B!; isotype, B!).

Tradescantia diuretica а mollis (Kunth) Seubert in
Martius, Flora Brasiliensis 3 (1) :251. 1855.

Tradescantia diuretica В glabriuscula Schultes ex Seu-
bert in Martius, Flora Brasiliensis 3(1) :251. 1855.
HOLOTYPE: Brazil Yrino, Sello 1309 (B!).

Tradescantia elongata B diuretica (Martius) Clarke in
DC., Monographiae Phanerogamarum 3:303. 1881.

Резсатвата diuretica Schlechtendal fide Hasskarl ex
Clarke in DC., Monographiae Phanerogamarum at
304. 1881, nom. mud. pro sym.

Tripogandra elongata forma diuretica (Martius)
Standley & Steyermark, Fieldiana: Bot. 24 (3) :38.
1952.

Illustrations: Bacigalupo, Darwiniana 14:407, 410.
1967; Figs. 4, БК, 1, as Tripogandra elongata.

Plants perennial, the base decumbent, rooting at the
nodes; stems to 11.5 dm long, flowering stems erect, rarely
branched; internodes to 12.1 cm long, with a complete or
partial line of uniseriate hairs extending down the side
from the sheath above, otherwise glabrous or rarely com-
pletely pilose or pilose distally. Leaves narrowly ovate to
ovate; blades to 14.2 ст long, to 2.6 ст wide, base oblique,
rounded on one side, cuneate on the other, dorsal and
ventral side glabrous to pilose, midvein with or without
a partial or complete line of uniseriate hairs ventrally,
margin ciliate; sheaths to 20.0 mm long, to 9.5 mm in
diam., villous at the orifice, with a line of uniseriate hairs
extending down the side opposite the blade, otherwise
glabrous to pilose. Inflorescences of 1-10 double cincinni
borne terminally and in the upper 1-4 leaf axils; peduncles
to 8.9 cm long, usually glabrous, rarely pilose or with a

1975] Tripogandra — Handlos 261

few scattered uniseriate hairs; buds, flowers and/or fruits
to 17 per double cincinnus; pedicels 3.5-9.0 mm long,
glabrous, reflexed in fruit; bracts at the base of each
pedicel glabrous or rarely pilose or with a few scattered
uniseriate hairs, margin entire, with a few uniseriate
hairs. or ciliate. Flowers pink; sepals narrowly ovate to
ovate, cymbiform, 4.5-7.0 mm long, 1.4-3.3 mm wide,
glabrous or with a few uniseriate hairs at the apex, rarely
pilose, margin hyaline, apex acute; petals more or less
elliptie, to 10.0 mm long, to 6.5 mm wide, base cuneate,
apex acute to rounded; stamens 3, opposite the sepals, fila-
ments 1.2-2.3 mm long, glabrous or with a few (2-3) hairs,
anthers 1.0-1.8 mm long, 0.6-1.1 mm wide, basifixed, anther
saes more or less parallel; staminodes 3, opposite the petals,
4.0-6.3 mm long, bearded in the upper third or fourth
with moniliform hairs, distal end bent in an S-shape, an-
thers 0.7-1.4 mm long, 0.8-1.5 mm wide, basifixed, with
parallel. yellow anther sacs, pollen sterile; ovary 0.7-1.4
mm iong, 0.7-1.0 mm in diam., glabrous, style 0.6-1.1 mm
long, stigma simple. Capsule globose or obovoid, 2.8-3.5 mm
long, 2.1-3.0 mm in diam., glabrous, brown; seeds 1-2 per
locule, triangular, the dorsal side convex, 1.2-1.8 mm long,
light gray, testa ribbed, areolate-foveate (Fig. 40), hilum
elliptic or punctiform (Fig. 41).

Chromosome number: 2n—64 (А. Sparrow, pers.
comm.).

Vernacular name: trepueraval, jupirava tupice fide
Martius; trepoerava fide Peckolt, Brazil.

Distribution and habitat: southern Brazil, Bolivia,
Paraguay, Argentina, and Uruguay; commonly in low wet
places near sea level to 1600 (-2200) m. This species in
Brazil seems adapted to moist swampy locations and is not
found in limestone outcrop areas as are many of the species
in México (Father Reitz, verb. comm.).

Flowering: Flowering occurs from December to July.

REPRESENTATIVE SPECIMENS

Bolivia. LA PAZ: San Bartolomé, near Calisaya, Basin of Rio Bopi,
province of S. Yungas, 750-900 m., 1-22 July 1939, Krukoff 10545

262 Ећодога [Уо]. 77

(GH, к). COCHABAMBA: vic. Cochabamba, 1891, Bang 1282 (BM,
E, F, G, GH, NY, US). Paraguay. Mbuveva, Apr. 1931, Jorgensen 4355
(Us). Brazil. PERNAMBUCO: Pesqueira, 20 Jan. 1943, Vellozo 561
(R). MATTO GROSSO: Smith 818 (R). MINAS GERAIS: Vicosa, road
from Canella to Repressa, Buraco Canella, 730 m., 7 Jan. 1930,
Mexia 5487 (BH, BM, Е, GH, LA, MICH, MO, NY, PH, UC, US). RIO DE
JANEIRO: Theresopolis, na Serra dos Orgãos, Jan. 1955, Vidal V-120
(R). FEDERAL DISTRICT: Botanical Garden and vicinity, Nov. 1915,
Curran 7 (US). SAO PAULO: Parque do Estado de Sao Paulo, 26
Mar. 1931, Hoehne 27392 (в, NY, SP). PARANA: Jaguariahyva, 28
Mar. 1916, Dusén 18037 (F, MO, NY, US). SANTA CATARINA: Mina
Velha, Garuva, S. Francisco do Sul, 10 m., 26 Mar. 1958, Reitz &
Klein 6618 (му, UC, US). RIO GRANDE DO SUL: Torres, Municipio
de Torres, Feb. 1939, Vidal (R). Uruguay. MONTEVIDEO: Miguelete,
10-20 m., May 1925, Herter 149 (B, BH, F, GH, M, MICH, MO, MSC,
NY, ©, UC, WIS). Argentina. MISSIONES: Dep. San Pedro, Loc.
Avellaneda, 8 Apr. 1949, Schwindt 1535 (BR, NY, US). BUENOS AIRES:
Partido de Delta, Paraya Mini, 18 May 1950, Cabrera 10635 (15).

The plants which I have included in Tripogandra diure-
tica generally have been considered a form of T. elongata
by taxonomists. The differences in the seeds, flowers, ratio
of style length to ovary length, leaf vesture when present,
and habitat are of a greater magnitude than those which
serve to distinguish forms. Furthermore, I consider T.
elongata sensu stricto to be composed of sterile material
which I include in 7. serrulata. The problem is further
discussed under that species.

Variation in leaf vesture is seen among the collections
of Tripogandra diuretica, but does not seem to follow any
discernible geographic pattern.

Herbarium material of this species is unusual because
a high percentage of specimens possesses open, dried flowers.
This is related to the fact that flowers on plants growing
in the greenhouse remain open all day and are longer lived
than those of other species.

7. Tripogandra encolea (Diels) Macbride, Revista Univ.
(Cuzco) 33(87) :142. 1944.

Tradescantia encolea Diels in Urban, Bot. Jahr. Syst.

37:381. 1906. TYPE: Perú. CAJAMARCA: San Pablo,

1975] Tripogandra — Handlos 263

2200-2400 m., 1906, Weberbauer 3855 (Holotype, B!;
isotype, G!).

Descantaria encolea (Diels) Briickner, Notizbl. Bot.
Gart. Berlin-Dahlem 10:56. 1927.

“Tripogandra encolea (Diels) Rohweder", Abh. Aus-
landsk., Reihe C, Naturwiss. 18:156. 1956.

Plants trailing, decumbent, stems branching and rooting
at the nodes; flowering stems upright, to 65 cm tall, 0.1-
0.4 em in diam. when dry, green; internodes 1.5-9.5 cm
long, always with a line of uniseriate hairs extending ina
line laterally down the internode from the sheath above,
otherwise glabrous basally and pilose distally in the erect
part of the plant. Leaves narrowly ovate; blade 1.9-7.4 cm
long, 0.9-2.6 em wide, reduced to sheathing bracts near the
top of the flowering stem, tapering or cordate at the base
on the lower part of the stem, usually amplexicaul at the
base below the inflorescence, glabrous ventrally, glabrous
dorsally or sometimes pilose near the sheath, hairs uni-
seriate, margin ciliate, apex acute; sheaths 0.4-2.9 cm long,
0.25-0.9 em in diam. when dry, villous at the orifice, other-
wise glabrous except for a line of uniseriate hairs laterally
opposite the leaf blade; sheathing bracts glabrous, pilose,
or with scattered capitate hairs. Inflorescences composed
of 1-4 double cincinni borne terminally and in the axils of
the upper leaves; peduncles 1.0-7.0 cm long, ca, 1 mm in
diam. when dried, glabrous or pilose at the base, densely
pilose near the грех, hairs capitate, brown or colorless;
double cincinni with 7-18 buds, flowers and/or fruits;
pedicels of flowers near anthesis 0.3-0.7 cm long, pilose
with capitate hairs; bracts at the base of each pedicel
glabrous or pilose with capitate hairs. Flowers white or
pink; sepals ovate-cymbiform, 4.5-6.2 mm long, 1.5-9.0 mm
wide, pilose with brown or colorless capitate hairs, apex
obtuse, margin entire; petals obovate, ca. 4-5.8 mm long,
ca. 2.5-4.5 mm wide, tapering at the base, apex obtuse;
stamens of outer whorl 3, shorter, filaments before anthesis
1.3-2.0 mm long, glabrous, anthers 0.9-1.5 mm long, 0.7-

264 Ећодога [Vol. 77

1.0 mm wide, white, dorsifixed, versatile, with parallel
anther sacs; stamens of inner whorl longer, filaments 3.0-
3.5 mm long, densely bearded with moniliform hairs in the
distal portion, anthers 0.8-1.0 mm long, 0.7-1.1 mm wide,
basifixed, versatile, with yellow anther sacs parallel or
diverging slightly, longer than the connective; ovary 0.9-
2.0 mm long, 0.8-1.2 mm in diam., glabrous, style 0.3-0.7
mm long, stigma simple or capitellate. Capsule brown,
glabrous, ca. 3 mm long, 2.5 mm in diam.; seeds 6, tri-
angular, 1.3-1.5 mm long, dark brown; hilum punctiform
to elliptical.

Distribution and habitat: Реги to Bolivia; in moist rocky
soil.

SPECIMENS EXAMINED

Perú. CAJAMARCA: San Pablo, 2200-2400 m., 1906, Weberbauer
3855 (в, G). LA LIBERTAD: Sammne-Casmiche, Prov. Otuzco, 1800 m.,
21 May 1952, López M. 0844 (us). LIMA: San Mateo, Prov. Huaro-
chiri, 3200 m., 24 Mar. 1952, Hutchison 815 [grown and collected
Univ. of Calif.-Berkeley, 19 Oct. 1957, С.В. Newcomb] (BH, F, С,
GH, UC). Matucana, ca. 8000 ft., 12 April-3 May 1922, Macbride &
Featherstone 350 (G, US); Prov. Cajabomba, Banos de Churin, 9000
ft., 7 Feb. 1946, Laudemer 5414 (к). Bolivia, COCHABAMBA: Ayo-
paya, Sailapata, 2700 m., Cárdenas 3029a (vs). Río de Guinllabamba,
10 June 1876, Andre 3608 (к, NY).

Diels’ original description of this species reported the
length of leaf sheaths to be 3-4 cm, the peduncles to be
1.5-2 ст, and petals to be 8-9 mm. My measurements of
the type specimens show leaf sheaths 0.7-2.9 cm long,
peduncles 1.0-4.5 ст long, and petals 5.8 mm. If additional
information was available to Diels he does not mention
that fact.

An examination of all the specimens available of this
rather rare species reveals that there are minor morpho-
logical differences among them which is not surprising if
one considers the limited amount of materia] available
and the ecological diversity of the mountains which this
species inhabits.

1975] Tripogandra — Handlos 265

8. Tripogandra glandulosa (Seubert) Rohweder, Abh.
Auslandsk., Reihe C, Naturwiss. 18:156. 1956.
Tradescantia glandulosa Seubert in Martius, Flora
Brasiliensis 3(1):253. 1855. HOLOTYPE: Brazil.
PARANA: Rio Negro, 7 Mar. 1823, Sello 995 (B!).
Tradescantia radiata Clarke in Chodat & Hassler, Bull.
Herb. Boissier 3:245. 1903. HOLOTYPE: Paraguay.
AMAMBAY: in regione cursus superioris fluminis Apa,

Feb. 1901-2, Hassler 8493 (G!).

Tradescantia pfanzii Brückner, Bot. Jahr. Syst. 61(1):
13. 1927 (‘1926’), mom, mud.

Descantaria glandulosa (Seubert) Brückner, Notizbl.
Bot. Gart. Berlin-Dahlem 10:56. 1927.

Descantaria pflanzii Brückner, Notizbl. Bot. Gart. Ber-
lin-Dahlem 10:57. 1927. HOLOTYPE: cult. in hort.
Berol. (В!).

Descantaria radiata (Clarke) Briickner, Notizbl. Bot.
Gart. Berlin-Dahlem 10:56. 1927.

Tripogandra pflanzii (Brückner) Rohweder, Abh. Aus-
landsk., Reihe C, Naturwiss. 18:156. 1956.

Tripogandra radiata (Clarke) Bacigalupo, Darwiniana
13:90. 1964.

Illustrations: Bacigalupo, Darwiniana 13:92, 93. 1964
(Figs. 1, 2); 14:404, 410. 1967 (Figs. 3, 5).

Plants perennial, rooting at nodes of decumbent stems;
flowering stems upright, to 39 ст tall, branching from
decumbent stems; internodes to 9.5 em long, glabrous or
with a partial line of uniseriate hairs extending down the
stem from the sheath above. Leaves narrowly ovate to
ovate; blades to 6.7 cm long, to 2.3 em wide, glabrous, base
cuneate on the lower part of the plant, rounded on the
upper portion of the plant, margin ciliate, apex acute;
sheaths to 11.0 mm long, to 8.6 mm in diam., a few long
hairs at the orifice, a line of uniseriate hairs extending
down the side of the sheath opposite the blade, otherwise
glabrous. Inflorescences of 1-10 (-23) double cincinni borne
terminally and in 1-4 of the upper leaf axils; peduncles

266 Ећодога [Vol. 77

to 3.1 cm long, variously glabrous or with 1-2 lines of uni-
seriate hairs, or glabrous proximally and pilose distally,
hairs capitate; double cincinni with up to 11 buds, flowers
and/or fruits; pedicels 2.3-6.0 mm long, recurved in fruit,
pilose with capitate hairs; bracts subtending each pedicel
glabrous to pilose with capitate hairs, margin denticulate,
entire, or ciliate. Flowers white or pink; sepals narrowly
ovate, 3.0-4.2 mm long, 1.2-1.9 mm wide, pilose, hairs capi-
tate, margin hyaline, apex acute; petals са. 3.5-5.0 mm
long; stamens 6, in two whorls, the outer shorter, filaments
to 1.5 mm long, with moniliform hairs borne on the mid-
dorsal portion, anthers 0.6-0.9 mm long, 0.4-0.8 mm wide,
basifixed, anther sacs parallel; stamens of the inner whorl
longer, filaments to 3.8 mm long, glabrous, ligulate, con-
cave in the upper third of their length, anthers 0.4-1.1 mm
long, 0.4-0.7 mm wide, basifixed, anther sacs parallel ;
ovary 0.5-1.0 mm long, 0.4-0.9 mm in diam., glabrous,
style 0.2-0.5 mm long, stigma simple or capitellate. Capsule
globose, to 2.7 mm long, to 3.1 mm in diam., glabrous, light
brown; seeds 2 per locule, rounded-triangular, 0.8-1.4 mm
long, light gray to brown, testa reticulate-foveate with
small prominent ribs radiating from the punctiform hilum
(Figs. 42, 43).

Chromosome number: n==8.

Distribution and habitat: southern Bolivia and Brazil,
Paraguay, Uruguay, and northeastern Argentina; in moist
places and along streams.

Flowering: Flowering of this species seems to occur
between September and March.

REPRESENTATIVE SPECIMENS

Brazil. PARANA: Sete Quedas, Mun. Guaira, 11 Dec. 1965, Hatsch-
bach, Lindeman & Haas 13339 (US). Paraguay. AMAMBAY: zwischen
Río Apa und Río Aquidaban, 1908-9, Fiebrig 4487 (RM, G, K, L).
SAN PEDRO: Alto Paraguay, Primavera, 13 Oct. 1955, Woolston 596
(C, NY). GUARIA: Villarrica, Cerro Peladu(?), Dec. 1930, Jorgensen
4112 (Ds, F, MO, NY, PH, US). Argentina. FORMOSA: Colonia Clorinda,
150 m., 30 Dec. 1926, Venturi 9163 (US). CATAMARCA: road 38 from
Tucumán to Catamarca, Cuesta de Totoral, km. 1384.7, Dept. Paclín,
900 m., 24 Mar. 1966, Hawkes, Hjerting & Rahn 3989 (C). TUCUMAN:

1975] Tripogandra — Handlos 267

El Duraguito, Dept. Capital, 550 m., 14 Jan. 1922, Venturi 1672
(ғ p.p. US, US p.p.) CHACO: Margarita Belén, Dept. Resistencia,
9 Dec. 1945, Aguilar 563 (с). SANTA FE: Lonteri, Estancia Bomaz-
zola, 1 Feb. 1936, Joh 1185 (05). CORRIENTES: Corrientes, 4 Sept.
1959, Mattson 3119 [grown in Copenhagen, collected by Pedersen in
1953] (С). ENTRE RIOS: Сопсерсїбп del Uruguay, 3 Dec. 1877,
Lorentz (GH). Uruguay. RIVERA: Cunapiru, 6-700 ft., 1928, Wright
(BM). CERRO LARGO: Rio Branco, 5-10 m., 27-29 Nov. 1947, Herter
2127 (MO, NY, US). TREINTA Y TRES: Vergara, 20 m., Dec. 1932,
Herter 1608 (vs).

Tripogandra glandulosa has very distinctive seeds and
this fact was recognized by the authors of the names
Tradescantia glandulosa, T. radiata, Descantaria pflanzii.
While the seeds are diagnostic, other statements in all of
the original descriptions are somewhat misleading. An
examination of the type specimens shows all the floral and
seed characters to be identical.

The combination Tripogandra pflanzii has been published
three times. The Gray Herbarium Card Index credits
Celarier (1955) with the earliest combination. This com-
bination was not validly published according to Article 33
of the International Code of Botanical Nomenclature
(1966). Burkart also combined the epithet in 1959, but
this publication was later than Rohweder's (1956) valid
publication of the name.

Bacigalupo (1964) was the first person to consider
Tripogandra radiata and T. pflanzii the same species but
he does not seem to have seen the type specimen of T.
glandulosa and recognized its identity.

Several collections of apparently sterile plants have
been made in Trinidad [Broadway 9145 (BM, UC); Fend-
ier 862 (BM, к): Grisebach 22 (K)], French Guiana
[Sag. 948 (BM)], and Para, Brazil [Huber 93 (BM)].
Pollen from an herbarium specimen mounted in aniline
blue-lactophenol did not stain and is interpreted as having
been sterile and non-functional when fresh. No specimens
have seeds though inflorescences are present.

These plants differ in the generally larger leaves (to 7.5
em long), the presence of one to several lines of hairs on

268 Ећодога [Vol. 77

the sheath, a line of hairs along the midrib on the lower
leaf surface, peduncles to 4.4 ст long, and all staminal
filaments glabrous. Broadway’s collections have small
stolon-like branches perforating the leaf sheaths.

The plants have been included in Tripogandra glandulosa
because the morphological similarities are greater than
with any other species. This decision is made reluctantly
because it violates the morphological and geographic unity
of the main body of collections. These plants seem to
represent disjunct outliers of the main group of popula-
tions of T. glandulosa and with the apparent sterility,
warrant further collection and study.

9. Tripogandra grandiflora (Donnell-Smith) Woodson,
Ann. Missouri Bot. Gard. 29:153. 1942.

Callisia grandifiora Donnell-Smith, Bot. Gaz. 31:125.
1901. TYPE: Guatemala. ALTA VERAPAZ: Cubilguitz,
350 m., Apr. 1901, von Tuerckheim 7684 (Lectotype,
US!; isolectotypes, GH!, Mol, US!).

Donnellia grandiflora (Donnell-Smith) Clarke in Don-
nell-Smith, Bot. Gaz. 33:261. 1902.

Neodonnellia grandiflora (Donnell-Smith) Rose, Proc.
Biol. Soe. Wash. 19:96, 1906.

Illustrations: Donnell-Smith, Bot. Gaz. 33: РІ. ХІ.
1902: Moore, Jr., Baileya 8:78-80, Figs. 24, 26. 1960.

Plant perennial; stem first erect and later trailing over
shrubs, to 3 m tall, to 1.0 cm in diam., unbranched or
branched: internodes to 17 em long, green, glabrous or
with a complete or partial line of uniseriate hairs extend-
ing down the internode from the sheath above, Leaves
two-ranked, narrowly ovate or elliptic, smaller apically and
reduced to sheaths in the inflorescence; blades to 15.2 cm
long, 4.4 ст wide, oblique at the base, cuneate on one side
and rounded on the other, glabrous dorsally, ventral sur-
face glabrous except for the midrib which is wholly or
partially covered with uniseriate hairs, margin ciliate or
entire with a few hairs near the sheath, apex acute; sheaths

1975] Tripogandra — Handlos 269

of vegetative leaves to 16 mm long, 8.0 mm in diam. when
dry, villous at the orifice, otherwise glabrous except for a
complete or partial line of uniserate hairs extending down
the side opposite the blade, sheaths of the cataphylls to
28 mm long, to 13 mm in diam., margin entire or ciliate.
Inflorescences of a single terminal double cincinnus or
paniculate, sometimes flexuous, composed of as many as
9 terminal and axillary double cincinni; peduncles to 4.0
cm long, or so reduced that cincinni appear sessile, green
or dark green with lighter green flecks, glabrous or with
1 or 2 lines of uniseriate hairs; double cincinni with up
to 13 buds, flowers and/or fruits; pedicels to 14 mm long
at anthesis, to 1.3 mm in diam., green or green with a
reddish tinge, glabrous, erect in fruit; bracts at the base
of each pedicel entire, glabrous or with a few marginal
uniseriate hairs. Flowers white; sepals narrowly ovate or
elliptie, to 8.5 mm long, to 4.0 mm wide at anthesis, green
or green with a pinkish base, glabrous, margin entire and
hyaline, apex acute or obtuse; petals ovate, elliptic, or
obovate, to 10.5 mm long, to 8.1 mm wide, cuneate at the
base, apex obtuse; stamens 3, opposite the sepals, filaments
1.0-2.5 mm long, white, glabrous (Fig. 9), anthers 0.9-1.3
mm long, 0.8-1.2 mm wide, whitish, dorsifixed, versatile,
open anther sacs white with a purple line around the edge
(Fig. 8), pollen white; staminodes 8, opposite the petals,
slightly epipetalous, filaments to 8.0 mm long, white, with
two patches of uniseriate hairs in the upper half, the
higher patch on the ventral side, the lower patch on the
dorsal side with a few scattered hairs below the patch,
filament bent in an open S-shape (Fig. 28), anthers dorsi-
fixed, not versatile, connective orange or yellow, C-shaped
with the anther sacs divergent, or connective elongate
with anther sacs parallel, pollen yellow, sterile; ovary 0.7-
1.6 mm long, 0.8-1.3 mm in diam., white or with a line of
pink between the carpels, glabrous, style to 1.1 mm long,
stigma simple or capitate and 3-lobed. Capsule elliptic,
green or brown, glabrous, to 6.5 mm long; seeds gray or
white and roughened, 1 or 2 per locule, elliptic (Fig. 77)

270 Ећодога [Уо1. 77

or triangular (Fig. 76), when elliptic with two sides up-
turned above the protuberant embryotega, to 5.0 mm long,
when triangular with two sides and one end upturned
forming an angular C-shaped ridge around the embryotega,
to 3.0 mm long, hilum linear (Fig. 78).

Chromosome number: n=8, 16.

Vernacular name: hoja de fluxion, Standley and Steyer-
mark, Guatemala, Fieldiana: Bot. 24(3):38. 1952.

Distribution and habitat: the Yucatan peninsula, south-
ern México, Guatemala and British Honduras; at low
elevations to 800-1500 m (7000-8800 ft, Nelson 3236c).

Flowering: Flowering occurs in August and September
in the area surrounding Tuxtla Gutierrez in México and
coincides with the rainy season in this location. In the
area of Yucatán flowering occurs later, January to March
and April. The fragrant flowers open sometime before
sunrise and remain open until noon.

REPRESENTATIVE SPECIMENS

México. CHIAPAS: km. 1026 of Hwy. 190, 24 km. west of Ocozo-
cuautla, 620 m., 13 Aug. 1967, Handlos 342 (BH). CAMPECHE: Mon-
terrey, 22 Jan. 1932, Lundell 1225 (ps, Е, MICH, US). British Hon-
duras. EL CAYO: Augustine, Mountain Pine Ridge, 1450 ft., 18 Mar.
1960, Hunt 399 (BM, MICH, MO, US). Guatemala. PETEN: Santa Cruz,
27-28 Mar. 1931, Bartlett 12390 (к, MICH, US). ALTA VERAPAZ: be-
tween Coban and Finca Chimoté, near Rubeltein, 800-1500 (300-
500) m., 25 Feb. 1942, Steyermark 44203 (F, MO). IZABAL: Rio
Chacón, 300 ft, 10 Feb. 1921, Johnson 1266 (Us).

Two different morphological forms can be distinguished
within Tripogandra grandiflora. One form, representing
the typical element, is more wide-spread and is charac-
terized by leaves with a ciliate margin, a paniculate
inflorescence, often with a flexuous axis and bearing 3-9
double cincinni, staminodes with a C-shaped connective and
divergent anther sacs and commonly one seed per locule.
Based on information on specimen labels the flowers are
very fragrant. This form is found in the eastern portions
of Guatemala, British Honduras, and southeastern México,
the Yucatán peninsula and Chiapas.

1975] Tripogandra — Handlos 271

The other form has leaves which lack the ciliate margin,
there being only a few hairs at the base of the blade near
the sheath. The inflorescence is composed of one to three
doubie cincinni, the staminodes have an elongate, straight
connective and the anther sacs are parallel. There are one
or two seeds per locule. The flowers have a weaker fra-
grance if Handlos 342 is typical, While the evidence is
limited there may also be a difference in a staminodial con-
nective color, this form being yellow while the former is
yellow or orange. The sterile pollen is more elongate in
the typical element and more similar to the fertile pollen
in Handlos 342. The second form is found in a restricted
area in Chiapas, México, centering around Tuxtla Gutierrez
(Ocozocuautla to San Cristóbal de las Casas).

At the present time the two forms seem to be undergoing
independent evolution. There may be significant inter-
actions and relationships between the pollinators and such
floral characters as form and color of staminodes, and
fragrance. If the two forms evolve in such a fashion that
two distinct sets of pollinators are involved in pollination
these two entities could retain their identity in the future
if the populations become sympatric. Geographical isola-
tion with some slight morphological and ecological differ-
entiation now seems to exist. With further increase in
differences these forms could be recognized as species but
at this point in time it seems premature to give any nomen-
clatural recognition to these forms.

The United States National Herbarium is the location
of two specimens which are held as the type material.
According to the International Code of Botanical Nomen-
clature, Article 7, Note 3, only one specimen can serve
as the holotype of the species. I designate the sheet (US
936917) which was annotated by C. V. Morton in August,
1940, as Neodonnellia grandiflora, as the lectotype with
the other sheet (US 936916) to serve as an isolectotype.
I have chosen this particular sheet because it holds both
flowering and fruiting branches.

272 Ећодога [Vol. 77

This species is distinguished by the revolute seed mar-
gin, the farinose seed surface, the two patches of hairs on
the inner filaments, the elongate shape of the cells of the
staminodial hairs and the sweetly scented flowers.

Reproduction in Tripogandra grandiflora is probably of
two sorts, sexually by seeds and vegetatively from sections
of fallen stem. Old stems fracture easily and in México
I have found young plants attached to pieces of stems
lying on the ground; the axillary bud at the base of the
internode apparently began to grow when conditions be-
came favorable, i.e., when sufficient moisture was available.
Dispersal may be aided by the fact that the plants grow
on hillsides and pieces of old stem could roll very easily
to a new location. In areas with abundant tropical showers,
pieces of stem could be carried and spread to new loca-
tions by water rushing down ravines and gullies,

10. Tripogandra guerrerensis Matuda, Anales Inst. Biol.
Nac. México 36:113. 1966 (‘1965’). TYPE: México.
Guerrero: Rincón de la Via, 775 m., 24 Sept. 1961,
Kruse 461 (Holotype, MEXU; isotype, MEXU!).

Illustration: Matuda, Anales Inst. Biol. Nac. México
36:113. 1966 (‘1965’).

Plants annual; stems erect, to 74 cm tall, 2-4 mm in
diam. when dry, unbranched or with 1-2 branches basally ;
internodes 1.8-9.7 em long, green, glabrous except for a
line of uniseriate hairs continuing down the internode
from the sheath on the side opposite the blade. Leaves nar-
rowly ovate; blades 2.9-10.7 ст long, 0.3-1.1 cm wide,
narrowed toward the base, smaller distally and leaves
reduced to sheathing bracts in the inflorescence, glabrous
ventrally, pilose dorsally, margin ciliate, apex acute;
sheaths 2.0-12 mm long, villous at the orifice, otherwise
glabrous except for a line of uniseriate hairs which con-
tinues down the side of the sheath opposite the blade.
Inflorescences paniculate, much-branched, terminal and in
the axils of the upper leaves, composed of 7-30 double cin-

1975] Tripogandra — Handlos 273

cinni, peduncles 7-23 mm long, 0.4-0.6 mm in diam., green,
glabrous; bracts within the inflorescence with short lamina
or reduced to a sheath near the apex; buds, flowers and/or
fruits 1-5 per double cincinnus; pedicels 1.4-3.5 mm long,
0.3-0.4 mm in diam., green, glabrous or with a few scattered
uniseriate or capitate hairs, erect in fruit; bracts sub-
tending the pedicels glabrous, margin entire. Flowers
white; sepals ovate-cymbiform, 2.3-3.2 mm long, 1.5-2.1 mm
wide, green with purple tips, glabrous or with a few scat-
tered capitate hairs, margin entire, apex acute or obtuse;
petals ovate-elliptic, 3.5-5.0 mm long, 2.3-3.5 mm. wide,
tapering at the base, apex obtuse; stamens 3, opposite the
sepals, filaments 1.0-1.1 mm long, white, glabrous basally
but bearded dorsally in the middle with a few uniseriate,
colorless hairs (Fig. 11), anthers 0.6-0.7 mm long, 0.4-0.5
mm wide, white, basifixed, versatile with anther sacs
parallel, longer than the gibbous connective, pollen white;
staminodes 8, opposite the petals, filaments slightly epipe-
talous, 3.5-4.5 mm long, white, glabrous, bent in an S-shape
in the upper half, expanded in the bent portion (Figs. 15,
17), anthers dorsifixed, versatile, filaments attached near
one side of the yellow, discoid connective and anther sacs
borne on the side opposite the filament, connective 0.6-0.8
mm long, 0.5-0.7 mm wide, pollen yellow, sterile; ovary
0.6-0.8 mm long, 0.6-0.8 mm in diam., light green, glabrous,
style 0.2-0.3 mm long, stigma simple, papillate, Capsule
obovoid, glabrous, light brown, 2.9-3.6 mm long, 2.2-2.5
mm in diam.; seeds 2 per locule (Fig. 49), the upper in
each locule longer (Figs. 70, 71), 1.6-1.9 mm, the lower
shorter, 1.1-1.4 mm (Figs. 68, 69), triangular to elliptical,
dark brown to black, testa areolate with ribs radiating
from the embryotega, hilum linear-elliptical (Figs. 69, 71).

Chromosome number: n=8,

Distribution and habitat: México in the states of Jalisco
and Guerrero: in rocky areas at low elevations, This species
is known from two locations which are about 455 km apart.
The intervening area is not easily accessible but it seems
highly probable that there are other suitable habitats be-

274 Ећодога [Vol. 77

tween the two known locations, and as the area is explored
new collections will almost certainly be made.

Flowering: Flowering occurs in September and October
in the two locations where this species has been collected.
Flowers in México and in the greenhouse at Ithaca, N.Y.,
open about 11:00 AM and close about 2:30 PM.

SPECIMENS EXAMINED

México. JALISCO: just before km. 148 on Hwy. 110 to Colima,
5.6 km. south of bridge over Río San Pedro, ca. 1050 m., 2 Oct. 1967,
Handlos 438 (BH); km. 147-148 on road between Colima and Ma-
zamitla, 1040 m., 10 Sept. 1961, Moore, Jr. & Bunting 8746 (BH, UC).

Tripogandra guerrerensis is unique among the species
I have seen in México because the petals are not sym-
metrically disposed at anthesis; rather, two petals bend
upward so that an angle of approximately 90? is formed
between the middle upright petal and the two lateral ones,
whereas in al! the other species the angle between petals
is about 120°,

The androecium bears a constant relationship to the
petals. In typical Tripogandra-fashion, the filaments of
two of the staminodes bend around two staminal filaments
and the three staminodes are then positioned in front of
the middle upright petal. The stamens surround the ovary
but because of the position of the petals and staminodes
the anthers can not be approached by a pollinator alighting
on the petals. From observations made of a natural popu-
lation in Jalisco, México, on October 2 and 3, 1967, the
disposition of the petals seems significant because bees are
restricted in their movements and are able to gather pollen
only by clinging to the staminodes. If the pollinator col-
lects sterile pollen, it hangs from the filaments of the
sterile stamens, with its abdomen pointed toward the
center of the flower. While gathering sterile pollen, the
abdomen is placed on the fertile anthers so that pollen is
deposited on the pollinator's abdomen. This seems to be
a device for insuring pollination. Insects are also able
to collect fertile pollen by clinging upside down to the

1975] Tripogandra — Handlos 275

staminodes with the head near the anthers. Further
critical studies are needed to determine how much cross-
pollination and how much self-pollination is effected, In
the greenhouse, abundant viable seed was set after insects
ceased entering from outdoors, so the plants are not self-
sterile.

11. Tripogandra kruseana Matuda, Anales Inst. Biol. Nac.
México 37:77, 78. 1967 (‘1966’). HOLOTYPE: México.
GUERRERO: Rincón de la Via, 735 m., 16 Oct. 1965,
Kruse 903 (MEXU!).

Illustration: E. Matuda, Anales Inst. Biol. Nac. México
37:70. 1967. C1966’).

Plants perennial?, rooting at the nodes; flowering stems
erect, to 61 cm tall, branched at the base; internodes to
9.5 ст long, with a partial line of uniseriate hairs extend-
ing down the side from the sheath above, otherwise
glabrous in the lower portion of the plant. Leaves nar-
rowly ovate; blades to 10.1 ст long, to 1.2 cm wide,
glabrous, margin ciliate, apex long acuminate; sheaths to
12 mm long, to 1.0 cm in diam., a few long hairs at the
orifice, with a line of uniseriate hairs on the side opposite
the blade, otherwise glabrous. Inflorescences borne termin-
ally and in the upper 8 leaf axils, composed of up to 9
double cincinni; peduncles to 1.8 em long, pilose with
capitate hairs; buds, flower and/or fruits 2-3 per double
cincinnus; pedicels 8-12 mm long, green, erect in fruit,
pilose with capitate hairs; bracts at the base of each
pedicel with margin entire or denticulate. Flowers pink
(white fide label) ; sepals ovate, 3.1 mm long, 1.6 mm wide
in flowers available (2 mm long, 1.2 mm wide fide desc.),
pilose with capitate hairs, margin red; petals not avail-
able; stamens 6, in two whorls, 3 shorter, 0.7 mm long,
bearded with white moniliform hairs; 3 larger, 2 mm
long, bearded with white moniliform hairs; ovary green,
glabrous, style short, stigma capitellate. Capsule globose,
2.5 mm long, 2.5 mm in diam. (immature).

276 Ећодога [Vol. 77

Distribution and habitat: This species is known only
from the type collection at Rincon de la Via, Guerrero,
México, in a rocky area at 735 m. altitude.

Flowering: The type specimen was in flower in mid-
October.

Matuda’s original description of 7. kruseana describes
the base of the leaf as amplexicaul but this is not readily
discernible in the holotype. If the plant has amplexicaul
leaves and is a perennial, it will key out to T. encolea. If
the plant lacks amplexicaul leaves, as the photograph of
the holotype seems to indicate, it will key near 7. saxicola
from which it may be distinguished by erect pedicels and
pilose peduncles.

12. Tripogandra montana Handlos, sp. nov. HOLOTYPE:
México. CHIAPAS: 3.2 miles N of junction of Hwy.
195 with road to El Bosque and Simojovel, 1770
(1690) m., 15 Aug. 1967, Handlos 355 (Us!).

Herba perennis; caulis decumbens, usque ad 2.6 m
longus, internodiis usque ad 12.8 ст longis, glabris vel
raro distaliter pilosis, linea unica pilorum instructis.
Folia angusto-ovata, laminis usque ad 14.5 ст longis,
usque ad 3.5 em latis, apice acutis, basi obliquis, glabris
vel pilis paucis dispersis instructis vel pilosis, margine
ciliatis vel medio nonnunquam eciliatis, vaginis usque ad
18.0 mm longis, usque ad 12.5 mm diam., linea unica
pilorum instructis, aliter glabris vel pilis dispersis in-
structis vel pilosis, orificio villosis vel pilis dispersis prae-
ditis. /Inflorescentiae terminales et in 1-4 axillis foliorum
summorum, ex 1-17 cincinnis duplicibus constantes;
pedunculi 0.7-12.7 em longi, glabri, pilosi vel 1-2 lineis
pilorum instructi; cincinni duplices omnes alabastra, flores,
vel fructus usque ad 17 gerentes; pedicelli usque ad 2.5-6.0
mm longi, 0.6-0.8 mm diam., maturitate reflexi, pilis capi-
tatis paucis instructi vel pilosi, bracteis basi pedicellorum
glabris vel pilis capitatis vel uniseriatis dispersis pubes-
centibus, margine ciliatis vel eciliatis. Flores rosei; sepala.

1975] Tripogandra — Handlos ДЕ

anguste ovata, elliptica, vel obovata, cymbiformia, 4.0-6.0
mm longa, 1.6-2.8 mm lata, pilosa vel pilis paucis, dispersis,
capitatis instructa, margine integra et hyalina, apice
obtusa; petala ovata, 5.5-11.0 mm longa, 3.5-6.8 mm lata,
basi cuneata, apice rotundata; stamina 6 in verticillis
duobus, 3 sepalis opposita filamentibus brevibus, 1.0-2.5
mm longis, subroseis, glabris vel raro pilis paucis praeditis
(Fig. 10), antheris 0.8-1.8 mm longis, 0.6-1.2 mm latis,
polline albido, 3 petalis opposita filamentibus epipetalis,
sigmoideis, 4.5-7.5 mm longis, medio pilis roseis monili-
formibus pubescentibus (Fig. 31), antheris 0.5-0.9 mm
longis, 0.5-1.1 mm latis, polline luteo; ovarium 1.2-1.7 mm
longum, 0.8-1.2 mm diam., glabrum, stylo 0.2-0.6 mm longo,
stigmate capitellato vel capitato. Capsula elliptica, 2.0-3.7
mm longa, 1.6-2.6 mm diam., glabra, seminibus triangulari-
bus, 1.1-1.7 mm longis, reticulatis (Figs. 44-46), hilo
punctiformi (Fig..45).

Chromosome number: n=21.
Vernacular name: та fide Standley, Guatemala.

Distribution and habitat: southern México, Guatemala,
Honduras, and El Salvador; in pine-oak forest areas at
higher elevations, ca. 1100-2500 m. The one undisturbed
location in which I found this species was in a black loam
soil in a pine woods with a scattering of oaks, Liquidambar
and tree ferns. The plants grew on a moist, east-facing
slope. An adjacent, drier, sunnier and steeper west-facing
slope had no plants of this species.

Flowering: In México flowering occurs between March
and November; in Guatemala between November and June;
in Honduras in February, March, April, July and Novem-
ber. The flowers open about 8:00 AM in their natural
habitat and close about 2:00 PM. The plants I found had
odorless flowers. Flowering seems to extend over a long
period of time because both young inflorescences and
mature seed were found in the location where Handlos
355 was collected.

278 Ећодога [Vol. 77

REPRESENTATIVE SPECIMENS

México. CHIAPAS: in the paraje of Kulak’tik, Municipio of Tene-
japa, 5500 ft., 25 Nov. 1965, Breedlove 14205 (Ds, F, MICH). Guate-
mala. HUEHUETENANGO: Canyon of Rio Chixoy near Malacatancito
about 20 km. southwest of Huehuetenango, 1600 m., 1 Dec. 1962,
Williams, Molina R. & Williams 22149 (F, NY). QUICHE: Cunen, 6000
ft., Apr. 1892, Heyde & Lux 3521 (GH, M, NY, US). ALTA VERAPAZ:
Chicoyonito, 4300 ft., Apr. 1889, Donnell-Smith 1643 (GH, NY, PH,
US). SAN MARCOS: slopes bordering Rio Malacate, barrancos 6 miles
south and west of town of Tajumulco, northwestern slopes of Volean
Tajumulco, 2300-2800 m., 26 Feb. 1940, Steyermark 36679 (F).
QUEZALTENANGO: mountains above Rio Samala, Sierra Madre Moun-
tains, 2 km. west of Zunil, 2300 m., 14 Dec. 1962, Williams, Molina R.
& Williams 23023 (к, NY). SOLOLA: steep slopes of Panajachel
water falls, road to Solalá, 2200 m., 12-23 Jan. 1966, Molina R.,
Burger & Wallenta 16233 (Е, NY). CHIMALTENANGO: near Río
Рахсауб, between Chimaltenango and San Martín Jilotepeque,
1650-1800 m., 3 Feb. 1939, Standley 64362 (F). SACATEPÉQUEZ :
slopes of Volcán de Agua, south of Santa María de Jesús, 1800-2100
m., 10 Dec. 1938, Standley 59449 (F). GUATEMALA: damp wooded
barranca 10 km. south of San Raimundo, about 1800 m., 18 Jan. 1939,
Standley 62882 (Е). EL PROGRESO: Montana Canahui, between Finca
San Miguel and summit of mountain, near upper limits of Finca
Caieta, 1600-2300 m., 10 Feb. 1942, Steyermark 43764 (F). ZACAPA:
along Rillito de! Volcán de Monos, Volcán de Monos, 1150-2100 m.,
10 Jan. 1942, Steyermark 42336 (F, MO). CHIQUIMULA: Montana
Nonojá, 3-5 miles east of Camotán, 600-1800 m., 11 Nov. 1939,
Steyermark 31687 (F). SUCHITEPÉQUEZ: southwestern lower slopes
of Volcán Zunil, in vicinity of Finca Montecristo, southeast of Santa
Maria de Jests, 1200-1300 m., 31 Jan. 1940, Steyermark 35221 (F).
Honduras. COPAN: 5 km. al S.O. de Santa Rosa de Copán, 1200 m.,
29 Mar. 1963, Molina R. 11675 (F). COMAYAGUA: vicinity of Sigua-
tepeque, 1080-1400 m., 14-27 Feb. 1928, Standley 56198 (F, Us).
INTIBUCA: Bafios de Esperanza, 1800 m., 27 Nov. 1958, ‘Hawkes,
Hjerting & Lester 2103 (c, к). El Salvador. Santo Tomás, 1922,
Calderón 1294 (мү, US).

The collections which I have included in Tripogandra
montana have usually been identified as T. elongata in the
past. On the basis of a distinct seed reticulation, a capitate
stigma, a short style, a different leaf texture, adaptation
to a relatively moist montane habitat, and a different and
distinct geographic range, I must recognize T. montana
as different from T. elongata. Tripogandra elongata is

1975] Tripogandra — Handlos 279

discussed further under T. serrulata. The long peduncles
and bright pink flowers of Т. elongata may indicate some
affinity with Т. montana but this is only speculation.

Variation in leaf vesture does occur within this taxon
but that seems to be a common occurrence within this
genus. The significance, function, and mode of inheritance
of this character should be investigated further.

13. Tripogandra multiflora (Swartz) Rafinesque, Flora
Telluriana 2:16. 1837 (‘1836’).

Tradescantia multiflora Swartz, Nova genera & species
plantarum seu prodromus . . . Indiam occidentalum
57. 1788; Flora Indiae occidentalis 1:599. 1797;
3:1972. 1806; Jacquin, Collectanea ad botanicam

. 8:226, 227. 1791. TYPE: Jamaica. Swartz
(Holotype, (B)?; isotype, M!).

Tradescantia parviflora Ruiz & Pavon, Florae Peru-
vianae et Chilensis prodromus, . . . 3:43. 1794.
TYPE: Perú. HUANUCO: Pozuzo, 1778-1788, Ruiz
& Pavon. (Holotype, MA; photograph, МА!; iso-
type, F!).

Tradescantia procumbens Willdenow, Species Planta-
rum 2:19 1799. HOLOTYPE: Jacquin (B); photo-
graph, B!.

Commelina floribunda HBK., Nova genera et species
plantarum 1:260. 1816. HOLOTYPE: Cumana, Bor-
dones & Quetepe, Humboldt & Bonpland (B) ; photo-
graph, B!.

Heminema multifiora (Swartz) Rafinesque, Flora
Telluriana 2:17. 1837 (‘1836’).

Aneilema floribunda (HBK) Hooker & Arnott, The
Botany of Captain Beechey’s Voyage 311. 1840.

Tradescantia cumanensis Kunth, Enumeratio Planta-
rum 4:96. 1843, based on Commelina floribunda
HBK non Tradescantia floribunda M. & G. (1842)
nec Т. floribunda Kunth (1843).

Tradescantia multiflora у linnaei Clarke in DC., Mono-
graphiae Phanerogamarum 3:306. 1881. TYPE:

280

Ећодога [Vol. 77

Colombia. in montibus juxta Bogotam, 6 Nov.
1852, Holton 127 (Holotype, K!; isotype, NY!).

Tradescantia multiflora B parviflora (Ruiz & Pavon)
Clarke in DC., Monographiae Phanerogamarum
3:306. 1881.

Descantaria procumbens Schlechtendal fide Hasskarl
ex Clarke in DC., Monographiae Phanerogamarum
3:305. 1881, nom. nud. pro syn.

Leptorhoeo floribunda (HBK) Baillon, Histoire des
Plantes 13:218. 1894 (‘1895’).

Tradescantia ionantha Diels in Urban, Bot. Jahrb.
Syst. 37:382. 1906. TYPE: Perú. PUNO: Sandia,
2100 m., 24 Mar. 1902, Weberbauer 588 (Holotype,
B!; isotype, G!).

Tradescantia multiflora var. tobagensis Urban, Sym-
bolae Antillanae 7:174. 1912. HOLOTYPE: Tobago.
juxta flumen “Bacolet” ad “Calder Hall", 31 Oct.
1889, Eggers 5693 (B!).

Descantaria multiflora (Swartz) Briickner, Notizbl.
Bot. Gart. Berlin-Dahlem 10:56. 1927.

Descantaria cumanensis (Kunth) Schlechtendal ex
Brückner, Notizbl. Bot. Gart. Berlin-Dahlem 10:56.
1927.

Descantaria ionantha (Diels) Briickner, Notizbl. Bot.
Gart. Berlin-Dahlem 10:56. 1927.

Tripogandra floribunda (HBK) Woodson, Ann. Mis-
sourl Bot. Gard. 29:152. 1942.

Tripogandra cumanensis (Kunth) Woodson, Ann. Mis-
souri Bot. Gard. 29:152. 1942.

Tripogandra ionantha (Diels) Macbride, Revista Univ.
(Cuzco) 33(87) :142, 1945.

Tripogandra parviflora (Ruiz & Pavon) Steyermark,
Phytologia 9:339. 1963 (‘1964’).

Tripogandra multiflora (Swartz) Woodson [fide Baci-
galupo] forma parviflora (Ruiz & Pavon) Baciga-
lupo, Darwiniana 14:398. 1967.

1975] Tripogandra — Handlos 281

Illustrations: Jacquin, Icones Plantarum rariorum 2:
355. 1790 (‘1781-1795’) ; Moore, Jr., Baileya 8:81, Fig. 27.
1960: Bacigalupo, Darwiniana 14:399, Fig. 1, 1967.

Plants perennial, trailing and rooting at the nodes or
caespitose; floriferous stems upright, to 83 cm tall, un-
branched or branched; internodes to 13.5 em long, the
uppermost 1-4 pilose (rarely glabrous), the lower glabrous
or glabrous proximally and pilose distally, in all speci-
mens with a line of uniseriate hairs extending down the
stem from the sheath above. Leaves narrowly ovate, ovate
or broadly ovate; blades to 8.8 ст long, to 2.8 ст wide,
glabrous or pilose dorsally, the ventral surface variously
glabrous, pilose and/or with a line or band of uniseriate
hairs near the margin, occasionally the base of the midvein
with a line of uniseriate hairs, base oblique, cuneate, or
rounded, margin ciliate, apex acute; sheaths to 14.2 mm
long, to 9.2 mm in diam., villous at the orifice, the upper-
most 1-7 pilose (rarely glabrous), the lower ones glabrous
or pilose, in all specimens with a line of uniseriate hairs
extending down the side opposite the blade. Inflorescences
of 1-26 double cincinni borne terminally and in the axils
of the upper 1-8 leaves; peduncles to 3.5 (-5.4) cm long,
densely pilose or rarely pilose distally and glabrous prox-
imally; double cincinni with up to 16 buds, flowers and/or
fruits; pedicels 1.4-4.0 mm long, reflexed in fruit, glabrous,
with scattered uniseriate hairs or pilose; bracts at the
base of each pedicel glabrous, with scattered uniseriate
hairs, or pilose, margin entire or long-ciliate. Flowers
white or pink; sepals ovate-cymbiform, 1.8-4.0 mm long,
1.0-1.8 mm wide, green, red, or green with a red tip and/or
base, glabrous, with scattered capitate hairs, or pilose,
margin hyaline, apex more or less acute; petals broadly
ovate(?), 1.6-4.0 mm long, 1.0-ca. 2.2 mm wide; stamens 6,
in two whorls, the outer shorter, filaments 0.5-1.3 mm long,
glabrous, anthers 0.3-0.6 mm long, 0.3-0.6 mm wide, pink,
basifixed, anther sacs parallel; stamens of the inner whorl
longer, 1.4-3.2 mm long, S-shaped, bearded with a ventral]

282 Кћодога [Vol. 77

tuft of moniliform hairs below the expanded distal end,
anthers 0.3-0.5 mm long, 0.2-0.8 mm wide, dorsifixed, con-
nective I- or C-shaped, yellow, anther sacs divergent or
parallel through bending of the connective; ovary 0.5-1.0
mm long, 0.4-0.8 mm in diam., glabrous, style 0.15-0.3 mm
long, stigma capitellate or capitate. Capsules 1.4-2.5 mm
long, 1.5-2.5 mm in diam., green or light brown, glabrous;
seeds 1-2 per locule, rounded-triangular, 0.7-1.0 (-1.4) mm
long, light or dark gray or brown, testa reticulate-foveate
(Figs. 32, 34), hilum punctiform (Figs. 33, 35).

Chromosome number: 2n=-64 (A. Sparrow, pers.
comm.)

Distribution and habitat: Jamaica, Trinidad, Tobago,
Costa Rica, Venezuela, Colombia, Pert, Bolivia, and Argen-
tina; on shaded or open rocky banks or slopes from sea
level to 2900 m.

Flowering: In Jamaica flowering occurs from June to
February; in Trinidad and Tobago in July and October;
in Costa Rica in November; in Venezuela from August to
January and in May and June; in Colombia from May to
February; in Perú in March, July, November, and Decem-
ber; in Bolivia from December to May.

REPRESENTATIVE SPECIMENS

Jamaica. Troy, 2000 ft., 16 Oct. 1917, Harris 12573 (F, NY, US).
MANCHESTER: vicinity of Mandeville, 15-26 Feb. 1910, Brown 84
(NY, PH). ST. ANDREW: off road from Kingston to Newscastle, 1250
ft, 17 June 1963, Crosby, Hespenheide & Anderson 178 (F, MICH,
MSC, UC). Tobago. Roxborough, 20 July 1914, Broadway 4906 (Us) ;
Mason Hall, 19 Oct. 1937, Sandwith 1865 (му). Trinidad. Point
opposite Melville Island, 25 Oct. 1925, F[reeman] & W [illiams] 11408
(му). Costa Rica. SAN JOSE: vicinity of El General, 1160 m., Nov.
1936, Skutch 2935 (GH, K, MO, NY, US). Venezuela. FALCON: Mea-
chiche, 18 Jan. 1942, Lasser 178 (US, VEN). LARA: Loma de León
Dto. Yribarren, 16 Sept. 1950, Tamayo 3726 (VEN). DISTRITO FEDERAL:
La Quesera, Upper Cotiza, near Caracas, above 1400 m., 18 Sept.
1921, Pittier 9821 (Us, VEN). MIRANDA: Los Mariches, on new road
to Sta. Lucia, 14 Dec. 1924, Pittier 11636 (US, VEN). MERIDA: 3
miles west; 2 miles south of Mérida Valley, 4500 ft., 26 Jan. 1981,
Reed 335 (vs). Colombia. MAGDALENA: Sierra Nevada de Santa
Marta, southeastern slopes; Hoya del Ric Donachuí; below the vil-

1975] Tripogandra — Handlos 283

lage Donachui near the river, 1350-1230 m., 24 Sept. 1959, Cuatre-
casas & Castaneda 24406 (US). NORTE DE SANTANDER: vicinity of
Mutiscua, 2900 m., 20-22 Feb. 1927, Killip & Smith 19669 (vs).
SANTANDER: vicinity of Charta, 2000-2600 m., 1-11 Feb. 1927, Killip
& Smith 17446 (GH, NY, US). BOYACA: Valle de la Uvita, Cordillera
Oriental, 2490-2560 m., 16 Sept. 1938, Cuatrecasas 1857 (05).
CUNDINAMARCA: Macizo de Bogotá, Quebrada de Chicó, 2650-2750
m., 1 June 1939, Cuatrecasas 5224 (US). HUILA: Cordillera Oriental,
east of Neiva, 800-1000 m., 31 July 1917, Rusby & Pennell 424 (NY).
Perü. SAN MARTIN: prope Tarapoto, 1855-6, Spruce 4198 (BM, BR,
E, NY). HUANUCO: west and above Puente Durand, north of Huanuco,
1900 m., 4 Nov. 1938, Stork & Horton 9589 (G, GH, K). LIMA: Lima,
1 July 1914, Rose & Rose 18551 (05). cusco: Potrero, Convención,
1300 m., 2 Mar. 1940, Vargas C. 1832 (GH). PUNO: Sandia, 2100
m., 24 Mar. 1902, Weberbauer 588 (B, с). Bolivia. LA PAZ: Apolo,
4800 ft, 10 Mar. 1902, Williams 55 (BM, US).

Tripogandra multiflora is a widespread species in South
America and a complex pattern of variation seems to exist
within the species as I conceive it. Collections below about
2,000 meters from Colombia, Venezuela and the Caribbean
islands form a fairly uniform group characterized by a
trailing habit, glabrous leaves with an oblique base, white
flowers and pilose sepals.

Collections from Bolivia in contrast are quite variable.
The leaf base is usually oblique and the base of the plant
seems to be decumbent. The flower color may be either
white or pink, sometimes even in the same collections, e.g.,
Buchtien 399. The leaf indument is variable — the leaves
may be (1) glabrous dorsally with a band of hairs near
the margin ventrally as in Williams 55; or (2) pilose on
both sides as in Rusby 1369, Buchtien 2457, 4183 (F, GH),
7190 (GH), Bang 603 (к, м, MICH); or (3) pilose dorsally
with a ventral marginal band of hairs as in Buchtien s.n.,
399 (US), 4183 (к), 7190 (му), Bang 603 (E, F, GH, MO,
PH, US). The sepals may be glabrous as in Williams 55,
Buchtien s.n., 399 (US), 4183 (F), 5350, 7190 (NY), Bang
603 (E, Е. GH, MO, NY, PH, US), or have scattered capitate
hairs as in Rusby 1369, Buchtien 399 (US), 2457 (мү),
4183 (GH), 7190 (GH), Bang 603 (M, MICH); or be pilose
as Buchtien 4183 (F, G). These plants must be studied in

284 Rhodora [Vol. 77

the field for a better understanding of the problems. I do
not know whether hybridization is occurring. It is possible
that collections have become mixed before being glued to
herbarium sheets; most collections were made by Buchtien
and Bang. It is possible, however, that the.species in
Bolivia is very polymorphic with some obvious traits
segregating within the populations.

Plants which grow in the highlands of Colombia, Peru
and Bolivia, above 2,000 meters elevation, tend to be caes-
pitose and the leaves are generally broadly ovate with a
cuneate to nearly amplexicaul base. Compared to lowland
plants of Т. multiflora there are few inflorescences and
these are partially covered by the upper one to five leaves;
the sepals, petals and pistil tend to be longer and the
stigma is broader, more nearly capitate than capitellate.
The flowers are always bright pink or magenta, I can
find no qualitative characters (aside, perhaps, from flower
color) which will allow one to separate the plants of this
group from the bulk of Т. multiflora. The presence of
plants which appear intermediate between the highland
and lowland forms leads me to consider the highland plants
as a mountain ecotype of T. multiflora. Intergradation as
seen from herbarium specimens may be more apparent
than real. It is possible that these forms are ecologically
separated and generally do not intergrade but are very
similar species. If one considers evolution to be a con-
tinuous and gradual process and not saltational, then it
is possible that the situation just described represents a
slight ecological divergence with no overwhelming morpho-
logical differences, a case of sibling or micro-species. These
essential questions can only be settled with further field
work.

Because of the pattern of morphological variations oc-
curring within Tripogandra multiflora, a number of names
have been applied to this species. These names are con-
sidered in detail here.

Willdenow (1799) named Tradescantia procumbens from
a collection with a procumbent stem from mainland South

1975] Tripogandra — Handlos 285

America. This name seems unnecessary for there appear
to be no other characters by which the island and the
mainland populations differ. Observations of greenhouse
grown plants and information on herbarium labels indi-
cate that, in fact, the island plants also tend to trail over
the ground.

Ruiz and Pavon’s Tradescantia parviflora does not seem
sufficiently distinct to be recognized as a species, the only
distinguishing character being the presence of pilosity on
the dorsal leaf surface. Clarke (1881) considered Т. parvi-
flora a variety (0) of Tradescantia multiflora. No other
collections that I have seen duplicate the distribution of
hairs found on the Ruiz and Pavon specimen. The other
collections cited under variety parviflora by Clarke have
additional hairs in a line near the margin on the ventral
surface which are not found in the Ruiz and Pavon speci-
men. Bacigalupo (1967) considered Tradescantia parvi-
flora a form of Tripogandra multiflora. , The specimen
which he cites from Argentina apparently has additional
indument on the ventral leaf surface not found on the
Ruiz and Pavon specimen. In terms of the distribution
of hairs on the leaves, the Ruiz and Pavon specimen seems,
therefore, to be an intermediate between the Bolivian, Ar-
gentinian and other Peruvian collections and the Colombian,
Venezuelan and Jamaican glabrous-leaved plants. Steyer-
mark (1963) cited specimens of Tripogandra parviflora
from Ecuador but apparently these are referable to a
species of Gibasis.

Urban’s variety tobagensis of Tradescantia multiflora
is represented by a very few collections. While the seeds
he observed were 0.6-0.7 mm in diameter, I find that seeds
from other collections on Trinidad and Tobago range from
0.7-1.0 mm in diameter. The plants do seem smaller but
a formal Latin designation seems unnecessary. The small
stature of the plants may represent only stunted growth
and no real genetic difference.

The mountain ecotype described earlier was named
Tradescantia multiflora variety y linnaei by Clarke (1881)

286 Ећодога [Vol. 77

and T. ionantha by Diels (in Urban, 1906). Judging from
herbarium specimens there seems to be no character ex-
cept, perhaps, the bright magenta flower color by which
these plants can be distinguished.

The names Tradescantia cumanensis and Tripogandro
cumanensis have been applied to specimens of Tripogandra
serrulata, but these usages seem to stem from an initial
misidentification of specimens by Clarke. Kunth’s original
description of Tradescantia cumanensis (as Commelina
floribunda) notes that the top of the stem and the peduncle
are “hirto-pilosis” while the sepals are “glanduloso-pilosis.”
These characters coincide with those found in T. multi-
flora which is common in Venezuela where the type of
T. cumanensis was collected, but these characters are not
found in Т. serrulata which seems to be rare or at least
not often collected in Venezuela.

Kunth’s concept of similarities may be considered as a
further line of evidence to demonstrate the identity of
Т. multiflora and Т. cumanensis. Kunth (1843) grouped
similar species in his treatment of the family and he noted
the affinities he believed existed. It can be seen that
Tradescantia schlechtendalii (a synonym for Tripogandra
serrulata) is number 41 in Kunth’s list while T. procum-
bens, T. parviflora, T. multiflora, and T. cumanensis are
numbers 44, 45, 46, and 47, respectively. Using this evi-
dence as a reflection of similarities, T. cumanensis stands
apart from T. schlechtendalii (= Ттродатата serrulata)
but close to T. multiflora. I have only seen a photograph of
the holotype of T. cumanensis but an examination of seeds
of this specimen would make a positive indentification
possible.

By application of the Internal Code of Botanical Nomen-
clature, Aneilema floribunda, Leptorhoeo floribunda and
Tripogandra floribunda based on Commelina floribunda
HBK must be cited in synonymy under Tripogandra multi-
flora for reasons which follow.

Hooker and Arnott published the combination Aneilema
floribunda based on Commelina floribunda HBK but the

1975] Tripogandra — Handlos 287

specimen to which they applied this name was a species
of Leptorhoeo, a genus in need of further study. Subse-
quent authors, Baillon (1894) and Woodson (1942), have
credited Hooker and Arnott as authors of the specific
epithet, floribunda, but Hooker and Arnott correctly
credited HBK and did not, in fact, publish a new species
description though they did apply the name incorrectly.

Rohweder (1956) published Tripogandra ionantha and
Steyermark (1964) published T. multiflora as new com-
binations. Both of these names were validly published
earlier so neither Rohweder nor Steyermark can be credited
as the correct authors.

14. Tripogandra neglecta Handlos, sp. nov. HOLOTYPE:
Brazil. MINAS GERAIS: ad Lagoa Sta., 27 Mar. 1864,
Warming 1069/1(c!).

Herba аппца (7) ; caulis erectus, usque ad 40 cm altus,
internodiis usque ad 10.5 cm longis, linea unica pilorum
instructis aliter glabris. Folia angusto-ovata, laminis usque
ad 11.3 cm longis, usque ad 2.1 em latis, apice acutis, basi
cuneatis, glabris, margine ciliatis, vaginis usque ad 2.3 ст
longis, usque ad 1.0 cm diam., linea unica pilorum instructis,
aliter glabris, orificio parce villosis. Inflorescentiae termi-
nales et in axillis foliorum summorum, ex 4-9 cincinnis
duplicibus constantes; pedunculi usque ad 4.0 em longi,
1-2 lineis pilorum instructi, cincinni duplices omnes ala-
bastra, flores, vel fructus usque ad 12 gerentes; ped icelli
usque ad 5.5 mm longi, glabri, maturitate reflexi, bracteis
basi pedicellorum margine integris, glabris, Flores albi?;
sepala ovata, cucullata, usque ad 4.0 mm longa, usque ad
2.3 mm lata, pilis paucis uniseriatis instructa, margine
integra et hyalina, apice + obtusa; petala ovata, apice
obtusa; stamina 6 in verticillis duobus, 3 sepalis opposita
Матеп из brevibus, usque ad 1.3 mm longis, glabris,
antheris usque ad 1.1 mm longis, usque ad 1.2 mm latis,
3 petalis opposita filamentibus longioribus, usque ad 3.2
mm longis, sigmoideis, barbatis distaliter, antheris usque
ad 0.8 mm longis, usque ad 1.2 mm latis, connectivo C-

288 Ећодога [Vol. 77

formi; ovarium usque ad 1.0 mm longum, usque ad 0.8 mm
diam.. glabrum, stylo usque ad 0.6 mm longo, stigmate
simplici. Capsula globosa, usque ad 3.0 mm longa, usque
ad 3.1 mm diam., glabra, seminibus triangularibus, usque
ad 1.6 mm longis, minute reticulatis, hilo punctiformi.
Distribution and habitat: Known only from one location
in Brazil.
Flowering: The one specimen available was in flower
and fruit in March.
SPECIMENS EXAMINED

Brazil. MINAS GERAIS: ad Lagoa Santa, 27 Mar. 1864, Warming
1069/2 (c); without precise location: Warming (C).

This rarely collected species is similar to Tripogandra
diuretica from which it differs in the shape of the leaf
base, the unique presence of a few uniseriate hairs at the
junction between adjacent sepals, the flower color, the
surface texture of the testa and duration (apparently).

15. Tripogandra palmeri (Rose) Woodson, Ann. Missouri

Bot. Gard. 29:153. 1942.

Tradescantia palmeri Rose, Contr. U.S. Nat. Herb.
1:113. 1891. TYPE: México. SONORA: Alamos, 16-
30 Sept. 1890, Palmer 737 (Holotype, US!; isotypes,
GH!, NY!).

Descantaria palmeri (Rose) Brückner, Notizbl. Bot.
Gart. Berlin-Dahlem 10:56. 1927.

Illustration: Matuda, Anales Inst. Biol. Nac. México
26:374. 1956 (‘1955’). The magnification is given as <1
but it is more nearly X1.

Plants annual; stems erect, to 45 em tall, to 4.0 mm in
diam. when dry, unbranched to branched at every node
(6 branches) ; internodes to 11.7 em. long, green, glabrous
to pilose. Leaves elliptie, narrowly ovate, or ovate; blades
0.7-9.5 ст long, 0.2-4.0 em wide, cuneate, rounded or trun-
cate at the base, dorsal surface glabrous or pilose, ventral
surface variously glabrous, glabrous with a few uniseriate
hairs along the base of the midvein, or pilose, margin

1975] Tripogandra — Handlos 289

ciliate, apex acute; sheaths 1.0-9.5 mm long, 0.6-9.5 mm in
diam. when dry, villous at the orifice, with a line of hairs
extending down the side opposite the blade, otherwise
glabrous or pilose. Inflorescences borne terminally and in
the axils of the upper leaves; peduncles to 5.4 ст long, or
so reduced that cincinni appear sessile, green, proximally
glabrous or pilose, hairs uniseriate, distally variously
glabrous, with a few scattered hairs, or pilose, hairs capi-
tate; cincinni with 2-21 buds, flowers and/or fruits; pedi-
cels 1.5-8.0 mm long at anthesis, 0.3-0.6 mm in diam.,
green, variously glabrous, with a few scattered capitate
hairs, or pilose, erect at anthesis and in fruit; bracts at
the base of the pedicels with an entire margin, glabrous
or with a few scattered capitate hairs. Flowers white;
sepals cymbiform, elliptic to ovate, 1.5-4.1 mm long, 1.0-
2.3 mm wide, green with the tip darker, pilose or with a
few scattered capitate hairs, margin entire and hyaline,
apex acute or obtuse; petals elliptic, ovate, or obovate, 2.0-
4.5 mm long, ca. 1.0-5.0 mm wide, cuneate at the base, apex
obtuse or rounded; stamens 8, opposite the sepals, shorter
than the staminodes, filaments 0.5-1.2 mm long, colorless
with a few (1-6) uniseriate hairs on the upper half of the
dorsal side (Fig. 13), anthers 0.3-1.0 mm long, 0.2-0.6 mm
wide, white, basifixed, with white anther sacs, pollen white;
staminodes 3, opposite the petals, longer, slightly epipeta-
lous, filaments 1.5-3.4 mm long, white, glabrous, slightly
bent in the middle (Fig. 21) or S-shaped above the middle
(Fig. 22), anthers 0.3-1.0 mm long, 0.4-0.9 mm wide,
broadly basifixed, not versatile, connective yellow, incon-
spicuous, broadened with anther sacs parallel, or C-shaped
with the anther sacs divergent, pollen white or yellow;
ovary 0.5-1.0 mm long, 0.4-0.9 mm in diam., white,
glabrous, style 0.2-0.4 mm long, stigma simple. Capsule
globose, 2.0-3.3 mm long, brown or green, glabrous; seeds
one per locule, ovate with the broad end and two sides
rolled inward (involute) which results in concavities on
three sides due to the inrolling, these concavities alter-
nating with three convexities where the edge is not rolled

290 Rhodora [Vol. 77

(turned), involute margins nearly touching and covering
the protruding embryotega (Fig. 72), testa black, areolate,
hilum linear (Fig. 78).

Chromosome number: n=16.

Distribution and habitat: México, in the states of So-
nora, Sinaloa, Jalisco, Michoacan, Guerrero, and Puebla ;
at low elevations, 50 to ca. 1400 m. The plants grow in
relatively dry areas with seasonal rainfall; the vegetation
is characterized as thorn scrub; candelabra cacti usually
are abundant. The soil in which these plants grow is
brown or black loam with abundant humus but in dry
stream beds it is loose due to the presence of sand and
gravel.

Flowering: Plants flower in August, September and
October. The flowers commonly open about 8:00 AM and
close about noon.

REPRESENTATIVE SPECIMENS

México. SONORA: Chorijoa, Río Mayo, 22 Aug. 1935, Gentry 1598
(ARIZ, F). SINALOA: El monte, Los Labrados, 50 m., 15 Oct. 1926,
Mexia 927 (vc, US). JALISCO: near Bolaños, Sept. 1897, Rose 2891
(GH, K, NY, US). MICHOACAN: Apatzingán, Dist. Apatzingán, 300 m.,
19 Aug. 1938, Hinton et al. 12051 (GH, K, NY). MEXICO: Cerro de
Los Capulines, Palmar Chico, 1100 m., 26 Aug. 1954, Matuda 31337
(MEXU). GUERRERO: Coyuca-Chacamerito, Dist. Coyuca, 19 Sept.
1934, Hinton et al. 6620 (GH, K, MICH, NY, US). PUEBLA: km. 232.5
of Hwy. 190, ca. 4 km. west of Río Atoyac, 1370 (1040) m., 9 Sept.
1967, Handlos 403 (вн).

Tripogandra palmeri is unique because the seeds are
borne one per locule; at anthesis there are two ovules in
each locule but the lower one always aborts. The seeds are
further unique in that the margins are involute on three
sides, rolling up in such a fashion that they nearly cover
the protuberant embryotega.

An interesting pattern of morphological variation can
be seen in the 20 collections I have examined. АП 15 col-
lections made north of Guerrero or north of 18? 30' N
latitude have narrowly ovate, nearly glabrous leaves, few
flowers per inflorescence, a narrow connective, parallel

1975] Tripogandra — Handlos 291

anther sacs and epipetalous filaments which are bent
slightly in the middle. The five collections made south of
18° 30’ N in the southern drainage area of the Rio Balsas-
Rio Mexcala have larger, more nearly ovate leaves which
are commonly pilose on both surfaces (though some plants
have leaves which are glabrous ventrally except for the mid-
vein), there are more flowers per inflorescence, the connec-
tive is C-shaped, anther sacs are divergent, and the epipe-
talous filaments are S-shaped. Both of these forms were col-
lected in the drainage basin of the Rio Balsas. As the plants
are annuals and self-pollination is probably the most com-
mon breeding pattern, variation in local populations would
be expected to be small. Since the two morphological types
occur in different geographical areas they might be recog-
nized as subspecies. Because so few collections have been
made. a more complex pattern of variation may emerge
in the future and I decline, therefore, to introduce a new
name. However, the linear distance covered by this limited
number of collections is rather extensive, 1140 and 360
kilometers for the northern and southern forms respec-
tively. Further studies and collections should be made
throughout the range of this species and especially in the
Guerrero-Michoacán border area in the region of Presa
del Infiernillo and to the east which is the area of pre-
sumed contact between the two forms.

16. Tripogandra purpurascens (Schauer) Handlos, Baileya
Pes oT.
Tradescantia purpurascens Schauer in Nees ab Esen-
beck and Schauer, Linnaea 19:700. 1847. HOLO-
TYPE: Mejico, Aschenborn 493(B!).

Plants annual; stem erect, to 73 em tall, branched or un-
branched, lower portion sometimes decumbent, then root-
ing at the nodes; internodes to 12.4 em long, glabrous or
with scattered capitate hairs, a line of uniseriate hairs
always present. Leaves narrowly ovate; blades to 8.0 em
long, to 2.7 cm wide, complicate, with base rounded or
sometimes cuneate, glabrous on both sides (rarely with a

292 Ећодога [Уо]. 77

few hairs dorsally), margin ciliate, apex acute; sheaths
to 12.5 mm long, to 8.0 mm in diam., villous or with a few
long hairs at the orifice, a line of uniseriate hairs extending
down the side opposite the blade, otherwise glabrous or
with scattered capitate hairs to 1 mm long. Inflorescences
of 1-5 double cincinni borne terminally and in the axils of
the upper leaves; peduncles to 14.1 cm long, always with
seattered capitate hairs and with or without 1-2 lines of
uniseriate hairs extending down the sides; double cincinni
with up to 26 buds, flowers and/or fruits; pedicels to 7.0
mm long, reflexed in fruit, pilose or with scattered capitate
hairs (Fig. 1); bracts at the base of each pedicel pilose
or with scattered capitate hairs, margin partly, wholly,
or not at all ciliate. Flowers pink or rarely white ;
sepals ovate-cymbiform, to 5.0 mm long, to 2.8 mm wide,
green or with a red tip and/or base and/or margin, pilose
or with scattered capitate hairs (Fig. 1) to 1 mm long,
margin hyaline, apex more or less acute; petals obovate,
to 6.0 (-8.6) mm long, to 4.8 mm wide, base cuneate, apex
acuminate or irregularly indented; stamens in two whorls,
the outer whorl shorter, filaments to 1.8 mm long, pink in
the middle, white on the ends, bearing a tuft of moniliform
hairs on the mid-portion of the dorsal side (Fig. 5), anthers
0.4-1.1 mm long, 0.4-1.2 mm wide, white with a purple
line around the oper sac, basifixed, anther sacs more or
less parallel, pollen white; inner whorl of stamens longer,
to 5.5 mm long, pink in the middle and white on both ends,
giabrous, bent in an open S-shape and slightly expanded
in the upper portion (Fig. 18), anthers 0.5-1.1 mm long,
0.5-1.4 mm wide, basifixed, connective inflated, bent, yellow,
anther sacs divergent or nearly parallel through bending
of the connective, pollen cream-colored; ovary 0.8-1.4 mm
long, 0.6-1.4 mm in diam., green, glabrous, style 0.3-0.5 mm
long, stigma simple, capitellate or capitate. Capsule obo-
vate, 2.2-3.5 mm long, 1.8-4.0 mm in diam., green to light
brown, glabrous; seeds two per locule, triangular, 1.2-1.8
(-2.1) mm long, brown, testa ribbed areolate (Figs. 50,
51, 54, 55), hilum punctiform (Figs. 51, 55).

1975] Tripogandra — Handlos 293

In the past, plants of this species were identified as
Tripogandra disgrega. As noted earlier, the holotype for
T. disgrega is unmistakeable so another name must be
applied to these plants. The next available name is Tra-
descantia purpurascens Schauer. The description accords
well with Aschenborn 493 in the Berlin herbarium and
because I have seen no other specimen I consider this to
be the holotype. The description does not mention the
stamens or seeds but dissection of the flowers proved the
specimen to be Tripogandra. This specimen was seen by
C. B. Clarke, annotated by him, and is cited apparently
as “ Атђетђеан 493”, under his description of Tradescantia
disgrega. Several specimens of T. purpurascens were listed
by him under T. amplexicaulis. He apparently did not
have clear concepts of the species T. amplexicaulis and T.
disgrega, He does not include the name T. purpurascens
in his list of synonyms for T. disgrega or any other species.
This omission is surprising considering the number of
manuscript names that are included as synonyms in the
various species he dealt with.

The separation of and differences between Tripogandra
disgrega and T. purpurascens are discussed under the
former species.

Tripogandra purpurascens is found in every country
from México to Panamá, is absent from most of South
America and occurs again in southern Bolivia and northern
Argentina. This outlier is morphologically recognizeable
by the presence of scattered capitate hairs on all the inter-
nodes. I recognize these plants as T. purpurascens sub-
species australis. In all other respects it seems to ђе
identical with the typical element of the species.

16a. Tripogandra purpurascens (Schauer) Handlos subsp.
purpurascens.
Tradescantia minuta Clarke in DC., Monographiae
Phanerogamarum 3:307. 1881. TYPE: México.
Uhde 141а (Holotype, B!; isotype, L!).

294 Ећодога [Vol. 77

Tradescantia parvula Brandegee, Univ. Calif. Publ.
Bot. 6:51. 1914. TYPE: México. PUEBLA: Boca del
Monte, Oct. 1913, Purpus 6486 (Holotype, UC!; iso-
types, BM!, F!, GH!, MO!, NY!).

Descantaria minuta (Clarke) Brückner, Notizbl. Bot.
Gart. Berlin-Dahlem 10:56. 1927.

Tripogandra minuta (Clarke) Woodson, Ann. Mis-
souri Bot. Gard. 29:153. 1942.

Lower internodes glabrous except for a line of uniseriate
hairs extending down one side, rarely the upper internodes
with scattered capitate hairs.

Chromosome number: 716.
Vernacular name: matlale fide Kerber, México.

Distribution and habitat: México, Guatemala, Honduras,
El Salvador, Nicaragua, Costa Rica, Panamá; in seasonal
stream beds or as a weed in cultivated fields at elevations
from 990 to 2500 meters.

Flowering: In México flowering occurs from August to
December while in Guatemala it occurs from September
to January and occasionally in April and June. In the
remainder of Central America flowering plants mav be
found from September to November in Honduras and El
Salvador, December to February in Nicaragua, July to
December in Costa Rica, and December to March in
Panamá.

Flowers open about 10:00 AM and close about 12:45
PM.

REPRESENTATIVE SPECIMENS

México. CHIHUAHUA: base of Sierra Madre, 3 Oct. 1888, Pringle
1680 (BM, BR, L, M, UC). SINALOA: km. 1184 of Hwy. 40, 1.7 km.
above Potrerillos, 1600 m., 15 Oct. 1967, Handlos 464 (BH). DURANGO:
at the city of Durango and vicinity, 6297 ft., Apr.-Nov. 1896, Palmer
646 (BM, C, F, GH, MO, NY, US). ZACATECAS: Sierra de los Morones,
near Plateado, Sept. 1897, Rose 2713 (мү, US). SAN LUIS POTOSI:
Alvarez, 5-10 Sept. 1902, Palmer 139 (сн, MO, NY, US). NAYARIT:
vicinity of Jalisco, 10 Nov. 1925, Ferris 5893 (DS, US). JALISCO:
about 11 miles southeast of Lagos de Moreno, near highway to León,

1975] Tripogandra — Handlos 295

1900 m., 7 Sept. 1952, McVaugh 12829 (MICH). HIDALGO: lower
slopes of Mt. Lena, west of Lena Station (FCNM), Mun. Nopala,
Dist. Huichapan, 2500 m., 10 Oct. 1946, Moore, Jr. 1441 (BH, GH).
VERACRUZ: La Luz pr. Cérdoba, 7 Oct. 1882, Kerber 90 (BM, C, GOET,
M, US). MICHOACAN: sparsely to densely vegetated slopes of lava
flow east of San Juan Nuevo, ca. 8 km. south of Uruapan, 6100 ft.,
11-15 Oct. 1961, King & Soderstrom 4719 (MICH, NY, UC, US). MEXICO:
Ixtapan, Dist. Temascaltepec, 1000 m., 19 Oct. 1932, Hinton 2281
(F, GH, MO, NY, US). DISTRITO FEDERAL: Mt. Guadalupe prés México,
24 Aug. 1865-66, Bourgeau 888 (BR, C, GH, K, L, M, NY, US). MORELOS:
Las Guacamayas, Cuernavaca, 1800 m., 5 Aug. 1966, Rebolledo V.
(MICH, MSC). PUEBLA: Laguna San Baltasar, vicinity of Puebla,
2135 m., 20 Sept. 1906, Arséne 327 (US). GUERRERO: Pilas, Dist.
Mina, 1000 m., 24 Sept. 1937, Hinton 10712 (BR, GH, NY, POM, US).
OAXACA: Santiago Huitzo, 5500 ft., 22 Oct. 1894, Smith 231 (GH).
CHIAPAS: Milpa on the north edge of San Cristóbal las Casas, Muni-
cipio of San Cristóbal, 7100 ft., 25 Sept. 1965, Breedlove 12430 (ps,
F, MICH). Guatemala. HUEHUETENANGO: 8 km. south of Huehue-
tenango, 1800 m., 30 Nov. 1962, Williams, Molina К. & Williams
29117 (F, NY). QUICHE: mts. east of Quiché, 2020 m., 20 Nov.-4 Dec.
1940, Grant 656 (F, GH p.p., MICH p.p.). SAN MARCOS: San Marcos,
2000 m., 17 June 1882, Lehmann 1609 (вм, US). QUEZALTENANGO:
Quezaltenango, Sept. 1876, Bernoulli & Сато 781 (GOET). SOLOLA:
mountain slopes above Lake Atitlán, about 3-5 km. west of Pana-
jachel, 2100 m., 6-7 Dec. 1963, Williams, Molina R. & Williams 25378
(F, NY). CHIMALTENANGO: near Finca La Alameda, near Chimal-
tenango, 1830 m., 7 Dec. 1938, Standley 59029 (F). GUATEMALA:
Finea Bretafia, road between Guatemala and Fiscal, 1200 m., 12
Dec. 1938, Standley 59667 (F). JALAPA: Laguna de Ayarza, 8000
ft., Sept. 1892, Heyde & Lux 3885 (GH, US). CHIQUIMULA: Volcán
Quezaltepeque, 3-4 miles northeast of Quezaltepeque, 1500-2000 m.,
8 Nov. 1939, Steyermark 31502 (F). SANTA ROSA: Laguna Los Pinos,
below Cerro Redondo, 25 Oct. 1942, Steyermark 52168 (F, MO).
Honduras. MORAZAN: Guamiles sobre las faldas noroeste de la Mt.
Uyuca, cerca de Las Flores, drainage of the Río Yeguare, 1600 m.,
20 Oct. 1948, Molina R. 1273 (F, GH, MO). El Salvador. LA LIBERTAD:
rim of Volcán San Salvador, 1800 m., 22 Sept. 1946, Williams &
Molina R. 10622 (GĦ, MICH, MO). Nicaragua. JINOTEGA: road to La
Fundadora, entering at km. 142 from Managua, region of Santa
María de Ostuma, 1400 m., 7 Dec. 1958, Hawkes, Hjerting & Lester
2197 (C, K). MATAGALPA: road to La Fundadora, cloud forest area
north of Sta. María de Ostuma, Cordillera Central de Nicaragua,
1300-1500 m., Feb. 1963, Williams, Molina R. & Williams 24949
(F, NY). Costa Rica. ALAJUELA: Clairiere au rancho de l'Achiote
(Poas), 2200 m., Nov. 1896, Tonduz 10765 (BR, US). SAN JOSE:

296 Rhodora [Vol. 77

А San José, 1135 m., Oct. 1890, Tonduz 3047 (BR, м, US). CARTAGO:
Cartago, 4250 ft., Nov. 1887, Cooper 5962 (GH, NY, US). Panama.
CHIRIQUI: Alto Lino, vicinity of E] Boquete, 990 m., 3 Feb.-15 Mar.
1938, Maurice 743 (us).

The species Tradescantia minuta has confused various
authors — Matuda (1956) for example, includes specimens
of Leptorhoeo and Murdannia in his circumscription of it.
Clarke's original description states that the staminal fila-
ments are glabrous. One assumes from this that all six
are glabrous and if one studies the dissected flower present
on the holotype (Uhde 141a), this seems correct. The
problem arises from the fact that the dissected flower is
an immature one and the hairs on the filaments have not
yet developed. An examination of flowers well past an-
thesis with nearly mature capsules reveals that the short
filaments of the outer whorl are bearded and the longer
filaments of the inner whorl are glabrous. The remainder
of the description seems correct. Uhde 141a seems to repre-
sent a very reduced (depauperate) form of Tripogandra
purpurascens subsp. purpurascens. The corrected obser-
vation of the presence of bearded filaments leaves no
character by which to distinguish this material as a
different species.

As Clarke erred in describing his new species with
glabrous filaments, so T. S. Brandegee erred in describing
the six filaments of his new species, Tradescantia parvula,
as bearded. Again, a closer examination of the specimens
reveals that the three short filaments are bearded but the
three longer ones are glabrous. Tradescantia parvula
represents another depauperate form of Tripogandra pur-
purascens subsp. purpurascens. Plants which have a
strictly annual habit seem to be able to flower and produce
seeds when of a very small size and with very few leaves.
On the other hand some plants seem capable of producing
long stems. In México, plants of Handlos 160 flowered at
a height of 12.5 cm, but when seeds of these plants were
grown in the greenhouse in Ithaca, N.Y., the seedlings
reached a height of over a meter before flowering.

1975] Tripogandra — Handlos 297

Throughout México this subspecies is relatively uniform
in size considering the plasticity of annuals. The presence
of one or two lines of uniseriate hairs on the peduncle is
a sporadic character over much of México but in southern
México and in Central America it becomes more common
and is used as a key character by Standley and Steyermark
in their Flora of Guatemala. There is still variability —
some plants lack the line — but the frequency of occur-
rence is much higher than in México north of Chiapas.

The colonizing and weedy propensities of Tripogandra
purpurascens subsp. purpurascens should be noted. Plants
of this subspecies probably existed in stream beds and
naturally disturbed areas such as those around volcanoes
prior to man’s invasion of the ecosystem. Often I have
seen T. purpurascens subsp. purpurascens in seasonal
stream beds and on August 19, 1965, on a trip to the
volcano of Paricutin, I found plants of subsp. purpur-
ascens growing within ten meters of the edge of a lava
flow. The area had been covered with a layer of volcanic
ash but the plants were growing, flowering and producing
seeds under these conditions. Volcanic activity has oc-
curred extensively and for a long period of time in México.
Clausen (1959) has designated the central, volcanically
active area of México as the Trans-Mexican Volcanic Belt.
The weedy habit in this species probably could have existed
before the advent of man and his disturbances of the
natural vegetation but man must be credited for opening
additional areas for colonization by this species and allow-
ing it to become the most common species of Tripogandra
in México.

16b. Tripogandra purpurascens (Schauer) Handlos subsp.
australis Handlos, subsp. nov. TYPE: Bolivia. Toldos
bei Bermejo, 1850 m., 26 Nov. 1903, Fiebrig 2221
(Holotype. GH!; isotypes, K!, L!, M!).

Illustrations: Bacigalupo, Darwiniana 13:402. 1967,
Figs. 2, 5a-g, as T. disgrega.

298 Ећодога [Vol. 77

Internodia pilis capitatis, dispersis pubescentia. In
Bolivia australi et Argentina boreali indigena.

Internodes all with scattered capitate hairs and a line
of uniseriate hairs.

Distribution and habitat: southern Bolivia and northern
Argentina; in wet wooded areas or as a weed in disturbed
places at elevations from 450 to 1850 meters.

Flowering: In January, February and March in Argen-
tina.

REPRESENTATIVE SPECIMENS

Bolivia. Fiebrig 2221 (GH, к, L, M). Argentina, SALTA: Dept.
Gen. M. Güemes, camino de La Cornisa, between Jujuy and Salta,
at km. 1648.9, 1480 m., 18 Mar. 1966, Hawkes, Hjerting & Rahn
8909 (с). TUCUMAN: Cuesta del Garabatal, Sierra de Ende, Jan.
1874, Lorentz & Hieronymus 875 (CORD, E, F, СОЕТ).

17. Tripogandra saxicola (Greenman) Woodson, Ann. Mis-
souri Bot. Gard. 29:154, 1942.

Tradescantia saxicola Greenman, Proc. Amer. Acad.
Arts 39:70. 1903. TYPE: México. GUERRERO: Iguala
Саћоп, 14 Sept. 1900, Pringle 9270 (Holotype, GH!;
isotype, vT!).

Descantaria saxicola (Greenman) Brückner, Notizbl.
Bot. Gart. Berlin-Dahlem 10:56. 1927.

Plants annual, erect or with the lower portion decumbent,
rooting at lower nodes; stems to 31 ст long, unbranched
to branched at every node; internodes 0.1-7.4 cm long,
green or purplish-tinged at the base of the plant, a band
of uniseriate hairs extending from the sheath above to the
node below, otherwise variously glabrous to pilose, Leaves
narrowly ovate to ovate; blades 1.0-7.4 cm long, 0.1-2.0
em wide, glabrous dorsally, ventrally variously glabrous,
pilose along the midvein, or pilose over the entire surface,
base cuneate or truncate, margin ciliate; sheaths 1.0-5.0
mm long, 1.2-5.0 mm in diam. when dry, villous at the orifice,
a line of uniseriate hairs extending down the side opposite
the blade, otherwise glabrous or with a few scattered uni-
seriate hairs. Inflorescences few, borne terminally and in

1975] Tripogandra — Handlos 299

the upper leaf axils; peduncles to 5.9 ст long, ог so
reduced that the double cincinni appear sessile, glabrous
except for one or two lines of uniseriate hairs extending
down the side; double cincinni with 1-10 flowers; pedicels
2.2-5.5 mm long, 0.3-0.6 mm in diam., pilose to densely
pilose (Fig, 2), reflexed in fruit; bracts at the base of
each pedicel usually ciliate. Flowers white; sepals ovate-
cymbiform, 2.0-4.6 mm long, 1.3-2.4 mm wide, green,
sparingly pilose to densely pilose, apex acute to obtuse,
margin entire, hyaline; petals elliptic-ovate, ca. 2.5-5.8
mm long, ca. 2.0-4.5 mm wide, tapering at the base, apex
obtuse; stamens 6, in two whorls, those of the outer whorl
shorter, filaments 1.0-1.6 mm long, glabrous (Fig. 12),
anthers 0.6-1.0 mm long, 0.3-0.7 mm wide, basifixed, versa-
tile with parallel white anther sacs which are longer than
the gibbous white connective, pollen white; stamens of the
inner whorl longer, slightly epipetalous, filaments 2.0-4.5
mm long, bearded on the dorsal distal surface, with white
moniliform hairs (Fig. 20), connective U-shaped, 0.3-0.7
mm long, 0.3-0.9 mm wide, yellow, dorsifixed, versatile,
with yellow divergent anther sacs, pollen yellow: ovary
0.7-1.1 mm long, 0.6-1.1 mm in diam., green, glabrous,
style 0.2-0.4 mm long, stigma capitellate, Capsule glabrous,
brown, 2.5-2.7 mm long; seeds usually 6, 0.9-1.3 mm long,
triangular or triangular with a notch in one end, with 2-3
furrows on the hilar side forming 3-4 lobes, brown to gray-
brown, surface areolate (Fig. 56), hilum punctiform (Fig.
57) to elliptical.

Chromosome number: n—21.

Distribution and habitat: in the states of Guerrero,
Morelos, and Puebla, México; in thin soil in rocky areas
or under shrubs at elevations from ca. 840 to 1500 m.

Flowering: Flowers open about 3:00 PM and close about
5:00 PM. Therefore, this species appears to be unique
among the Mexican Tripogandra species because it is the
only one in which the flowers open in the afternoon; all
other species flower in the morning even though blossoms
may not close until afternoon.

300 Ећодога [Vol. 77

REPRESENTATIVE SPECIMENS

México. MORELOS: Yautepec, near Cuernavaca, 22 Oct. 1902,
Pringle (ут). PUEBLA: km. 216 on road to Oaxaca, Hwy. 190, about
16 km. SE of Izúcar de Matamoros, 30 Aug. 1965, Handlos 215
(BH). GUERRERO; Сайоп del Mano, along railroad tracks north of
Iguala, ca. 3 km. north of El Naranjo, са. 840 m., 13 Sept. 1967,
Handlos 418 (BH).

The origin of the annual habit may be illustrated by the
growth cycle of Tripogandra saxicola. In México this
species acts as an annual. All collections seem to be of the
current season’s growth judging from the presence of
juvenile leaves at the base of the plant and the absence
of old trailing stems and interconnections between plants.
In the greenhouse, however, plants of Handlos 215 have
functioned as perennials. The plants lose most of their
leaves during the winter, though flowering continues, and
some of the stems remain green. In May and June, vege-
tative shoots are produced, the plants grow vigorously,
rooting at the nodes, and vegetative reproduction is very
easy. However, not all collections react in this fashion.
Plants of Handlos 415 acted as annuals in the greenhouse
and died completely after flowering and producing seeds
in 1967 and 1968. This behavior seems significant as it
indicates that both the annual and perennial habit may
exist within some species. With the extinction of all the
perennial plants the species would appear strictly annual.
The presence of annual and perennial populations also
occurs in Tripogandra serrulata and may indicate that
the annual habit has arisen independently several times
within the genus and that some annual species may be of
recent origin.

The collections Handlos 198 and 419 merit mention be-
cause these plants were collected in the same area in two
different years. In 1965, when Handlos 198 was collected,
the area was covered with large shrubs and small trees
and had received enough rain so the ground was moist
and water dripped from the ledges. Plants reached a
maximum height of 31 centimeters and were found only

1975] Tripogandra — Handlos 301

after crawling under and through the bushes. Two years
later I found the whole area cut over, apparently for fire-
wood, with very little brush left. It was then possible to
walk and climb over the rocks with little interference
from the remaining woody vegetation. The plants col-
lected as Handlos 419 were wilted, what little soil was left
was exceedingly dry, and the few individuals which were
found reached a maximum height of only 15 centimeters.
This information not only gives an impression of how
much variation may occur in an area from year to year
but it also gives us a glimpse of the fate of the plants in
this area. First, there is a diminution of plant size; second,
there is a reduction in population size, and lastly, extinc-
tion is probable as the soil is gradually washed away from
the rocky ledges and weedy plants which are better com-
petitors grow in the alluvium deposited at the cliff bases.

18. Tripogandra serrulata (Vahl) Handlos, Baileya 17:33.
1970.

Commelina serrulata Vahl, Eclogae Americanae 2:4.
1798. HOLOTYPE: Ryan (C!).

Tradescantia elongata Meyer, Primitae Florae Es-
sequeboensis . . . 146. 1818. HOLOTYPE: apparently
lost.

Tradescantia congesta Martens & Galeotti, Bull. Acad.
Bruxelles 4(2) :377. 1842. HOLOTYPE: México.
VERACRUZ: Mirador, 3000 ft., June-Oct. 1840, Gale-
otti 4949 (BR!).

Tradescantia balbisii Kunth, Enumeratio Plantarum
4:97. 1843. TYPE: Portorico, 1834, Balbis (Holo-
type, B!; isotype, B!).

Tradescantia schlechtendalii Kunth, Enumeratio Plan-
tarum 4:94. 1843. TYPE: México. Hacienda de la
Laguna, Oct. 1828, Schiede 972 (Holotype, B!; iso-
type, HAL!).

Tradescantia schomburgkiana Kunth, Enumeratio
Plantarum 4:663. 1843. TYPE: Guiana Angl., 1842,
Schomburgk 189 (Holotype, B!; isotype, K!).

302 Rhodora [Vol. 77

Tradescantia guianensis Miquel, Linnaea 18:374. 1845
(‘1844’). TYPE: Surinam. Focke 400 (Holotype,
U!; isotype, GH!).

Descantaria ?balbisit Hasskarl ex Clarke in ЮС,
Monographiae Phanerogamarum 8:808. 1881, nom.
nud. pro sun,

Descantaria schlechtendalii Hasskarl ex Clarke in
DC., Monographiae Phanerogamarum 8:804. 1881,
nom. nud. pro syn.

Heterachtia gaudichaudiana Hasskarl ex Clarke in
DC., Monographiae Phanerogamarum 3:303. 1881,
nom. nud. pro syn.

Tradescantia cumanensis B glabrior Clarke in DC.,
Monographiae Phanerogamarum 3:306. 1881.
TYPE: Nicaragua. Chontales, 1867-8, Tate 452
(Holotype, K!; isotype, BM!).

Tradescantia elongata ё schlechtendalii (Kunth)
Clarke in DC., Monographiae Phanerogamarum
3:304. 1881.

?Descantaria elongata (Meyer) Brückner, Notizbl.
Bot. Gart. Berlin-Dahlem 10:56. 1927.

?Tripogandra elongata (Meyer) Woodson, Ann. Mis-
souri Bot. Gard. 29:152. 1942.

Tripogandra cumanensis f. glabrior (Clarke) Stand-
ley E Steyermark, Fieldiana: Bot. 24(3) :36. 1952.

Illustrations: Standley and Steyermark, Fieldiana: Bot.
24(3) :39, 1952, Fig. 9, a line drawing of a flower and
upper part of the plant. The position of the stamens is
incorrectly shown in the open flower and misplaced in the
flower drawing.

Plants perennial, decumbent at the base, rooting at
nodes, flowering stems erect; stems to 1.14 т long,
branching irregularly; internodes to 14.1 сш long, glab-
rous except for a line of uniseriate hairs extending
down the side from the sheath above (Fig. 4). Leaves
narrowly ovate, occasionally ovate; blades to 13.2 ст long,

1975] Tripogandra — Handlos 303

to 2.7 ст wide, glabrous to pilose dorsally, rarely with a
line of hairs along the midvein, glabrous to sparingly pilose
ventrally, often with a line of uniseriate hairs along the
midvein, base oblique, margin ciliate, apex acute; sheaths
to 2.1 em long, fo 1.2 cm in diam,, villous at the orifice, a
line of uniseriate hairs extending down the side opposite
the blade (Fig. 4), otherwise variously glabrous or the
uppermost occasionally with scattered uniseriate hairs or
pilose. Inflorescences composed of 1-13 double cincinni
borne terminally and in the axils of the upper 1-4 leaves;
peduncles 0.4-5.1 (-6.7) ст long, glabrous or with 1-2 lines
of uniseriate hairs extending down the sides; double cin-
cinni with up to 17 buds, flowers and/or fruits; pedicels
0.7-5.0 mm long, 0.5-0.7 mm in diam., reflexed in fruit,
glabrous to sparingly pilose (Fig. 3), hairs capitate; bracts
at the base of each pedicel glabrous, margin entire, ciliate,
or with a few scattered hairs. Flowers white or pink
(Fig. 16) ; sepals ovate, 2.5-4.6 mm long, 1.3-2.7 mm wide,
glabrous to sparingly pilose (Fig. 3), hairs capitate, mar-
gin hyaline, apex acute to obtuse; petals ovate-elliptic,
3.5-6.2 mm long, 2.3-4.3 mm wide, base cuneate, apex
rounded; stamens 6 in two whorls, the outer shorter, fila-
ments 1.0-1.5 mm long, white, glabrous or with a few (1-3)
moniliform hairs, anthers 0.5-1.0 mm long, 0.4-1.0 mm
wide, anther sacs parallel, connective inconspicuous, pollen
white; stamens of the inner whorl longer, filaments 2.7-
4.5 mm long, white, bent in an open S-shape, bearded with
moniliform hairs in the upper half (Fig. 19), anthers
0.4-0.8 mm long, 0.5-1.0 mm wide, basifixed, connective
elongate and narrow, anther sacs divergent, yellow, pollen
yellow; ovary 0.6-1.3 mm long, 0.5-1.0 mm in diam., white,
glabrous, style 0.1-0.8 mm long, stigma capitate, weakly
3-lobed. Capsule elliptical, 2.0-2.8 mm long, 1.5-2.5 mm in
diam., light brown, glabrous; seeds usually 2 per locule,
rounded triangular, 0.9-1.5 mm long, gray or brownish-
gray, testa reticulate, reticulations lighter in color (Fig.
38), hilum punctiform (Fig. 39).
Chromosome number: 716, 24.

304 Ећодога [Vol. 77

Vernacular names: suelda fide Bro. Daniel, Colombia ;
siempre viva fide Archer, Colombia; palm grass (Bar-
badian) fide Standley, Panamá ; matalin, shit fide Martínez-
Calderón, México; yerva del poyo fide Curtiss, México.

Distribution and habitat: central México, south to
Panamá, Colombia, Ecuador, Perü, Venezuela, Guyana,
Surinam, and the Caribbean islands; from sea level to
about 1500 m. Most specimens with altitudinal informa-
tion are from intermediate to low elevations. Two records
are exceptions. Matuda reports 2500 meters for one loca-
tion in México, while Heyde and Lux reported 8000 feet
for a collection in Guatemala. These reports should be
investigated further.

Flowering: in México from January to December; in
Guatemala from August to January and March, May, and
July; in British Honduras from October to February and
April; in Honduras from November to March and May,
June, August, and September; in El Salvador from Decem-
ber to April and July; in Nicaragua in April and De-
cember; in Costa Rica in every month except January; in
Panamá in every month of the year; in Colombia in
February, March, April, May, June, August, October,
November. and December ; in Ecuador in May and August;
and in Perü in April.

Flowers open in the morning about 9:00 AM and close
about 2:30 PM.

REPRESENTATIVE SPECIMENS

México. SAN LUIS POTOSI: Tamazunchale, 250 #., 5 Aug. 1987,
Fisher (MO, NY, US). HIDALGO: wooded slopes between Calnali and
Huazalingo, Dist. Huejutla, 29 May 1947, Moore, Jr. 3016 (BH).
VERACRUZ: Orizaba, Mt. Orizaba, 4000 ft., 29 Aug. 1891, Seaton 29
(F, GH, NY, US). DISTRITO FEDERAL: Monte Guadalupe prés México,
24 Aug. 1865, Bourgeau 888 (GH). MORELOS: Cuernavaca, 29 Oct.
1903, Holway 5261 (GH). PUEBLA: near Metlaltoyuca, 800 ft., 31
Jan. 1898, Goldman 49 (Us). OAXACA: Yaveo, Arroyo del Perrico,
Dist. Choapam, 475 m., 15 Mar. 1938, Mexia 9153 (F, GH, MO, NY,
0, UC, US). CHIAPAS: Escuintla, Nov.-Dec. 1937, Matuda 2176 (F,
GH, MICH, NY, UC). Guatemala. ALTA VERAPAZ: Finca Mocca, 3200
ft., 5 Jan. 1920, Johnson 169 (NY, US). IZABAL: vicinity of Quirigua,

1975] Tripogandra — Handlos 305

75-225 m., 15-31 May 1922, Standley 24165 (GH, МУ, US). SAN
MARCOS: river 5 mi. W of Malacatan, 400 m., 20 Nov. 1940, Grant
566 (F, GH). QUEZALTENANGO: Finca Pireneos, below Santa María
de Jesús, 1350-1380 m., 11 Mar. 1939, Standley 68347 (Е, MICH).
SACATEPEQUEZ: Ciudad Vieja, Mar. 1915, Tejada 343 (US). GUATE-
MALA: Brefias y seta cerca de Guatemala, 1400 m., July 1921, Tonduz
669 (US). JALAPA: Laguna de Ayarza, 8000 ft., Oct. 1892, Heyde &
Lux 3882 (GH, K, M, US). ZACAPA: Gualán, 420 ft., 20 Jan. 1905,
Deam 404 (GH, MICH). RETALHULEU: бап Felipe, 13 Jan. 1917,
Holway 710 (US). SUCHITEPEQUEZ: Chojoja p. Mazatenango, Sept.
1867, Bernoulli 469 (BR, NY). ESCUINTLA: Escuintla, 1100 ft., Mar.
1890, Donnell-Smith 2220 (GH, M, US). SANTA ROSA: Rio María
Linda, 3000 ft., Sept. 1893, Heyde & Lux 6251 (GH, US). SAN MARCOS:
Cangutz, 1140 m., 31 July 1922, Galas 11 (vs). British Honduras.
Gracie Rock, Sibun River, 15 April 1935, Gentle 1594 (F, GH, MICH,
MO, NY, US). Honduras. SANTA BARBARA: Río Permejo, 600 ft., Dec.
1888, Thieme 5532 (GH, US). CORTES: in ravine near Lake Yojoa,
Agua Azul, 630 m., 28 Dec. 1946, Williams & Molina К. 11411 (BH,
GH, MICH, MO, UC). COMAYAGUA: Rittenhouse’s hacienda near Sigua-
tepeque, 1050 m., 30 June 1936, Yuncker, Dawson & Youse 5527
(F, GH, MICH, MO, U). ATLANTIDA: Ceiba, 26 Sept. 1916, Dyer A94
(us). voRo: Farm 39 of the Tela Railroad Company, Guaymas
Dist., 30 m., 2 Feb. 1928, Standley 55489 (US). OLANCHO: a la orilla
de la quebrada cerca de El Plomo, Valle Catacamas, 300 m., 19 Nov.
1963, Molina R. 13290 (G, NY). LEMPIRA: faldas de Montana Puca
cerca de Los Cuábanos, 1300 m., 25 Sept. 1963, Molina К. 12956
(F, NY). MORAZAN: along Santa Clara Creek, drainage of the Rio
Yeguare, 850 m., 6 Aug. 1949, Williams & Molina R. 15865 (GH, US).
EL PARAISO: Montana entre Cifuentes у El Urraco, 900 m., 15 Mar.
1963, Molina R. 11428 (Е, мү, US). El Salvador. AHUACHAPAN:
vicinity of Ahuachapán, 800-1000 m., 9-27 Jan. 1922, Standley 19858
(GH, NY, US). SONSONATE: Finca Chilata, 26, 27 Dec. 1921, Standley
19311 (GH, NY, US). LA LIBERTAD: vicinity of Santa Tecla, 790-950
m., 10 Apr. 1922, Standley 23054 (US). SAN SALVADOR: San Salvador,
Tuly 1922, Calderón 913 (GH, MO, NY, US). LA PAZ: Zacatecoluca,
Mar. 1922, Calderón 303 (GH, NY, US). SAN VICENTE: vicinity of
San Vicente, 350-500 m., 2-11 Mar. 1922, Standley 21727 (GH, мо,
NY, US). Nicaragua. JINOTEGA: road to La Fundadora, entering at
km. 142 from Managua, region of Santa María de Ostuma, 1400 m.,
7 Dec. 1958. Hawkes, Hjerting & Lester 2198 (C). CHONTALES:
slopes of Mt. Mombacho, near Grenada, 460 m., 18 Dec. 1940-9 Feb.
1941, Grant 787 (F, GH, MICH). ZELAYA: vicinity of El Recreo, on
Río Mico, са. 30 m. 23 Apr.14 May 1949, Standley 19091 (F).
Costa Rica. ALAJUELA: Villa Quesada, Canton San Carlos, 825 m.,
10 Mar. 1940, Smith p2574 (F, MICH, MO). SAN JOSE: vicinity of

806 Ећодога [Vol. 77

El General, 915 m., Feb. 1936, Skutch 2602 (GH, MICH, MO, NY, US).
CARTAGO: Angostura, 19 June 1874, Kuntze 2050 (NY). LIMON:
Jiménez, Llanos de Santa Clara, 650 ft, Apr. 1894, Donnell-Smith
4976 (GH, K, US). Panamá. BOCAS DEL TORO: Bocas del Toro, 6 Nov.
1920, Carleton 71 (GH, NY, US). CHIRIQUI: Boquete, Boquete Dist.,
3800 ft., 17 May 1938, Davidson 665 (F, GH, MO, US). COCLE: Lower
Río Anton, vicinity of El Valle De Anton, 800-1000 (600) m., 30
Dec. 1936, Allen 111 (GH, MO, PH). CANAL ZONE: ruins of fort, Fort
San Lorenzo, Fort Sherman Military Reservation, 14 June 1923,
Maxon & Valentine 7018 (c, GH, US). PANAMA: Tumba Muerto
Road, near Panamá, 6 Jan. 1924, Standley 29717 (C, US). DARIEN:
vicinity of Boca de Сире, са. 40 m., 5 Oct. 1938, Allen 890 (Е, GH,
MO, NY, US). SAN BLAS: Perme, 24 Apr. 1933, Cooper III 253 (NY,
US). Venezuela. CARABOBO: Represa del acueducto de San Esteban,
Pto. Cabello, 7 Jan. 1965, Aristeguieta 5421 (VEN). ARAGUA: entre
Guamilas y Rancho Grande, P.N., 850 m., 6 Oct. 1938, Williams
10382 (VEN). FEDERAL DISTRICT: Caracas and vicinity, 3000-3500 ft.,
9 Jan. 1921, Bailey & Bailey 811 (NY, US). MIRANDA: bosque de Los
Guayabitos, arriba de Baruta, Nov. 1964, Aristeguieta 5395 (VEN).
MERIDA: 3!4 miles west of city of Mérida, 5000 ft., 24 Jan. 1931,
Reed 288 (us). BOLIVAR: San José, Ciudad Bolivar and vicinity, on
the Orinoco, about 200 ft., 9 Jan. 1921, Bailey & Bailey 811 (BH).
Colombia. MAGDALENA: Minca road, Santa Marta, 1200 ft., 28 Nov.
1898-1899, Smith 2280 (BM, BR, E, F, GH, L, MICH, MO, NY, PH, U, UC,
Us, VT, WIS). BOLIVAR: Los Hurtados, on Río Sinu, 40-70 m., 4 Feb.
1918, Pennell 4153 (Ny, US). CHOCO: Andagoya, 70-100 m., 20-30
Apr. 1939, Killip 35075 (BM, мо, US). ANTIOQUIA: vicinity of Medel-
Ип, 10 Mar. 1927, Toro 37 (му, US). SANTANDER: Puerto Wilches
and vicinity, 100 m., 28 Nov.-2 Dec. 1926, Killip & Smith 14773 (NY).
VALLE DEL CAUCA: Estero de Congrejo, north shore of Buenaventura
Bay, near sea level, 8 June 1944, Killip & Cuatrecasas 38730 (Е, US).
CALDAS: Santa Cecilia, Cordillera Occidental, Vertiente Occidental,
800 m., 16 Feb. 1945, von Sneidern 5181 (US). TOLIMA: “La Trini-
dad," Libano, 1000-1200 m., 21-25 Dec. 1917, Pennell 3363 (NY).
Ecuador. PICHINCHA: entre Santo Domingo у la Heda. Lelia, Sec-
éion Occidental, 400-800, 950-1100 m., 11 Aug. 1945, Acosta-Solis
10937 (Е, US). IMBABURA: entre El Pajón y Сасћасо, 600, 740 m.,
30 May-12 June 1949, Acosta-Solís 12728 (к, US). Pert. TUMBES:
a 8 km. al sur de Tumbes, 15-20 m., 24 Apr. 1949, Ferreyra 6002
(US). LORETO: near km. 194 below Divisoria on road from Tingo
Maria to Aguaytia, Prov. Coronel Portillo, ca. 1400 m., 22 Dec. 1960,
Moore, Jr., Salazar C. & Smith 8632 (вн). JUNIN: La Merced,
ca. 700 m., 29 May-4 June 1929, Killip & Smith 23416 (Ny, US).
Guyana. POMEROON DISTRICT: Mora Landing, Moruka River, 21-23
Aug. 1922, De La Cruz 1844 (BH, F, GH, MO, NY, US). Surinam.

1975] Tripogandra — Handlos 307

Suriname R. near Gansee, 15 Nov. 1933, Lanjouw 1307 (Ny, U, US).
Dominican Republic. Madre Vieja, Nagua, Prov. Maria Trinidad
Sánchez, 20 Dec. 1964, Jimenez 5108 (мү). Puerto Rico, San Juan,
14 km. 5 on Mil. Road, 2 Mar. 1899, Heller & Heller 662 (к, му, US).
Guadeloupe. 1895, Duss 3619 (F, NY, US). Dominica. prope Wotten
Waven, 200 m., Dec. 1887, Eggers 690 (BR, CORD, GOET, L, M, UC).
Martinique. La riviere du Galion (Trinite) et de la riviere du Carbet,
Oct. 1888, Duss 1024 (мү). St. Vincent. 1000 ft., March 1890, Smith
& Smith 1660 (BM). Trinidad. North Range, roadside forest, Arima
Valley Road, 500 m., 24 Mar. 1959, Cowan & Simmonds 1180 (Nv,
Us):

Tripogandra serrulata may be easily distinguished from
other species by its gray or gray-brown seeds with lighter
colored reticulations which produce a cobweb pattern over
the surface.

Within Tripogandra serrulata there is variation in leaf
shape as well as in the vesture of leaves, peduncles, pedicels,
and sepals. I have not seen any consistency or pattern in
this variation. This does not of necessity mean that none
exists, however, only that my study has not been precise
enough and that the information available to me has not
been complete.

Many of the specimens associated with this species have
been identified in the past as Tripogandra cumanensis.
This seems to be the result of a misidentification of speci-
mens by Clarke. The name which has priority is Com-
melina serrulata. Tradescantia cumanensis is cited as a
synonym of Tripogandra multiflora and is discussed under
that species.

Vahl’s description of Commelina serrulata agrees well
with the type specimen, but does not mention the stamens
or the seeds. It would be difficult to know from the descrip-
tion alone to which species or even genus this specimen
should be referred. The specimen has nearly mature seeds
which have a reticulation matching that on plants which
recent taxonomists have called Tripogandra cumanensis.
The stamens follow the diagnostic Tripogandra pattern;
they are dimorphic with the shorter ones opposite the

308 Ећодога [Vol. 77

sepals, It was probably because of the dimorphic stamens
that Vahl placed this species in the genus Commelina.
The name Commelina serrulata has not been used in recent
years but the type specimen is undoubtedly a Tripogandra.
Kunth (1843) included this species in the genus Commelina
in a section called “Species valde dubiae," indicating that
he was not certain what the plant was. His decription was
taken from Vahl and in parts is nearly a word-for-word
copy, implying that he had probably not seen a specimen.

The problem of the identity of Tradescantia elongata
has plagued taxonomists for many years. I have not seen
the type specimen; it should be a Rodshied collection
(Stearn and Williams, 1957) preserved at Gottingen but
was not located among the specimens obtained on loan.
If T. elongata does actually apply to Tripogandra it can
only apply to the one taxon which seems to occur in
the vicinity of the Essequibo River in Guyana. The plants
from this area are of great interest and should be studied
further. I have examined sixty-one sheets of specimens
from this area; all sheets have several inflorescences but
none of them have fruits in any stage of development. In
addition Т mounted pollen in aniline blue-lactophenol from
some of the plants identified as T. elongata from through-
out its range. The pollen does not stain and on that basis
is considered to have been inviable when fresh. From per-
sonal experience with T. serrulata in México and knowledge
of the existence of sterile plants within these populations,
I have treated all these sterile collections as part of the
species T. serrulata, but only after studying and describing
that species from fertile specimens. The pattern and range
of variation within the sterile collections falls within that
of fertile plants of T. serrulata, except for the peduncle
length of three collections from Guyana. The longest pe-
duncles were 5.7, 6.4, and 6.7 ст. while the longest peduncle
on fertile T. serrulata was 5.1 ст. I do not consider this
problem solved. Тће apparent sterility of these plants
must be studied in the field to see if it is characteristic
or whether my sample by some quirk is very biased.

1975] Tripogandra — Handlos 309

Tripogandra elongata has been interpreted broadly in
the past and three taxa have been included within it. These
are T. diuretica of southern Brazil, Bolivia, Paraguay,
Agentina, and Uruguay, Т. montana of Central America,
and T. elongata sensu stricto of eastern Venezuela, Guyana
and Surinam. These three taxa are not sympatric in any
part of their ranges and are morphologically distinguish-
able. Therefore, I prefer to recognize T. diuretica and
T. montana as separate and distinct species, while the
sterile plants of Т. elongata are included within T. ser-
rulata.

Kunth (1843) proposed the name Tradescantia schlech-
tendalli based on Schiede 972, which Schlechtendal had
misidentified and called Commelina mexicana Presl. Presl’s
original description contains several points indicating that
the plants were different: “Petala caerulea, . . . Stamina
tra. Filamenta erecta fertilia hirsuta, duo pistillo breviora,
tertium pistillo longius. Antherae . . . apice poro dehis-
centibus polliniferia . . . Stylus simplex curvatus. Stigma
emarginatum.” The Schiede specimen has white petals,
six stamens, three longer and three shorter, and a capitate
stigma. Kunth proceeded, however, to recognize Com-
melina mexicana Presl as a species of Tradescantia. I
believe that this name may apply to a species of Com-
melina though I am not certain of this, In any case it
certainly is not a species of Tradescantia or Tripogandra,
on the basis of the description given by Presl.

Clarke’s variety 8 glabrior of Tradescantia cumanensis
does not seem to differ from 7. congesta. Clarke states
that the pedicels and sepals are glabrous when, in fact,
there are a few hairs to be found on the type specimen
though some structures are glabrous. These two collec-
tions are certainly conspecific but Clarke considered gla-
brior to be a variety of T. cumanensis while T. congesta
was made a synonym of T. elongata.

Plants of Tripogandra serrulata have been collected from
several Caribbean islands. АП of these collections lack
seeds except one from Hispaniola. The seeds seem identical

310 Кћодога [Vol. 77

with those of mainland Т. serrulata. If this plant repre-
sents the native and not a recently introduced Tripogandra,
this gives me further confidence in considering these plants,
which have been called Т. elongata, as conspecific with
T. serrulata.

19. Tripogandra silvatica Handlos, sp. nov. TYPE: México.
VERACRUZ: Montepio, 19 km. al E de Catemaco, 19
Mar. 1965, Gonzales Quintero 2239 (Holotype, MICH!;
isotype, MSC!).

Herba perennis(?); caulis decumbens, usque ad 30 cm
longus; internodiis usque ad 5.5 cm longis, linea unica
pilorum instructis, aliter glabris. Folia ovata, laminis
usque ad 2.9 ст longis, usque ad 1.4 ст latis, basi obliquis,
apice acutis, dorsaliter glabris, ventraliter glabris vel pilis
dispersis instructis praeter lineam partialem pilorum secus
costam, margine ciliatis, vaginis usque ad 5.6 mm longis,
usque ad 3.0 mm diam., orificio villosis vel pilis dispersis
praeditis, linea unica pilorum instructis, aliter glabris.
Inflorescentiae terminales, ex 1 (-3) cincinnis duplicibus
constantes; pedunculi usque ad 2.3 ст longi, glabri vel
pilis capitatis paucis dispersis instructi; cincinni duplices
omnes alabastra, flores, vel fructus usque ad 13 gerentes;
pedicelli usque ad 4.5 mm longi, maturitate reflexi, gla-
briusculi ad pilosi, bracteis basi pedicellorum glabris,
margine erosis, nonnunquam ciliatis. Flores albi; sepala
ovata, cucullata, usque ad 3.7 mm longa, usque ad 1.6 mm
lata, pilis capitatis pilosa, margine hyalina, apice = ob-
tusa; petala usque ad 4 mm longa; stamina 6 in verticillis
duobus, 3 sepalis opposita Матеп из brevibus, usque ad
1.8 mm longis, glabris vel ?pilis paucis moniliformibus
praeditis, antheris usque ad 0.5 mm longis, usque ad 0.6
mm latis, 3 petalis opposita filamentibus longioribus, usque
ad 2.7 mm longis, glabris, distaliter sigmoideis, antheris
usque ad 0.7 mm longis, usque ad 0.6 mm latis; ovarium
usque ad 0.7 mm longum, glabrum, stylo usque ad 0.4 mm
longo, stigmate capitellato. Capsula obovoidea, usque ad
2.3 mm longa, usque ad 1.5 mm diam., glabra, basi stipitata,

1975] Tripogandra — Handlos Bil

seminibus triangularibus, usque ad 1.2 mm longis, reticu-
latis, hilo punctiformi.

Distribution and habitat: in the state of Veracruz,
México, in the area around Colipa-Misantla and Catemaco;
in wet forest lowlands.

Flowering: March and April.

SPECIMENS EXAMINED

México. VERACRUZ: Misantla, Mar. 1841, Liebmann (C); inter
Colipa et Misantla ad Palenque, Маг. 1841, Liebmann (c); Colipa,
Mar. 1841, Liebmann 350 (c); Colipa, Mar. 1841, Liebmann (С);
Jalapa, 4000 ft., З Apr. 1899, Pringle 7810 (vT).

Dried specimens of this species resemble Leiandra
eordifolia superficially — usually bearing only one terminal
double cincinnus. Dissection of the flower is necessary to
reveal the dimorphic nature of the stamens.

20. Tripogandra warmingiana (Seubert) Handlos, comb.
nov.

Tradescantia warmingiana Seubert in Warming,
Vidensk. Meddel. Dansk Naturhist. Foren. Kjoben-
havn 126. 1872. HOLOTYPE: Brazil. MINAS GERAIS:
Lagoa Santa, 6 Mar. 1866, Warming 1046 (c!).

Plants annual(?) ; stem decumbent at the base and root-
ing at the nodes, to 19 ст long, branched; internodes to
5.6 em long, glabrous except for a line of uniseriate hairs
extending down the side from the sheath above. Leaves
narrowly ovate to ovate; blades to 4.4 em long, to 1.85 em
wide, with the base always oblique but narrowed and sub-
petiolate on the lower part of the plant, glabrous dorsally,
glabrous ventrally except for a line of hairs at the proximal
end of the midvein, margin ciliate, apex acute and slightly
acuminate; sheaths to 4.4 mm long, to 3.1 mm in diam.,
sparingly villous at the orifice, otherwise glabrous except
for a line of uniseriate hairs extending down the side
opposite the blade. Inflorescences composed of 1-3 double
cincinni borne terminally and in the upper leaf axils;
peduncles to 1.2 ст long, green, glabrous; double cincinni

312 Ећодога [Vol. 77

with up to 8 buds, flowers and/or fruits, pedicels to 4.5
mm long, green, with a few uniseriate hairs near the
distal end, reflexed in fruit; bracts at the base of each
pedicel glabrous, margin erose. Flowers white; sepals
ovate, to 3.4 mm long, to 1.7 mm wide, pilose along the
midvein, the remainder with scattered uniseriate hairs,
margin hyaline, apex acute; petals ovate-elliptic (fide
Warming), to 5.0 mm long, to 2.3 mm wide, apex obtuse
(fide Warming); stamens 6, in two whorls, the outer
shorter, filaments to 1.2 mm long, glabrous, anthers to 0.7
mm long, to 0.5 mm wide, anther sacs parallel; stamens
of the inner whorl longer, filaments to 3.7 mm long, bent
in an S-shape (fide sketch, presumably by Warming),
bearded in the lower portion of the upper half with monili-
form hairs, anthers to 0.5 mm long, to 1.2 mm wide, con-
nective elongate and anther sacs divergent; ovary to 0.9
mm long, 0.8 mm in diam., glabrous, style to 0.5 mm long,
stigma minutely capitellate. Capsule globose, to 3.0 mm
long, to 2.8 mm in diam., light brown, glabrous; seeds 2
per locule, rounded-triangular, gray or brown, testa reticu-
late-foveate (Fig. 61), hilum punctiform.

Distribution: known only from three collections at Lagoa
Santa, Brazil.

SPECIMENS EXAMINED
Brazil. MINAS GERAIS: Lagoa Santa, 10 Mar. 1864, Warming
1070 (778) (с); Lagoa Santa, Warming (F, US р.р.).

I know this species from only three collections. Accord-
ing to the original description it was found in a wooded
area above a rocky calcareous site near Lagoa Santa,
Brazil and flowered from January to March.

The original description accords well with the type
specimen. One discrepancy exists concerning the shorter
stamens. The original description states: “Stamina tria
interiora minora, . . ." In fact, the shorter stamens are
opposite the sepals and are the outer whorl of stamens.

C. B. Clarke considered this species as part of his
Tradescantia elongata. The habit and size of the plant.

1975] Tripogandra — Handlos 313

shape of the laminar base, seeds, flower size and color, and
stigma show this species to be distinct.

EXCLUDED SPECIES

Tripogandra cordifolia (Swartz) Aristeguieta, Bol. Acad.
Ci. Fis. 25:125. 1965.
Tradescantia cordifolia Swartz, Nova genera & species
plantarum seu prodromus . . . 57. 1788.

Leiandra cordifolia (Swartz) Rafinesque, Flora Tellu-

мапа 2:16. 1837 (‘1836’).

Callisia cordifolia (Swartz) Anderson & Woodson, Contr.

Arnold Arbor. 9:117. 1935.
Phyodina cordifolia (Swartz) Rohweder, Abh. Aus-
landsk., Reihe C, Naturwiss. 18:151. 1956.

I have excluded Tripogandra cordifolia because it has
six more or less similar, glabrous stamens. The genus
Phyodina as interpreted by Rohweder is composed of di-
verse elements which show little affinity. This problem
has been discussed to some extent by Moore (1963) but
requires much more study in conjunction with a considera-
tion of Tradescantia gracilis, T. debilis, and Leptorhoeo
filiformis.

Tripogandra lundellii (Standley) Woodson, Ann. Missouri
Bot. Gard. 29:153. 1942.

Tradescantia lundellii Standley, Publ. Field Mus. Nat.
Hist. Bot. Ser. 22:5. 1940. TYPE: Lundell 7098 (Holo-
type F!; isotypes, MICH !, NY!).

Gibasis sp.

Standley's original description of Tradescantia lundellii
described stamens of two lengths. This species cannot ђе
considered one of Tripogandra because at anthesis the
plants have six essentially similar stamens which are borne
on flowers in a single cincinnus.

Tripogandra rosea (Ventenat) Woodson, Ann. Missouri
Bot. Gard. 29:153, 1942.
Tradescantia rosea Ventenat, Jard. Cels 24. 1800. HOLO-
TYPE: Michaux P; photograph, GH!.

314 Ећодога [Vol. 77

Cuthbertia rosea (Ventenat) Small, Flora of the South-
eastern United States 287. 1903.

Phyodina rosea (Ventenat) Rohweder, Abh. Auslandsk.,
Reihe С, Naturwiss. 18:151. 1956.

Tripogandra rosea lacks the dimorphic stamens which
characterize Tripogandra sensu stricto. Rohweder’s inclu-
sion of this species in Phyodina should be investigated
further.

Tripogandra stenophylla (Brandegee) Matuda, Anales
Inst. Biol. Univ, Nac. México 26:369. 1956 (‘1955’).
Tradescantia stenophylla Brandegee, Univ. Calif. Publ.
Bot. 3:377. 1909. TYPE: Purpus 3352 (Holotype,
uc!; isotype, NY!).
Tripogandra stenophylla is properly placed in the genus
Tradescantia because this plant has six similar stamens
and two foliaceous bracts subtending the double cincinnus.

Tripogandra warscewicziana (Kunth & Bouche) Woodson,

Ann. Missouri Bot. Gard. 29:154. 1942.

Tradescantia warszewicziana Kunth & Bouche, Ind.
Sem. Hort. Berol. 11. 1847.

Spironema warszewiczianum (Kunth & Bouche) Brück-
ner, Notizbl. Bot. Gart. Berlin-Dahlem 10:56. 1927.

Phyodina warscewicziana (Kunth & Bouche) Rohweder,
Abh. Auslandsk., Веће С, Naturwiss. 18:151. 1956.

Hadrodemas warszewicziana (Kunth & Bouche) Moore,
Baileya 10:134. 1963 (‘1962’).

Tripogandra warscewieziana must be excluded because
of its inflorescence structure and a lack of dimorphic
stamens. Moore (1963) has more fully discussed the place-
ment of this unusual species.

Descantaria laxiflora (Clarke) Briickner, Notizbl. Bot.
Gart, Berlin-Dahlem 10:56. 1927.

Tradescantia laxiflora Clarke in DC, Monographiae
Phanerogamarum 3:307. 1881. TYPE: Andrieux 51
(Holotype, K!; isotype, M!).

Gibasis sp.

1975] Tripogandra — Handlos SE

This species has a single cincinnus so it cannot ђе con-
sidered a species of Tripogandra.

ACKNOWLEDGMENTS

The author expresses his appreciation and gratitude to
Drs. H. E. Moore, Jr., H. T. Stinson, and R. B. Root,
Special Committee members, for their advice, criticism,
and assistance. The writer is indebted to Drs. C. H. Uhl,
М. W. Uhl, С. Eickwort, and У. J. Dress, who have pro-
vided assistance and encouragement with special aspects
of this work. Deepest appreciation is due Dr. Moore for
initial inspiration for this project and for the many hours
he devoted to guidance and advice.

The writer wishes to thank the curators of the following
herbaria for the loan of specimens: ARIZ, B, BAB, BH, BM,
BR, C, CORD, DS, Е, Е, С. GH, СОЕТ, HAL, К, L, LA, M, MA, MICH,
MO, MSC, NY, PH, R, RSA, P, SP, TENN, U, UC, US, VEN, VT,
WIS. The staff at the Instituto de Biología, Universidad
Nacional Autonoma de México (MEXU), have been espe-
cially kind and helpful during the author's trips to México.

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BOTANY DEPARTMENT
UNIVERSITY OF ZAMBIA
LUSAKA, ZAMBIA

320 Кћодога [Vol. 77

Fig. 1. Upper pedicel and lower calyx of Tripogandra purpur-
ascens bearing capitate hairs. X 17.

Fig. 2. Upper pedicel and lower calyx of T. saxicola bearing
uniseriate hairs. X 17.

Fig. 3. Upper pedicel and lower calyx of T. serrulata. X 17.

Fig. 4. Sheathing leaf base and node of Т. serrulata showing a
continuous line of uniseriate hairs on the sheath and internode.

x 11.

1975] Tripogandra — Handlos 321

322

Rhodora

[Vol. 77

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Outer stamen of Tripogandra purpurascens.

Stamen of 7.
Stamen of 7.
Stamen of T.
Stamen of T.
Outer stamen
Stamen of 7.

Outer stamen of T. saxicola. X 22.

Stamen of 7.

amplexans. X 11.
amplexicaulis. X 11.
grandiflora. X 11.
grandiflora. X 11.

of T. montana. X 11.
guerrerensis. X 11.

palmeri. X 22.

Stamen of T. angustifolia. X 22.

X

11.

Tripogandra — Handlos

1975]

по А "E У
Мр

»

324

Rhodora

[Vol. 77

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Androecium of Tripogandra guerrerensis.

Flower of T.
Staminode of
Inner stamen
Inner stamen
Inner stamen
Staminode of
Staminode of

serrulata.

х 7.5.

T. guerrerensis. X 11.

of Т. purpurascens. X 11.

of T. serrulata. 11.
of T. saxicola. X 11.

T. palmeri.
T. palmeri.

х 11.
х 11.

x 11.

Tripogandra — Handlos

1975]

Rhodora [Vol. 77

Fig. 23.

Distal portion of staminode of Tripogandra amplexicaulis

showing the inflated filament. X 11.

Fig. 24.

Germinating seed of T. purpurascens showing the emerg-

ing radicle and capping embryotega. X 34.5.

Fig. 25.

Androecium and gynoecium of T. amplerans showing

the relative positions of stamens and staminodes. Х 11.

Fig.
Fig.
x 22.
Fig.
Fiz.
X 11.
Fig.
Fig.

26.
27.

28.
29.

30.
31.

Staminode of T. amplexans, lateral view. X 11.
Distal portion of filament and anther of T. angustifolia.

Distal portion of staminode of T. grandiflora. X 11.
Staminode of T. amplexicaulis-T. amplexans intermediate.

Staminode of T. angustifolia. X 16.
Distal portion of inner stamen of T. montana. X 11.

1975]

Tripogandra — Handlos

328 Кћодога [Vol. 77
Fig. 32. Seed of Tripogandra multiflora, dorsal surface.
Fig. 33. Seed of T. multiflora, ventral surface showing puncti-
form hilum.
Fig. 34. Seed of Т. multiflora, dorsal surface.

Fig. 35. Seed

form hilum.

Fig. 36. Seed

Fig. 37. Seed
hilum.

Fig. 38. Seed

Fig. 39. Seed
form hilum.

Fig. 40. Seed

Fig. 41. Seed

form hilum.

Fig. 42. Seed

Fig. 43. Seed
form hilum.
Fig. 44. Seed
Fig. 45. Seed
form hilum.
Fig. 46. Seed
Fig. 47. Seed
Fig. 48. Seed
hilum.
Fig. 49.
upper
carpel.

The

of T. multiflora, ventral surface showing puncti-

of Tradescantia triandra, dorsal surface.
of T. triandra, ventral surface showing punctiform

of Tripogandra serrulata, dorsal surface.
of T. serrulata, ventral surface showing puncti-

of T. diuretica, dorsal surface.
of T. diuretica, ventral surface showing puncti-

of T. glandulosa, dorsal surface.
of T. glandulosa, ventral surface showing puncti-

of T. montana, dorsal surface.
of T. montana, ventral surface showing puncti-

of T. montana, lateral surface.
of T. brasiliensis, dorsal surface.
of T. brasiliensis, ventral surface showing elliptical

Cleared ovary of T. guerrerensis showing two seeds, the
larger and the lower smaller, and the dorsal bundle of one

scale represents 1 mm.

Handlos 329

1975] Tripogandra

330 Ећодога [Vol. 77

Fig. 50. Seed of Tripogandra purpurascens subsp. purpurascens,
dorsal surface.

Fig. 51. Seed of T. purpurascens subsp. purpurascens, ventral
surface showing punctiform hilum.

Fig. 52. Seed of T. disgrega, dorsal surface.

Fig. 53. Seed of T. disgrega, ventral suface showing punctiform
hilum.

Fig. 54. Seed of T. purpurascens subsp. australis, dorsal surface.

Fig. 55. Seed of T. purpurascens subsp. australis, ventral sur-
face showing punctiform hilum.

Fig. 56. Seed of T. saxicola, dorsal surface.

Fig. 57. Seed of T. saxicola, ventral surface showing punctiform
hilum.

Fig. 58. Seed of T. amplexans, dorsal surface.

Fig. 59. Seed of T. amplexans, dorsal surface.

Fig. 60. Seed of T. amplexans, ventral surface showing linear
hilum.

Fig. 61. Seed of T. warmingiana, dorsal surface.

Fig. 62. Seed of T. amplexicaulis-T. amplexans intermediate,
dorsal surface.
` Fig. 63. Seed of T. amplexicaulis-T. amplexans intermediate,
ventral surface.

Fig. 64. Seed of T. amplexicaulis, dorsal side showing alveolate
surface.

Fig. 65. Seed of Т. amplexicaulis, ventral side showing linear
hilum and alveolate surface.

Fig. 66. Seed of 7. amplexicaulis, dorsal surface.

Fig. 67. Seed of T. amplexicaulis, ventral surface showing linear
hilum.

The scale represents 1 mm.

1975] Tripogandra

Handlos 33

332 Rhodora [Vol. 77
Fig. 68. Seed of Tripogandra guerrerensis, dorsal surface, lower

seed of the locule.

Fig. 69. Seed
eal hilum, lower

Fig. 70. Seed
the locule.

Fig. 71. Seed
cal hilum, upper

Fig. 72. Seed

Fig. 73. Seed
hilum.

Fig. 74. Seed

Fig. 75. Seed

form hilum.

Fig. 76. Seed

Fig. 77. Seed

Fig. 78. Seed
hilum.

of T. guerrerensis, ventral surface showing ellipti-
seed of the locule.

of T. guerrerensis, dorsal surface, upper seed of

of T. guerrerensis, ventral surface showing ellipti-
seed of the locule.

of T. palmeri, dorsa! surface.

of T. palmeri, ventral surface showing linear
of T. angustifolia, dorsal surface.

of T. angustifolia, ventral surface showing puncti-

of T. grandiflora, dorsal surface.
of T. grandiflora, dorsal surface.
of T. grandiflora, ventral surface showing linear

The scale represents 1 mm.

1975] Tripogandra — Handlos 333

THE FLORA ОЕ CONCORD': A REVIEW

The greater Boston area has a long history of botanical
publications, beginning with Bigelow’s Florula Bostoniensis
of 1814. A “Flora of Middlesex County” by L. L. Dame
and F. 5. Collins was published in 1888. R. J. Eaton’s Flora
of Concord is in this tradition. Based on his own knowl-
edge, the herbarium of the New England Botanical Club and
the Gray Herbarium — which holds the most of Thoreau’s
herbarium — he has synthesized an annotated catalog which
is both ecological and historical.

The Flora of Concord has been a labor of love. It is for-
tunate that the Museum of Comparative Zoology of Harvard
University, having a Field Station in Concord, should have
undertaken the publication of so useful a book. One hopes
that this will stimulate the production of other local floras
which are, after all, the stuff on which more synoptic works
must eventually be based.

GORDON P. DE WOLF, JR.
ARNOLD ARBORETUM
JAMAICA PLAIN, MASS. 02130

!'Eaton, К. J. — A Flora of Concord; from Thoreau’s Time to the
Present Day. Cambridge, Mass.: Museum of Comparative Zoology:
1974.

334

REPORT ON ENDANGERED AND THREATENED
PLANT SPECIES OF THE UNITED STATES

The Smithsonian Institution’s Report on Endangered and
Threatened Plant Species of the United States has been
presented to Congress and published by the Government
Printing Office. Contained in the 200-page Report are lists
of endangered, threatened, commercially exploited and re-
cently extinct species of the United States (including Alaska
and Hawaii), as well as recommendations for the preserva-
tion and protection of these species, Listings of endangered
and threatened species arranged alphabetically by States
are included. A limited number of complimentary copies
of the Report are available, upon written request, from:
Endangered Flora Project, Department of Botany, Smith-
sonian Institution, Washington, D. C. 20560.

EDWARD 5. AYENSU
DEPARTMENT OF BOTANY
SMITHSONIAN INSTITUTION
WASHINGTON, D. C. 20560

335

APPEAL FOR SUPPORT FOR THE
INDEX HOLMENSIS PROJECT

The INDEX HOLMENSIS is an index of plant distribu-
tion maps with world-wide coverage. It is the only interna-
tional bibliography on distribution of vascular plants in
area and vegetation maps.

We have so far published four volumes, i.e. volume I,
covering vascular cryptogams and gymnosperms, volume II
containing Monocotyledoneae A-I, volume III Monocotyle-
doneae J-Z, and finally volume IV covering Dicotyledoneae
A-B, together more than 1000 pages. We intend to continue
to publish one volume each year. The total number of
distribution maps so far published is estimated at about
400,000, all of which will finally be listed in the index
or its supplement. Although the main work is at present
done at the Swedish Museum of Natural History in Stock-
holm, the indexing work is served by an international
editorial board. Members of this board to some extent
guarantee completeness of the file for their particular area.

Still, the number of area and vegetation maps published
annually is growing rapidly owing to the increased im-
portance that is felt for the geographic compound of plant
taxa. Consequently, not only are there wide areas all over
the world where the entire flora is mapped systematically,
but mapping has become a common feature in monographs
in different fields, as for instance economic botany, palaeo-
botany, vegetational history, palynology, and last but not
least phytocoenology.

In order to keep the file for the INDEX HOLMENSIS
and its planned supplement volumes up-to-date we here-
with ask our fellow botanists to inform us about their
published maps and/or to send reprints of their publica-
tions. Needless to say, we shall also continue to supply all
information on distribution maps so far not published in
the INDEX HOLMENSIS to colleagues on request.

АП correspondence should be addressed to: Hans Tralau,
The Swedish Museum of Natural History, S — 104 05
Stockholm 50.

Volume 77, Мо. 810, including pages 165-336, was issued Sept. 30, 1975

336

Cover illustration
Rhododendron canadense (L.) Torr.
from
Curtis’ Botanical Magazine t.474

RHODORA June, 1975 Vol. 77, No.

CONTENTS

De Plantis Toxicariis E Mundo Novo Tropicale Commentationes
XII. Notes on Biodynamic Piperaceous Plants
опата Еуста SNohulles /........ 5. eerte

Chromosome Counts of Compositae from the United States,
Mexico, and Guatemala
David J. Keil and Tod Е. Stuessy ................ PRE ИНЧИ Men

А Vegetational Survey of the Vascular Plants of the Kent
Island Group, Grand Manan, New Brunswick
ЈОНИ e MODI EE otro EUN

A Red-petioled Form of Thalia geniculata L. from Central
Florida
Allen С. Shuey .... ове 4 e

The Taxonomy of Tripogandra (Commelinaceae)
"adu D. ОНООВ iier чш pira nh sott

The Flora of Concord: A Review
Gale Драч. а ТЕ МИН illie син

Report on Endangered and Threatened Plant Species of the
United States
КОМО EON чуо.» TE АД ios uic

Appeal for Support for the Index Holmensis Project
Hans Tralau ... ОСОН ЫРА РА МАНИ ОНА

810

165

171

196

ока,

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Vol. 77 September, 1975 No. 811

= m

a x "d aX true Ma

"e " EN. Ju

~

A

The Nem Tugend ESIA PH e

Botanical Museum, Oxford Street, Cambridge, Mass. | 02138 -

Conducted and published for the Club, by

ALFRED LINN BOGLE, Editor-in-Chief
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Rbodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 77 September, 1975 No. 811

A REVISION OF THE
NORTH AMERICAN GENUS AMORPHA
(LEGUMINOSAE-PSORALEAE)

ROBERT L. WILBUR!

Amorpha is a small North American genus of papilionate
legumes belonging to the tribe Psoraleae. It has been the
subject of several taxonomic revisions (Schneider, 1907;
Rydberg, 1919; and Palmer, 1981) as well 25 numerous
regional treatments and one might suppose that such atten-
tion would have resulted in a rather satisfactory taxonomic
understanding at the present time. However, such a favor-
able supposition would be completely unjustified and my
presentation below is offered as an alternative. Although
it is the result of an intensive if somewhat sporadic study
timewise for more than fifteen years and an examination
of more than twelve thousand specimens, I am not so naive
as to believe that my account of the genus presented below
is the ultimate answer either. Naturally I hope that future
studies will show that the revision presented Fere is a
closer approximation of the biological reality than the
earlier accounts, but I would be less than candid if I were
not to admit that I have found the genus to be extremely
difficult and not especially tractable to the usual investiga-
tory procedures of a ‘‘non-experimental” taxonomist.

Grateful acknowledgment is hereby made for support given by the
National Science Foundation to this research under NSF Grants
5636 and GB-13815.

987

388 Ећодога [Vol. 77

The difficulty encountered with Amorpha as with any
taxonomically perplexing taxon is not that there is too
much variability or too little but that the variability that
seemed so discrete when a relatively small number of speci-
mens was examined becomes ever so much more like a
continuum when thousands of specimens are studied. En-
vironmentally induced plasticity of at least certain members
of the genus is sometimes spectacular, and it certainly
would prove disconcerting to those who have recognized
numerous taxa in what I refer to as the fruticosa-complex
to see the number of specimens that clearly possess the
characteristics of one “taxon” in the growth produced
early in the season and that of another “species” or “уа-
riety” in the later growth. The extraordinary morphologi-
cal diversity exhibited by certain species of Amorpha, due
both to apparently great genetical flexibility (i.e. numerous
biotypes) and environmental plasticity, has resulted in the
recognition of numerous taxa. For example, Rydberg
(1919) recognized 23 species while Palmer (1931) ac-
cepted 20 species, 11 additional varieties and 7 forms for
а total of 38 named taxa. The treatment presented here
recognizes 15 species (including one new species, A. oua-
chitensis Wilbur) and 3 additional varieties.

Any definitive statement concerning the interrelation-
ships of the genus Amorpha with its tribal neighbors in
the Psoraleae must await more complete understanding of
those genera as well as of Атотрћа itself. Morphologically
Amorpha seemingly is most closely related to the ditypic
Parryella Т. & С. ex A. Gray of Arizona and New Mexico
and to the more widespread but still western genus Eysen-
hardtia H.B.K., which ranges from central Texas west to
California and south into Guatemala. Parryella differs
principally from Amorpha in that it is without at least all
outward evidence of petals while Eysenhardtia has all 5
petals. The fruits of all three genera are 1(2)-seeded but,
although all are indehiscent, those of Parryella and Amor-
pha are far more similar in texture, shape, and all other
respects. With the facts available to us at the present time,

1975] Amorpha — Wilbur 339

future research may support the hypothesis that Amorpha,
Parryella, and Eysenhardtia were derived from a common
ancestral complex and that Amorpha and Parryella are
more closely related to one another than either is to Eysen-
hardtia.

My gratitude to the curators of the herbaria listed below
is far greater than is even usually the case in similar ac-
knowledgments for they have been extremely tolerant in
permitting me to keep loans for far longer than is custom-
ary and without their understanding the study could not
have been completed.

A, ARIZ, CAS, COLO, CS, DES, DS, DUKE, F, FLAS, FSU, GA,
GH, IA, ISC, JEPS, KANU, KSC, MICH, MINN, MO, NCU, NDA,
ND-GREENE, NEB, NSC, NY, OKL, OKLA, PHIL, POM, RSA, SDC,
SDU, SMU, TENN, TEX, UARK, UC, UMO, US, USF.

Amorpha L., Sp. Pl. 743. 1753; Gen. Pl. ed. 5. 319. 1754.

Bonfidia Necker, Elem. Bot. 3: 46. 1790. (Art. 20(2)
of the LC.B.N. clearly states that this is not to be
regarded as a generic name.)

Suffrutescent low shrubs to large bushy-topped shrubs.
Leaves odd-pinnately compound with setaceous to linear,
caducous stipules and short- to long-petiolate; leaflets (77)
11-29(45), entire to crenulate, epunctate to very densely
and conspicuously glandular punctate, petiolulate, stipel-
late on the upper side. Inflorescence a spike-like, terminal
raceme or the racemes clustered and then appearing panicu-
late; flowers pedicellate from the axil of a setaceous to
linear, caducous bract. Calyx persistent, obconic, funnel-
form or even shortly campanulate, almost imperceptibly to
strongly 5-lobed. Corolla reduced to a single petal, the
banner, and it erect, enveloping the internal floral series,
clawed, obovate to obcordate, often eventually the apical
lobes adaxially recurved, purple, blue, violet or white, entire
to erose. Stamens 10, at least basally monadelphous and
occasionally the staminal tube exserted beyond the calyx;
the filaments distinct above and exserted beyond the сајух

840 Ећодога [Vol. 77

and usually the petal; anthers versatile, small, oblong,
longitudinally dehiscent. Ovary ovoid, slightly compressed,
2-ovulate; style slender, elongate, exserted beyond the calyx
and often the petal; stigma terminal, capitate. Fruit a
1-seeded, indehiscent pod, + oblique, straight to strongly
curved, compressed, slightly to very noticeably longer than
the calyx, eglandular to conspicuously punctate glandular;
seeds ovoid to oblong, laterally compressed.

Typification: Amorpha fruticosa L.; the genus was com-
prised of only one species when originally established.

Distribution: a North American genus of about 15 spe-
cies ranging from southern Canada south to southern
Florida and west to California.

KEY TO THE SPECIES OF AMORPHA

1. Petioles and the rachises of leaves beset with spine-like

glands; plants of Arizona, California and Baja Cali-
fornia.

2. Vexillum reddish-purple, claw indistinct but about
1 mm long; filaments 6-7 mm long, united for the
basal 2-3.2 mm; fruit densely to moderately pilosu-
lose throughout (except in one area north of San
Francisco Bay); plants of Arizona and northern
California south into northern Baja California. ..
TTC 1. A. californica.

2. Vexilum white, claw 2-3 mm long; filaments 8-12
mm long, united for the basal 1-1.5 mm; fruit gla-
brous or sparingly pilosulose only near the apex;
plants known only from the Sierra San Pedro Martir
of northern Baja California. ...... 2. А. apiculata.

1. Petioles and the rachises of leaves lacking spine-like
glands.

3. Shrubs usually less than 1 m tall; leaves shortly
petiolate with the petiole typically shorter than the
width of the lowest leaflet.

1975] Amorpha — Wilbur 841

4. Midvein of the leaflets included or but barely
exserted and then its tip distinctly swollen, or if
noticeably exserted, then the tip of mucro con-
spicuously swollen; plants of the southeastern
United States.

5. Calyx-tube densely puberulent, or, if not, at
least the lobes externally sparsely puberulent ;
stem, rachises (of both leaf and inflorescence),
and leaflets densely pubescent, or, if not, usu-
ally at least somewhat puberulent; leaflets
entire or but inconspicuously crenulate; peti-
olules usually less than 1.5 mm long and
densely to sparsely pubescent; ranging from
North Carolina south into peninsular Flor-

stem, rachises (of both leaf and inflorescence),
and leaflets glabrous or but very sparsely
puberulent or short-pubescent; leaflets usually
conspicuously crenulate; petiolules usually
more than 1.5 mm long and glabrous (or
rarely very sparingly puberulent); known
only from Dade County, Florida. ..........
е а Ds ee е 4. A. crenulata.
4. Midvein of the leaflets exserted into a slender
mucro with a tapered tip; plants of the south-
eastern United States and much of the central
part of the continent.

6. Adaxial calyx-lobes 0.4-0.8(1.2) mm long,
the abaxial lobe (1)1.2-1.6(1.8) mm long;
plants of the coastal plain of the southeastern
United States (the Carolinas and Georgia).
рО ИК uu сог. 5. A. georgiana.

6. Adaxial calyx-lobes (0.6) 1-1.5 (1.8) mm long,
the abaxial lobe (1.2) 1.5-2.2 (2.5) mm long;
plants of the central United States (southern
Canada south into Texas and from Indiana
west into New Mexico and Wyoming).

342

Rhodora [Vol. 77

7. Foliage and/or calyces conspicuously pu-
bescent to the unaided eye and often ca-
nescent. .............. 6. A. canescens.

7. Foliage and calyces glabrous or nearly so,
or at least not conspicuously pubescent to
the unaided eye and never canescent.

8. Leaflets appearing epunctate or at

least the punctate glands on the lower
surface of the leaflets not discernible
without magnification; racemes usually
clustered and mostly in groups of 5-10.
Lee eee ee ee eee ae 6. A. canescens.

Leaflets conspicuously punctate and
the glands readily visible without mag-
nification; racemes solitary. .......
Lee eee eee eee eee eae 7. A. nana.

3. Shrubs usually more than (1)1.2 m tall; leaves with
petioles longer than the width of the lowermost

leaflet.

9. Calyx-lobes nearly obsolete, all less than 0.8 mm
long and never narrowly acute. .. 8. A. glabra.

9. Calyx-lobes or at least some well developed and

longer than 0.8 mm and some narrowly acute or

acuminate,

10. Calyx-lobes from about half as long to as
long as the calyx-tube, all acute or acumi-
nate; the lateral lobes more than 1 mm long.

11.

11.

Secondary venation slightly to moder-
ately elevated beneath but the leaflets
never conspicuously reticulate; abaxial
calyx-lobes (1.8) 2.5-3.5 mm Jong; plants
of the Carolinas, Georgia, and Alabama.
Lee ee eee eee teen 9. A. schwerinii.
Secondary venation conspicuously еје-
vated beneath often to the 3rd and 4th
degree of branching and thus forming a
striking reticulum; abaxial calyx-lobes

Amorpha — Wilbur 343

(1.2) 1.5-2.0 mm long; plants of Louisi-
ana and eastern Texas. ....... i
SK PHP: 10. A. paniculata.

10. Calyx-lobes or at least some of them less
than half as long as the calyx-tube, at least
the adaxial usually rounded; the lateral
lobes less than 1 mm long.

12.

Foliage, calyces and pedicels blackening
or at least darkening upon drying;
calyx-tube eglandular to sparingly
glandular with very small and incon-
spicuous, slightly elevated punctate
одак p E 11, A. nitens.

Foliage, calyces апа pedicels never

blackening and rarely even darkening

upon drying; calyx-tube usually glandu-
lar with often large, conspicuous, ele-
vated punctate glands.

13. Petiolules conspicuously pustulate-
glandular; leaflets typically emargi-
nate and often with a globose tip
to the midvein; plants of Texas or
Oklahoma and/or Arkansas.

14. Petals present on specimen.
15. Petals bright blue to deep
violet; stipules glabrous;
plants of eastern Texas
апа Oklahoma i тр,

RM Nu 12. A. laevigata.

15. Petals purplish; plants of
central Texas; stipules
pubescent; eastern Okla-
homa and/or west central

Arkansas.

16. Abaxial calyx-lobes
(0.8) 1.0-1.2(1.4) mm
long; central Texas
on the Edwards Pla-

844

Ећодога [Vol. 77

teau. ..............
.. 18. А. roemeriana.
16. Abaxial calyx-lobes
about 0.6-0.9 (1.2)
mm long; Ouachita
Mts. of southeastern
Oklahoma and west
central Arkansas.
. 14. A. ouachitensis.

14. Petals lacking on specimen.

17.

17.

Calyx-tube beset with
large conspicuous pustu-
late glands throughout
the upper 1/2-2/3 its
length, glabrous; eastern

Texas and southeastern

Oklahoma. ............

e, 12. A. laevigata.

Calyx-tube sparingly to

moderately supplied with

often inconspicuous pus-
tulate glands in the up-
per third of its length;
glabrous to moderately
spreading to appressed
short-pubescent; central

Texas or the Ouachita

Mts. of southeastern Ok-

lahoma and west central

Arkansas.

18. Abaxial calyx-lobes
(0.8) 1.0-1.2(1.4) mm
long; fruit about 6-7
mm long and 2.5-3.5
mm wide; central
Texas on the Ed-
wards Plateau.

. 18. А. roemeriana.

1975] Атогрћа — Wilbur 345

18. Abaxial calyx-lobes

0.6-0.9(1.2) mm long;

fruit about 7-9 mm

long and 3-4 mm

wide; Ouachita Mts.

of southeastern Okla-

homa and west cen-

iral gen и:

. 14. A. ouachitensis.

13. Petiolules eglandular or with only
inconspicuous glands; leaflets acute

to broadly rounded, only rarely
emarginate and the midvein only
rarely terminating in a globose or
swollen tip; plants of almost the
entire range of the genus. .......

15. A. fruticosa.

1. Amorpha californica Nutt. in Torr, & Gray, Fl. N. Am. 1:
306. 1858.

Erect, bushy shrub (0.7) 1-3(4) m tall. Current season's
growth sparingly to more typically moderately puberulent
or crinkly-pilosulose bearing few to numerous, usually con-
spicuous, often apically pointed or sharply hooked, as well
as basally swollen, resiniferous, amber-colored glands.
Leaves ascendant to spreading, (0.5) 1-1.5 (2.0) dm long.
Petioles (0.7) 1-1.5 (2.5) em long, usually equaling or longer
than the width of the lowermost leaflet, densely crinkly-
pilosulose or puberulent to glabrous with several to numer-
ous, rounded and pointed, resiniferous, pustulate glands.
Stipules caducous, oblong-lanceolate to lanceolate or linear,
eglandular, reddish-brown, sparingly to densely, appressed
tawny-pilosulose, (2)4-6 mm long. Rachis of leaf slender,
about 1 mm in diameter, densely crinkly-pilosulose or
puberulent to glabrous, moderately beset with both sharp-
pointed and rounded, amber-colored, resiniferous, sessile
glands. Leaflets (11)13-19(25), oblong to elliptic-oblong,
(0.8) 1.5-2.5 (4.2) em long, (0.6) 1-1.5 (2.0) em wide, typi-

846 Ећодога [Vol. 77

cally 1.5-2.2 times as long аз wide, opposite or more typi-
cally alternate, symmetrical, more or less broadly rounded
to subcordate basally, typically broadly rounded to obtuse
and emarginate apically, entire to inconspicuously crenu-
late; secondary veins very slightly, if at all, elevated be-
neath. Midvein usually terminating in a swollen, globose
tip flush with the margin or very rarely slightly exserted
or very rarely somewhat tapering. Lower surface of leaf-
lets moderately pilosulose throughout but densely so along
the midvein to glabrous or very nearly so, conspicuously
glandular-punctate below with light to dark brown, resin-
ous glands of apparently 2 sizes; upper surface typically
moderately pilosulose but occasionally glabrous, eglandu-
lar. Petiolule 0.7-1.5(1.8) mm long, densely spreading
pilosulose to glabrous, sparingly to moderately pustulate-
glandular, Racemes erect, usually solitary but rarely few
together, (0.5)1-1.8(2.5) dm long; rachis of inflorescence
moderately spreading-pilosulose, hirsutulous or puberulent
to glabrous, sparingly glandular pustulate with both
rounded and sharp-pointed glands. Pedicels 0.3-1.2 mm
long, densely spreading- to appressed-pilosulose hirsutulous
or puberulent to glabrous, eglandular; bracts lanceolate to
narrowly linear, 1.5-4 mm long, externally moderately
spreading- to appressed-pilosulose, eglandular to moder-
ately pustulate-glandular, internally glabrous, caducous.
Calyx-tube narrowly cylindric or somewhat funnelform,
2.0-2.8 mm long, moderately spreading- to appressed-pilosu-
lose or puberulent throughout to glabrous, abundantly to
sparingly pustulate-glandular in upper half. Calyx-lobes
usually tipped by a conspicuous pustulate gland and also
frequently sparingly to moderately glandular-pustulate,
densely to moderately pilosulose or glabrous both externally
and internally. Adaxial calyx-lobes broadly to narrowly
triangular-dentate, acute, (0.4) 0.8-1.5(2) mm long; lateral
calyx-lobes slightly longer, acute to acuminate, (0.5) 1-2
(2.2) mm long; abaxial lobe narrowly lanceolate, acumi-
nate, (0.8)1.5-2.5 mm long. Vexillum 5-7 mm long, about
2.5-4 mm wide, broadly obovate, gradually tapering to the

1975] Amorpha — Wilbur 347

short, indistinct claw about 1 mm long, moderately to
strongly arching, apically more or less truncate to emargi-
nate and often with a central apiculation, reddish-purple
and with the claw and blade both strongly enveloping the
filaments. Filaments 6-7 mm long, united into a sheath for
the basal 2.0-3.2 mm, glabrous; anthers about 0.5-0.7 mm
long, pale yellow. Style moderately ascending-pilose; ovary
either densely pilosulose or very sparingly so and then only
apically. Fruit 6-8 mm long, 2.5-3.5 mm wide, broadest
above the middle, tapering somewhat basally, with the
adaxial margin straight or slightly outwardly outcurved
above and the abaxial strongly outwardly bowed, densely
to moderately pilosulose throughout or only sparingly
strigillose apically, pustulate-glandular in the upper half
or two-thirds. Seeds smooth, plump, 2.5-3.2 mm long, light-
to olive-brown.

Distribution: Northern California (approximately 41°
N) south into the Sierra San Pedro Martir of northern
Baja California (approximately 31° N) and sporadically
eastward as far as southeastern Arizona.

Two rather strikingly distinct varieties have been recog-
nized within this species. Their distribution as shown by
our rather conventional but crude mapping does not sug-
gest that they are strictly allopatric but a more refined
appraisal of their relationship should wait at least careful
field observations. The two may be separated by the fol-
lowing key.

A) Upper surface of leaflets, rachises and petiolules moder-
ately to densely spreading-pilosulose or puberulent;
calyx-tube moderately spreading-pilosulose or occasion-
ally appressed-pilosulose; calyx-lobes all, or at least the
longest, over 1 mm long. ........ la. var. californica.

A) Upper surface of leaflets, rachises and petiolules gla-
brous or very nearly so; pubescence on lower surface
of leaflet lacking or largely restricted to the midvein;
calyx-tube glabrous or nearly so or sparingly to rarely

348 Ећодога [Vol. 77

moderately strigillose; calyx-lobes all 1 mm long or less.
1b. var. napensis.

e 9 е ө ө ө 9 ө 9 ө ө ө ө 9 ө ө э ө ө ө ] 2 е э ө ө ө ө ө э ө ө ө

la. A. californica var. californica
Amorpha californica Nutt. in Torr. & Gray, Fl. N. Am.
1: 306. 1838. TYPIFICATION: *St. Barbara, Califor-
nia, near the coast," Nuttall s.n. (holotype(?), му !).
A. hispidula Greene, Fl. Fran. 14. 1891. TYPIFICA-
TION: California: MONTEREY CO., Hickman s.n.
(lectotype, ND-G!).
A. californica var. hispidula (Greene) Palmer, Jour.
Arnold Arb. 12: 163. 1931.

Branchlets, rachises, petioles and petiolules moderately
spreading pilosulose or puberulent; leaflets spreading- to
appressed-pilosulose above and below. Calyx-tube usually
moderately to densely spreading- or appressed-pilosulose or
puberulent, eglandular or sparingly and inconspicuously
pustulate-glandular. Calyx-lobes usually more than 1 mm
long. Pod moderately to densely more or less appressed,
short-pubescent.

Distribution: Coastal Ranges north of San Francisco to
about 38°30’ N and along the Coastal Ranges south of San
Francisco from Monterey Co. (about 36°30’ N) into north-
ern Baja California in the Sierra San Pedro Martir (about
30°30’ N); also in an area south of Mt. Shasta (about
41° N) and on the western slopes of the Sierra Nevada
(about 39° N); disjunctly occurring in central and south-
eastern Arizona, (Map 2.)

1b. A. californica var. napensis Jepson, Man. Fl. PI. Calif.
556. 1925. TYPIFICATION: California: Howell МЕ,
Napa Range, Jepson 6835 (lectotype, JEPS!). The
lectotype, designated by Jepson (ЕІ. Calif. 2: 331.
1936), is a sterile specimen and apparently a crown
sprout while his number 6834 collected on the same
day and the same place, is the fruiting specimen from
which most of the diagnostic information was ob-
tained.

1975] Amorpha — Wilbur 349

MAPS 1-2. Map 1. Amorpha californica var. napensis (dots
north of San Francisco Bay) and A. apiculata (dots with horizontal
line in Baja California). Map 2. A. californica var. californica.

Branchlets, rachises, petioles and petiolules very spar-
ingly pilosulose or more typically glabrous; leaflets gla-
brous above and either glabrous beneath or the pubescence
primarily restricted to the midvein. Calyx-tube glabrous
or sparingly strigillose above or rarely moderately strigil-
lose throughout. Calyx-lobes usually all 1 mm long or less
and ranging from 0.4-1.0 mm long. Fruit sparingly to
moderately strigillose apically or throughout.

Distribution: endemic to the area north of San Francisco
Bay in Marin, Napa and Sonoma Counties. (Map 1, in
part.)

Both recent California floras (Munz, 1959, and Abrams,
1944) indicate the species is divisible into two varieties: a
northern one ranging from the Shasta area of southern
California south to Monterey and a southern variety rang-
ing from the Santa Lucia Mountains just south of Mon-
terey south to the Santa Ana and Santa Rosa Mountains

850 Ећодога [Vol. 77

of extreme Southern California. The range of the varieties
presented by these authors is in obvious conflict with those
found in this study although the morphologica] features
characterizing the taxa in all three studies seem to be
identical.

2. Amorpha apiculata Wiggins, Contr. Dudley Herb. 1: 171.
1933. TYPIFICATION: Mexico: banks of the Rio Santo
Domingo about 5 miles above Mission Santo Domingo,
Baja California, Wiggins & Demaree 4776 (holotype,
DS, not seen; isotypes, ARIZ!, DS!; F!; NY!, РОМ !, RSA!).

Erect, slender shrub 2-5 m tall. Current season’s growth
moderately to sparsely spreading-pilosulose bearing few to
numerous, rounded, resiniferous, amber-colored, elliptic
glands and a few, apically pointed, basally rounded, spine-
like glands. Leaves ascendant to spreading, 1-2 dm long.
Petioles 1-2.5 cm long, usually equaling or longer than the
width of the lowermost leaflet, sparingly to moderately
spreading-pilosulose, with several to numerous, usually
rounded but occasionally pointed, amber-colored glands.
Stipules caducous to tardily deciduous, linear to linear-
setaceous, glandular, sparingly pilosulose, 2-3 mm long.
Rachis of leaf slender, about 1 mm in diameter, sparingly
to moderately pilosulose, moderately beset with rounded or
pointed, amber-colored, resiniferous, sessile glands. Leaf-
lets (9)13-19(25), elliptic to oblong-elliptic, mostly 1.5-
3.0(3.8) ст long апа (0.5)1.0-1.5 em wide, typically 2.2-
2.9 times as long as wide, usually alternate and symmetri-
cal, tapering either both basally and apically and often
acute to more or less rounded; secondary veins very slightly
elevated bencath. Midvein exserted, 0.2-0.8 mm long,
slightly to conspicuously swollen at the tip. Lower surface
sparsely pilosulose throughout or more or less restricted to
the midvein where tending to be wide-spreading, "light
green, slightly glaucous," conspicuously glandular-punctate
below with dark, resinous glands of apparently 2 sizes;
upper surface glabrous, “bright green," moderately to
sparingly punctate-glandular. Petiolules 1.5-2.2 mm long,

1975] Amorpha — Wilbur 351

sparingly to moderately spreading to somewhat appressed
pilosulose, moderately pustulate-glandular. Racemes soli-
tary to several, 1-7 in number, 1-3 dm long, rachis mod-
erately spreading-pilosulose; pedicels 0.8-1.2 mm long,
moderately spreading to appressed pilosulose; bracts nar-
rowly linear, 2.5-3.0 mm long, sparingly pilosulose, pustu-
late-glandular, caducous. Calyx-tube narrowly funnelform,
2.5-3.0 mm long, moderately spreading-pilosulose through-
out, conspicuously pustulate-glandular in upper fifth. Calyx-
lobes usually tipped by a resinous, pustulate gland and
otherwise eglandular, moderately to sparingly pilosulose
or puberulent both externally and internally. Adaxial caly-
cine lobe triangular-dentate, acute to less frequently acumi-
nate, 0.5-1 mm long; lateral lobes longer and more nar-
rowly lanceolate, acuminate, 0.8-1 mm long; abaxial lobe
narrowly lanceolate, acuminate, 1.2-2 mm long. Vexillum
5-7 mm long, about 4 mm wide, broadly obovate with a
slender claw and a broad, truncate apical notch with a
central, triangular-dentate apiculation about 0.1-0.2 mm
long, white; the claw enveloping the filaments and the
blade either plane or reflexed away from the filaments.
Filaments greatly elongate, 8-12 mm long, united for the
basal 1-1.5 mm, glabrous; anthers about 0.5 mm long, ap-
parently pale yellow. Style moderately ascending-pilose;
ovary glabrous except for the uppermost pilosulose region
near the style. Fruit (apparently still immature) about
6 mm long and 2.5 mm wide, broadest above the middle,
tapering to base, with the adaxial margin very nearly
straight and the abaxial margin gradually outwardly
bowed, glabrous or sparingly pilosulose only near the top,
conspicuously glandular-pustulate.

Distribution: Known only from the Sierra San Pedro
Martir of northern Baja California, Mexico at approxi-
mately 31° N. (Map 1, in part.)

This species is clearly very closely related to Amorpha
californica and especially to that species’s var. californica
which apparently also reaches its southern limit in the

852, Ећодога [Vol. 77

Sierra San Pedro Martir. I have examined only eleven
different collections of either species from this area. These
two species have been confused both in the field and the
herbaria by collectors and identifiers and at the present
time one cannot form any realistic picture of their ecologic
or spatial arrangement to one another. It does not seem
that the two are strictly altitudinally separated from one
another but if anything A. californica tends to occupy the
higher elevations.

The two species can seemingly be readily distinguished
from each other vegetatively and either in flower or fruit.
Additional collections and field observations from this re-
gion are certainly much to be desired.

Although A. apiculata seems in some respects to be less
strikingly different from A. californica var. californica
than is A. californica var. napensis, nothing would seem to
be gained by attempting to surmise now the results of the
needed research. The three taxa form a most distinctive
complex within the genus.

3. Amorpha herbacea Walt., Fl. Car. 179. 1788.

A shrub mostly (0.3)0.6-1.2(1.5) m high arising from a
thick, woody, often horizontal rootstock reportedly up to
2 m in length and 2 cm in diameter. Current season's
growth dull reddish-purple, olivaceous to light brown,
slender, about (1)2-4 mm in diameter, with numerous fine
longitudinal grooves and ridges and sprinkled throughout
with few to numerous, usually inconspicuous, small punc-
tate glands, mostly densely short-pubescent or puberulent
and then often appearing canescent but occasionally (espe-
cially in Florida) only very sparingly pubescent. Leaves
numerous, spreading to ascendant, mostly about (0.6) 0.8-
1.8(2.4) dm long. Petioles mostly densely short-pubescent,
puberulent or strigillose but occasionally glabrous or nearly
so and with few to numerous postulate glands especially
abundant near the base, typically shorter than the width
of the lowest leaflet but occasionally as long as 1.5 times
the width, about (0.5)1-10(13) mm long. Stipules incon-

1975 Amorpha — Wilbur 355
spicuous, pigmented like the stem and rachis, glabrous to
sparingly pubescent, caducous or persisting as a withered
remnant, appressed, acicular to setaceous, about (1) 1.2-
2.5(3) mm long. Rachis of leaf slender, about 0.5-1 mm in
diameter, usually densely short-pubescent or puberulent
throughout (or if differentially so, then more densely
pubescent above in and about the channeled groove) to less
commonly sparsely short-pilose, strigillose or even glabrous
or nearly so, sparingly glandular throughout. Leaflets (15)
23-45(63) in number, about (0.7) 1.0-2.5 (8.2) cm long and
(3)4-10(15) mm wide, mostly (1.2)2-3(4) times as long as
broad, usually symmetrical but rarely asymmetrical, oppo-
site or alternate, the interval between petiolules on the
same side of the rachis about (0.3) 0.6-1.2 (1.8) ст long.
Blades of leaflets mostly broadly to narrowly oblong or
occasionally oblong-elliptic to elliptic, or rarely ovate-ob-
long, ovate, obovate, obcordate or almost orbicular with the
base usually broadly rounded to obtuse, occasionally sub-
cordate, truncate or nearly so, oblique or rarely tapering
and then more or less acute; the apex obtuse to broadly
rounded to almost truncate and occasionally shallowly
emarginate, with the midvein terminating in a sessile or
subsessile, distinctly swollen, often globose, glandular knob,
or, when exserted, the swollen tip on a stalk about 0.2-
0.5(0.8) mm long. Texture of leaflets coriaceous to sub-
coriaceous when dry, finely reticulate above from the very
slightly elevated venation excepting the scarcely depressed
midvein; the margin usually slightly to conspicuously revo-
lute and very inconspicuously crenulate to entire, or nearly
so. Midvein prominently elevated beneath and the second-
ary veins but slightly raised. Both surfaces of the leaflets
usually very densely pubescent, but rarely the upper or
even both sides glabrous or very nearly so, and when most
nearly glabrous, then often strigillose beneath along the
principal veins; pubescence of the upper surface usually
short-pilose, puberulent, or even strigillose and that of the
lower surface usually short-villous, pilose, puberulent, or
rarely strigillose; punctate glands on the lower surface

854 Rhodora [Vol. 77

usually conspicuous and readily apparent to the unaided
eye, appearing to be of approximately two size classes,
usually numerous but occasionally few, ranging in number
from about (40) 75-225 (425) glands per half-leaflet. Petio-
lule about (0.7)1-2 mm long, usually densely puberulent,
occasionally strigillose, less commonly sparsely puberulent
or strigillose and rarely glabrous, with few to numerous
pustulate glands. Stipels acicular to setaceous, mostly (0.4)
0.8-2 mm long, rarely tardily deciduous, glabrous to pu-
berulent, especially near the base. Racemes usually few to
several or occasionally numerous, appearing terminally
clustered or occasionally solitary, about (1)4-12(20) in
number, mostly appearing slender and elongate, about
(0.3) 1-1.8(4) dm long and (0.8) 1.2-1.8(2) ст in diameter,
sessile, subsessile, or on a peduncle 1-4(7) em long and
with a usually densely but occasionally sparsely puberulent,
conspicuously ridged and grooved, sparingly pustulate
glandular rachis bearing numerous, usually densely clus-
tered, mostly densely short-pubescent to puberulent or oc-
casionally glabrous pedicels about (0.2) 0.4-1.2(1.8) mm
long, subtended by a caducous, sparsely glandular-punctate,
usually sparingly puberulent to short-pubescent, brownish
to amber colored, linear-subulate to setaceous bract about
(1.2) 1.8-2.5(3) mm long. Calyx-tube turbinate to nar-
rowly campanulate or cylindrical, mostly about 1.5-2.5(3)
mm high with the abaxial side slightly longer than the
adaxial, usually either very densely puberulent or short-
pilose throughout, and with the pubescence curly and
spreading or, rarely, densely to sparingly minutely strigil-
lose throughout, or the strigillose pubescence restricted to
the upper third or half, or (occasionally in Florida) the
tube completely glabrous or the lower half or two-thirds
of the tube glabrous and the upper portion variously pubes-
cent, bearing few to numerous conspicuous pustular glands
scattered throughout the upper one-third to two-thirds of
the tube. Calyx-lobes externally usually densely puberulent
or short-pubescent and hence the margins not appearing
strikingly ciliate or occasionally (especially in the more

1975] Amorpha — Wilbur 355

glabrous Floridian forms) merely sparsely puberulent and
then the margins appearing densely short-ciliate, punctate-
glandular, internally the lobes and occasionally the very
uppermost portion of the tube densely puberulent to short-
pubescent (the tube otherwise glabrous within). Adaxial
calyx-lobes usually triangular-dentate and often narrowly
so, mostly acute to acuminate but occasionally (most com-
monly in the Floridian variety with a glabrous calyx-tube)
oblong and then obtuse, about (0.4)0.5-1(1.2) mm long ;
lateral calyx-lobes broadly to narrowly triangular, mostly
longer than the adaxial lobes but shorter than the abaxial,
acute to acuminate and mostly (0.6)0.8-1.2(1.5) mm long;
the abaxial lobe usually narrowly triangular, acute to
aeuminate and mostly (0.8)1-1.5(2) mm long. Vexillum
about (4)5-6(7) mm long and (2) 2.5-3.5 mm wide, broadly
obeordate and tapering into a narrow claw, strongly arch-
ing and incurved laterally, and thus enveloping the inner
floral parts with an entire to finely erose margin, blue-
violet, purplish-violet to white. Filaments about 6-8 mm
long, glabrous. Anthers about (0.3)0.4-0.6(0.8) mm long,
yellow to yellowish orange. Pistil about 6 mm long with a
sparsely ciliate or puberulent to glabrous ovary about 1
mm high, and broadest at or about the middle and tapering
to either end and with a densely antrorsely pubescent style
about 5 mm long terminated by a small, truncate, stigmatic
tip. Fruit about 4-6 mm long and about 1.8-2.5 mm wide,
broadest at or near the middle and tapering to the 1-2 mm
stipe-like base, conspicuously pustular-punctate in the up-
per half or two-thirds, densely short-pubescent to glabrous
or nearly so, exceeding the calyx-tube by about 2.5-4 mm,
laterally compressed, obliquely obovate to very slightly,
crescent-shaped, the adaxial side straight to slightly arched
concavely away from the rachis, the abaxial side strongly
outwardly bowed, terminated by the often 0.5-0.5 mm per-
sistent base of the style.

Distribution: Dry, open woods, sandhills, or savannahs,
typically of the Coastal Plain but occasionally in the Pied-
mont or mountains from North Carolina south into Florida.

856 Ећодога [Vol. 77

This species, the most widespread of the southeastern
dwarf taxa is not morphologically uniform throughout its
extensive area. Its variability is most pronounced in
Florida.

A striking extreme within the species as interpreted here
is that element which was formerly designated A. floridana.
My earlier survey of the southeastern dwarf species
(Rhodora 56: 261-265. 1954) indicated that A. floridana
represented a distinct species endemic to the west coast of
Florida from about the Apalachicola region south at least
to Manatee and Highlands counties. The suggested differ-
ence in curvature of the dorsal suture of the fruit employed
by Small (Man. SE. Flora 688. 1933.) to distinguish the
two taxa is not tenable. The dorsal suture of most fruiting
specimens of the floridana type is straight, or very nearly
so, just as in А. herbacea. Only rarely is it as curved as
shown by Palmer (Jour. Arn. Arb. 12: 194. f. 5. 1931).
Examination of many more specimens of A. herbacea s.s.
and of A. floridana has convinced me that the difference
between these two taxa is merely one of degree of pubes-
cence. However, there are also a small number of speci-
mens which seem at least partially intermediate between
these extremes. Macroscopically they appear much closer
to extreme A. floridana but microscopically the calyx-tubes
are pubescent to a varying degree. Some of these tubes
are sparsely puberulent or short-villous throughout, while
others are sparingly to densely strigillose throughout.
These intermediates might indicate introgression from A.
herbacea, since they are often found in areas in which that
taxon is known and are often collected with it. Except for
the apparently isolated area of the glabrous extreme about
Apalachicola in Franklin County, the range of the so-called
A. floridana coincides with part of that of A. herbacea in
western peninsular Florida. I tentatively have distin-
guished these two less than ideally separable taxa by the
following key.

1975] Amorpha — Wilbur 357

KEY TO THE VARIETIES OF A. HERBACEA

Upper portion of the plant including the leaves usually
conspicuously pubescent; calyx-tube densely puberulent
to short-pilose; fruit densely to sparsely puberulent or
rarely glabrate. ........ 3a. А. herbacea var. herbacea.

Upper portion of the plant and the leaves appearing gla-
brous or nearly so, or at least not densely pubescent;
calyx-tube glabrous, sparingly puberulent or sparsely to
densely minutely strigillose; fruit glabrous. ......-..-.
Куз por ura e dea arn m 3b. А. ћетђасеа var. floridana.

3a. A. herbacea var. herbacea. TYPIFICATION : Doubtfully

present in Walter’s Herbarium (BM) as Dr. Carroll
E. Wood, Jr. did not see a photograph of a specimen
of this distinctive species in the GH photocopy of
Walter’s Herbarium. A neotype should therefore per-
haps be designated.

A. pubescens Willd., Berlin Baumz. 17. 1796.

A. pumila Michx., Fl. Bor. Am. 2: 64. 1805.

(nom. illegit., Intern. Code Art. 63). (TYPIFICATION :
holotype, P, not seen; phototype, GH 1)

А. cyanostachya М. А. Curtis, Boston Jour. Nat. Hist.
1: 140. 1835. TYPIFICATION: North Carolina: Wil-
mington, М. A. Curtis s.n. (lectotype, NY!).

A. herbacea var. а typica Schneider, Bot. Gaz. 43: 299.
1907.

A. herbacea var. В Boyntoni Schneider, Bot. Gaz. 43:
300. 1907. TYPIFICATION: Florida: PASCO CO., pine
barrens, Richland, Curtiss 6664 (lectotype, MO!;
isolectotypes, CU!, GA!, GH!, ISC!, MIN], NEB!, NY!,
PHIL!, UC!, US!)

Usually at least the upper portion of the stem densely
short-pubescent or puberulent, but occasionally only
sparsely pubescent and rarely almost glabrous. Rachis of
leaf usually densely short-pubescent or puberulent through-

358 Rhodora [Vol. 77

out or rarely sparsely short-pilose, puberulent or strigillose.
Rachis of the inflorescence usually densely short-pubescent
or puberulent. Calyx-tube very densely puberulent or
short-pilose throughout, and with the pubescence curling
and spreading and not at all strigillose. Fruit densely to
sparsely puberulent or short-pubescent throughout or in
part glabrous.

Typification: Probably not in Walter's Herbarium (BM).
Supposedly from the area about Walter's plantation in
present day Berkeley County, South Carolina, where it is
common.

Distribution: Dry, open woods, sandhills, or savannahs,
typically of the Coastal Plain but occasionally in the Pied-
mont and mountains from North Carolina south into pen-
insular Florida. (Map 3.)

Schneider did not treat his two proposed varieties as
distinct species since they were "apparently connected . . .
by some intermediate forms." The extremes in leaflet size
and number are striking but the transition from one
morphological type to the other is complete. The more
northern specimens are often separable from those from
the southern portion of the range, but exceptions are fre-
quent.

3b. A. herbacea var, floridana (Rydb.) Wilbur, Jour. Elisha
Mitchell Sci. Soc. 80: 55. 1964.

A. floridana Rydb., N. Am. F]. 24: 31. 1919. TYPI-
FICATION: Florida: [no other locality given],
Chapman s.n. (МҮ!).

Upper portion of the stem glabrous or nearly so to
sparsely short-pilose or puberulent, but occasionally notice-
ably puberulent or strigillose. Rachis glabrous or nearly
so to more or less densely puberulent or short-pubescent,
especially above. Rachis of the inflorescence sparsely to

1975] Amorpha — Wilbur 359

MAPS 3-5. Map 3. Amorpha herbacea var. herbacea. Map 4.
A. herbacea var. floridana. Map 5. A. georgiana var. georgiana (dots
in the Carolinas and Georgia), A. georgiana var. confusa (triangles)
and A. erenulata (dots with vertical and horizontal lines in southern
Florida).

densely puberulent. Calyx-tube glabrous or nearly so to
sparsely short-puberulent or minutely strigillose through-
out, and the pubescence, when present, often restricted to
the upper portion of the tube or at least sparsely dis-
tributed below, or, if more or less equally dispersed and
dense, then the pubescence strigillose.

360 Ећодога [Vol. 77

Distribution: Known definitely only from western Flor-
ida from Franklin County to Manatee County in dry pine
lands and fields, sandy roadsides. (Specimens of this spe-
cies collected by Leavenworth [GH, PH], and others [NY]
are labeled “East Florida.") (Map 4.)

4. Amorpha crenulata Rydb., N. Am. Fl. 24: 30, 1919.
TYPIFICATION: Florida: DADE CO., in hummocks, be-
tween Coconut Grove and Cutler, Small & Wilson
1898 (holotype, NY!; a presumed isotype, Е!).

A low, apparently rhizomatose shrub about 0.4-1.0(1.5)
m high. Current season's growth usually dull reddish-
purple or more rarely olivaceous, usually slender, 1-2(3)
mm in diameter and but sparsely puberulent to glabrous,
or very nearly so, with numerous fine longitudinal] grooves
and ridges and sprinkled throughout with few to numerous
small glandular blistered areas; older branches gray to
light brownish and less conspicuously lined. Leaves numer-
ous, spreading to ascendant, (0.8)1.5-2.5(3) dm long.
Petioles very sparingly puberulent to glabrous and usually
longer than the breadth of the lowermost leaflet, but oc-
casionally equaling or exceeded by the breadth of the lower-
most leaflet, sparingly glandular throughout and densely
so near the base, (0.3)0.8-1.5(1.8) em long. Stipules in-
conspicuous and pigmented like the stem and rachis, gla-
brous, usually caducous, appressed, setaceous to acicular,
1.2-2.2 mm long. Rachis slender, about 0.5-1 mm in diam-
eter, glabrous to sparsely puberulent, usually sparingly
glandular, channeled above. Leaflets (19)23-33(41) in
number, (0.7) 1.2-2.5(4.2) cm long, (2.5)5-9(11) mm wide,
mostly (1.2)2-4(5) times as long as broad, remote, usually
well separated from the nearest leaflet on the same side of
the rachis, the interval between those petiolules about (5)
7-17(20) mm long. Blades of leaflets mostly narrowly to
broadly oblong or elliptic but varying from almost orbicular
to ovate-oblong or obovate, with the base commonly broadly
rounded to almost truncate or gradually tapering and

1975] Amorpha — Wilbur 361

sometimes acute; the apex broadly rounded to almost trun-
cate and commonly emarginate or occasionally somewhat
acute, with the midvein terminating in a sessile or sub-
sessile globose glandular knob or exserted 0.2-0.5 mm as a
gland-tipped mucro. Texture of leaflets coriaceous to sub-
coriaceous when dry, finely reticulate above from the
slightly elevated venation although the midvein depressed,
the margin flat or more typically slightly to conspicuously
revolute, usually very noticeably crenate or crenulate.
Midvein prominently elevated beneath, the secondary veins
but slightly raised. Both surfaces of the leaflets glabrous
or rarely very sparsely pubescent beneath along the prin-
cipal veins; punctate glands on the lower surface usually
conspicuous and readily apparent to the unaided eye, ap-
pearing roughly of two sizes, varying from dense and very
numerous to relatively sparsely scattered and few, ranging
mostly between 40-180 punctate glands per half-leaflet.
Petiolule glabrous or rarely sparsely pubescent, pustulate-
glandular, (1)1.5-2(3) mm long. Stipels acicular, 0.4-1.5
mm long, glabrous, often not long-persisting and the lower
often not developing. Racemes solitary or very commonly
few to several closely associated, slender and elongate,
mostly (0.5)1-2.5(3) dm long and about 0.7-1.8 cm in
diameter, sessile or nearly so, or on a peduncle about 1-2
(4) em long, with a conspicuously ridged and grooved gla-
brous to very sparingly puberulent rachis bearing numer-
ous densely clustered to more loosely arranged glabrous to
very sparsely puberulent pedicels (0.5) 1-1.8 (2.2) mm long,
subtended by a caducous, gland-dotted, brownish to amber-
colored, sparsely puberulent, linear-setaceous or narrowly
spatulate bract nearly 1.2-2.5 mm long. Calyx-tube turbi-
nate to narrowly campanulate, mostly (2)2.2-3.2 (3.5) mm
high with the abaxial side slightly longer than the adaxial,
glabrous or very nearly so throughout, with the few to
numerous punctate glands mostly restricted to the upper
half of the tube. Calyx-lobes punctate-glandular, glabrous
or very nearly so externally with a densely short-ciliate
margin; internally the lobes and uppermost portion of the

362 Ећодога [Vol. 77

tube densely matted-puberulent, the lower portion of the
tube glabrous internally. Adaxial lobes triangular-dentate
and acute to acuminate to oblong and obtusely rounded and
about 0.5-1 mm long; lateral lobes broadly to narrowly
triangular and acute to acuminate and mostly about 0.8-
1.2 mm long; abaxial lobe usually narrowly triangular and
acute or more commonly acuminate, mostly (1)1.2-1.8(2)
mm long. Vexillum about (4.5)5-6(7) mm long and 3-4.5
mm wide, very broadly obcordate and tapering abruptly to
a slender claw, strongly arched and incurved and envelop-
ing the inner floral parts, the margin irregularly finely
erose to almost entire, white to violet-blue. Filaments about
6-9 mm long, glabrous. Anthers about 0.5-0.7 mm long,
yellowish. Pistil about 6 mm long with a glabrous ovary
about 1 mm high, an antrorsely pubescent style about 5 mm
long and a small, capitate, terminal stigma. Fruit about
4-6.5 mm long and 1.8-2.5 mm wide with the base tapered
below into a stipe about 1-2 mm long, glabrous, conspicu-
ously punctate-glandular in the upper two-thirds, exceed-
ing the calyx-tube by about 2.5-4 mm, laterally compressed,
obliquely obovate, the adaxial side straight or nearly so,
the abaxial side strongly outwardly bowed, terminated by
the often 0.5 mm long, persistent base of the style.

Distribution: Apparently restricted to Dade County,
Florida. (Map 5, in part.)

This appears to be a weakly differentiated species en-
demic to the southern tip of Florida. It seems very closely
related to A. herbacea and particularly to the more glabrate
variant of that species, var. floridana.

5. Amorpha georgiana Wilbur, Rhodora 56: 261. 1954.
[10 Jan. 1955]

A low shrub 0.8-1 m high. Current season's growth
olivaceous to dull reddish purple, about 1-3 mm in diameter,
very sparsely puberulent or more commonly glabrous with
numerous fine longitudinal grooves and ridges and sparsely
sprinkled with small punctate glands; the older growth

1975] Amorpha — Wilbur 363

glabrous, dull reddish purple or brownish and less notice-
ably ridged and occasionally, especially in the southern por-
tion of its range, persisting for one or perhaps more years
and then darker and often grayish or black. Leaves numer-
ous, spreading to ascendant, mostly (3)6-15 (18) em long.
Petioles glabrous or sparsely to moderately puberulent or
short-pubescent, ranging in length from practically absent
to two or more times longer than the breadth of the lower-
most leaflet, varying from 1-15(20) mm long, sparingly
glandular to densely so especially at or near the base.
Stipules inconspicuous, glabrous, caducous to semipersist-
ent, appressed, linear-subulate or more commonly setaceous,
mostly about (1)1.5-2(2.5) mm long. Rachis slender, about
0.5-1 mm in diameter, glabrous to sparingly strigillose,
puberulent or short-pubescent below and the channeled
groove similar or rather densely short-puberulent, spar-
ingly glandular throughout. Leaflets varying from about
(11) 15-43 (47) in number, (0.3) 0.6-2.5 (3.2) ст long and
(0.2) 0.6-1.2(1.6) em wide, usually about (1)1.5-2.5(9)
times as long as wide, opposite or nearly so to conspicu-
ously alternate, approximate and often imbricately over-
lapping with an interval between petiolules of (0.2) 0.5-
0.5(0.6) em to rather widely separated with the interval
between petiolules on the same side of the rachis about
(0.5) 0.8-1.4(1.8) cm long. Blades of the leaflets mostly
broadly to narrowly oblong or occasionally oblong-elliptic
to elliptic and rarely ovate-oblong, or the terminal some-
times obovate, obcordate or even nearly orbicular, with
the base broadly rounded or almost truncate to subcordate
and the apex usually obtuse to broadly rounded and com-
monly slightly emarginate with the midvein exserted as a
slender mucro about (0.2)0.4-1 mm long and tapering
gradually to the not at all or rarely very indistinctly swol-
len, globose or knob-like tip. Texture of leaflets coriaceous
to subcoriaceous when dried; inconspicuously reticulate
above from the scarcely elevated venation except for the
sometimes but slightly depressed midvein; the margin
usually slightly to conspicuously revolute although occa-

364 Ећодога [Vol. 77

sionally flat or nearly so; entire or inconspicuously crenu-
late and with the midvein prominently and the secondary
veins slightly elevated beneath; at maturity both surfaces
glabrous or nearly so, or with the lower surfaces sparsely
to moderately strigillose, especially along the principal
veins, or rarely moderately spreading, short-pubescent, the
punctate glands on the lower surface usually inconspicuous
but still noticeable to the unaided eye and of uniform size,
or at least not conspicuously of two size-classes, mostly
about (30)50-120(160) glands per half-leaflet. Petiolule
glabrous or sparsely to densely strigillose, puberulent or
pilosulose with few to numerous conspicuous to very in-
conspicuous pustulate glands, 0.7-1.8(2) mm long. Stipels
acicular or setaceous, glabrous, usually persistent, mostly
0.8-1.8 mm long. Racemes usually with densely clustered
flowers or less commonly with the flowers more loosely ar-
ranged, usually appearing terminally clustered and arising
from the naked upper portion of the stem or from the axils
of the upper leaves or occasionally solitary, varying in
number from (1)4-11(15), mostly (0.2)0.5-2.0(3.0) dm
long and about (1.0)1.2-1.6 ст in diameter, sessile or
nearly so or on a peduncle up to about 8 cm long, the rachis
conspicuously ridged and grooved, glabrous or nearly so
to sparsely strigillose or puberulent or less commonly
densely puberulent, sparingly glandular, bearing numerous,
glabrous to sparingly strigillose or puberulent pedicels
about 0.4-1 mm long subtended by a caducous, sparingly if
at all glandular-punctate, sparsely short-pubescent and
often merely ciliate, brownish to amber-colored, linear to
linear-subulate bract mostly 1.5-2.5(8) mm long. Calyx-
tube turbinate, narrowly campanulate or cylindrical, about
1.7-2.2 mm high with the abaxial side slightly longer than
the adaxial, glabrous or rarely inconspicuously and very
sparingly pilosulose, bearing few to numerous more or less
inconspicuous pustular glands on the upper third to two-
thirds of the tube, Calyx-lobes externally glabrous to
puberulent (if the latter, usually sparingly so), and usually

1975 Amorpha — Wilbur 365

punctate glandular and with densely white-ciliate margins;
internally the lobes and occasionally even the uppermost
portion of the tube densely puberulent to short-pubescent
(with the tube otherwise glabrous within). Adaxial calyx-
lobes either triangular-dentate and acute to acuminate or
oblong, and then obtuse to rounded, mostly 0.4-0.8 (1.3) mm
long; lateral lobes usually longer than the adaxial] lobes
and shorter than the abaxial, usually triangular-dentate
and acute to acuminate but occasionally oblong and obtuse
to rounded, mostly (0.6) 0.8-1.2 (1.5) mm long; abaxial lobe
usually narrowly triangular to lance-subulate, acute or
more typically acuminate, mostly (1.0) 1.2-1.6(1.8) mm
long. Vexillum about (4)5-6 mm long and 3-3.5(4) mm
wide, broadly obovate to obcordate, tapering abruptly into
a narrow claw, strongly arching and incurving laterally
and thus enveloping the inner flora] parts, strikingly to
inconspicuously emarginate, the margins entire to very
finely erose, purplish-blue, bluish or violet in color, Fila-
ments about 5-8 mm long, glabrous. Anthers about 0.4-0.6
(0.7) mm long, exserted, yellowish prior to pollen dis-
charge. Pistil about 6 mm long with a glabrous ovary about
0.8-1 mm high, including the somewhat stipitate base,
broadest above the middle, with an antrorsely pubescent
style about 4-5 mm long terminated by a small, truncate,
stigmatic tip. Fruit about 4-5.5 mm long and 2-2.5 mm
wide, broadest at or near the apex and tapering to the
1-2 mm stipe-like base, glabrous, conspicuously punctate-
glandular in the upper half or two-thirds, exceeding the
calyx-tube by about 2.2-3.5 mm, laterally compressed,
obliquely obovate, the adaxial side straight or nearly so, the
abaxial side strongly outwardly bowed, usually terminated
by the 0.2-0.5 mm long, persistent base of the style.

Distribution: Sandy wire-grass savannahs, pine woods,
and thickets in the Coastal Plain from southeastern North
Carolina southward into eastern Georgia.

This species seems to be represented by two varieties
which are distinguished in the following key:

366 Rhodora [Vol. 77

KEY TO THE VARIETIES OF A. GEORGIANA

Lateral leaflets (3)6-10(15) mm long and (2)3-5(8) mm
wide; petioles 1-3(5) mm long; racemes typically solitary
but rarely with 1-3 short, additional racemes, mostly
(2)3-5(6) ст long; vexillum reddish-purple. ........
к —— Ба. var. georgiana.

Lateral leaflets typically (10) 15-25 (35) mm long and (7)
9-15(18) mm wide; petioles usually (6)8-15(20) mm
long; racemes in clusters of (1)3-5(8), 10-20(30) cm
long; vexillum light to more typically an intense bright
blue. .............................. 5b. var. confusa.

Ба. A. georgiana var. georgiana. TYPIFICATION: Georgia:
TELFAIR CO., sandy wire-grass savannah about 3
miles northwest of Lumber City, Wilbur 3158 (holo-
type, GH!; isotypes, DUKE!, GA!, MICH!, MO!, NSC!,
NY!, US!).

Distribution: Sandy flats bordering streams and low-
land woods from the central Coastal Plain of North Caro-
lina south into central coastal Georgia. (Map 5, in part.)

5b. A. georgiana var. confusa Wilbur, Jour. Elisha Mitchell
Sci. Soc. 80: 58. 1964. TYPIFICATION: North Caro-
lina: BRUNSWICK CO., grassy savannah, 7 miles south-
west of Wilmington, Godfrey & Shunk 4122 (holo-
type, GH!; isotype, US!).

A. glabra in the sense of Boynton, Bot. Gaz. 25: 279.
1898 and in Small’s Fl. SE. U.S. 626. 1903, but
not of Poiret, Encycl. Méth. Suppl. 1: 330. 1810.

A. caroliniana in the sense of Т. & С. Fl. N. Am. 1:
305. 1838 in part; Schneider, Bot. Gaz. 43: 302.
1907 and Ill. Handb. Laubh. 2: 74. 1907; Rydb.,
N. Am. FI. 24: 29. 1919, but not of Croom, Am.
Jour. Sei. 25: 74. 1834.

A. cyanostachya in the sense of Palmer, Jour. Arn.
Arb. 12: 169. 1931, and of Small, Man. SE. ЕІ.
639. 1933, but not of M. A. Curtis, Boston Jour.
Nat. Hist. 1: 140. 1835.

1975] Amorpha — Wilbur 367

Distribution: Pine woods, sandy ridges, savannahs, and
sandy roadbanks of southeastern North Carolina (Bladen,
Brunswick, and Columbus Counties). (Map 5, in part.)

The confused basis of the previous names applied to this
taxon has been discussed rather fully in an earlier paper
(Jour. Elisha Mitchell Sci. Soc. 80: 58-61. 1964) and there
is no need to repeat that discussion here. Although, I
remain skeptical that the present arrangement of the
georgiana and confusa taxa will prove to be satisfactory
when more is learned about them, I refrain from further
nomenclatural shifts at the present time. I suspect future
investigation may well demonstrate that the two taxa are
specifically distinct. Perhaps I am overly impressed by the
strikingly different colors of their petals.

6. Amorpha canescens Pursh, Fl. Am. Sept. 467. 1814.
Typification: Not seen. Perhaps in Pursh's col-
lection but Nuttall (Gen. N. Am. Pl. 2: 92. 1818)
states that “Мг. Pursh places his usual mark of v.v.
to this species, although he has never seen a flower-
ing specimen except in my herbarium.” “Оп the
banks of the Missouri and Mississippi," Pursh; and
“From the banks of Fox River and the Quisconsin
[Wisconsin River] to the Mississippi; around St.
Louis, Louisiana, and on the banks of the Missouri
probably to the Mountains," according to Nuttall.

A.canescens Nutt., Fraser's Cat. no. 4. 1815. nom.
nud.

A. canescens var. B leptostachya A. Gray, Mem. Am.
Acad. П. 4: 31. 1849. nom. nud.

A. canescens var. glabrata A. Gray, Smithson. Contr.
Knowl. 3(Art. 5, Pl. Wright): 49. 1852. TYPIFI-
CATION: eastern Texas, Charles Wright, not seen.

A. canescens var. « typica C. K. Schneid, Bot. Gaz.
43: 300. 1907.

A. brachycarpa Palmer, Jour. Arnold Arb. 12: 171.
1931. TYPIFICATION: Missouri: STONE CO. dry

868 Ећодога [Vol. 77

slopes and ledges, rocky hills near Galena, Е. J.
Palmer 19197 (holotype, A!; isotypes, KANU!,
MIN!, us!).

A. canescens f. glabrata (A. Gray) Fassett, Rhodora
38: 191. 1936.

A low, ascendant to erect, apparently rhizomatose, usu-
ally canescent, shrub mostly about (3)5-8(10) ат high.
Current season's growth light brown to olivaceous and
slender (about (1)2-3 mm in diameter) and typically very
densely grayish tomentose or occasionally but moderately
so or very rarely glabrous or nearly so, with numerous very
fine longitudinal striations and apparently eglandular or at
most with very few minute, amber-colored, very incon-
spicuous glands; older branches mostly light gray to brown-
ish and very slightly, if at all, lined, glabrous or very
nearly so. Leaves numerous, spreading, about (3.0)6.0-
12.0(15.0) em long. Petioles whitish to pale gray, moder-
ately to very densely tangled tomentose, villous or even-
spreading short-pubescent or very rarely moderately pu-
berulent to spreading short-pilose, sometimes equaling but
usually very much shorter than the breadth of the lower-
most leaflet, apparently always lacking pustulate glands,
mostly (0.5)1-3(5) mm long. Stipules inconspicuous, ca-
ducous, densely canescent-pilose externally, glabrous, dark
reddish and amber-colored within, initially appressed but
spreading and often becoming reflexed with age, linear-
lanceolate, linear to setaceous, mostly (1.2)2-3(3.5 mm
long, pustular glands apparently always lacking. Rachis of
leaf slender, mostly about (0.2)0.5-1.0(1.2) mm in diam-
eter, usually very densely grayish, tangled-tomentose but
occasionally varying to sparingly so to even strigillose or
spreading puberulent, channeled above, pustular glands
apparently always absent. Leaflets mostly (11)27-41 (47)
in number, usually about (0.3)1.0-1.8(2.5) ст long and
(2.0)4.0-7.0(12.0) mm wide, usually about (1.8)2.0-3.0
(3.8) times as long as broad, typically crowded and com-
monly overlapping one another, the interval between peti-

1975] Amorpha — Wilbur 369

olules on the same side of the rachis characteristically
about (2.0)3.0-8.0(10.0) mm long. Blades of leaflets mostly
ovate-oblong, oblong-elliptic, oblong, elliptic, or even ovate
with the terminal rarely obcordate and those of the dwarf
suckers often almost suborbicular to obovate, with the base
obtuse to broadly rounded, and more rarely subcordate or
even somewhat truncate and the apex obtuse to broadly
rounded but rarely acute or even emarginate with the mid-
vein exserted into a slender tapering mucro (rarely slightly
swollen at tip on a few leaflets) usually about (0.2) 0.4-
0.8(1.2) mm long. Texture of leaflet subcoriaceous to
thick-emembranous when dry, venation often completely
obscured or nearly so by the dense pubescence, but when
discernible usually only finely and inconspicuously reticu-
late above from the slightly elevated secondary and tertiary
venation (the midvein somewhat depressed) ; the margin
flat to very slightly revolute and entire; the midvein
markedly elevated beneath and the secondary and tertiary
venation often forming an inconspicuous net (which is
usually completely obscured or nearly so by the dense pu-
bescence); both surfaces usually very densely grayish
tomentose and then appearing canescent to less commonly
moderately short-pubescent or very rarely nearly glabrous
with pubescence restricted to the midvein and sometimes
principal veins and to the margin, and the leaves then
greenish (the pubescence usually both denser and longer
below than above) ; pustular-punctate glands usually pres-
ent beneath, although completely obscured from view by
the tangled pubescence, when not hidden by pubescence still
relatively inconspicuous, light olive-green or brown in color,
more or less of uniform size, very small. Petiolules very
densely spreading pilose to short-pubescent but rarely
sparingly short-pubescent, usually appearing canescent,
mostly (0.5)0.8-1.2 mm long, usually lacking punctate
glands or glands, if present, very inconspicuous. Stipels
acicular, about (0.5)0.8-1.2(1.5) mm long, mostly slightly
exceeding but sometimes almost twice as long as the peti-
olule, glabrous to sparsely short-pubescent, long-persistent,

370 Rhodora [Vol. 77

dark reddish brown, the lower rarely developing beyond
the swollen base. Racemes usually numerous in the axils
of the uppermost leaves and often forming a dense com-
pound cluster, mostly (1)5-20(30) or even more in number,
very densely flowered, mostly (2.0)7.0-15 (25) ст long and
about 1-1.5 cm thick; peduncle short, mostly about 1-5 (8)
em long; rachis strongly ridged and grooved, very densely
spreading-pilose to short-pubescent, sparingly and very
inconspicuously glandular-punctate and bearing numerous,
very densely arranged, moderately to (more typically)
densely spreading, wavy, short-pubescent or pilose pedicels
appearing short and stout with a truncate apex, about
(0.4)0.6-1.2(1.5) mm long, subtended by a caducous,
sparsely glandular-pustulate, amber-colored, moderately to
densely pilose externally and glabrous within, linear-
setaceous to narrowly lanceolate (with a long tapering
apex) bract mostly (2.5)3.0-4.0(4.5) mm long. Calyx-tube
turbinate to obconic-campanulate, mostly (1.5) 1.8-2 (2.5)
mm high, slightly asymmetrical with the abaxial slightly
longer than the adaxial, usually very densely canescent
throughout, but very rarely sparingly pilose to almost
glabrous (the trichomes mostly spreading short- to long-
pilose), usually rather inconspicuously (due to the pubes-
cence) punctate-glandular, internally glabrous except for
possibly the uppermost fringe. Calyx-lobes all narrowly
triangular-lanceolate, obscurely punctate-glandular and
acute; adaxial lobes (0.6) 1-1.4(1.6) mm long; lateral lobes
(1.0) 1.2-1.5(1.8) mm long; abaxial lobe (1.2) 1.5-2.2 (2.5)
mm long, all very densely canescent, short- to long-pilose
externally and internally more sparingly pilose. Vexillum
about 4.5-6.0 mm long and 2.5-4 mm wide, very broadly
obcordate and abruptly tapering into a slender claw (about
1-1.5 mm long), strongly arched and incurved, enveloping
the inner floral parts, its apex usually somewhat emargi-
nate but occasionally also obscurely apiculate, its margin
entire to irregularly erose, bright violet. Filaments golden-
brownish to purplish, about 6-8 mm long, glabrous, united
into a tube for the lowest 1.5-2.0 mm. Anthers about 0.4-

1975] Атогрһа — Wilbur 311

MAP 6. Атотрћа canescens.

0.8 mm long, yellowish to golden brown. Pistil about 5.0-
7.0 mm long with densely pilose ovary about 1 mm high, a
densely antrorsely pubescent style about 4-6 mm long and
a small, capitate, terminal stigma. Fruit about (3.0)3.5-
4.5(5.0) mm long and (1.5)1.8-2.2 mm wide, tapering to
the somewhat stipe-like base, usually densely to moderately
villous-canescent but very rarely glabrous, punctate-glandu-
lar in the upper half or two-thirds, exceeding the calyx-
tube by 2-3.5 mm, with the adaxial straight or very nearly
so and the abaxial side strongly outwardly bowed, termi-
nated by the 0.5-1.5 mm long, persistent base of the style.

372 Rhodora [Vol. 77

Distribution: Roadsides, fields, prairies, hillsides, and
open woodlands from Indiana west into the Dakotas, Wyo-
ming, and New Mexico, and southern Canada south into
eastern Texas. (Map 6.)

The reasons for recognizing neither A. canescens var.
glabrata A. Gray nor А. brachycarpa have been presented
in an earlier paper (Jour. Elisha Mitchell Sci. Soc. 80:
62-63. 1964) and will not be repeated here.

An apparently rarely encountered hybrid between A.
canescens and A. fruticosa may be better discussed here
than elsewhere since the overall aspect of the plant is such
that it most likely would be taken as an aberrant, oversized
A. canescens. The rarity of the supposed hybrid certainly
attests to the effectiveness of the isolating mechanisms
between the two species for they occupy the same general
geographical area over a very extensive region of the
central United States.

Palmer (Rhodora 55: 158-159. 1953) described the
hybrid of these two species as Amorpha X notha from a
single plant found on a “low rocky bank between upland
prairie and alluvial valley of Center Creek, about 1 mile
north of Webb City, Jasper Co., Missouri.” Both species
are common in southwestern Missouri and yet Palmer, who
was extremely familiar with the plants of that area and
who had a particular interest in Amorpha, knew only one
example of the hybrid. In addition to the two collection
numbers made by Palmer from the type plant (Palmer
52047, 27 May 1951, in flower; isotypes, F, MO, SMU, ОМО);
and Palmer 52930, 9 August 1951, in fruit (F, UMO).
Palmer made an additional collection (#56605) from the
same area, if not from the same plant, two years later.
Another collection which I take to represent this hybrid
was collected by Palmer, #25305, 4 June 1924: dry rocky
ledges, limestone bluffs of Spring River, near Melugin,
Jasper Co., Missouri (A, MIN, ОМО) and was cited, mis-
takenly, I believe, (Jour. Arnold Arb. 12: 167. 1931) as
A. canescens var. glabrata A. Gray.

1975] Amorpha — Wilbur 378

The hybrid differs from A. canescens with which it has
the greatest overall affinity in its taller habit, much shorter
calyx-lobes and particularly in its dorsally strongly curved
fruit. From its A. fruticosa parentage, it differs in densely
canescent, pilosulose calyx-tube as well as the finely canes-
cent leaves and young stems. The practice of giving bi-
nomials to hybrids of this sort has, I believe, very little to
recommend itself as a botanical custom.

7. Amorpha nana Nutt., Fraser's Cat, 1813. TYPIFICATION:
“Collected near the Mandan towns, 1600 miles up
the Missourie," Nuttall, not seen.

A. microphylla Pursh. Fl Am. Sept. 466. 1814.
nom. illegit., Art. 63. TYPIFICATION: not seen;
Pursh stated that he based name on a specimen
presented to him by Meriwether Lewis of the
Lewis and Clark Expedition.

A. punctata. Raf., New Fl. 3: 14. 1887. TYPIFICA-
TION: Specimen not seen. “Discovered by Brad-
bury in the upper Missouri."

A low, erect, apparently rhizomatose shrub aobut 8-6
(9) dm high. Branches of the current season light reddish-
brown to pale olivaceous; clustered near the top of the
stem; slender (about 1-2(3) mm in diameter), sparsely
or, more typically, moderately strigillose but becoming
glabrate below with age and with numerous very fine,
longitudinal grooves and ridges and a sparse to moderate
sprinkling of small inconspicuous punctate-pustular, amber-
colored glands; older branches usually light gray or pale
brownish, only slightly, if at all, lined. Leaves numerous,
spreading to ascendant, (1.5)3-7(10) cm long. Petioles
sparsely to densely strigillose or somewhat finely spreading-
puberulent, becoming glabrous or nearly so, equaling or
usually longer than the breadth of the lowermost leaflet
(rarely the breadth of leaflet is greater than length of
petiole), about (2)4-8(10) mm long, very sparingly pustu-
late-glandular. Stipules inconspicuous, pigmented like the

374 Rhodora [Vol. 77

young stem and petiole, caducous, short-pubescent along
margins and outer surface and often with a moderately
long tuft of trichomes near tip, appressed to spreading,
setaceous to linear, (2)3-5 (6.5) mm long, occasionally
sparingly pustulate-glandular. Rachis of leaf very slender,
mostly about 0.3-0.7 mm in diameter, glabrous or nearly
so to densely very short-puberulent or strigillose and
sparingly to moderately beset with minute pustular-glands,
channeled above. Leaflets mostly (7)13-27 (31) in number,
(2)6-13(18) mm long, (2)3-6(7.5) mm wide, usually (1)
1.8-2.9(3.5) times as long as broad, rarely overlapping to
more commonly remote, usually the interval between peti-
olules on the same side of the rachis about (3)4-7(9) mm
long. Blade of leaflet mostly narrowly to broadly oblong or
even somewhat elliptical but occasionally varying to ovate
or obovate to almost orbicular, with the base usually
rounded or occasionally with margins gradually tapering
and with the base then cuneate, the apex usually broadly
rounded or even truncate and commonly emarginate but
occasionally acute, with the midvein exserted into a slender,
tapering mucro usually about (0.6)0.8-1.2(1.5) mm long.
Texture of leaflet coriaceous to subcoriaceous when dry,
finely and inconspicuously reticulate above from the some-
what elevated secondary and tertiary venation (the mid-
vein somewhat depressed), the margin flat or more typi-
cally somewhat revolute and entire to inconspicuously cren-
ulate; the midvein conspicuously elevated beneath and with
the secondary and sometimes the tertiary veins somewhat
elevated and sometimes forming an inconspicuous net;
surface glabrous on both sides but the midvein beneath and
the margin usually sparingly short-pubescent, puberulent
or somewhat strigillose or rarely glabrous or nearly so;
the punctate glands conspicuous and readily visible to the
unaided eye on the lower surface, appearing to be of uni-
form size or at least not falling mostly into two more or
less distinct size classes, usually with about (10) 20-35 (55)
punctate glands рег half-leaflet, Petiolule densely to spar-
ingly short-pubescent or strigillose and sometimes becoming

1975] Amorpha — Wilbur 375

glabrate with usually few to several small pustulate glands
and mostly about (0.5)0.7-1 mm long. Stipels acicular,
about (1.0) 1.5-2.5 (3) mm long, often 2 times or more as
long as the petiolule, glabrous, long-persistent. Racemes
solitary at the tips of the current season’s growth, con-
tracted and very densely flowered, mostly (2.0)3-7(9) cm
long and about 1-1.5 cm in diameter, subsessile or the
peduncle about 0.3-1.5 (2.0) ст long; rachis conspicuously
ridged and grooved, moderately to very densely puberulent
or occasionally strigillose; pedicels bearing numerous very
densely clustered, moderately to densely puberulent or
strigillose (often becoming sparsely so or even glabrate in
fruit), often long-persistent (and with a truncate, some-
what expanded apex), about 1.0-2.0(2.5) mm long, sub-
tended by a caducous, sparsely glandular-pustulate, brown-
ish to amber-colored, moderately to densely puberulent,
linear-setaceous to narrowly spatulate bract mostly (2.5)
3.0-4.0(5.0) mm long with a long, tapering apex. Calyx-
tube turbinate, mostly 1.8-2.2 mm high with the abaxial
side slightly longer than the adaxial, glabrous throughout
with few to more usually numerous punctate glands mostly
arranged in rows and restricted to the upper half of the
tube. Calyx-lobes all narrowly triangular-lanceolate and
acute, punctate-glandular, glabrous or very sparsely pu-
berulent externally but with densely short-ciliate margins;
internally the lobes and the very uppermost portion of the
tube densely matted short-puberulent and the lower portion
of the tube internally (as well as externally) glabrous;
adaxial lobes about (0.8) 1.0-1.5 (1.8) mm long; lateral lobes
mostly (1.0)1.2-1.8(2.0) mm long; abaxial lobe (1.5) 1.8-
2.0(2.2) mm long. Vexillum about 4.5-6.0 mm long and
3.5-4.5 mm wide, very broadly obcordate, abruptly taper-
ing into a slender claw, strongly arched and incurved and
enveloping the inner floral parts, its margin emarginate
and usually finely erose, dark purple. Filaments purplish,
about 6-8 mm long, glabrous; anthers about 0.4-0.6 mm
long, purplish. Pisti] about 4.0-6.0 mm long with a gla-
brous ovary about 0.8 mm high, an antrorsely pubescent

[Уо]. 77

Rhodora

376

Amorpha nana.

MAP 7.

1975] Amorpha — Wilbur | 377

style about 3.5-5 mm long and a small, capitate, terminal
stigma. Fruit about 4.5-5.5 mm long and 2.0-2.8 mm wide,
oblongish but tapering to a stipe-like base of about 0.5 mm
long, glabrous, conspicuously punctate-glandular in the
upper two-thirds, exceeding the calyx-tube by about 2.0-
3.5 mm, the adaxial side straight or nearly so, the abaxial
side strongly outwardly bowed, terminated by persistent
base of the style, this about 0.5 mm long.

Distribution: Prairies, hillsides, plains, and buttes from
Iowa and Minnesota west into the Dakotas and Colorado
and from southern Manitoba and Saskatchewan south into
New Mexico; primarily a plant of the northern plains.
(Map 7.)

8. Amorpha glabra Poir., Encycl, Meth. Suppl. 1: 330. 1810.
TYPIFICATION: type not seen, perhaps at P; based on
a specimen grown in cultivation in Paris.

24. glabra Desf., Tabl. Ecole Bot. Paris 192. 1804,
nom. nud.

?A. glabra Pers., Syn. Pl. 2: 295. 1807. nom semi-
nud.

A. montana Boynt., Biltmore Bot. Stud. 1: 158. 1902.
TYPIFICATION : North Carolina: rocky woods, Bilt-
more, Biltmore Herb. 14 (lectotype, Us!; fruiting
specimen collected 29 August 1896 on sheet num-
bered 331562; isolectotypes, A!, CU!, GH!, MIN!,
MO!, ND-Greene!, NY!, RM!, us!).

ФА. fruticosa var. glabra Bean, Trees and Shrubs
Brit. Isles 1: 198. 1914.

A much-branched, erect shrub mostly 1-2 m high. Cur-
rent season's growth light to dark purplish or light-brown
to olivaceous, slender to moderately thickened, about 1-2
(3) mm in diameter, usually completely glabrous or at most
very sparsely and obscurely strigillose to puberulent, spar-
ingly, if at all, beset with inconspicuous, minute, amber-
colored pustulate glands; branches of the preceding year

378 Rhodora [Vol. 77

mostly grayish to blackish. Leaves wide-spreading, mostly
(1.0) 1.4-2.2(3.0) dm long. Petioles mostly (1.6) 2.0-4.0
(5.7) ст long, typically much longer than the width of the
lowermost leaflet, glabrous or very nearly so or very spar-
ingly puberulent, sparsely beset, if at all, with amber-col-
ored pustulate-glands. Stipules inconspicuous, appressed
when present, caducous, thin, sparsely to moderately pu-
berulent to short-pubescent on margins and outer surface,
dark reddish-brown, typically linear to linear-lanceolate,
about 2.5-4.5 mm long, apparently eglandular. Rachis of
leaf about 0.5-1.0 mm in diameter, usually greenish-oliva-
ceous, glabrous or rarely at most exceedingly sparingly
puberulent or strigillose, usually sparingly glandular pus-
tulate throughout, interval between petiolules on the same
side of the rachis about (0.8) 1.5-3.0 (3.6) ст long. Leaflets
mostly (7)11-15(21), either opposite or alternate, about
(1.4) 2.2-4.6(7.4) em long and (1.0) 1.4-2.8(3.6) cm wide,
mostly (1.2) 1.4-2.2(2.7) times as long as broad, character-
istically widely spaced and not overlapping, usually broadly
oblong, oblong-elliptic to elliptic or less commonly ovate-
oblong to ovate or rarely nearly orbicular, basally typically
very broadly rounded to truncate or nearly so or even
subcordate but rarely gradually tapering to even acute,
apically very broadly rounded to strongly obtuse and almost
invariably conspicuously emarginate. Midvein usually ter-
minating in a swollen, globose, glandular tip either sessile
or exserted on a stalk about 0.2-0.4 mm long. Both surfaces
with venation scarcely either elevated or depressed on
either surface except for the conspicuously elevated mid-
vein and the primary and secondary veins obscurely reticu-
late, glabrous or rarely sparsely and inconspicuously stri-
gillose beneath along the midvein and principal lateral
veins, usually conspicuously glandular-punctate; punctate
glands variable in size, usually numerous, mostly ranging
in number from about (30)50-150(300) glands per half-
leaflet. Margin entire, inconspicuously and finely undulate
to conspicuously crenulate, slightly revolute to more or less
flush-margined. Petiolules about 2.0-4.0(5.5) mm long,

1975] Amorpha — Wilbur 319

usually glabrous but occasionally sparingly to moderately
puberulent or strigillose, mostly sparsely glandular-pustu-
ate, wrinkled when dried. Stipels acicular to setaceous,
mostly 1.2-2.0 mm long (but the tips quickly caducous and
thus appearing shorter), glabrous, usually long-persistent,
dark reddish-brown. Racemes often solitary and terminal
or few and terminally clustered, 1-3(4) in number, with
densely clustered flowers, mostly about (0.5) 1.0-1.8 (2.8)
dm long and 1.2-2.0 cm in diameter, sessile or nearly so or
with a peduncle 1-6 cm long; rachis sparsely glandular-
punctate, glabrous or very sparingly strigillose or puberu-
lent. Pedicels glabrous or at most sparsely strigillose or
puberulent, usually eglandular, about (0.8) 1.0-2.5 (5.0) mm
long; bracts caducous, apparently eglandular, golden-
brown, usually puberulent or ciliate, linear-oblong or linear-
lanceolate, about (0.5) 0.8-1.5 (2.0) mm long. Calyx-tube
broadly turbinate or campanulate, mostly (2.0) 2.2-5.0
(3.2) mm high, somewhat asymmetrical with the abaxial
side slightly higher than the adaxial, glabrous externally
and internally throughout excepting the densely short-
ciliate rim, bearing 1 or 2 rows of small, inconspicuous,
pustulate-glands mostly restricted to the upper third of the
tube. Calyx-lobes often obsolete or nearly so and the top
of the calyx then appearing truncate or the lobes low and
depressed with the calyx-rim appearing weakly undulate
but when noticeably developed usually broader than high,
mostly rounded or broadly triangular-dentate ; adaxial lobes
often rounded and shorter than the lateral lobes but fre-
quently triangular-dentate and then occasionally longer
than the lateral lobes, when developed mostly (0.1) 0.2-0.8
(0.5) mm long; lateral lobes often broadly rounded but
commonly broadly triangular-dentate and when developed
beyond mere undulations then mostly (0.1)0.2-0.4(6.5) mm
long; adaxial lobe typically triangular and acute although
oecasionally rounded, mostly (0.1)0.2-0.6(0.8) mm long.
Vexillum about (4.5) 6.0-8.0 (8.5) mm long and 4.0-6.0 mm
wide, broadly obcordate, abruptly to gradually tapering
into a narrow, short claw about 1 mm long, entire to erose,

380 Ећодога [Vol. 77

MAPS 8-10. Map 8. Amorpha glabra. Map 9. А. schwerinii
Map 10. А. paniculata.

usually moderately to strongly emarginate, bright reddish-
purple. Filaments yellowish to golden, about 8-11 mm long,
glabrous; the monadelphous tube noticeably exserted at
maturity from the calyx-tube for about (0.5)1.0-2.0 mm,
the united portion of the filaments mostly about 3.0-4.5 mm
long; anthers about 0.5-1.0 mm long, golden-yellow to
orange. Pistil about 7.0-9.0 mm long with a glabrous, obo-
vate ovary about 1 mm high and an antrorsely short-
pubescent style terminated by a truncate stigmatic tip.
Fruit about (6.5)7.5-9.0(10.5) mm long and (2.5)3.0-

1975] Amorpha — Wilbur 381

4.0(4.5) mm wide, broadest at or above the middle and
tapering to the slender, conical base, elabrous throughout,
the upper two-thirds sparsely to moderately covered with
small but usually conspicuously pustulate glands, obliquely
obovate with the adaxial valve straight or nearly so and
the abaxial valve very strongly outwardly bowed above,
often terminated by the persistent, 0.2-0.5 mm long base of
the style.

Distribution: Endemic to the southern Appalachian Mts.
of western N. and S. Carolina, northeastern Ga. and east-
ern Tennessee. (Map 8.)

The attribution of this species to Arkansas and Okla-
homa (Jour. Arnold Arb. 12: 174-175. 1931, ard Proc.
Oklahoma Acad. За. 27: 69. 1947.) is, I believe, an error.
Specimens so annotated in the past seem to me to be A.
ouachitensis whose closest relative appears to be the central
Texan A. roermeriana (=A. texana) and not the southern
Appalachian endemic.

9. Amorpha schwerinii Schneider, Bot. Gaz. 43: 301. 1907.
[as Sehwerini]. TYPIFICATION: North Carolina:
ROWAN CO. Dunn’s Mountain, alt. 1200 ft. 7. К.
Small s.n. (holotype, Mo!; isotypes, F! му!).

A. densiflora Е. E. Boynton ex Small, Fl. SE. U.S. ed.
2. 1342. 1913. TYPIFICATION: North Carolina:
GASTON CO., slopes of Crowder's Mountain, Bilt-
more Herb. 14756b (lectotype, NY!; isolectotypes,
MIN! NY! RM!).

A bushy, widely branched, erect shrub mostly (1) 1.5-
25 m high. Current season's growth dull olive-brown to
reddish-brown or dull grayish to almost black and typically
slender, about 1.0-2.0 mm in diameter, mostly densely pu-
berulent to curly short-pubescent, usually with a moderate
to dense scattering of inconspicuous, small, amber-colored,
pustular glands; older branches dark grayish to blackish
but becoming dark reddish-brown with numerous, incon-
spicuous longitudinal fissures and more or less orbicular,

382 Ећодога [Vol. 77

slightly elevated, lenticels, becoming progressively glabrate.
Buds globose to ovoid, scales densely puberulent to short-
pilose. Leaves wide-spreading, about (0.5)0.8-1.2(2.2) dm
long. Petioles mostly (0.7) 1.0-1.8 (2.6) ст long, mostly
equaling or exceeding the length of the lowermost leaflet
but occasionally shorter, moderately to densely pubescent
with spreading hyaline trichomes (up to 0.7 mm long),
bearing none or but few pustular, amber-colored glands.
Stipules inconspicuous, usua!ly appressed while present but
soon caducous, copiously tawny puberulent to curly short-
pubescent externally and glabrous or nearly so within, dark
reddish-brown, typically linear to linear-lanceolaie, about
3.0-4.5 mm long, eglandular. Rachis of leaves slender,
mostly 0.5-0.7 mm in diameter, typically brownish to gray-
ish, moderately to densely puberulent to spreading short-
pubescent with the trichomes up to about 0.5 mm long,
eglandular or the pustulate glands sparse and inconspicu-
ous. Leaflets mostly (7)19-27 (29) in number, usually about
(0.5) 1.5-3.0(4.0) em long and (0.4)0.8-1.5(1.9) ст wide,
typically about (1.3)2.0-3.0(3.8) times as long as broad,
rarely overlapping, the interval between petiolules on the
same side of the rachis mostly about (0.6) 0.8-1.5 (1.8) ст
long; usually narrowly to broadly oblong but varying from
elliptic to elliptic-oblong and rarely from ovate-oblong to
ovate or even almost orbicular in unusual terminal leaflets,
typically with the base broadly rounded but varying to
somewhat truncate or even slightly subcordate and the apex
mostly obtuse to broadly rounded or occasionally emar-
ginate; venation slightly elevated above and more pro-
nouncedly so below. Midvein usually swollen terminally,
usually exserted for 0.2-0.5(0.8) mm. Margin entire to very
obscurely crenulate and inconspicuously revolute. Lower
surface densely pilose with soft ascendant or spreading hy-
aline, tapering trichomes mostly 0.3-0.6 mm long particu-
larly pubescent along the midvein, moderately to densely
beset with olivaceous to amber, punctate glands beneath
(about 30-120 glands /half-leaflet) ; upper surface minutely
and obscurely but copiously short-pubescent with hyaline,

1975] Amorpha — Wilbur 383

appressed to spreading trichomes about 0.1-0.3 mm long.
Petiolules usually (0.5)1-2 mm long, densely spreading
pilosulose to short-pubescent with hyaline trichomes mostly
0.2-0.6 mm long, indistinctly glandular-pustulate. Stipels
acicular, about 1.0-2.2 mm long, often equaling the petio-
lules in length, typically long-persistent, dark reddish-brown,
pilosulose and often densely so at base and glabrous or
nearly so above, sparingly pustulate. Racemes 1 or some-
times 2 and the principal one usually terminating strong
shoots and the second when present smaller and arising at
or near base of the terminal cluster, mostly (2.0)4.0-8.0(12.0)
em long and 1.0-1.5(1.8) em in diameter; rachis of inflores-
cence densely pilosulose to short-pubescent, eglandular or
nearly so. Pedicels puberulent to pilosulose, eglandular,
about (0.8) 1.0-1.5(2.0) mm long; bracts caducous, sparingly
glandular-pustulate, moderately to densely pilosulose ex-
ternally and glabrous within, linear to linear-lanceolate,
about 2.0-3.5 mm long. Calyx-tube turbinate to obconic-
cylindrical, mostly 1.8-2.5 mm long, somewhat asym-
metrical with the adaxial side slightly longer than the
abaxial, externally sparingly to densely spreading pilosulose
throughout, moderately to densely beset with amber-col-
ored, pustulate glands in the upper one-fifth to one-third
but eglandular below. Calyx-lobes mostly linear to linear-
lanceolate (very rarely the four paired lobes triangular-
dentate), acute, sparingly to moderately glandular-punctate,
short-pilose both externally and internally, mostly equaling
or exceeding the length of the tube. Adaxial lobes about
(1.2)2.0-3.0 mm long; lateral lobes about (1.5)2.0-3.2 mm
long; abaxial lobe (1.8)2.5-3.5 mm long. Vexillum about
4.5-6.5 mm long, 3.0-4.2 mm wide, very broadly obovate to
obcordate, tapering into the slender claw of about 1.5 mm
long, apically emarginate and sometimes also inconspicu-
ously apiculate, margin entire to irregularly erose, purplish.
Filaments yellowish to golden, about 6-8 mm long, glabrous,
united at base into a tube 1.5-2.0 mm long; anthers about
0.5-0.8 mm long, golden-yellow. Pistil about 5-7 mm long
with a densely antrorsely pubescent style about 4-6 mm

384 Ећодога [Vol. 77

high and a capitate stigma. Fruit about 5-6.5 mm long and
1.8-2.2 mm wide, tapering to base, usually densely short-
pubescent or rarely glabrous or nearly so, usually conspicu-
ously glandular-pustulate in upper half, the adaxial valve
straight or nearly so and the abaxial very strongly out-
wardly bowed above, terminated by the persistent base of
the style.

Distribution: River banks and open slopes from the inner
North Carolina Piedmont southwest into Alabama, (Map 9.)

This is morphologically one of the most distinctive and
clearcut species within the genus.

10. Amorpha paniculata T. & G., Fl. N. Am. 1: 306. 1838.
TYPIFICATION: Texas: Т. Drummond 461 (lectotype,
NY!; isolectotype, GH!).

Erect coarse suffrutescent herb or shrub 1-3 m tall and
relatively unbranched. Current season’s growth usually
densely puberulent to spreading short-pubescent or glabrate
in age, often somewhat canescent and lacking pustulate
glands. Buds obovoid, densely appressed short-pubescent,
canescent. Leaves wide-spreading, mostly (1)2-4 dm long.
Petioles (2)3-6(9) ст long, usually noticeably longer than
the width of the lowermost leaflet, stout, canescent, densely
puberulent. Stipules caducous, setaceous, pubescent, about
3 mm long. Rachis of leaf about 2.5 mm in diameter,
densely puberulent throughout and more or less canescent.
Leaflets drying coriaceous, mostly 11-19, oblong to oblong-
elliptic, usually (1.5)3-6(8) ст long and 1.5-3(5.3) ст
wide, typically (1.4) 1.8-2.2 (2.6) times as long as wide,
opposite or more commonly alternate, usually symmetrical,
typically broadly rounded both apically and basally, con-
spicuously reticulate below with the veins elevated often to
the fourth degree of branching. Midvein exserted up to
1.8 mm or ending at the margin, usually somewhat taper-
ing but rarely swollen. Lower surface usually moderately
to densely covered with spreading, crisped, hyaline to
tawny, pilosulose trichomes but rarely sparingly to mod-

1975] Amorpha — Wilbur 385

erately short-pubescent or even glabrate; upper surface
glabrate or sparingly short-pubescent to puberulent espe-
cially along the principal veins, usually inconspicuously
and sparingly to moderately beset beneath with small, pus-
tulate, punctate glands. Petiolules (0.2)0.4-1.0 cm long,
moderately to more typically densely spreading pilosulose,
somewhat canescent, sparingly and inconspieuously pustu-
late-glandular. Racemes several to numerous, (1)5-12 in
number, about (0.5) 1.5-3.0 (4.0) dm long; rachis of inflor-
escence finely pilosulose, canescent. Pedicels slender, about
1-2 mm long, pilosulose; bracts very slender, setaceous,
pilosulose, about 2.95 mm long. Calyx-tube funnel-form,
(1.8)2(2.2) mm long, moderately to densely pilosulose with
fine hyaline trichomes or rarely glabrate in age, sparingly
to moderately and inconspicuously glandular-pustulate
above. Calyx-lobes inconspicuously glandular-pustulate,
densely but finely pilosulose ; adaxial lobes broadly to nar-
rowly triangular-dentate or ovate, acute, 0.8-1.0(1.2) mm
long; lateral lobes longer and usually more narrowly lanceo-
late, acute to acuminate, (1.0)1.2-1.4 mm long; abaxial
lobe (1.2) 1.5-2.0 mm long, linear to linear-lanceolate. Vex-
illum 5-7 mm long, 3-4 mm wide, entire to slightly erose
apically, purple. Filaments about 5-6 mm long, united for
about half their length, glabrous; anthers about 0.6-0.8 mm
long, orange. Style densely villous; ovary glabrous, Fruit
4-6(8) mm long and 2.95 mm wide, broadest somewhat
above the middle, tapering to the base, with the adaxial
margin strongly outwardly bent in the upper quarter and
the abaxial margin strongly outwardly bowed, glabrous,
conspicuously glandular-pustulate. Seed reddish-brown,
about 3.5 mm long and 1.8 mm wide, smooth.

Distribution: Thickets, bogs, swampy woods and ditches
of southwestern Arkansas, west central Louisiana and
eastern Texas. (Map 10.)

The distinctiveness and relative uniformity of this spe-
cies is suggested by its lack of synonyms. The only syno-
nym sometimes associated with it has been A. roemeriana

386 Rhodora [Vol. 77

Scheele, but I believe this binomial properly belongs to the
central Texan plant previously referred to as A. texana
Buckl.

11. Amorpha nitens Boynton, Biltmore Bot. Stud. 1: 139.
1902. TYPIFICATION: Georgia: swamp near Waynes-
boro, Biltmore Herbarium s.n. (holotype: not seen,
designated as at the Biltmore Herbarium and hence
presumably at the us).

A. nitens var. leucodermis E. J. Palmer, Jour. Arnold
Arb. 12: 177. 1931. TYPIFICATION: Georgia:
thicket along lake, Augusta, Boynton 7035 (holo-
type, aH !).

An erect, branching shrub 1-3 m tall. Current season's
growth usually blackening or at least darkening upon dry-
ing, glabrous to sparingly or even moderately puberulent
to pilosulose, usually eglandular or bearing only a few
scattered, small inconspicuous glands. Buds compressed
and broadly oblong or ovoid, blackish when dried, sparingly
pilosulose to glabrate except for the shortly puberulent
margins of the outer scales. Leaves more or less wide-
spreading to ascendant, mostly 1-1.8(2.2) dm long, usually
conspicuously darkening or blackening upon drying. Peti-
oles glabrous to moderately crispy short-pilose, eglandular
or with a few, inconspicuous, small, pustulate glands, typi-
cally longer than the width of the lowermost leaflet, mostly
(1.5) 2-3.5(5) ст long. Stipules caducous, linear to linear-
setaceous, eglandular, drying blackish, usually glabrous
except externally with an apical tuft of tawny pubescence,
mostly 3-5 mm long. Rachis of leaf about 0.6-1.0 mm in
diameter, glabrous to moderately crispy-pilosulose, usually
eglandular but rarely very sparingly and inconspicuously
glandular. Leaflets (7)9-15(19) in number, oblong to el-
liptic-oblong or occasionally somewhat ovate, mostly 2-4 (7)
ст long and (1.0) 1.5-3.5 (4.5) em wide, typically 1.5-2.5
times as long as wide, opposite or more typically alter-
nate, usually widely spaced, symmetrical, usually broadly

1975] Amorpha — Wilbur 387

rounded basally, apically obtuse to more typically broadly
rounded and often emarginate, entire; secondary venation
usually little, if at all, elevated beneath. Midvein typically
shortly exserted and but little tapered. Lower surface
of leaflets glabrous to moderately spreading pilosulose
throughout or the pubescence sometimes restricted to the
midvein, eglandular or inconspicuously and often rather
sparingly beset with small punctate glands; upper surface
eglandular and glabrous and often shiny at maturity but
occasionally sparsely beset with extremely minute and
fine, appressed short-pubescence. Petiolules about 2-3.2 mm
long, glabrous to moderately spreading pilosulose, eglandu-
lar to inconspicuously and very sparingly pustulate gland-
ular. Racemes erect, solitary or with several clustered
together, (0.5) 0.8-1.5 (2.5) dm long; rachis of inflorescence
glabrous to moderately pilosulose or puberulent, eglandular
or very sparingly and inconspicuously pustulate-glandular.
Pedicels 1-2.2 mm long, glabrous to moderately pilosulose
or crispy pubescent, slender, eglandular, blackening upon
drying; bracts usually caducous, basally attached on pedi-
cel or distally as much as midway towards the tip, very
narrowly linear to setaceous, 1.5-3(4) mm long, glabrous
to externally tangled tawny-pilosulose especially along the
margins, usually eglandular but occasionally inconspicu-
ously and very sparingly pustulate glandular, typically
blackening upon drying. Calyx-tube slenderly obconic to
broadly funnelform to rarely even roundedly campanulate,
mostly 2-2.5 mm long, glabrous to moderately pilosulose,
often completely eglandular but occasionally very sparingly
beset with inconspicuous, minute pustular glands in the
upper third. Calyx-lobes with a ciliate fringe and with the
outer surface glabrous to pilosulose, eglandular; adaxial
calyx-lobes usually broadly rounded and obtuse but occa-
sionally varying to triangular, and acute, about 0.2-0.6 mm
long; lateral calyx-lobes broadly rounded to more typically
triangular, about 0.1-1.0 mm long; abaxial lobe triangular
and acute and often narrowly so, 0.6-1.4 mm long. Vexil-
lum 4.5-6 mm long, 3-4.8 mm wide, broadly obovate, grad-

388 Rhodora [Vol. 77

ually tapering into an indistinct claw about 1-1.5 mm long,
slightly to strongly arched, apically broadly rounded and
entire to irregularly erose, apparently reddish purple
[“blue-purple fide Palmer"] and with the claw and blade
both enveloping the filaments. Filaments 5-7 mm long,
united into a sheath for the lower 2-2.5 mm, glabrous;
anthers 0.5-0.8 mm long, yellow. Style moderately ascend-
ingly pilose, 5-7 mm long; ovary glabrous. Fruit 6-8 mm
long and 2.5-3.5 mm wide, broadest above the middle and
tapering basally with the adaxial suture straight or slightly
curved and the abaxial strongly outwardly bowed, glabrous,
moderately pustulate glandular distally. Seeds smooth,
dark reddish-brown to blackish, 3-4.2 mm long.

Distribution. Thickets and moist woods along creeks,
rivers and bottomlands from southern Illinois southeast
into Georgia and west into Louisiana and eastern Okla-
homa. (Map 1i.)

12. Amorpha laevigata Nutt. in Torr. & Gray, Fl. N. Am.
1: 306. 1838. TYPIFICATION: “Banks of the Arkan-
sas, near Salt River,” Nuttall s.n. (holotype, BM, not
seen; phototype: A!)

A. laevigata var. typica Schneider, Illustr. Handb.
Laubh. 2: 74. 1907.

Shrub 1-2(3) m tall. Current season's growth sparingly to
moderately strigillose and glandular-pustulate, light brown
to more characteristically deep reddish-purple. Buds sub-
orbicular to ovoid, compressed; the scales sparingly short-
pubescent. Leaves wide-spreading to moderately ascendant,
mostly (0.5)0.8-1.5(2.4) dm long. Petioles (0.6) 1.0-2.5
(3.4) em long, about 1 mm in diameter, very sparingly
strigillose or glabrous, sparingly to moderately glandular-
pustulate, usually longer than the width of the lowermost
leaflet. Stipules caducous, linear-subulate to setaceous,
glabrous, dark reddish-purple, about 2-3 mm long. Rachis
of leaf 0.5-1 mm in diameter, glabrous or very sparingly

1975] Amorpha — Wilbur 389

MAPS 11-14. Map 11. Amorpha nitens. Map 12. A. laevigata.
Map 13. A. roemeriana. Map. 14. A. ouachitensis.

strigillose, sparingly to moderately glandular-pustulate.
Leaflets (9)13-19(25), oblong or oblong-elliptie to rarely
obovate, usually (1.0)1.6-3.2(4.2) cm long and (0.4) 1.0-
1.5(2.2) em wide, typically (1.5)1.8-2.4(2.8) times as long
as wide, usually alternate and symmetrical, characteristi-
cally basally rounded but rarely acutely tapering, apically
usually broadly rounded and often emarginate; venation
little elevated beneath except for the midvein; margin
entire to slightly erenulate. Midvein either exserted as a
blunt or slightly tapering mucro or more characteristically
terminating in a swollen knob exserted not more than 0.2

390 Ећодога [Vol. 77

mm. Lower surface glabrous or very sparingly strigillose
especially along the midvein, moderately to densely beset
with conspicuous punctate glands often appearing to be of
two size classes or rarely nearly epunctate; upper surface
glabrous. Petiolules (1)2-3(4) mm long, glabrous or more
typically sparingly to moderately strigillose, conspicuously
and densely beset with markedly elevated pustulate glands;
stipels usually persisting, dark reddish or blackish, about
1-2 mm long. Racemes solitary or loosely clustered, 1-3(8),
about (0.5) 1.0-2.0 (3.0) dm long; rachis of inflorescence
very sparingly strigillose, eglandular or sparingly glandu-
lar-pustulate. Pedicels usually 0.6-1.2 mm long and spar-
ingly to moderately strigillose, eglandular; bracts caducous,
setaceous to narrowly lanceolate with a long-tapering apex,
sparingly to moderately strigillose, 1.5-8 mm long, glandu-
lar-pustulate. Calyx-tube funnelform to somewhat cam-
panulate, about 1.5-3 mm high, glabrous or sparingly to
moderately strigillose, moderately to more typically densely
beset with pustulate glands in upper two-thirds. Calyx-
lobes sparingly to densely strigillose with a dense fringe of
white trichomes, glandular-pustulate; adaxial lobes broadly
rounded to triangular-dentate and acute, about 0.2-0.5 mm
long; lateral lobes usually triangular dentate and acute to
acuminate but occasionally broadly rounded, 0.5-0.8 mm
long; adaxial lobe narrowly triangular-dentate, 0.8-1.2 mm
long. Vexillum about 4-6 mm long and about 4.0-4.5 mm
wide, entire to finely erose, bright blue to deep violet blue.
Filaments to 9 mm long, united for about half their length,
glabrous; anthers about 0.5-0.7 mm long, orange. Ovary
glabrous; style antrorsely pubescent. Fruit about 4.5-6 mm
long and 2-2.5 mm wide, with the adaxial suture straight
or slightly incurved near the apex and the abaxial suture
strongly outwardly bowed, glabrous or rarely sparingly
strigillose, conspicuously glandular-pustulate on the distal
three-fourths.

Distribution: A rare species of prairies, open woods and
creek banks of eastern Texas and Oklahoma. (Map 12.)

1975] Amorpha — Wilbur 391

13. Amorpha roemeriana Scheele, Linnaea 21: 461. 1848.
TYPIFICATION: “In margine rivulorum prope, Aus-
tin," Е. Roemer s.n. (holotype: not seen by me or by
anyone else who has commented upon it.)

А. fruticosa var. [1] subglabra A. Gray, Boston
Jour. Nat. Hist. 6: 174. 1850. TYPIFICATION :
Texas: on a ereek near Fredericksburg, June 1847,
Lindheimer s.n. (holotype, GH !). It is doubtful if
Gray intended or actually did publish this varietal
name. The type-style is different than that em-
ployed in this paper for varieties and its punctua-
tion is also different. The next entry in the
Plantae Lindheimerianae is also listed by Gray
as A. fruticosa var. subglabra.

A. laevigata var. pubescens А. Gray, Smithsonian
Contr. Knowledge 3: 49. 1852. TYPIFICATION :
“Eastern” Texas, Charles Wright s.n. (holotype,
GH!).

A. texana Buckley, Proc. Acad. Nat. Sci. Philadel-
phia 1861: 452. 1862. TYPIFICATION: Texas: “Оп
the Pierdenalis River," S. B. Buckley s.n. (holo-
type, PHIL !).

A. subglabra (A. Gray) Heller, Contr. Herb. Frank-
lin & Marshall Coll. 1: 48. 1895.

A. texana [хат] mollis Boynton, Biltmore Bot. Stud.
1: 139. 1902. A needlessly created “nom. nov."
based on “А. laevigata pubescens А. Gray."

A. laevigata var. pubescens f. mollis (Boynton) C.
К. Schneider, Bot. Gaz. 43: 307. 1907.

A. texana var. glabrescens E. J. Palmer, Jour. Arn-
old Arb. 12: 180. 1931. TYPIFICATION: Texas:
[COMAL со.] Comanche Spring; New Braunfels
etc., Lindheimer 743 (holotype, A!; isotypes, ARIZ l,
F!, GH!, MO!, NY!, OKL!, PHIL!, UC!, US).

Shrub 1-3 m tall. Current season's growth sparingly to
densely puberulent or strigillose or rarely glabrous, occa-

392, Ећодога [Vol. 77

sionally glabrate in age, sparingly to moderately glandular-
pustulate. Buds ovoid, compressed, the scales often some-
what keeled, sparingly to moderately puberulent. Leaves
strongly divergent, (0.5)1-1.5(2) dm long. Petioles (0.5)
1.5-3.0 cm long, about 1 mm in diameter, very sparingly to
occasionally densely puberulent or strigillose, sparingly to
moderately glandular-pustulate, usually equaling or longer
than the width of the lowermost leaflet. Stipules caducous
and rarely seen, linear, tawny puberulent, ca. 2 mm long.
Rachis of leaf slender, 1-1.6 mm in diameter or less, very
sparingly to moderately strigillose or puberulent or rarely
densely puberulent. Leaflets (7)9-11(15), oblong or more
typically broadly oblong to oblong-elliptic or rarely obovate
to suborbicular, usually (1.0)2.5-4.0(5.2) ст long and
(0.7) 1.5-2.5 (3.8) ст wide, typically (1.0) 1.3-2.9 (3.0) times
as long as wide, opposite or more commonly alternate,
typically symmetrical, usually broadly rounded basally and
apically either broadly rounded or more characteristically
emarginate and often conspicuously so; venation little ele-
vated beneath other than the midvein and to a slight degree
the secondary veins; margins entire to conspicuously cren-
ulate. Midvein rarely exserted as much as 0.8 mm long as
a tapering mucro but more typically terminating in a
slightly swollen knob at the margin or tapering less than
0.2 mm beyond. Lower surface usually sparingly to mod-
erately pilosulose or puberulent with fine, hyaline trichomes
or rarely glabrous or nearly so, usually inconspicuously
and moderately beset with small punctate glands; upper
surface glabrous or at least glabrate to moderately pilosu-
lose or finely puberulent especially along the principal
veins, Petiolules usually 2-5(7) mm long, glabrous to
densely puberulent or pilosulose, often becoming wrinkled
upon drying, usually sparingly to moderately glandular-
pustulate but rarely eglandular. Stipels inconspicuous,
often caducous, dark reddish, about 1.2-2 mm long. Ra-
cemes solitary or loosely clustered, 1-3(6) in number, about
(4)6-12(20) cm long; rachis usually sparingly to moder-
ately strigillose or puberulent, rarely densely tawny or

1975] Amorpha — Wilbur 393

hyaline puberulent or short-pubescent, sparingly glandular-
pustulate; pedicels mostly 1-1.5 mm long, usually sparingly
to moderately puberulent or strigillose; bracts caducous,
setaceous to narrowly lanceolate and then often with a
long-tapering apex, puberulent or pilosulose, 1.2-2.2 mm
long. Calyx-tube funnelform, usually 2.5-3.8 mm long, very
sparingly to moderately strigillose to pilosulose with hya-
line trichomes, moderately glandular-pustulate in upper
third; lobes moderately to more characteristically densely
short-pubescent above with a conspicuous fringe of white
trichomes, glandular-pustulate; adaxial lobes broadly
rounded to broadly triangular-dentate and acute, (0.2)
0.4-0.6(0.8) mm long; lateral lobes broadly rounded to more
characteristically acutely triangular-dentate, 0.5-0.8 (1.0)
mm long; abaxial lobe usually narrowly triangular-dentate,
about (0.8)1.0-1.2(1.4) mm long. Vexillum about 5-7 mm
long, 5-6 mm wide, emarginate and occasionally apiculate,
entire to slightly erose, purple. Filaments at maturity to
10 mm long, united for up to two-thirds their length, gla-
brous; anthers about 0.6-0.8 mm long. Style villous; ovary
glabrous. Fruit about 6-7 mm long and 2.5-3.5 mm wide,
plump at maturity, with the adaxial margin straight or
somewhat inwardly bent in the upper third and the abaxial
margin strongly outwardly bowed, glabrous to moderately
strigillose, conspicuously glandular-pustulate above. Seed
bright brown, about 3 mm long and 2 mm wide.

Distribution: All creek bed and stream banks of the “Hill
Country" of centra] Texas in the eastern portion of the
Edwards Plateau. (Map 13.)

There is no record in the literature that anyone has ex-
amined authentic material of A. roemeriana Scheele. А.
Gray stated (Boston Jour. Nat. Hist. 6: 175. 1850.) that
A. roemeriana was “doubtless a form of A. fruticosa or of
A. paniculata.” Later Gray (Proc. Acad, Nat. Sci. Phil.
1862: 162) indicated that A. roemeriana was apparently
the same as A. texana Buckl. which he synonymized with
A. laevigata var. pubescens. Watson (Smithsonian Misc.

894 Ећодога [Vol. 77

Coll. 258. 188. 1878), Schneider (Bot. Gaz. 43: 307. 1907)
Rydberg (N. Am. Fl. 24: 27. 1919) and Palmer (Jour.
Arnold Arb. 12: 179. 1931) have all listed Scheele’s A.
roemeriana as a synonym of A. paniculata T. & G.

Unfortunately for the sake of stability, it would seem
most unlikely that A. roemeriana can be considered a
synonym of A. paniculata. A. paniculata is a largely East
Texas species of thickets and low swampy or marshy
ground. A. roemeriana was originally collected along a
creek bend near Austin which is its eastern range limit.
Amorpha texana has been collected in Travis Co. while
А. paniculata is not known from the vicinity. The original
description itself is far more suggestive of the central
Texan species than it is of more easternly ranging A.
paniculata. I have no doubt that A. roemeriana is an
earlier name for the species recently known as А. texana.

Palmer (1931) recognized a glabrous or subglabrate and
a spreading pubescent variety of this species. There seems
to be no geographic segregation of these pubescence types
and actually there seems to be as much of a continuum in
vestiture as one could expect in so small a number of col-
lections as are available of this species. I concur with
Turner (The Legumes of Texas, p. 143, 1959) and Correll
and Johnston (Man. Vasc. Pl. Tex., p. 818, 1970) who did
not recognize the varieties either. Both of these taxonomic
works suggest that A. laevigata is not specifically distinct
from A. texana, Turner suggesting that A. laevigata “per-
haps is best treated as a narrow-leaved eastern variety of
A. texana" while Correll and Johnston indicated that it was
*perhaps only a form of A. texana.” I believe the differ-
ences in petal color, calyx shape, texture and glandularity
etc. all indicate considerable morphological divergence be-
tween the two taxa.

14. Amorpha ouachitensis Wilbur, sp, nov. TYPIFICATION:
Oklahoma: LE FLORE CO., dry, rocky (sandstone)
slopes of Black Fork Mountain, near Page, Palmer
20572 (holotype, A!; isotypes, GA!, NY!, Us!).

1975] Amorpha — Wilbur 395

Frutex erectus 1-2 m altus. Foliola (7)9-13(17),
lateralia oblonga vel oblongo-elliptica, opposita, basi
et apice rotundata vel emarginata, (1.2) 2.5-4.0 (7.0)
cm longa et (0.9) 1.5-2.5 (3.6) cm lata. Tubus calycis
28.3.9 mm altus, + glaber, glanduloso-punctatus
in quarta parte superiore. Lobi calycis glabri vel
pubescentes; lobi abaxiales 0.4-0.6 mm longi; lobi
laterales 0.5-0.7 mm longi; lobi adaxiales 0.6-0.9
(1.2) mm longi. Vexillum 5-7.5 mm longum, rubi-
cundo-purpureum.

Shrub 1-2 m tall. Current season's growth glabrous or
more typically very sparingly to moderately pilosulose and
sparingly pustulate glandular with inconspicuous lenticular
glands. Buds ovoid to almost globose with the scales mod-
erately appressed pubescent on the outer surface and api-
cally densely ciliate with tawny-villous trichomes. Leaves
strongly divergent, 7-23 cm long. Petioles 1.2-2(2.4) ст
long and about 0.5-1.5(2) mm in diameter, moderately
puberulent or short-pilose with hyaline to tawny trichomes,
sparingly to moderately glandular-pustulate, usually equal-
ling or longer than the width of the lowermost leaflet.
Stipules caducous, lanceolate, densely tawny villous api-
cally, ca. 3-4 mm long including the apical tuft but only
about 2 mm long excluding the apical villosity. Rachis of
leaf mostly 1 mm in diameter or less, very sparingly
pilosulose to even glabrous, sparingly glandular-pustulate.
Leaflets (7)9-13(17), typically broadly oblong to oblong-
elliptic or oblong, mostly (1.2) 2.5-4.0(7.0) ст long and
(0.9) 1.5-2.5 (3.6) ст wide, typically (1.4) 1.5-2 (2.4) times
as long as wide, usually opposite, typically symmetrical,
usually broadly rounded basally and apically character-
istically conspicuously emarginate to broadly rounded;
venation but little elevated beneath other than the mid-
vein; margins entire to inconspicuously undulate-crenate.
Midvein exserted either as a tapering mucro about 0.2-
0.6(1.0) mm or only slightly exserted and terminating in a
swollen knob. Lower surface of the leaflets glabrous or

396 Rhodora [Vol. 77

glabrate to sparingly or even moderately appressed pilosu-
lose or less commonly spreading pilosulose or short pubes-
cent with fine, hyaline to pale tawny, slender trichomes
and usually conspicuously beset with numerous pustulate,
amber-colored glands; upper surface glabrous or glabrate
to moderately pilosulose with inconspicuous, slender tri-
chomes and lacking pustulate glands. Petiolules mostly 2-3
(4) mm long, glabrous or glabrate to moderately short-
pubescent or pilosulose, sparingly to moderately glandular-
pustulate or rarely eglandular. Stipels inconspicuous, some-
times caducous, drying dull dark reddish-brown, mostly
1.2-2.2 mm long, glabrous or sparingly appressed pilosulose.
Racemes solitary or in loose clusters of 2-4, mostly (8)
10-20 em long; rachis of inflorescence glabrous vo sparingly
or even moderately spreading short-pubescent with hyaline
or very rarely tawny trichomes and eglandular or very
sparingly and inconspicuously pustulate-glandular ; pedi-
cels about 1-1.5 mm long, glabrous to sparingly puberulent;
bracts caducous, narrowly lance-oblong, about 1-1.5 mm
long, marginally short-pubescent with stiff, hyaline tri-
chomes. Calyx-tube narrowly conical to funnelform, about
2.8.3.2 mm long, usually glabrous or nearly so to moder-
ately puberulent or spreading short-pubescent, sparingly
glandular pustulate in the upper quarter; lobes usually
glabrous but occasionally moderately appressed pubescent
on the external surface and densely margined with a con-
spicuous fringe of white trichomes, eglandular to sparingly
pustulate glandular; adaxial lobes broadly rounded, mostly
0.4-0.6 mm long; lateral lobes broadly rounded or perhaps
more characteristically broadly triangular-dentate, about
0.5-0.7 mm long; abaxial lobe narrowly triangular-dentate
with an acute to acuminate apex, about 0.6-0.9(1.2) mm
long. Vexilum about 5-7.5 mm long and 4.5-6 mm wide,
strongly emarginate but otherwise entire, purple. Fila-
ments at maturity 8-10 mm long, united for up to 3/5 their
length, glabrous; anthers about 0.6-0.8 mm long. Style
antrorsely pilosulose; ovary usually glabrous. Fruit about
1-9 mm long and 3-4 mm wide and with the adaxial margin

1975] Amorpha — Wilbur 397

either straight or more typically mostly straight but with
the upper portion bent sharply abaxially and the abaxial
margin strongly outwardly bowed, glabrous or occasionally
somewhat appressed, short-pubescent and conspicuously
pustulate-glandular. Seed solitary, dark brownish to black-
ish, about 4-5 mm long and 2.5-3 mm wide.

Distribution: Ouachita Mountains of west-central Arkan-
sas and southeastern Oklahoma. (Map 14.)

This species has previously been included within the
concept of other species. Palmer cited specimens of it as
А. glabra Ројт. (Jour. Arnold Arb. 12: 175. 1931) and also
A. virgata (Jour. Arnold Arb. 12: 182. 1931) which I have
treated as a synonym of A. fruticosa. Amorpha glabra 15
in my opinion a southern Appalachian endemic while the
range of A. fruticosa s.lat. spans the continent. I believe
the morphology of A. owachitensis indicates a closer rela-
tionship with the rare central Texan endemic A. roemeriand
(= A. terana) than with either A. glabra or A. fruticosa.

Representative Specimens: Arkansas: CONWAY CO., rocky moun-
tainsides, Petit Jean Mt., Demaree 37161 (SMU); GARLAND CO., shale
outcrop on banks of Ouachita River near Hot Springs, Palmer 24252
(А); LOGAN CO. rocky glades, top of Magazine Mt., Palmer 24187
(A, MO, UARK) ; MONTGOMERY CO., Mount Ida, open banks of Ouachita
River below bridge, Demaree 57965 (SMU); POLK CO., Rich Mt., rich
woods E of lodge near Lover's Leap, T'ucker 12285 (DUKE). Okla-
homa: LE FLORE CO., Ouachita National Forest, Goodman 2545 (GH,
ISC, MO, NY, OKL), rocky slopes of Blackfort Mt., near Page, Palmer
20913 (A); MCCURTAIN CO.: rocky sandy soil in woods 37 mi. N of
Broken Bow, Stratton 1146 (OKL); PUSHMATAHA CO., rocky stream
bank, 4 mi. W of Albion, Stratton 4161 (LL, OKL).

15. Amorpha fruticosa L., Sp. Pl. 2: 713. 1753.

A. fruticosa а vulgaris Pursh, Fl. Am. Sept. 2: 466.
1814.

A. fruticosa В emarginata Pursh, Fl. Am. Sept. 2:
466. 1814.

A. fruticosa y angustifolia Pursh, Fl. Am. Sept. 2:
466. 1814.

398

Rhodora [Vol. 77

A. croceolanata Wats., Dendr. Brit. 2: t. 139. 1825
[as crocealanata].

A. emarginata (Pursh) Sweet, Hort. Brit. 121. 1827.

A. fragrans Sweet, Brit. Fl. Gard. 3: t. 241. 1828.

A. caroliniana Croom, Amer. Jour. Sci. 25: 74. 1834.
TYPIFICATION: North Carolina: near New Bern,
Loomis s.n. (lectotype, NY!).

A. fruticosa var. 5 coerulea Loudon, Arb. Brit. 607.
1838.

A. humilis Tausch, Flora 21: 750. 1838. TYPIFICA-
tion: undetermined; based on cultivated plants
grown in Prague.

A. fruticosa var. 4 Lewisii Loudon, Arb. Brit. 2: 607.
1838.

A.tennessensis Shuttleworth ex Kunze in Delect.
Sem. Hort. Lips. 1848: 1. 1848; Linnaea 24: 191.
1851. TYPIFICATION: Tennessee: Ad rivulos prope
Dandridge, Rugel s.n., June 1842 (isolectotype,
GH!).

A. fruticosa var. caroliniana (Croom) 5. Wats.,
Smithsonian Misc. Coll. 258. 188. 1878.

A. fruticosa var. стара Kirchn., Arb. Musc. 370.
1864.

А. pendula Carr., Rev. Hort. 1870-71: 378.

A. fruticosa var. pendula (Carr.) Dipp., Laubh. 3:
691. 1893.

A. fruticosa f. albiflora Sheldon, Bull. Geol. & Nat.
Hist. Surv. Minnesota 9: 72. 1894. TYPIFICATION:
Minnesota: MILLE LACS СО., on the banks of the
Rum River, 3 miles north of Milaca, Sheldon
s.n. (holotype, MIN!).

A. virgata Small, Bull. Torrey Club 21: 17. pl. 171.
1894. TYPIFICATION: Georgia: DE KALB CO., NW.
slope of Stone Mountain, alt. 1000-1400 ft., Small
s.n. 3 July 1893 (lectotype, NY!; isolectotypes,
GA!, GH!, ISC!, MIN!, vol uc!, Us!).

A. fruticosa var. croceolanata (Wats.) Mouillefort,
Arb. Arbriss. 1: 577. 1894.

1975]

Amorpha — Wilbur 399

A. angustifolia (Pursh) Boynton, Biltmore Вог.
Stud. 1: 139. .1902.

A. fruticosa f. aureo-variegata Schwerin, Mitteil.
Deutsch Dendr. Ges. 16: 255. 1907.

A. fruticosa var. typica Schneider, Bot. Gaz. 43:
304. 1907.

A. fruticosa f. crispa (Kirchn.) C. К. Schneider,
Illustr. Hand. Laubh. 2: 72. 1907.

A. fruticosa f. pendula (Carr.) C. К. Schneider,
Illustr. Handb. Laubh. 2: 75. 1907.

A. fruticosa var. humilis (Tausch) Schneider, Bot.
Gaz. 43: 305. 1907.

A. occidentalis Abrams, М. Y. Bot. Gard. Bull. 6:
394. 1910. TYPIFICATION: California: San Diego
River, near Old San Diego Mission, Abrams 3425
(holotype, Ds, not seen; isotypes, A!, DS!, GH !, MO},
NY !, РОМ !, UC!, US!)

A. Bushii Rydb., N. Am. Fl. 24: 31. 1919. TYPIFICA-
TION: Florida: uncommon near spring, Chatta-
ћоосћее River, Bush 13 (holotype, NY!; isotype:
AT)

A. Curtissii Rydb., N. Am. Fl. 24: 30. 1919. TYPIFI-
CATION: Florida: DUVAL CO. low ground near
river, Jacksonville, Curtiss 4703 (holotype, му!;
isotypes, KANU!, US!)

A. arizonica Rydb., N. Am. Fl. 24: 33. 1919. TYPI-
FICATION: Arizona: COCHISE CO., along streams,
Ramsey Canyon, Huachuca Mountains, Gooding
136 (holotype, NY!; isotypes: ARIZ!, GH}, NEB],
NY!,RM!, uc!).

A. occidentalis var. arizonica (Rydb.) E. J. Palmer,
Jour. Arnold Arb. 12: 185. 1951.

A. fruticosa f. humilis (Tausch) E. J. Palmer, Jour.
Arnold Arb. 12: 189. 1931.

A. fruticosa f. coerulea (Loud.) E. J. Palmer, Jour.
Arnold Arb. 12: 189. 1981.

400

Ећодога [Vol. 77

А. fruticosa var. angustifolia f. glabrata Е. J.
Palmer, Jour. Arnold Arb. 12: 191. 1931. TYPIFI-
CATION: Texas: BRAZOS CO., shores of lake, Kurten,
Palmer 13479 (holotype: A!; isotype, мо!).

A. fruticosa var. tennesseensis (Shuttleworth ех
Kunze) E. J. Palmer, Jour. Arnold Arb. 12: 192.
1931.

A. fruticosa var. oblongifolia Е. J. Palmer, Jour.
Arnold Arb, 12: 192. 1931. TYPIFICATION: Arkan-
sas: PHILLIPS CO., low alluvial ground, along Mis-
sissippi River, near Helena, Palmer 26628 (holo-
type, A!; isotypes, DS!, MO!, UARK!).

A. occidentalis var. emarginata Е. J. Palmer, Jour.
Arnold Arb. 12: 185. 1931. TYPIFICATION: Ari-
zona: [MARICOPA CO.] Fish Creek, Apache Trail,
Eastwood 8745 (holotype, A!; isotype, CAS!).

A. emarginata Eastwood, Proc. Calif. Acad, Sci. 20:
148. 1931, not A. emarginata (Pursh) Sweet,
1827. TYPIFICATION: Arizona: [MARICOPA Col
Fish Creek, Apache Trail, Eastwood 8745 (holo-
type, CAS!; isotype: A!).

А. DeWinkeleri Small, Man. Se. Fl. 689. 1933.
TYPIFICATION: Florida: LEE CO., prairies near Ft.
Shackleford, Big Cypress, Small 8349 (lectotype,
NY!; isolectotypes, GH!, MO!, US!).

A. fruticosa var. augustifolia f. latior Fassett, Rho-
dora 38: 190. 1936. TYPIFICATION: Wisconsin:
ST. CROIX CO., stony shore of Lake St. Croix, 10
miles south of Hudson, Fassett 17014 (holotype,
WIS, not seen; isotypes, GH!, MO!, NY!).

A. fruticosa var. occidentalis (Abrams) Kearney &
Peebles, Jour. Washington Acad. Sci. 29: 483.
1939.

An erect shrub (1)2-3(4) m tall with one to several
stems arising from the base and often branching above
sufficiently so as to appear bush-topped. Current season’s
growth rarely glabrous or densely to moderately pilosulose

1975] Amorpha — Wilbur 401

to puberulent or strigillose, occasionally becoming glabrate;
eglandular or very sparingly and inconspicuously pustulate-
glandular, grayish to dark reddish-brown, occasionally
somewhat glaucescent. Buds ovoid to subglobose, light
brownish to dark reddish-brown, usually more or less gla-
brous except for the ciliate margins of the scales or occa-
sionally pilosulose. Leaves wide-spreading to somewhat
ascendant, mostly 1-2.8 dm long, usually remaining green
but occasionally turning brownish upon drying but not
blackening. Petioles moderately puberulent to crispy pilo-
sulose and often becoming glabrate, rarely glabrous, eg-
landular or sparingly pustulate with small, inconspicuous
glands, typically longer than the width of lowermost leaflet,
mostly 1-4 cm long. Stipules caducous, narrowly linear,
eglandular or very sparingly and inconspicuously pustulate,
reddish-brown, externally and especially apically tawny to
hyaline pilosulose, usually 2-4 mm long. Rachis of leaf
about 0.5-1 mm in diameter, rarely glabrous or more typi-
cally sparingly fo densely puberulent to crispy pilosulose
or glabrate, eglandular or very sparingly and inconspicu-
ously pustulate-glandular. Leaflets 9-21 (31), oblong to
elliptic-oblong or elliptic or even occasionally ovate, (1) 2-
4(5) em long and (0.5) 1-2(2.7) ст wide, mostly 2-3 (6)
times as long as wide, usually opposite or subopposite,
basally symmetrical ог rarely asymmetrical, acute to
broadly rounded basally, rounded to acute or rarely emargi-
nate apically, entire or nearly so; secondary venation mod-
erately elevated beneath. Midvein usually slender, exserted,
about 0.5-1.5 mm long and tapering, but occasionally only
very shortly exserted and little, if at all, tapering. Lower
surface of leaflets sparsely to densely spreading puberulent
to crispy pilosulose (occasionally so densely so as to appear
velvety) when young or rarely either glabrous or some-
times glabrate in fruit, often strigose or strigillose or oc-
casionally even short-spiculate with the trichomes either
hyaline, or ashy or even tawny; eglandular or inconspicu-
ously beset with small pustulate glands or very rarely
rather conspicuously glandular-punetate; upper surface

402 Rhodora [Vol. 77

usually very short-puberulent to glabrous when young and
typically glabrate when mature but occasionally densely
pilosulose, eglandular, Petiolules about (1.5)2-4 mm long,
sparingly to densely puberulent or more typically spread-
ing pilosulose, occasionally becoming glabrate, or very
rarely glabrous, usually inconspicuously pustulate-glandu-
lar. Stipels slenderly cylindric, setaceous mostly 2-4 mm
long, glabrous to moderately puberulent. Racemes erect,
solitary or more typically with several to many clustered
together, usually densely flowered, (0.5)1-2(2.5) dm long;
rachis of inflorescence usually moderately to densely pu-
berulent or pilosulose, sometimes becoming glabrate in age,
eglandular or very sparingly and inconspicuously pustulate-
glandular. Pedicels mostly 1-2.0 mm long, sparsely to
densely spreading puberulent or more typically crispy pilo-
sulose or short-spiculate or even glabrous, eglandular or
very sparingly and inconspicuously pustulate-glandular;
bracts caducous, borne basally on the pedicels, very nar-
rowly linear to setaceous, mostly 1.5-3 mm long, sparingly
to densely pilosulose externally, usually eglandular, typi-
cally dull reddish-brown. Calyx-tube usually obconic but
varying to funnelform or even somewhat campanulate,
mostly 2-3(4) mm long, glabrous or spiculate, or sparingly
to densely puberulent or pilosulose, sparsely to moderately
pustulate-glandular on the upper third of tube with small
and inconspicuous to large and conspicuous amber-colored
glands but occasionally eglandular; lobes with a hyaline,
ciliate-fringed margin and with the external surface spar-
ingly to moderately pilosulose and either eglandular or very
sparingly beset with inconspicuous pustulate glands; adax-
ial lobes broadly rounded to broadly triangular-dentate,
about 0.2-0.5 mm long; lateral lobes usually acute and tri-
angular-dentate but occasionally broadly rounded, about
0.3-0.8(1.0) mm long; abaxial lobe narrowly triangular-
dentate, acute, (0.5)0.8-1.2 mm long. Vexillum about 5-6
mm long about 3.5-4.2 mm wide, broadly obovate, gradually
tapering to a rather indistinct claw of about 1-1.5 mm long,
moderately to strongly arched, apically broadly rounded

1975] Amorpha — Wilbur 403

and entire or indistinctly emarginate to irregularly erose,
appearing dark reddish-purple either fresh or dried and
with the claw and blade enfolding and enveloping the fila-
ments and style. Filaments 6-8 mm long, united into a
sheath for the lower 1-2(3) mm, glabrous; anthers 0.4-0.6
mm long, yellow. Style moderately ascending pilose, 5-7
mm long; ovary glabrous or rarely pubescent. Fruit 5-9
mm long and (2.0)3-4.5 mm wide, basally tapering, straight
to strongly curved along the abaxial suture and the abaxial
suture strongly outwardly bowed, glabrous to densely ap-
pressed short-pubescent, conspicuously pustulate-glandular
or eglandular. Seeds smooth, reddish-brown, 3.5-4.5 mm
long.

Distribution: Southern Quebec south into Florida and
west into northern Mexico, southern California, and Wyo-
ming. (Map 15.)

This wide-ranging species is, as one might infer from its
lengthy synonymy, extremely variable. Numerous species
and/or varieties have been segregated from it in the past
and many have been recognized in most recent floristic
treatments. The striking variability 18 such that it might
seem reassuring indeed to have a formally designated sys-
tem by which one could categorize the very dissimilar
appearing plants. Unfortunately, however, the examination
of many hundreds of specimens soon demonstrates the
inadequacy of even the numerous described segregates to
convey the extent of the variability encountered not to
mention the not inconsiderable number of specimens which
clearly match the characteristics of one described taxon
in its older growth and another in its younger and more
recently formed portions. Clearly then what might be con-
veniently referred to as the fruticosa-complex is ап ех-
tremely diverse assemblage of populations whose variability
is due both to environmentally induced plasticity and also
to the presence of a great number of biotypes. I have found
it taxonomically unclarifying to attempt to categorize this
variability formally by the recognition of the previously

[Уо]. 77

Rhodora

404

"овог утај оуалошү

ст dVW

1975] Amorpha — Wilbur 405

segregated species, varieties and forms and have concluded
that the variability could best be treated as representing
one extremely polymorphic taxon. The alternate approach
is exemplified by Steyermark’s treatment in the flora of
Missouri where as many as four of the five varieties recog-
nized as occurring in Missouri are known from a single
county. No indication of ecological preference is suggested
to account for at least partial isolation of these populations
and they are in fact stated to be “intergrading.”

TAXA EXCLUDED FROM THE GENUS
AND BINOMIALS AND TRINOMIALS
OF UNCERTAIN POSITION

Certain binomials or trinomials have in the interests of
completeness been included in the following list which per-
haps could have been excluded since they were not validly
published. Names published without a diagnosis or descrip-
tion or without a reference to one that was effectively pub-
lished are nomina nuda and hence technically could be
ignored (ICBN Art. 32(3)). Also binomials or trinomials
which were originally published in synonymy are not
validly published (ICBN Art. 34(4)) and hence could be
ignored (Art. 12).

Amorpha arborea Hort. ex Schkuhr, Bot. Handb. Deutschl.
Gew. 2: 333. 1796. [Publication not seen by me; this
binomial was reportedly published as a synonym.]

Amorpha canescens [f.] glabrescens Zabel in Beissner et al.,
Handb. Laubh. Ben. 268. 1903. [Publication not seen by
me; reportedly this bionomial was published as a nom.
nud. and a synonym of A. canescens f. glabrata (A. Gray)
N. С. Fassett.]

Amorpha colorata Raf., Aut. Bot. 81. 1840. [Appalachian
Mts.] Rafinesque himself questioningly suggested that it
might be А. caroliniana Croom, and Merrill (Index
Rafimesquianus, p. 142. 1949) suggested, again with a
question mark, its identity with A. nitens Boynton. I
am unable to identify it with any certainty.

406 Ећодога [Vol. 77

Amorpha crocea Hort. ex Lavallée, Arb. Segrez. 60. 1877.
[Publication not seen by me but this binomial reportedly
appeared as a nom. nud. ]

Amorpha dealbata Hort. ex Lavallée, Arb. Segrez. 60. 1877.
[Publication not seen by me but this binomial reportedly
appeared as a nom. nud. |

Amorpha discolor Raf., Aut. Bot. 79. 1840. [ Alabama. ]
Merrill (Index Rafinesquianus, р. 142. 1949.) synony-
mized this binomial with A. fruticosa L.

Amorpha elata Hayne, Dendr. Fl. 134. 1822. [Publication
not seen by me. Rydberg cited this binomial in the syn-
onymy of A. fruticosa L.]

Amorpha elatior Hort. ex Lavallée, Arb. Segrez. 60. 1877.
[Publication not seen by me but this binomial is report-
edly a nom. nud.]

Amorpha flexuosa Raf., Aut. Bot. 80. 1840. [Unaka Mts. of
Carolina.] Perhaps a synonym of A. fruticosa L. as sug-
gested by Merrill (Index Rafinesquianus, p. 142. 1949).

Amorpha fruticosa var. fragans Bean, Trees and Shrubs
Brit. Isl. 1: 198. 1914. [Publication not seen by me;
Palmer states that the brief description would apparently
exclude it from synonymy of A. fragrans Sweet but that
its identity is undeterminable.]

Amorpha gaertnerii Hort. ex К. Koch, Dendr. 1: 70. 1869.
[Publieation not seen by me; reportedly a nom. nud. ]
Amorpha gardnerii Hort. ex. Kirchner, Arb. Muse. 370.
1864. [Publication not seen by me; reportedly a nom.

nud.]

Amorpha glandulosa Blanco, Flora de Filipinas. ed. 1. 555.
1837. [Philippines.] = Dalaea glandulosa (Blanco)
Merr., Dept. of Interior, Manila. 37. 1905. = Thornbera
dalea (L.) Rydb., Fl. N. Am. 24(2) : 120. 1920. = Dalea
annua (Mill) Kuntze, Rev. Gen. 178. 1891.

Amorpha glauca Raf., Aut. Bot. 80. 1840. [Missouri.] A
synonym of A. fruticosa L. s.lat. as suggested by Merrill
(Index Rafinesquianus, p. 142. 1949.)

1975] Amorpha — Wilbur 407

Amorpha Lewisii Loddiges ex Loudon, Arb. Brit. 2: 607.
1838. [Publication not seen by me; reportedly published
in synonymy. |

Amorpha ludoviciana Hort. ex Lavallée, Arb. Segrez. 60.
1877. [Publication not seen by me but this binomial is
reportedly a nom. nud. |

Amorpha Ludwigii Hort. ex К. Koch, Dendr. 1: 70. 1869.
[Publication not seen by me; reportedly published in
synonymy. |

Amorpha ? lutea Raf., Fl. Ludov. 105. 1817. [ Louisiana. ]
Clearly not an Amorpha but its identity has not vet been
established.

Amorpha macrophylla Raf., Aut. Bot. 79. 1840. [ Florida. ]
Probably another synonym of А. fruticosa L. s.lat. as
suggested by Merrill (Index Rafinesquianus, p. 142.
1949.). The identity of this name is totally impossible to
surmise if one were following Palmer's treatment with
its numerous accepted segregates from the Amorpha
fruticosa complex.

Amorpha marginata Hort. ex Lavallée, Arb. Segrez. 60.
1877. [Publication not seen by me but reportedly the
binomial appeared as a nom. nud.]

Amorpha non-perforata Schkuhr, Bot. Handb. Deutschl.
Gew. 2: 333. 1796. [Publication not seen. Usually listed
in the synonymy of A. fruticosa L.]

Amorpha ornata Wenderoth, Ind. Sem. Hort. Marburg.
1835. [Publication not seen by me; reportedly a nom.
nud. |

Amorpha отапа М. Е. Jones, Contr. West. Bot. по. 16:
32. 1930. Typification: Arizona: Miller Canyon, Hua-
ећиса Mts., М. E. Jones 25027 (holotype, not seen; iso-
types, CAS!, мо!, OKLA!, RM!, UC!) = Indigofera sphae-
rocarpa A, Gray.

Amorpha pedalis Blanco, Flora de Filipinas, ed. 1. 553.
1837. [Philippines.] According to Merrill (Dept. of In-
terior, Manila. 19. 1905) this is probably Solominia
oblongifolia DC. (Polygalaceae).

408 Ећодога [Vol. 77

Amorpha perforata Schkuhr, Bot. Handb. Deutschl. Gew. 2:
333. 1796. [Publication not seen. Usually listed in the
synonymy of A. fruticosa L.]

Amorpha rabiae Lexarza, Nov. Veg. Desc. fasc. I. 22. 1824.
[Michoacan, Mexico.] Palmer (Jour. Arn. Arb. 12: 197.
1931) presents a resumé of the original description. It
does not possess the characters of an Amorpha but its
identity is otherwise uncertain.

Amorpha retusa Raf., Aut. Bot. 80. 1840. [Appalachian
Mts.] Probably a snyonym of A. fruticosa L.; Rafines-
que's description mentioning its tomentose leaflets and
acute calyx-lobes would certainly exclude the possibility
of its being A. glabra as was suggested by Merrill (Index
Rafinesquianus, p. 142. 1949) and Rehder (Bibliogr.
Cult. Trees and Shrubs. 369. 1949).

Amorpha tomentosa ? Raf., Fl. Ludov. 105. 1817. [Louisi-
ana.] Identity uncertain as all that Rafinesque states
about it is “а doubtful species, of which Robin only men-
tions it being tomentose...” Rydberg (N. Am. Fl. 24
(1): 27. 1919? questioningly placed it in the synonymy of
A. paniculata but the description is far too incomplete
to allow anyone to identify the plant being described. It
is not even certain that it is an Amorpha.

Amorpha tomentosa Raf., Aut. Bot. 81. 1840. not Raf., FI.
Ludov. 105. 1817. [“Оп Wabash and West Kentucky."]
Merrill (Index Rafinesquianus, p. 142. 1949) question-
ingly suggests that it might be a synonym of A. canes-
cens Pursh but that species is unknown to me from
“West Kentucky" one of the two cited localities.

Amorpha verrucosa Raf. Aut. Bot. 80. 1840. [Alabama.]
Merrill (Index Rafinesquianus, p. 142. 1949) suggested
this binomial was a synonym of A, fruticosa L. and I
would agree that the original description is vague enough
to fit the very broad concept I have of the species, al-
though it would be impossible to tel] to which of the
segregates recognized by Palmer from the A. fruticosa
complex that Rafinesque's species belongs.

1975] Amorpha — Wilbur 409

LITERATURE CITED

ABRAMS, L. 1944. Illustrated Flora of the Pacific States. Amorpha
in 2: 555. Stanford Univ. Press.

MUNZ, P. A. 1959. A California Flora. Amorpha on p. 852. Univ.
of California Press.

PALMER, E. J. 1931. Conspectus of the Genus Amorpha. Jour.
Arnold Arb. 12: 157-197.

RYDBERG, P. A. 1919. [Treatment of Amorpha] in the N. Amer.
Fl. 24: 26-34.

SCHNEIDER, C. K. 1907. Conspectus Generis Amorphae. Bot. Gaz.

13: 297-307.
DEPARTMENT OF BOTANY

DUKE UNIVERSITY
DURHAM, N.C. 27706

PAPAVER IN NORTH AMERICA
NORTH OF MEXICO!

ROBERT W. KIGER*

The following diagnostic treatment includes the native
and introduced species of Papaver found in North America
north of Mexico. It is primarily synthetic in nature and
is based in large part on the works cited under “Litera-
ture,” as well as on the collection of specimens in the
United States National Herbarium, Smithsonian Institu-
tion. This treatment is not offered as a definitive revision,
the achievement of which will require extensive study in
the field, herbarium and laboratory. In some cases, exist-
ing collections are very inadequate, and this is especially
true of the problematic scapose poppies. In view of the
present confused situation with regard to the latter in
North America, a broad species concept is here adopted
for them. In this arctic and alpine complex many species
of dubious merit have been proposed, some on the basis
of only one or a few specimens and with little regard for
the overall generic context. Much importance has been
accorded chromosome numbers in some instances, but until
the complex cytological patterns in Papaver are fully in-
vestigated and their relationships to morphological patterns
analyzed, such data contribute little to sound taxonomic
circumscriptions and dispositions,

When the native North American species of Papaver
are better known, it is probable that some taxa herein
submerged in broadly conceived species will be recognized
separately at the specific and varietal levels. Until such
time, however, practicality and fidelity to what is actually

‘Based on a study conducted for the Flora North America Program,
supported by National Science Foundation Contract C-757 with the
Smithsonian Institution.

?Present address: Hunt Institute for Botanical Documentation,
Carnegie Mellon University, Pittsburgh, Pennsylvania 15213.

410

1975] Papaver — Kiger 411

known about the native poppies seem best served by a
synoptic approach utilizing these broad taxa. I have made
no attempt to relate the introduced poppies to the many
infraspecific taxa that have been proposed for those species
in their native ranges. This would be virtually impossible
on the basis of the morphologies of most individual plants
found in North America. Any such correlations which
could be made would be essentially meaningless in the
North American context, within which, due to geographic-
ally random introductions and subsequent hybridizations,
there is no biologically significant pattern of variation in
evidence.

The sectional nomenclature utilized herein follows that
presented in my recent review of the subject (Kiger, 1973).
The circumscriptions and descriptions of sections generally
follow the traditional ones of existing revisions, particu-
larly that by Fedde (1909). Only characters of species
found in North America north of Mexico are included in
the sectional descriptions. The descriptions of all taxa
follow the outline and glossary developed for Flora North
America (Porter, et al., 1975).

Papaver L., Sp. Pl., 506. 1753; Gen. Pl ed. 5, 224. 1754.

Plants annual or perennial, herbaceous, caulescent, sub-
scapose or scapose, monoclinous, sap gummy, opaque. Main
stems eramous or ramose. Leaves alternate, sometimes
basally rosulate; blades entire to bipinnatipartite. Flowers
solitary, pedicellate or scapose, actinomorphic, complete,
synearpous, apostemonous. Sepals 2, free, caducous, entire.
Petals 4, entire, aestivation corrugate, Stamens numerous.
Carpels 3-18; styles absent; stigmas 3-18, sessile, radiate
on a dise, interstigmatie membrane usually present, often
conspicuous. Fruits capsular, dehiscence poricidal, pores
subapical, 3-18, locules 3-18.

412 Rhodora [Vol. 77

KEY TO SPECIES

a. Plants caulescent or rarely subscapose, at least a few
cauline leaves present. b. |
b. Upper саш пе leaves amplexicaulous. ............
e, 1. P. somniferum.
b. Upper cauline leaves not amplexicaulous. с.
с. Ovaries and capsules setose. .. 5. P. hybridum.
c. Ovaries and capsules glabrous. d.
d. Plants perennial; stems eramous; petals 5 ст
or more long. ............ 6. P. orientale.
d. Plants annual; stems ramose; petals less than
5 em long. e.
e. Stigmatic dise depressed-conic, usually
umbonate. ......... 4. P. californicum.
e. Stigmatic disc essentially plane. f.

f. Pedicels markedly patent hispid dis-
tally; capsules less than twice as long
as broad. ............. 2. P. rhoeas.

f. Pedicels strongly appressed hispid
distally; capsules twice as long as
broad. ............... 3. P. dubium.

a. Plants scapose, leaves all basal. g.
g. Leaves entire to 3 (rarely 5)-lobed, essentially gla-
brous; capsules obovoid-obconic. ... 9. P. walpolei.
g. Leaves pinnatipartite or bipinnatipartite with more
than 5 lobes, variously vestite, at least when young;
capsules narrowly clavate to globose. h.
h. Setae on ovaries and capsules basally tubercu-

late. 1.
i. Petals salmon pink to yellow, paler basally;
Rocky Mountains. ......... 7. P. alpinum.
i. Petals white to rose with yellow basal spot;
Alaska. ................ 8. P. alboroseum.
h. Setae on ovaries and capsules not basally tuber-
culate. j.

j. Stigmatic disce distinctly conic and/or um-
bonate. k.

о

1975] Papaver — Kiger 413

k. Leaves mostly bipinnatipartite; capsule
obovoid. ............ 11. P. meconnellit.
k. Leaves mostly pinnatipartite; capsule nar-
rowly oblong-cylindric to clavate. ......
10. P. тасонттп.

i. Stigmatic dise essentially plane. 1.

l. Mature plants over 25 ст tall; petals 4-6
em long; scapes glabrate to sparsely ves-
tite, ................. 13. P. nudicaule.

|l Mature plants less than 25 ст tall; petals
to 4 (rarely to 5) em long; scapes moder-
ately to densely vestite. ................

12. P. lapponicum ssp. occidentale.

SECT. PAPAVER

Plants annual, caulescent, glaucous, glabrate. Leaf
blades simple or pinnatipartite. Basal leaves petiolate. Cau-
line leaves sessile, upper amplexicaulous. Filaments cla-
vate. Stigmatic dise essentially plane. Fruits globose or
subglobose, glaucous, glabrous.

1. Р. somniferum L., Sp. Pl., 508. 1758.

Plants 8-15 dm tall. Main stems distally cauliramous,
stout; branches few. Leaf blades simple and coarsely bi-
dentate, or pinnatipartite with lobes irregularly dentate.
Pedicels glabrate or sparsely pale setose distally. Petals
white, pink, red or purple, darker spotted basally, 3.5-6.0
em long. Filaments white; anthers pale yellow. Stigmas
5-18. disc margin deeply lobed, membrane conspicuous.
Fruits usually stipitate, 3-9 cm long, not ribbed.

Adventive locally throughout the United States and
Canada as far north as Newfoundland, introduced from
Europe, native of Eurasia; fields, roadsides and waste
places, especially about habitations; flowering June-Septem-
ber.

414 Rhodora [Vol. 77

SECT. RHOEADES Bernh.

Plants annual, caulescent or subscapose, glabrous, pilose,
hirsute, hispid or setose. Leaf blades pinnatipartite or
bipinnatipartite. Basal leaves petiolate. Cauline leaves ses-
sile, not amplexicaulous. Filaments filiform, Stigmatic disc
essentially plane, sometimes umbonate, margin lobed, lobes
rounded to obtuse. Fruits glabrous.

2. P. rhoeas L., Sp. Pl., 507. 1753.

Plants caulescent, hispid or setulose, 2.5-9.0 dm tall. Main
stems distally cauliramous, slender to stout. Leaf blades
pinnatipartite or bipinnatipartite, lobes acuminate, coarsely
dentate. Cauline leaves often somewhat clustered beneath
the pedicels. Pedicels sparsely to moderately patent hispid
or setulose. Petals pink to red, sometimes darker spotted
basally, 2.0-4.5 em long. Filaments purple; anthers bluish.
Stigmas 5-18, disc essentially plane, broader than capsule
apex, membrane conspicuous. Fruits slightly stipitate or
sessile, broadly obovoid or subglobose, 1-2 cm long, ob-
scurely ribbed, glaucous.

Adventive locally throughout the United States and
Canada as far north as Alaska, introduced from Europe;
fields, roadsides and disturbed sites, especially about habi-
tations; flowering March-October.

3. P. dubium L., Sp. Pl, 1196. 1753.

Plants caulescent or subscapose, 2-6 dm tall. Main stems
basicauliramous, hirsute to hispid. Basal leaf blades pin-
natipartite, glaucescent, hispid. Cauline leaf blades bipin-
natipartite, glaucescent, hispid. Pedicels distally appressed
hispid, proximally patent hispid. Petals red, sometimes
darker spotted basally. Filaments purple; anthers violet.
Stigmas 7-9, disc essentially plane. Fruits sessile or slightly
stipitate, obovoid, L:W::2:1, usually distinctly ribbed,
glaucous.

1975] Papaver — Kiger 415

Naturalized locally throughout the contiguous United
States, introduced from Europe; fields, roadsides, thickets,
disturbed sites, especially near habitations; flowering May-
August.

4. P. californicum A. Gray, Proc. Amer. Acad. 22: 313.
1887.

Р. lemmonii Greene

Plants caulescent, glabrous or sparsely pilose, 3-6 dm
tall. Main stems distally cauliramous, slender. Leaf blades
pinnatipartite or bipinnatipartite, lobes sometimes dentate,
Petals red, greenish spotted basally, 1-2 cm long. Fila-
ments greenish yellow; anthers yellow. Stigmas 5-8, disc
plane to depressed-conic, usually umbonate, not broader
than capsule apex, membrane conspicuous. Fruits sessile,
ellipsoid to obovoid-turbinate, 1-2 ст long.

Native, endemic to western California; chaparral and
oak woodlands in the mountains at elevations below 800 m,
especially in clearings, burns and other disturbed sites;
flowering April-May.

SECT. ARGEMONIDIUM Spach

Plants annual, eaulescent, pilose, villous or hispid. Leaf
blades pinnatipartite or bipinnatipartite. Basal leaves pe-
tiolate, Cauline leaves sessile, not amplexicaulous. Fila-
ments clavate. Stigmatic dise convex and vaulted, lobes
obtuse. Fruits setose.

5. P. hybridum L., Sp. Pl, 506. 1758.
P. apulum Ten. var. micranthum (Boreau) Fedde
misappl.

Plants 1-5 dm tall. Main stems basiramous and cauli-
ramous. Pedicels moderately to densely appressed to
spreading hispid. Petals red, darker spotted basally. Fila-
ments dark violet: anthers pale blue. Stigmas 4-8, disc
deciduous at maturity. Fruits sessile, obovoid to subglobose,

416 Ећодога [Уо]. 77

1.0-1.5 ст long, obscurely to distinctly ribbed, vestiture
recurved-arcuate, spreading or ascending.

Naturalized locally in California, introduced from Eura-
sia; fields, vineyards and disturbed sites; flowering March-
May.

Specimens attributed to Р. apulum var. micranthum in
herbaria and reported as such in Munz and Keck (1959)
and in Munz (1968) belong here.

Sect. ОХУТОМА Bernh.

Plants perennial, robust, caulescent, hispid. Leaf blades
pinnatipartite. Basal leaves petiolate. Cauline leaves ses-
sile, not amplexicaulous, Flowers very large. Filaments
clavate. Stigmatic dise plane. Fruits subglobose, glaucous,
glabrous.

6. P. orientale L., Sp. Pl., 508. 1753.

Plants 6-10 dm tall. Main stems eramous. Leaf lobes
aristate. Pedicels moderately to densely appressed pale
hispid. Petals pale orange, sometimes pale spotted basally,
5-8 em long. Filaments purple; anthers violet. Stigmas
13-18. Fruits sessile, to 3.5 cm long.

Adventive locally in the middle-Atlantic United States,
escaping from cultivation, introduced from Europe, native
of southwest Asia; fields and disturbed sites; flowering in
May.

Papaver bracteatum Lindl., which some consider distinct
frora P. orientale, has deep red petals and has foliaceous
bracts subtending the sepals. It is occasionally cultivated
and might be found as an escape.

SECT. LASIOTRACHYPHYLLA Bernh.

Plants perennial, scapose. Main stems eramous. Basal
leaves petiolate, rosulate, blades simple, pinnatipartite or
bipinnatipartite. Cauline leaves absent. Stigmatic disc
plane to conic, sometimes vaulted, sometimes umbonate.
Fruits setose.

1975] Papaver — Kiger 411

7. P. alpinum L., Sp. Pl, 507. 1755.

P. ридтавит Rydb.
P. nudicaule L. ssp. radicatum (Rottb.) Fedde var.
pseudocorydalifoliwm Fedde

Plants eespitose. Leaf blades pinnatipartite or bipin-
natipartite. glabrate or sparsely hispid. Scapes erect, to
10 em tall, sparsely hirsute. Petals yellow to salmon pink,
paler spotted basally, to 1 em long. Stigmas usually 5, disc
vaulted. Fruits obovoid-cylindrie or ellipsoid, whitish
setose, trichomes basally tuberculate.

Native to Montana, Alberta and British Columbia; talus
slopes in the Rocky Mountains from 1500 to 2700 m; flower-
ing July-August.

For discussion of the relationship of these plants to
European and Asian members of the complex see D. Love
(1969).

8. P. alboroseum Hult., Fl. Kamtchatka 2, Sv. Vet.-Akad.
Handi ser. 3, 5, no. 2: 141, t. 8, f. c. 1928.

Plants cespitose, caudex short. Leaf blades bipinnati-
partite. pale setose above and beneath, primary lobes 2-5-
partite. Scapes arcuate-ascending, to 15 ст tall, setose.
Petals white to rose, yellow spotted basally. Stigmas 5-6,
dise essentially plane. Fruits ovoid to globose, ribbed,
whitish to brown setose, trichomes basally tuberculate.

Native to the Kenai Peninsula, Alaska, also to Kamt-
chatka, U. S. S. R.; sandy and gravelly soils at low eleva-
tions.

9. P. walpolei Pors., Rhodora 41: 231. 1939.

Plants densely cespitose. Leaf blades entire or pinnati-
partite. subcoriaceous, glabrous, lobes (when present) 8,
rarely 5, broadly obtuse, margins revolute, petiole bases
conspieuously persistent. Scapes erect, to 16 cm tall, hir-
sute or hirtellous distally. Petals pale yellow or cream and

418 Кћодога [Vol. 77

yellow spotted basally, to 2.5 em long. Stigmas usually 5,
dise conic, broader than capsule. Fruits obovoid-obconic,
1.5 cm long, pale yellowish setose or setulose, trichomes
slender, basally tuberculate, sometimes only obscurely so.

Native to the Seward Peninsula, Alaska, also to north-
eastern Asia; tundra from sea level to ca. 800 m on moun-
tain slopes in gravelly loam and solifluction soils, often of
limestone origin; flowering May-August.

10. P. macounii Greene, Pittonia 3: 247. 1897.

keelei Pors.

alaskanum Hult. var. macranthum Hult.
scammianum D. Love

hultenti Knaben

microcarpum DC., misappl.

nudicaule L., misappl.

VDDD

Plants solitary to densely cespitose. Leaf blades pinnati-
partite or rarely bipinnatipartite, glabrate to pilose. Scapes
erect, to 4 dm tall, moderately pilose. Petals yellow.
Stigmas 3-5, disc conic, umbonate, sometimes vaulted.
Fruits narrowly oblong or clavate, L:W::2:1, setose, tri-
chomes not basally tuberculate.

Native and widespread in Alaska, Yukon Terr., North-
west Terr. and British Columbia; sandy and gravelly soils,
heaths, thickets, meadows, often on slopes, to over 2100 m.

11. P. meconnellii Hult., Fl. Alaska & Yukon 5, Lunds Univ.
Arssk. N. F. Avd. 2, 41, no. 1: 803, 7. 7. 1945.

Plants cespitose, to 15 ст tall. Leaf blades bipinnati-
partite, rarely pinnatipartite, glaucous, sparsely pilose
above and beneath. Scapes erect, pilose. Petals yellow,
Stamens usually shorter than mature ovary. Stigmatic
dise convex, distinctly apiculate-umbonate, membrane соп-
spicuous. Fruits obovoid, pale setose, trichomes not basally
tuberculate.

Endemic to Yukon Terr. ; sandy and gravelly soils.

1975] Papaver — Kiger 419

12. P.lapponicum (Tolm.) Nordh. ssp. occidentale (Lundstr.)
Кпађеп, Op. Bot. 2, 3: 55. 1959.

P. radicatum Rottb. ssp. lapponicum Tolm.

P. radicatum ssp. occidentale Lundstr.

P. lapponicum ssp. porsildii Knaben

P. nudicaule L. ssp. radicatum (Rottb.) Fedde var.
coloradense Fedde

nudicaule ssp. radicatum var. columbianum Fedde
kluanensis D. Love

freedmanianum D. Love

nigroflavum D. Love

cornwallisensis D. Love

alaskanum Hult. var. alaskanum

denalii Gjaerevoll

radicatum Rottb. ssp. radicatum, misappl.

kb

Plants cespitose. Caudex short to elongate. Leaf blades
mostly bipinnatipartite, pilose ; petiole bases usually per-
sistent, sometimes conspicuously so, pale to dark brown.
Scapes ascending to erect, to 20 (rarely 25) ст tall, sparse-
ly to moderately appressed to patent hispid or pilose. Petals
yellow or rarely white. Stigmas 5-8, disc plane to slightly
convex, sometimes vaulted. Fruits subglobose, ellipsoid or
obovoid, setose, trichomes not basally tuberculate.

Native and widespread in arctic North America from
Greenland to Alaska, extending southward in disjunct
populations in the Rocky Mountains to Colorado and north-
ern New Mexico at high elevations, also in Lapland; sandy
and gravelly soils, often on talus, arctic and alpine tundra
to са. 3000 m.

For discussions of the proper typification of P. radica-
tum Rottb. and its bearing on the nomenclature in this
complex see A. Lóve (1962a, 1962b), Knaben (1958) and
Knaben and Hylander (1970). The latter interpretation 15
here accepted. See D. Lóve (1969) for discussion of the
Rocky Mountain plants of this complex, held separate by
her.

420 Rhodora [Vol. 77

18. P. nudicaule L., Sp. Pl., 507. 1753.

Plants cespitose, Leaf blades pinnatipartite, less fre-
quently bipinnatipartite, glabrate or setose. Scapes erect,
25-50 ст tall, stout, glabrate or sparsely hispid. Petals
yellow, red or white. Stigmas 4-6, disc plane, membrane
ohsolescent. Fruits clavate or obovoid, setose, trichomes
not basally tuberculate.

Adventive locally in Alaska and Yukon Terr., escaping
from cultivation, native to arctic and alpine Asia; road-
sides and disturbed sites, especially about habitations.

LITERATURE

ABRAMS, L. 1944. Illustrated flora of the Расте States, Washing-
ton, Oregon and California. Vol. 2. Stanford.

ANDERSON, J. P. 1959. Flora of Alaska and adjacent parts of
Canada. Ames.

BócHER, T. W., ET AL. 1966. Grønlands flora, ed. 2. Copenhagen.

Ernst, W. R. 1962. The genera of Papaveraceae and Fumaria-
ceae in the southeastern United States. Jour. Arnold Arb. 43:
315-343.

FABERGE, A. C. 1944. Genetics of the Scapiflora section of Papaver.
III. Interspecifie hybrids and genetic homology. Jour. Genet.
46: 125-149.

FEDDE, F. 1909. Papaver in ENGLER, A., ED. das Pflanzenreich
40 (4, 104) : 288-3806.

FERNALD, M. L. 1950. Gray's manual of botany, ed. 8. New York.

GLEASON, Н. А. 1963. The new Britton and Brown illustrated flora
of the northeastern United States and adjacent Canada. Vol. 2.
New York.

HULTÉN, E. 1945. Flora of Alaska and Yukon, 5. Lunds Univ.
Arssk. N. F. Avd. 2, 41, no. 1.

19683. Comments on the Flora of Alaska and Yukon.

Ark. Bot. 7, 1.

1968b. Flora of Alaska and neighboring territories. A
manual of the vascular plants. Stanford.

JEPSON, W. L. 1922. A flora of California. Vol. 1, Part 7. Berke-
ley.

Касек, К. У. 1973. Sectional nomenclature in Papaver L. Taxon
22: 579-582.

——

1975] Papaver — Kiger 421

KNABEN, С. 1958. Papaver-studier, med et forsvar for P. radi-
catum Rottb. som en islandsk-skandinavisk art. Blyttia 16: 61-
80.

————————. 1959a. On the evolution of the radicatum-group of the
Scapiflera papavers as studied in 70 and 56 chromosome species.
Part A. Cytotaxonomical aspects. Op. Bot. 2, 3.

1959b. On the evolution of the radicatum-group of the
Scapiflora papavers as studied in 70 and 56 chromosome species.
Part B. Experimental studies. Op. Bot. 3, 3.

, & М. HyLANDER. 1970. On the typification of Papaver
radicatum Rottb. and its nomenclatural consequences. Bot. Not.
123: 338-345.

Love, A. 1962a. Typification of Papaver radicatum — а nomen-
clatural detective story. Bot. Not. 115: 115-136.

1962b. Nomenclature of North Atlantic Papavers.
Taxon 11: 182-138.

LóvE, D. 1969. Papaver at high altitudes in the Rocky Mountains.
Brittonia 21: 1-10.

‚ & N. J. FREEDMAN. 1956. A plant collection from SW
Yukon. Bot. Not. 109: 153-211.

Mowat, A. B., & S. M. WALTERS. 1964. Papaver in TUTIN, T. G.,
ET AL., EDS., Flora Europaea. Vol. 1, 247-250. Cambridge.
MvNZ, P. A. 1968. Supplement to a California flora. Berkeley.

, in collaboration with D. D. Кеск. 1959. A California
flora. Berkeley.

PoLUNIN, N. 1940. Botany of the Canadian eastern Arctic. Part I.
Pteridophyta and Spermatophyta. Nat. Mus. Canada Bull. 92.

1959. Circumpolar arctic flora. Oxford.

Ророу, M. С. 1937. Papaver in Komarov, V. L., ED., Flora S. S.
S. R. Vol. 7, 598-646. Moscow.

Porsinp, A. E. 1939. Contributions to the flora of Alaska (con-
tinued). Rhodora 41: 199-254.

1951. Botany of southeastern Yukon adjacent to the
Canol Road. Nat. Mus. Canada Bull. 121.

-———— 1955. The vascular plants of the western Canadian
Arctic Archipelago. Nat. Mus. Canada Bull. 135.

1964. Illustrated flora of the Canadian Arctic Archi-
pelago. Nat. Mus. Canada Bull. 146.

PORTER, D. M., ET AL. 1973. A guide for contributors to Flora
North America. Part II. An outline and glossary of terms for
morphclogical and habitat description (provisional edition). ЕМА
Report 66. Washington.

422 Rhodora [Vol. 77

RYDBERG, P. A. 1922. Flora of the Rocky Mountains and adjacent
plains, ed. 2. New York.

SMALL, J. K. 1933. Manual of the southeastern flora. New York.

WiGGINS, I. L, & J. Н. THoMas. 1962. А flora of the Alaskan
arctic slope. Arctic Inst. N. Amer. Special Publ. 4.

DEPARTMENT OF BOTANY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. 20560

THE NORTH AMERICAN SPECIES,
BOERHAVIA SPICATA (NYCTAGINACEAE),
IN NORTHWESTERN ARGENTINA

PHILIP D. CANTINO

Boerhavia spicata Choisy is a morphologically variable
herb widespread in the Sonoran and Chihuahuan deserts
of the southwestern United States and northern Mexico,
and heretofore known only from that region. During the
course of field study as a worker in the IBP Structure of
Ecosystems Program, I found an isolated population of
this species in Catamarca Province, Argentina. This con-
stitutes the first known occurrence of the species in South
America, and is an addition to the already sizable list of
species known to have range disjunctions between the
desert regions of southwestern North America and north-
ern Argentina and Chile (Bray, 1900; Johnston, 1940;
Raven. 1963: Solbrig, 1972).

Plants of Boerhavia spicata oceur abundantly on a small
group of sand dunes located 35 km west of the town of
Andalgalá, at the north edge of the Bolsón de Pipanaco.
In spite of extensive field work in the Andalgalá region
over an eight-month period in 1972-73, no other popula-
tions, or even isolated individuals, of the plant were found.
Several Argentine botanists familiar with the flora of the
region were consulted, and none had seen the species pre-
viously. In addition, a literature search revealed no refer-
ences to the occurrence of B, spicata or other species with
similar characteristics in South America.

I considered the possibility that the population might
represent an undescribed species. An inspection of the
worldwide collection of Boerhavia at the Harvard Univer-
sity Herbaria revealed a strong resemblance between the
Argentine specimens and members of the В. spicata
complex (explained below). No marked resemblance was
found to any other species of Boerhavia.

423

424 Rhodora [Vol. 77

The Boerhavia spicata complex has been considered by
a number of authors (Standley, 1909; Tidestrom and Kit-
tell, 1941; Kearney and Peebles, 1960; Ferris, 1964) to
consist of three to four distinct species: В. spicata Choisy,
B. watsonii Standley, B. coulteri (Hook. f.) S. Watson, and
B. torreyana (S. Watson) Standley. More recently it has
been interpreted as a single morphologically variable
species, B. spicata Choisy (Reed, 1969), Upon examina-
tion of the Harvard Herbaria collection of the complex
(50 specimens), T tend to agree with Reed's interpretation
that we are dealing with a single variable species. The
charaeters used by the above-mentioned authors to dis-
tinguish the four species (e.g., degree of crowding of the
flowers in the inflorescence; shape of the ridges and furrows
of the anthocarp; degree of rugosity of the anthocarp;
presence or absence of glandular pubescence on the stems
and glandular dots on the foliage) all show gradual varia-
tion between the extreme character states. However, the
species should, perhaps, be segregated into two varieties
-— one, with a center of distribution in the Sonoran Desert,
having a tendency towards closely spaced flowers, acute
to acuminate leaf-apices, and densely glandular-villous
stems; and the other, with a center of distribution in the
Chihuahuan Desert, having a tendency towards loosely
spaced flowers, obtuse leaf apices, and puberulent stems
with few or no glands. The Argentine population clearly
has affinities with the latter variety.

The above analysis is intended as a suggestion only. I
do not feel justified in dividing the species into varieties
based solely on the very limited herbarium study that I
nave done. Crossing experiments in the laboratory and a
field study of ecotypic variation would help greatly to
clarify the taxonomic relationships within this difficult
complex. Meanwhile, for the purposes of identifying the
Argentine population, I am following Reed in his place-
ment of the entire complex in the species Boerhavia spicata
Choisy. The Argentine plants are clearly members of the
complex; thus until someone does a definitive revision of
the genus, they should be placed in B. spicata Choisy.

1975] Воегһау1а — Сап по 425

The following description is patterned after Reed’s de-
scription of Boerhavia spicata but has been modified so as
to describe the Argentine population rather than the
species as a whole:

Erect or decumbent annual from a slender taproot, usu-
ally with many branches spreading from the base; stems
2.6 dm long, much branched, often tinged with pink, white-
puberulent on the lower parts (occasionally sparsely glan-
dular-hirtellous at the nodes only), glabrous on the upper
parts; leaves opposite, the petioles puberulent, 0.2-5.0 em
long, the leaf-blades oval to ovate or ovate-lanceolate, 0.8-
4.0 cm long, 0.4-2.5 em wide, truncate to obtuse or slightly
ohlique at base, obtuse to acuminate at the apex, the margins
entire to sinuate, green above, paler beneath, brown-punc-
tate or not, sparsely puberulent to glabrate; inflorescence
eymose-paniculate, spreadingly much-branched, the branches
slender, glabrous; flowers on short pedicels 0.8 to 1.0 mm
long, loosely spaced on the ultimate branches of the inflores-
сепсе; bracts minute, deciduous, ovate to lanceolate, pink-
ish, ciliolate; perianth pale pink, 1.5 mm long, puberulent;
stamens 2, short-exserted: anthocarp narrowly obovoid,
3 mm long, 1 mm wide, rounded at apex, acute at the
base, 5-angulate, the sulci nearly closed, strongly rugose.

Specimens (Cantino 664, 724) are deposited in the her-
baria of the following institutions in the United States
and Argentina: Harvard University, University of Michi-
gan, University of Arizona, New Mexico State University,
University of Texas, Instituto Miguel Lillo (Tucumán),
Universidad Nacional de Córdoba, and Instituto de Botán-
ica Darwinion (San Isidro).

ACKNOWLEDGMENTS

I wish to express my appreciation to Dr. Otto Т. Solbrig,
Dr. Reed C. Rollins, Dr. Carroll E. Wood, and Dr. Rolla
M. Tryon for their advice and encouragement in the prepa-
ration of the manuscript.

426 Ећодога [Vol. 77

LITERATURE CITED

Bray, W. L. 1900. The relations of the North American flora to
that of South America. Science II. 12: 709-716.

Ferris, R. S. 1964. Nyctaginaceae. Pp. 476-482. Iw Е. SHREVE &
I. L. WIGGINS, Vegetation and flora of the Sonoran Desert. Vol. 1.
Stanford University Press, Stanford, Calif.

JOHNSTON, I. M. 1940. The floristic significance of shrubs common
to North and South American deserts. Jour. Arnold Arb. 21:
356-863.

KEARNEY, T. H., & R. Н. PEEBLES. 1960. Arizona Flora. Univ. of
Calif. Press, Berkeley & Los Angeles. (Boerhavia, pp. 275-277)

RAVEN, Р. Н. 1963. Amphitropical relationships in the floras of
North and South America. Quart. Rev. Biol. 38: 151-177.

REED, С. К. 1969. Nyctaginaceae. Pp. 203-220. In С. L. LUNDELL,
Flora of Texas. Vol. 2. Texas Research Foundation, Renner,
Texas.

SoLBRIG, O. T. 1972. New approaches to the study of disjunctions
with special emphasis on the American amphitropical desert dis-
junctions. Pp. 85-100. In D. VALENTINE, ED. Taxonomy, phyto-
geography, and evolution. London.

1972. The floristic disjunctions between the “Monte”
in Argentina and the “Sonoran Desert” in Mexico and the United
States. Ann. Missouri Bot. Gard. 59: 218-223.

STANDLEY, P. C. 1909. "The Allioniaceae of the United States with
notes on Mexican species. Contr. U.S. Natl. Herb. 12: 303-389.

TIDESTROM, I., & T. KITTELL. 1941. А flora of Arizona and New
Mexico. Catholic University of America Press, Washington,
D.C. (Boerhavia, pp. 652-655)

GRAY HERBARIUM
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS 02138

GEOGRAPHICAL DISTRIBUTION OF THE
CEDAR GLADE ENDEMIC VIOLA EGGLESTONII

JERRY M. BASKIN AND CAROL С. BASKIN

Viola eqglestonii Brainerd is a perennial, stemless, blue
violet that heretofore was thought to be endemic to the
cedar glades of central Tennessee. Distribution maps pub-
lished by Russell (1958, 1965) show the species as occurring
in four counties, and Baskin and Baskin (1970) list five
additional counties making а total of nine in the Central
vasin of Tennessee. These counties include Bedford,
Cannon, Davidson, Marshall, Maury, Rutherford, Smith,
Sumner and Wilson. Viola egglestonii previously has been
reported from Warren (Brainerd, 1921) and Hart (Braun,
1943) counties in south-central Kentucky, but Russell
(1965) in his extensive treatment of the violets of central
and eastern United States could not find specimens to
verify its occurrence in Kentucky. The purpose of this
note is to report the discovery of populations of V. eggle-
stonii in cedar (limestone) glades in Kentucky, Georgia
and Alabama.

On 15 April 1972 we discovered a population of Viola
egglestonii in a cedar glade area in Bullitt Co., Kentucky,
along Co. Rd. 1604, 0.3 miles south of St. Rd. 480 (Baskin
& Baskin 1191). Two additional populations were found
on 23 April 1972 in cedar glades in Bullitt Co. The loca-
tions of these two populations are: Woodsdale Rd., 1.9
miles east of Deatsville Rd. (Baskin & Baskin 1225);
Bells Mill Rd., 2.6 miles east of U.S. 61 (Baskin & Baskin
1234). An additional population was found in Bullitt Co.
on 1 April 1973 on Clarks Lane, 0.7 miles north of St. Kd.
480 (Baskin & Baskin 1639).

Populations of Viola egglestonii were found in cedar
glade areas in Walker and Catoosa counties, Georgia, on
26 May 1972. The locations of these populations are as
follows: Walker Co., along U.S. 27, 5.1 miles south of St.
Rd. 2 (Baskin & Baskin 1301) ; Walker Co., along U. 5. 27,

427

428 Rhodora [Vol. 77

Fig. 1. Geographical distribution of Viola egglestonii. Each dot
represents a county where the species is known to occur.

4.8 miles south of St. Rd. 2 (Baskin & Baskin 1306); Catoosa
Co., Chickamauga National Park, 2 miles east of Ft. Ogle-
thorpe on Co. Rd. 2 (Baskin & Baskin 1308).

On 27 May 1972 a population of Viola egglestonii was
found in a cedar glade in Franklin Со. Alabama. This
population occurs along Co. Rd. 83, 0.6 miles north of St.
Rd. 24 (Baskin & Baskin 1328).

The present known geographical distribution of Viola
egglestonii is shown in Fig. 1. Herbarium specimens from

1975] Viola — Baskin & Baskin 429

the new locations reported in this note have been sent to
the herbarium at Vanderbilt University (VDB).

LITERATURE CITED

RussELL, N. H. 1958. The violets of Tennessee. Castanea 23:
63-76.

———————. 1965. Violets (Viola) of central and eastern United
States. Sida 2: 1-113.

BASKIN, J. M., & C. C. BaskIN. New county records for the cedar
glade endemic Viola egylestonii. Jour. Tenn. Acad. Sci. 45:
60-61.

BRAINERD, E. 1921. Violets of North America. Vt. Agri. Exp.
Sta. Bull. 224.

Braun, E. L. 1943. An annotated catalogue of spermatophytes of
Kentucky. 161 pp. Published by the author, Cincinnati, Ohio.

SCHOOL OF BIOLOGICAL SCIENCES
UNIVERSITY OF KENTUCKY
LEXINGTON, KENTUCKY 40506

RECENT ADDITIONS TO THE MARINE ALGAL
FLORA OF NAHANT, MASSACHUSETTS!

E. E. WEBBER

The only comprehensive volume to date treating the
benthic marine algae of northeastern North America is
that of Taylor (1957). In addition to the systematics and
species descriptions of relevant plants, Taylor also presents
a brief historical summary of marine phycology from
the late 1800’s to approximately the mid-1950’s.

Since that time, however, studies of benthic marine algae
in New England have progressed rapidly. A summation of
recent research dealing with algae of the sublittoral, with
those of estuaries and adjacent salt marshes, and with the
autecology of several species, has been published (Webber,
1975). This paper also contains a preliminary tabulation
of 107 benthic algae specifically from the vicinity of the
Marine Science Institute at Nahant.

In the early summer of 1973 nineteen additional species
were collected, mostly from the Nahant rocky shore, and a
few from a nearby salt marsh. These are listed below.

References used in determinations of the species were:
Farlow (1882), Koster (1955), Taylor (1957), Bliding
(1963), Cardinal (1964), Wilce (1965), and Drouet (1968).
Extended comments on these species will be reserved for
forthcoming publications.

LIST OF SPECIES
Cyanophycophyta

Oscillatoria lutea Ag. — Forming a 2 mm thick mat on
deeply shaded and moist rocks just above high tide
mark; mixed with Microcoleus vaginatus (Vauch.) Gom.

Rivularia atra Roth — epilithic in 2 small rock pools in
spray zone.

Tolypothrix sp. and Cylindrospermum sp. — collected from
salt marsh soil, mid-littoral.

!Contribution No. 14 from the Marine Science Institute, Nahant,
Massachusetts 01908.

430

1975] Nahant Algae — Webber 431

Chlorophycophyta

Tetraspora sp. — Located at the bases of the marsh grass,
Spartina patens Ait. (Muhl.) upper littoral.

Pringsheimiella scutata (Reinke) Schmidt et Petrak — col-
lected once, attached to the operculum of Littorina
obtusata.

Blidingia marginata (J. Ag.) Dangeard ssp. marginata —
commonly epilithic at and just above high tide level.
B. marginata ssp. subsalsa (Kjell.) Bliding — located im-

mediately above B. marginata.

Enteromorpha compressa (L.) Grev. var. compressa —
occurs in quantity in a splash zone pool at the top of
the very exposed East Point rocks.

E. ahlneriana Bliding — common in tide pools at high tide
level.

E. clathrata (Roth) Grev. (type I) — attached to small
stones in the lower littoral zone of Lynn Harbor.

E. clathrata (Roth) Grev. (type II) — entangled with
Cladophora sp. in the lower littoral zone of Lynn Harbor.

E. flexuosa (Wulfen ex Roth) J. Ag. ssp. flexuosa — grow-
ing on shells in the lower littoral zone of Lynn Harbor.

Urospora collabens (C. Ag.) Holmes et Batters — com-
monly ере in the mid-littoral zone at Nahant, often
mixed with Codiolum gregarium A. Braun.

Rhizoclonium riparium (Roth) Harv. f. riparium — while
R. tortuosum is the common species of this genus at
Nahant, R. riparium was located in one splash-zone tide
pool at the East Point rocks.

Phaeophycophyta

Ectocarpus fasciculatus var. refractus (Kutz.) Ата. —
common on Laminaria stipes; previously reported by
Taylor (1957) only from Maine, as an epiphyte on
Porphyra.

Myriotrichia filiformis Нату. — epiphytic on Chordaria
flagelliformis in a lower littoral zone tide pool.

432 Ећодога [Vol. 77

Saccorhiza dermatodea (De la Pyl.) J. Ag. — epilithic at
8 to 20 foot depths, mixed with Laminaria saccharina
(Wilce, 1965) and L. digitata. Published accounts of
S. dermatodea along the northeast coast suggest that this
species is limited to coastal waters north of Cape Ann,
Massachusetts (Farlow, 1882; Lamb and Zimmermann,
1964; Stone, et. al., 1970; Mathieson and Fralick, 1972).
However, records for the distribution for Saccorhiza in
New England (Taylor, personal communication) reveal
that plants of this species extend southward to the Cape
Cod (Sandwich, Massachusetts) area. Indeed, S. derma-
todea is known from the sublittoral at both ends of the
Cape Cod Canal (Wilce, personal communication).

Rhodophycophyta

Nemalion multifidum (Weber et Mohr) J. Ag. — epilithic
in the upper sublittoral, uncommon, only female plants
located.

ACKNOWLEDGMENTS

Appreciation is expressed to Dr. W. Randolph Taylor
for sending me collection records of Saccorhiza dermatodea
from his personal herbarium. Plants of this species were
collected simultaneously at Nahant (July 14, 1973) by
Jane Menge and Barry Spracklin.

in addition, Myriotrichia filiformis, Nemalion multifidum,
and Rivularia atra were collected initially by Jane Menge,
while Prigsheimiella scutata was located by Barry
Spracklin.

LITERATURE CITED

BLIDING, С. 1963. А critical survey of European taxa in Ulvales.
I. Opera Botanica (Suppl. Bot. Not.). 8: 1-160. Lund.

CARDINAL, A. 1964. Etude sur les ectocarpacées de la Manche. Nov.
Hedw. 15: 1-86. 41 figs.

DROUET, F. 1968. Revision of the classification of the Oscilla-
toriaceae. Monogr. 15. Acad. Nat. Sci. Philadelphia.

1975] Nahant Algae — Webber 438

FaRLow. W. С. 1882. The marine algae of New England. U.S.
Comm. Fish and Fisheries. Commissioners Report, 1879. Wash-
ington: Government Printing Office. Pp. 1-210.

Koster, J. T. 1955. The genus Rhizoclonium in the Netherlands.
Pub. Sta. Zool. Napoli. 27: 335-357.

Lame, I. M, & M. H. ZIMMERMANN. 1964. Marine vegetation of
Cape Ann, Essex County, Massachusetts. Rhodora 66: 217-254.

MATHIESON, А. C., & R. A. FRALICK. 1972. Investigations of New
England marine algae. V. The algal vegetation of the Hampton-
Seabrook estuary and the open coast near Hampton, N.H. Rho-
dora 74: 406-435.

$тохкЕ, В. A., E. J. Henre, J. M. Conway, & A. C. MATHIESON.
1970. A preliminary checklist of the marine algae of Campo-
bello Island, New Brunswick, Canada. Rhodora 72: 313-338.

TAYLOR, W. R. 1957. Marine Algae of the Northeastern Coast of
North America. (Rev. ed.) Univ. Mich. Press. Ann Arbor. 509
pp., 60 pls.

Werrer, E. E. 1975. Phycological studies from the Marine Science
Institute, Nahant, Massachusetts. I. Introduction and prelimi-
пату tabulation of species. Rhodora 77: 149-158.

WiLcr, R. T. 1965. Studies in the genus Laminaria. Bot. Gotho-
burgensia. III. Proe. Fifth Mar. Biol. Symp. Góteborg. 247-256.

DEFT. OF BIOLOGY
KEUKA COLLEGE
KREUKA PARK, N.Y. 14478

LACTUCA MURALIS IN MAINE
JAMES P. POOLE

In a letter dated November 15, 1969, Mrs. Roberta G.
Poland of South Deerfield, Mass. reported finding Lactuca
muralis (L.) Gaertn. on Mt. Desert Island in Salisbury
Cove, Hancock County, Maine on August 16, 1969. It was
collected from a small group of plants that were still in
good condition when checked again in 1970 and 1971.
Voucher specimens have been deposited in the Jesup Her-
barium at Dartmouth College (HNH), in the University
of New Hampshire Herbarium (NHA), in the Pringle
Herbarium at the University of Vermont (VT), and in
the herbarium of the University of Massachusetts at Am-
herst, Mass. (MASS). Collection of this species in Cornish,
Sullivan County, N.H. was reported in Rhodora 68:774,
1966. At that time Professor Albion R. Hodgdon at the
University of New Hampshire wrote that the Cornish dis-
covery was the first report of the species in New England,
and Frank C. Seymour’s Flora of New England published
in 1969 lists Cornish as the only New England station.
Mrs. Poland’s discovery represents a really disjunctive
jump.

JESUP HERBARIUM
DARTMOUTH COLLEGE
HANOVER, N.H. 03755

484

LACTUCA MURALIS IN BRITISH COLUMBIA
JoHN T. WITHERSPOON AND THOMAS J. WATSON, JR.

Lactuca muralis (L.) Gaertn. was recently collected in
British Columbia (KOA campground, ca. 10 mi W of Hope,
2 Sept. 1973, Watson 1048). The specimen is deposited at
MONTU. This is the most inland known record in British
Columbia as well as the farthest north in the foothills of
the Cascade Mountains. The locality, latitude 49°20'N
longitude 121°40’W, is approximately the same longitude
as a reported locality in the Cascade foothills of Marion
County, Oregon (Peck, 1961).

Lactuca muralis is an introduction from northern Europe
and is sparingly adventive in North America. In eastern
North America it is known from western Quebec, eastern
New York and west to Michigan (Fernald, 1950; Gleason
and Cronquist, 1963). It was recently reported from New
Hampshire (Poole, 1966).

It is apparently more common in the eastern parts of
North America and previous to this report it was known
only from the Oregon locality above, Clallam County and
San Juan Island, Washington, and from Lake Cowichan,
Vancouver Island, British Columbia in western North
America (Hitchcock, et al., 1955). It seems to be migrating
slowly inland, both eastward and northward in mainland
British Columbia.

LITERATURE CITED

FERNALD, М. L. 1950. Gray’s Manual of Botany. 8th ed. Van
Nostrand Reinhold Co., New York. lxiv + 1682 pp.

GLEASON, H. A. and A. CRONQUIST. 1963. Manual of Vascular Plants
of Northeastern United States and Adjacent Canada. D. Van
Nostrand Co., Inc., Princeton. li + 810 pp.

Нітснсоск, C. L., A. CRONQUIST, M. OWNBEY and J. W. THOMPSON.
1955. Vascular Plants of the Pacific Northwest. Part 5, Com-
positae, by A. Cronquist. Univ. of Washington Press, Seattle.
343 pp.

Poore, J. P. 1966. Lactuca muralis in New England. Rhodora
68: 138.

DEPT. OF BOTANY
UNIVERSITY OF MONTANA
MISSOULA, MT 59801

435

CHIMAPHILA MACULATA IN NEW HAMPSHIRE
JAMES P. POOLE

In December, 1966, Е. 5. Page, Emeritus Professor of
Botany at Dartmouth College, collected two specimens of
Chimaphila maculata (L.) Pursh on a wooded hill two
miles south of George Pond and about % mile west of
the dirt road that borders the west shore of the pond.
The plants bore neither flowers nor fruit, but the leaves
were sufficiently characteristic for identification. Since in
Gray’s Manual, 8th edition, this species is listed as growing
in southern New Hampshire only, the collection was ac-
cessioned in Jesup Herbarium as voucher specimens for
what appeared to be a range extension for the species. The
following summer, in August, Professor Page guided Pro-
fessor Carl Wilson and me to the area where the plants
had been collected, but careful search over the whole area
by the three of us failed to turn up another specimen. In
light of our experience it is interesting to quote from the
article covering the distribution of this species in Maine
and New Hampshire by Albion R. Hodgdon and Lesley M.
Eastman in Rhodora 75:162-165 (1973). “In northern
New England Chimaphila maculata tends to occur as dis-
junct colonies of a few individuals or sometimes single
plants, in this respect resembling some species of orchids.
Even with careful searching it is usually impossible to find
any more plants after having found a few shoots at any
one station. While the stems are of low stature and stand
but a few inches above the forest floor, the leaves are
conspicuous and easy to detect.” In the same article a
number of records extending the range of this species in
New Hampshire are listed. One collection is listed for
Tuftonboro, Carroll County, on Sentinel Mountain, ap-
proximately ten miles farther north than George Pond in
Grafton County, but I find no record of an earlier collection
in Grafton County, I am listing this station as in Enfield,
Grafton County, but it is difficult to locate exactly on the

436

о

1975] Chimaphila maculata — Poole 437

topographic map, so difficult that it could actually be spotted
on either side of the boundary separating Enfield from
Grantham in Sullivan County. The only other New Hamp-
shire specimen in Jesup Herbarium was collected on a
wooded hill in Kensington, Rockingham County, in July
1896, and bears a printed label reading “Ex Herb. Alvah
A. Eaton.”

JESEP HERBARIUM

DARTMOUTH COLLEGE
HANOVER, N.H. 03755

BOOK REVIEW:
WINTER KEYS TO WOODY PLANTS OF MAINE!

When one is faced with the task of teaching a field ori-
ented course in introductory taxonomy in the more north-
ern regions of this continent one must, of necessity, give
due consideration to those plants that are available during
the winter season — our woody vascular plants. I enthusi-
astically welcome this excellent field manual for the identi-
fication of woody plants in winter condition. The authors
of this carefully constructed, excellently illustrated work
have succeeded in providing a thorough treatment of the
woody plants of Maine, native, naturalized and adventive,
with nearly 250 species included. The work goes beyond
the "clearly" woody species and provides a means for iden-
tifying such marginally woody species as Arceuthobium
pusillum, Mitchella repens, Potentilla tridentata, Aralia
hispida, A. nudicaulis and Decodon verticillatus. The keys
are well constructed, strictly dichotomous, and usually
employ a minimum of 2-3 characters per lead. Frequently
additional information regarding persistent fruits or veg-
etative features is provided in the leads to further facilitate
the identification process. Habitat data are also incorpor-
ated in keys where helpful. The value of the manual is
greatly enhanced by the superb set of plates of original
line drawings, made from fresh material. In addition to
drawings of twigs and of critical leaf scar or bud features,
the illustrator has provided drawings of leaves and fruits
of certain species in which they are frequently found per-
sistent. It is no understatement when the authors claim it
to be fully illustrated.

The manual is organized into a “Кеу to Genera and Aber-
rant Species" followed by a “Кеу to Species" with genera

'Campbell, Christopher S. and Fay Hyland. 1975. Winter Keys
to Woody Plants of Maine. Illustrated by Mary L. F. Campbell.
University of Maine Press, Orono, Maine. 52 pp + 63 plates. $3.00.
Available from: Secretary, Department of Botany, Deering Hall,
University of Maine, Orono, Maine 04473.

438

Crow 439

1975] Winter Keys

arranged alphabetically. Additional notes are included at
the end of each set of keys providing information regard-
ing infraspecific taxa. Nomenclature follows Fernald,
Gray’s Manual of Botany edition 8. A clear, concise and
complete glossary is also provided. The very thorough
index not only includes the scientific name, but also lists
every conceivable combination of common names (e.g.,
Pyrus americana; American Mountain Ash; Ash, American
Mountain; Mountain Ash, American). The back inside
cover provides a map of the counties of Maine and a metric-
English scale.

The publication is appropriately dedicated to the mem-
ory of Professor Merritt Lyndon Fernald, whose long term
interest in the plants of New England and northeastern
North America found its stimulus in the plants of his boy-
hood haunts in the vicinity of Orono, Maine. This fine piece
of work should be of interest and value to students, ama-
teur and professional botanists, ecologists, horticulturalists,
foresters and naturalists of not only Maine, but of her
adjacent states and provinces. At $3.00, this is a welcome
bargain.

GARRETT Е. CROW

DEPARTMENT OF BOTANY & PLANT PATHOLOGY
UNIVERSITY OF NEW HAMPSHIRE

DURHAM, NEW HAMPSHIRE 03824

Volume 77, No. 811, including pages 332-439, was issued Dec. 15, 1975

Cover illustration
Rhododendron canadense (L.) Torr.
from
Curtis’ Botanical Magazine t.474

RHODORA September, 1975 Vol. 77, No.

CONTENTS

A Revision of the North American Genus Amorpha (Leguminosae-
Psoraleae).
е О. Шон н ТОРИНО. ~ А Iiis

Papaver in North America North of Mexico.
Robert W. Kiger Wa

The North American Species Boerhavia spicata (Nyctaginaceae),
in Northwestern Argentina.
а D. CHUNG Wee EE УОИ

Geographical Distribution of the Cedar Glade Endemic Viola
egglestonii.
Jerry M. Baskin and Carol C. Baskin

Recent Additions to the Marine Algal Flora of Nahant, Massa-
chusetts.
О АНАН EE CR

Lactuca muralis in Maine.
Мо ха es a ids taba Er dead

Lactuca muralis in British Columbia.
John T. Witherspoon and Thomas J. Watson, Jr. ....................

Chimaphila maculata in New Hampshire.
James P. Poole

Book Review: Winter Keys to Woody Plants of Maine.
Garrett E. Crow ..

811

337

410

423

421

480

434

435

436

. 438

Rhodora

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Vol. 77 December, 1975 No. 812

The No Englmd Botanical Club, Ine.

Botanical Museum, Oxford Street, Cambridge, Mass. 02138

Conducted and published for the Club, by

ALFRED LINN BOGLE, Editor-in-Chief
ROLLA MILTON TRYON
STEPHEN ALAN SPONGBERG
GERALD JOSEPH GASTONY
RICHARD EDWIN WEAVER

Associate Editors

_—

RHODORA.— A quarterly journal of botany, devoted primarily to
the flora of North America and floristically related areas. Price
$20.00 per year, net, postpaid, in funds payable at par in the United
States currency at Boston. Some back volumes, and single copies
are available. For information and prices write RHODORA at
address given below..

Scientific papers and notes, relating directly or indirectly to the
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mittee for publication. Articles concerned with systematic botany
and cytotaxonomy in their broader implications are equally accept-
able. All manuscripts should be submitted in duplicate, and should
be double-spaced throughout. Please conform to the style of recent
issues of the journal. Illustrations can be used only if the cost of
engraver's blocks is met through the author or his institution.
Forms may be closed five weeks in advance of publication. Ex-
tracted reprints, if ordered in advance, will be furnished at cost.

Address manuscripts and proofs to:
Dr. A. Linn Bogle
Dept. of Botany and Plant Pathology,
University of New Hampshire,
Durham, New Hampshire 03824

Subscriptions and orders for back issues (making all remittances
payable to RHODORA) should be sent to RHODORA, Botanical
Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive
the next number of RHODORA, changes of address must be received
prior to the first day of March, June, September or December.

Second Class Postage Paid at Boston, Mass.

MANUFACTURED BY
THE LEXINGTON PRESS, INC.
LEXINGTON, MASSACHUSETTS

Cover illustration
Rhododendron canadense (L.) Torr.
from

Curtis’ Botanical Magazine t.474

Rhodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 77 December, 1975 No. 812

HYBRIDS IN AMERICAN CYATHEACEAE

DAVID S. CONANT

Several small tree ferns collected in a ravine near Cerro
de Punta, Puerto Rico, are proposed as à hybrid between
Alsophila dryopteroides (Maxon) Tryon and Nephelea
portoricensis (Kuhn) Tryon, both of which were growing
abundantly at the site. This is the first unequivocal report
of a hybrid in the family.

In addition, six other hybrids are proposed, one on the
basis of substantial evidence, and the others on the basis
of inference. Four previous reports suggest that hybridiza-
tion may occur in the Cyatheaceae, but definitive analyses
of the parental and hybrid characters were not presented.
Но ши and Sen (1961) and Holttum (1963, 1974) report
that Cyathea alternans (Wall ex Hook.) Presl of Malaya,
Sumatra and Borneo might be a series of hybrids between
Cyathea moluccana R. Br. and С. squamulata (Bl.) Copel.
on the basis of its indusium and leaf architecture. Gastony
(1973) treats Nephelea concinna (Kuhn) Tryon as a spe-
cies but notes the possibility that it is a hybrid between
N. pubescens (Kuhn) Tryon and М. Tussacii (Desv.)
Tryon on the basis of intermediate characters of the lamina
architecture and partially contracted fertile pinnules.

441

442 Ећодога [Vol. 77

DEFINITE HYBRIDS

1. Alsophila dryopteroides (Maxon) Tryon X Nephelea
portoricensis (Kuhn) Tryon

Plants of this hybrid (Fig. 1) resembled large plants of
Alsophila dryopteroides at a distance, but on closer inspec-
tion they were seen to have the black squaminate spines of
Nephelea on the petiole. Species of Cyatheaceae growing
nearby were Lophosoria quadripinnata (Gmel) C. Chr.,
Cyathea arborea (L.) Sm., C. furfuracea Baker, Trichip-
teris armata (Sw.) Tryon, T. borinquena (Maxon) Tryon,
Alsophila bryophila Tryon, A. dryopteroides (Maxon)
Tryon, and Nephelea portoricensis (Kuhn) Tryon.

The black squaminate spines on the petiole and rachis
(Fig. 10) are a generie character of Nephelea (Fig. 12).
They firmly establish one parent as N. portoricensis since
it is the only member of the genus in Puerto Rico. The
setate scales on the lamina of the hybrid restrict the second
parent to a species of Alsophila of which there are three
in Puerto Rico; A. Brooksii (Maxon) Tryon, A. bryophila,
and A. dryopteroides. Alsophila Brooksii is not known
from the central mountains where the hybrids grew, it
lacks the dark lanceolate type of scale present in the hybrid,
and its long petiole is inconsistent with the requirements
for the second parental species. Similarly, A. bryophila
may be eliminated as a possible parent because it also lacks
the dark lanceolate type of scale, and a parent with a short
rather than a tall trunk is required. Alsophila dryopter-
oides remains as the second possible parent and all of the
characters of the hybrid are consistent with this choice.

A detailed comparison of the hybrid and parental spe-
cies is presented in Table 1. Some of the important char-
acters of the hybrid are in the leaves, stems, and lamina
scales.

Characters which are intermediate are lamina shape and
dissection, stem, petiole and rachis spininess, number of
leaf scars per unit of stem length, and leaf scar diameter.
The lance-ovate lamina of the hybrid is intermediate be-

1975] Cyatheaceae — Conant 443

Figs. 1-2. Alsopliia dryopteroides X Nephelea portoricensis:

1, plant, stem ca. 75 cm. tall, leaves ca. 1.0 m. long, Conant 679;

2, spore, 1500, Conant 687. Figs. 3-4. Cnemidaria horrida X
Cyathea arborea: 3, plant, stem ca. 2.0 m. tall: leaves: са 2.9. m.

long. Conant 626; 4, bicolorous scales on crozier, Х 1/3, Conant 546.
, 3 3 b ,

(all specimens GH).

444 Ећодога [Vol. 77

Alsophila

Li Pu ЛИЛЛИАН
444 МАУ А АН ЧИ,

Figs. 5-6. Leaf tracings and pinnae of Nephelea portoricensis,
Hybrid, and Alsophila dryopteroides: 5, leaf tracings, A, №. por-
toricensis, B, from Conant 679 and C, from Conant 680, Hybrid,
D, from Conant 681, E, from Conant 690 and F, from Conant 682,
A. dryopteroides, X 1/30; 6, pinnae, A, Conant 2002, N. portori-
consis, B, Conant 1980, Hybrid, C, Conant 1852, A. dryopteroides,
X 1/5. (al specimens GH).

1975] Cyatheaceae — Conant 445

tween the oblanceolate lamina of A. dryopteroides and the
ovate lamina of N. portoricensis (Fig. 5). The lamina
dissection of the hybrid is bipinnate-lobed which is inter-
mediate between the pinnate-pinnatifid condition of A.
dryopteroides and the bipinnate-pinnatifid condition of М.
portoricensis (Fig. 6). The stem, petiole and rachis of the
hybrid have a few spines reaching 2.5 mm. long (Figs. 9
and 10), a condition which is intermediate between the
spineless A. dryopteroides (Figs. 7 and 8) and the numer-
ous spines reaching 5.5 mm. Jong in N. portoricensis (Figs.
11 and 12). The hybrid has from 18-25 leaf scars 1.5 cm.
in diameter, per 10 em. of stem (Fig. 9). Alsophila dry-
opteroides has about 43 leaf scars 1.0 em. in diameter, per
10 em. of stem (Fig. 7) while N. portoricensis has about
13 leaf sears 2.0 ст. in diameter, per 10 cm. of stem
(Fig. 11).

One character of the hybrid is similar to that of Al-
sophila dryopteroides. The dark lanceolate laminar scales
with a single apical seta found in the hybrid (Fig. 14)
closely resemble the most prominent laminar scale of A.
dryopteroides (Fig. 13).

Other characters of the hybrid are like those of Nephelea
portoricensis. The acaroid type of laminar scale occurs in
both the hybrid (Fig. 15) and in N. portoricensis (Fig. 17)
and the marginally setate type of scale also occurs in the
hybrid (Fig. 16) and in N. portoricensis (Fig. 18).

The spores of the hybrid plants are well developed
(Fig 2). A study of their viability has been planned but
until it is completed it is not possible to suggest the number
of F, hybrids probably involved in the five plants seen.

Trunk erect, occasionally short spiny, to 2 m. high, leaf
scars 1.5 em. in diameter, 18-25 per 10 cm. of stem length.
Leaves 1.0-1.5 m. long, petiole to 10 cm., armed with short
black squaminate spines ; lamina lance-ovate, gradually nar-
rowed at base (basal pinnae to !4 as long as the median),
to 60 em. broad at the middle, acuminate, bipinnate-lobed
to occasionally bipinnate-pinnatifid, leaf tissue glabrous,

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Figs. 7-18. Alsophila dryopteroides, Hybrid, and Nephelea porto-
ricensis. Figs. 7-8. A. dryopteroides: 7, stem, Х 2/3, Conant 682,

8, petiole, х 2, Conant 690. Figs. 9-10. Hybrid: 9, stem, X 2/3,
Conant 680, 10, rachis, X 2, Conant 680. Figs. 11-12. N. portori-

9

censis: 11, stem, X 2/3, Tryon & Tryon 6950, 12, petiole, X 2/3,
Conant 412. Fig. 13. A. dryopteroides, lanceolate type seale, х 100,
Conant 598. Figs. 14-16. Hybrid: 14, lanceolate type scale, X 100,
Conant 687, 15, acaroid scale, X 100, Conant 687, 16, marginally
seiate scale, X 100, Conant 687. Figs. 17-18. М. portoricensis: 17,
acaroid scale, X 100, Conant 188, 18, marginally setate scale, X 100,
Coaant 188. (all specimens GH).

448 Еһодога [Vol. 77

herbaceous, glaucescent beneath, dark green above; rachis
dull castaneous, occasionally with short spines at the base,
glabrescent to squamulose beneath, invested with stiff,
brown, antrorse trichomes and a few long filiform scales
above; pinnae to 25, spreading, sessile, the largest 30 cm.
long, 7 cm. broad near base, becoming pinnatifid towards
the apex; pinna-rachis beneath invested with marginally
setate scales, acaroid scales, and dark bodied, light mar-
gined, lanceolate scales with a single, dark, stout apical
seta, and with intermediate scales ranging from oblong to
lanceolate, uniformly brown to dark bodied, and many
setate to singly and stoutly setate, these latter types be-
coming sub-bullate towards the apex; undersurface of pin-
na-rachis becoming sparsely pubescent apically; pinnules
18-24 pairs, 19-34 mm. long, 7-10 mm. broad, deeply lobed
to occasionally pinnatifid at the base; costa beneath often
with acaroid scales at the base, and beyond with sub-
bullate scales with few to many setae, these progressing
to bullate scales at the pinnule apex, occasionally with a
few stiff trichomes apically; costules with marginally setate
sub-bullate scales at the base, these grading into squamules
apically; veins 4-5 pairs, sori 2-3 on each lobe, closer to the
costule than to the margin; indusia deeply cyathiform, tan,
membranous, glabrous; spores well developed, light tan,
trilete.

SPECIMENS EXAMINED;

Puerto Rico: Five plants from the vicinity of Cerro de Punta,
40 miles w.s.w. of San Juan, ravine on north side of road, Rt. 143,
km. 18.5 e. of jct. with Rt. 10, Conant 678 (GH, NY, US), Conant
679 (GH, NY, US), Conant 680 (GH, му, US), Conant 687 (F, GH, IJ,
NY, RPPR, US, USD, Herb. El Verde Field Station, Puerto Rico);
ravine on north side of road, Rt. 143, km. 22.2 e. of jet. with Rt.
10, Conant 599 (GH).

2. Cnemidaria horrida (L.) Presl X Cyathea arborea (L.)
Sm. Hemitelia Wilsonii Hook. in Hook. & Baker, Syn.
Fil. 30. 1865. TYPE: Jamaica, Wilson 781 (к). (See
Maxon, Contrib. U.S. Nat. Herb. 17: t. 18. 1914.)

1975] Cyatheaceae — Conant 449

The status of Hemitelia Wilsonii (Fig. 3) as a species is
untenable because well developed spores have not been
found in its sporangia. Other factors indicating a hybrid
status are the highly variable leaf architecture and the
variable venation.

I have seen Hemitelia Wilsonii growing at two sites in
the Luquillo Forest, 20 miles e.s.e. of San Juan, Puerto
Rico (Rt. 186, km. 19.0 s. of jct. with Rt. 3, Municipio de
Rio Grande, and Sabana Rd., km. 3.0 e. of ject. МИАВ Rt LIL,
Municipio de Rio Grande). In both places Cnemidaria
horrida and Cyathea arborea were growing nearby.

The hybrid is intermediate between the parental species
in its venation, indusia, and leaf architecture. The venation
of the hybrid is usually free but occasionally anastomoses
or forms loops or dead-ends or both (Fig. 22). This con-
dition is intermediate between the free venation of Cyathea
arborea (Fig. 24) and the anastomosing venation of Cne-
midaria horrida. (Fig. 20). The indusia of the hybrid range
from hemiteloid to sub-cyathiform (Fig. 21), a condition
intermediate between the hemiteloid indusia of Cn. horrida
(Fig. 19) and the cyathiform indusia of C. arborea, (Fig.
23). The leaf architecture of the hybrid ranges from
bipinnate-lobed to bipinnate-pinnatifid. This is intermedi-
ate between the pinnate-pinnatifid condition of Cn. horrida,
and the bipinnate-pinnatifid to tripinnate architecture of
C. arborea.

Characters of the hybrid which are closest to Cnemidaria
horrida are the shape of the pinna and leaf apex and the
bicolorous scales on the croziers and petioles (Fig. 4).
One character of the hybrid which is found in Cyathea
arborea but not in Cn. horrida is the small whitish scales
on the costa undersurface.

The apparent absence of viable spores, the combination
of characters, and the documented occurrence of Hemitelia
Wilsonii in the vicinity of both parental species, support
the proposal of Н. Wilsonii as a hybrid between Cnem idaria
horrida and Cyathea arborea.

450 Rhodora [Vol. 77

Figs. 19-24. Cnemidaria horrida, Hybrid, and Cyathea arborea.
Figs. 19-20. Cn. horrida: 19, indusia, X 20, Gastony et al. 655,
20, venation, X 715, Scamman 8117. Figs. 21-22. Hybrid: 21,
indusia, upper arrow, subcyathiform indusium, lower arrow, hemi-
teloid indusium, х 20, Gastony et al. 654, 22, venation, upper arrow,
anastomosing veins, middle arrow, space between free veins (note
loop in lower vein), lower arrow, dead-end in vein, X 714, Conant
546. Figs. 28-24. С. arborea: 23, indusia, X 20, Howard & Nevling
15776, 24, venation, X 715, Gastony 10. (all specimens GH).

1975] Cyatheaceae — Conant 451

SPECIMENS EXAMINED:

Puerto Rico: Conant 546, 626, 627 (GH), Hess 371 (му), Hioram
182, 804 (хү), Kepler, Sabana Rd., El Yunque Rd., 5 Mar., 1970,
Herb, El Verde Field Station, Municipio de Rio Grande. Dominican
Republic: Abbott 2660 (GH, NY, us), Ekman 14752 (NY), 15014
(GH, NY), Gastony, Jones & Norris 654 (сн). Haiti: Ekman H4835
(NY, S), H4846 (s, US). Jamaica: Jenman (NY), Proctor 18419 (A).

PUTATIVE HYBRIDS

The establishment of the Alsophila-Nephelea intergeneric
hybrid with a bipinnate-lobed leaf architecture helps to
interpret other rare species and unusual collections with a
similar architecture within the Alsophila-Nephelea evolu-
tionary line. These are discussed below in order of the
certainty of their parental species. All of them combine
characters of species with very different leaf architecture
and (or) lamina indument. АП are very rare, known
mostly from one collection, and do not fit within the vari-
ation of the species proposed as parents. Nephelea con-
cinna may be a hybrid, perhaps between N. pubescens and
N. Tussacii as suggested by Gastony (1973) but a detailed
analysis of it has not been made.

3. Alsophila Brooksii (Maxon) Tryon ~ Nephelea portori-
censis (Kuhn) Tryon

A collection from Indiera Fria, near Maricao, Puerto
Rico (Britton, Cowell, & Brown 4520 (NY) ) resembles the
Alsophila dryopteroides — Nephelea portoricensis hybrid
quite closely in leaf dissection and in width to length pro-
portions of the pinnae and pinnules. It differs in that it
lacks the dark bodied, light margined, lanceolate scales
with a single, dark, stout apical seta of A. dryopteroides.
In addition the lamina base is abruptly narrowed, and the
petiole is up to 25 cm. long. The complexity of the lamina,
the abundance of acaroid scales, and the pubescence of the
veins and costules of this collection are all characters
attributable to N. portoricensis as pointed out by Gastony
(1973). Th» unusually long and narrow pinnae suggest
that this may be a hybrid involving N. portoricensis and

452, Еһодога [Vol. 77

an Alsophila species with a less dissected lamina and long
narrow pinnae. Alsophila Brooksii is well known from the
Maricao Forest about 10 km. to the west of Indiera Fria.
It is the only species of Alsophila occurring in the western
end of the island, and it has a pinnate-pinnatifid lamina
abruptly reduced at the base. It has long narrow pinnae
and its petiole reaches 78 cm. in length. Its characters,
combined with those of N. portoricensis, would result in a
plant very similar to the Britton et al. 4520.

4. Nephelea balanocarpa (D. C. Eaton) Tryon X Nephelea
woodwardioides (Kaulf.) Gastony

Cuban material with bipinnate-lobed lamina architecture
such as Leon, Clement, & Roca 10533 (NY) and Hioram &
Clement 6377 (US) seems to represent hybrids between
pinnate-pinnatifid and bipinnate-pinnatifid species, One
parent is probably the bipinnate-pinnatifid Nephelea wood-
wardioides. It is not clear whether var. woodwardioides or
var. cubensis (Maxon) Gastony is involved. Nephelea
balanocarpa is probably the second, pinnate-pinnatifid
parent. It is far more common than A. Brooksii, also pin-
nate-pinnatifid, which is known in Cuba only from the type
collection. The abundance of tiny antrorse squamules cov-
ering the veins in the hybrid is in agreement with the in-
dument of N. balanocarpa.

5. Alsophila hotteana (C.Chr. & Ekman) Tryon X Nephe-
lea sp. Cyathea confinis C. Chr. Kungl. Svensk. Vetens.-
акаа. Hand. ser. 3, 16:13. 1937. TYPE: Haiti, Massif de
la Hotte, Jeremie, Ekman 10382 (holotype, s; isotype,
us!).

Cyathea confinis, a Haitian species known only from the
type collection at Massif de la Hotte, has a bipinnate-lobed
lamina, suggesting that it is probably of hybrid origin.
It is described as having an aculeate trunk to 2 m. high, a
short petiole armed with short black spines, and a grad-
ually reduced lamina base. A relationship to the genus

1975] Cyatheaceae — Conant 453

Nephelea is evident in the aculeate trunk and the black
petiole spines. Three species of the genus occur in Haiti,
N. crassa (Maxon) Tryon, N. fulgens (C.Chr.) Gastony,
and М. woodwardioides (Kaulf.) Gastony var. Hieronymi
(Brause) Gastony. АП are tall and have a bipinnate-
pinnatifid lamina making it difficult or impossible to deter-
mine from herbarium specimens which of the species is
involved in the cross.

Alsophila hotteana is probably the pinnate-pinnatifid
parent. Its short petiole, gradually narrowed lamina base,
and tall stem are consistent with the characters required
of the second parent, and it is the only Alsophila known to
occur in the Massif de la Hotte. Other species of Alsophila
in Haiti are restricted to the Massif du Nord.

PUTATIVE HYBRIDS INVOLVING UNDETERMINED SPECIES
OF ALSOPHILA AND NEPHELEA

6. Cyathea irregularis Brause Urban Symb. Ant. 7. 155.
1911. TYPE: Dominican Republic, Santo Domingo, Con-
stanza, Turckheim 3212 (holotype, B; isotype, NY!).

Cyathea irregularis, known only from the type collection,
is another species with a bipinnate-lobed lamina. It is evi-
dently a hybrid of Alsophila апа Nephelea since all species
of Alsophila in the Dominican Republic are pinnate-pin-
natifid and all species of Nephelea are bipinnate-pinnatifid.
The single specimen of a midportion of the lamina, how-
ever, provides too few characters to choose between the
four Alsophilas and three Nepheleas known to grow in the
country.

Another specimen from the Dominican Republic, with a
similar leaf morphology, Abbott 2031 (Us, pro parte) is
probably an Alsophila X Nephelea hybrid but may not
have the same parents as Cyathea irregularis,

7. Cyathea jamaicensis Jenm. Jour. Bot. 20: 323. 1882.
TYPE: Jamaica, Mansfield, near Bath, Wilson 686 (holo-
type, BM; isotype, GH!).

454 Rhodora [Vol. 77

The bipinnate-lobed Cyathea jamaicensis is known only
from the type collection. It is similar to Nephelea con-
cinna, but may be separated from it by its pinna rachis
indument and by its indusium. Cyathea jamaicensis has
the undersurfaces of its pinna-rachises nearly without
scales and trichomes and it has a meniscoid indusium.
Nephelea concinna has its pinna-rachis undersurfaces quite
well invested with scales and trichomes and it has a cyathi-
form to urceolate indusium. Also, C. jamaicensis is de-
scribed as having an unarmed stem whereas N. concinna
has a spiny stem.

It seems probable that Cyathea jamaicensis is a hybrid
because of its lamina architecture and its rarity. The lack
of spines on its trunk implies that it could not be an intra-
generic cross involving two species of Nephelea. The de-
gree of lamina dissection of C. jamaicensis suggests that a
bipinnate-pinnatifid species of Nephelea has crossed with
a pinnate-pinnatifid species of Alsophila.

CONCLUSIONS

Two certain and five putative hybrids have been pro-
posed. All combine characters of species with very differ-
ent leaf architecture, lamina indument, and (or) indusia.
The implication is that other species with similar leaf
architecture, lamina indument, and (or) indusia should
also cross. These hybrids would be very difficult or im-
possible to detect. Some polymorphic species in current
taxonomie treatments may require reassessment because
hybrids may have been included in the taxa. Careful field
observations of species that are growing together, when-
ever tree fern collections are made, will help to solve this
problem,

The presence of well developed spores in the Alsophila
dryopteroides X Nephelea portoricensis hybrid suggests
that these genera are closely related. The apparent absence
of well developed spores in the Cnemidaria horrida X
Cyathea arborea hybrid implies a more distant relation of
the parents.

1975] Cyatheaceae — Conant 455

ACKNOWLEDGEMENTS

The writer wishes to express sincere thanks to the Puerto
Rico Nuclear Center, U.S. Atomic Energy Commission,
Rio Piedras, Puerto Rico, and particularly to Dr. Richard
С. Clements, Director, Division of Terrestrial Ecology, for
making Student Research Grants available during 1972
and 1973 when collections and field studies were made.
Dr. Rolla M. Tryon has contributed to the organization and
presentation of ideas and conclusions embodied in this
paper. Thanks are due to Dr. Alice F. Tryon and Paulo G.
Windisch for help in preparing the plates and for percep-
tive discussion. I am indebted to Bruce Tiffney and to
Dr. Elso Barghoorn for assistance in the use of the photo-
graphic equipment of the Paleobotanieal Laboratory at
Harvard. Many thanks go to Jose and Elvira Colon, and
to Michael Burke for assistance in making collections and
in pressing specimens.

LITERATURE CITED

GASTONY, G. J. 1973. А revision of the fern genus Nephelea.
опр. Gray Herb. 203: 81-148.
HorrrUM, К. E. 1963. Cyatheaceae. Flora Malesiana II, 1(2):
65-176.
1974. The tree ferns of the genus Cyathea in Borneo.
Garden’s Bull. 27: 167-182.
‚ & U. SEN. 1961. Morphology and classification of the
tree ferns. Phytomorphology 11: 406-420.

BIOLOGICAL LABORATORIES
HARVARD UNIVERSITY
CAMBRIDGE, MASS.

ZOSTERA MARINA L., ITS GROWTH
AND DISTRIBUTION IN THE
GREAT BAY ESTUARY, NEW HAMPSHIRE!

STANLEY А. Bees, JR. AND RICHARD A. FRALICK*

Protected coastal and estuarine waters of New England
often contain dense meadows of the marine Angiosperm
Zostera marina L. Little was known of the ecology of
Zostera, or eel grass as it is commonly called, until a wast-
ing disease attributed to the parasitic marine fungus Laby-
rinthula almost exterminated the Zostera population in
New England waters (Jepps, 1931; Renn, 1934, 1935,
1936a, 1936b; Young, 1937, 1943).

The present paper compares the seasonal growth (bio-
mass), reproductive phenology, and local distribution of
Zostera with seasonal variations in temperature and salin-
ity. The results of vertical and horizontal transplants are
discussed as well as the possibility of ecotypical adaptation.

METHODS AND MATERIALS

Bimonthly observations and measurements of Zostera
marina populations were made at eight stations within the
Great Bay estuary system during 1972 (Table 1). Meas-
urements of surface water temperature and salinity were
made at mean low water (MLW). Biomass was also re-
corded at MLW since the majority of Zostera plants oc-
curred in this zone.

1Jackson Estuarine Laboratory Scientific Contribution Number 27.

2Present address: Department of Natural Science, Plymouth State
College of the University of New Hampshire, Plymouth, N.H. 03264.

456

1975]

Zostera — Riggs & Fralick

457

Table 1. Characteristics of eight stations within the
Great Bay Esturary.
Stations/ Temperature Salinity Substrate
nautical °C EAT
miles inland
1. Jaffrey Point avg. 10.2 avg. 30.3 rocky outcrops
(0.0) max. 23.0 max. 32.5 with some sand
min. 1.0 min. 26.0
2. Pierce Island avg. 10.5 avg. 27.0 broken rocks, mud
(2.0) max. 21.5 max. 32.0 and silt
min 0.0 min. 19.0
3. Newington Town avg. ПЛ avg. 24.5 small rocks, mud
Landing (5.5) max. 24.0 max. 231.0 and silt
min. 0.4 min. 16.0
4. Dover Point avg. 12.5 avg. 28.7 large rock outcrop-
(7.0) тах. 25.0 max. 32.0 pings, mud and silt
min. 3.0 min. 15.0
5. Cedar Point ave. 13.6 avg. 22.1 mud and silt
(8.3) max. 27.0 max. 32.0
min. 0.0 min. 6.0
6. Adams Point avg. 18.8 avg. 21.0 rock outcroppings,
(10.7) max. 28.0 max. 31.5 shale, cobble,
min. 0.5 min. 8.0 mud and silt
7. Chapmans avg. 14.2 avg. 5.7 mostly mud,
Landing (15.2) max. 26.5 max. 22.0 some small
min 0.0 min. 0.0 rocks and boulders
8. Exeter avg. 14.4 ave. 3.4 mud and silt
(19.1) max. 28.1 max. 20.0
min 0.0 min 0.0

458 Ећодога [Уо1. 77

Growth studies were initiated during March, 1972.
Tagged plants (approximately 100) were established at
Stations 2 through 6, and the blade length was measured
according to the method of McRoy (1970). Horizontal and
vertical transplants of Zostera were also initiated in March,
and the plants were subsequently observed and measured
for a period of 10 months. Each of the transplants was
duplicated, one using the substrate of the original station
and the other using the substrate of the new station.

McRoy (1970) showed that a rapid estimate of the stand-
ing crop of Zostera is possible because of a relationship
between blade length and dry weight. By incorporating
both the width and the length of the blades we derived and
untilized the following equation for conversion to dry
weight (biomass) without harvesting or damaging the
plants.

Formula Example:

Y = (0.308 X 107) (x) — 48.2 X 10+
Y — dry wt. (grams)
x—L*xW (mm)

blade width — 5 mm

blade length — 100 mm

L X W = 500

Y = (0.308 X 10%) (500) — 48.2 X 10+

predicted — 10.6 X 103 g dry wt.

actual — 10.8 X 103 g dry wt.

A linear regression was performed between the predicted
dry weight and the actual dry weight, and a correlation co-
efficient of 0.99 was obtained.

Temperature was recorded with a laboratory grade, sub-
mersible, mercury thermometer. Salinity was determined
with a sot of hydrometers (G.M. Manufacturing Co., N.Y.)
and the readings were corrected to 15°C.

1975] Zostera — Riggs & Fralick 459

Hydrographic factors: Figure 1 shows the seasonal vari-
ation in temperature at two representative stations for 12
months. Stations 1 and 2 exhibited similar temperature
values with maximum temperatures of 28.0° and 21.5° С
occurring in July and minimum temperatures 1° and 0° C
occurring in January, February, and December. Stations
3 and 4 exhibited maximum temperature values of 25.0°
to 26.0° C in July with minimum values of 0° C during the
winter months. Stations 6, 7, and 8 showed the greatest
ranges in temperature with maximum values of 27.5° C
in July and minimum values of 0° C in January, February,
and December.

FIG. 1
SH i-initial Growth STATION 2 ———

а : STATION 6
Ь-Віотаѕѕ Мах. +

TEMPERATURE (С)

О
F ME Г ТИ ЕЕ cua хемо

MONTHS (1972)

=

460 Ећодога [Vol. 77

Figure 2 shows the seasonal variation of salinity at four
representative stations for 12 months. The values are con-
sistent with the proximity of each station to the open coast.
Thus, Station 1 had a seasonal salinity range of 26.0 to
32.5 о/оо.; Station 2 showed a range of 19.0 to 32.0 о/оо.;
Station 6 had a range of 8.00 to 21.0 o/oo., while Station 8
had the lowest salinities within a range of 0.0 to 20.0 0/oo.

304

257

20-

SALINITY (%)
л

i-Initial Growth
f-Flowering STATION 2 ——
b-Biomass Max. STATION ЗЕ ——

[| Т

F M А M J J A
MONTHS

T T т 1
о М D J

Un ~

Seasonal growth: Figure 3 indicates that active growth
of Zostera was initiated during March and April at Sta-
tions 2, 3, 5, and 6. The plants at Stations 2 and 3 exhibited
а conspicuous elongation of the blades (1 to 3 cm) in
March. The plants at Stations 5 and 6 did not initiate
growth until the end of April. Growth at Stations 2, 3, 5,
and 6 declined rapidly during November and December
and no growth was recorded in January and February. No

1975] Zostera — Riggs & Fralick 461

30 | FIG. 2b
AN
^ "а b
5f IN A 6b: э... =
HN spe
254 (EN s А ,
t | Ыб 4 HI 4
\ t ii , L6 2 SS
INDE ` Le ~“ /
LA ; E
a2 ГА V P is ud Ce |:
A 7! к” „Му
20 ; xi UN | / ! А
SEN H ! 23
oe ЈЕ ! E
= | |
ў C?
= 154 BO уља
= | *5i a
< PAK
л : 9.
FECE
ө;
y
DG SE
i-Initial Growth
f-Flowering STATION 5 «——-
b-Biomass Max. STATION (6/6: -
О T T T | T Џ T T T T
F M A M J J A 5 О N D J

MONTHS (197 2)

growth was observed at Stations 1, 7, and 8 throughout the
season. The growth at Station 4 is not illustrated because

the colony was considered subtidal, while all other stations
were intertidal.

Standing crop: Figure 3 illustrates the seasonal varia-
tion in standing crop of Zostera at several stations. Mini-
mum values were apparent from December through March.
Maximum standing crop values were recorded in August
at Station 5. The plants at Station 6 reached their maxi-
mum biomass in late September. Two peaks were evident
at Stations 2 and 3; the first occurred in mid-June and the
second in early August. The initial peak in biomass was
associated with flowering, while a decrease in early July
was attributed to the abscission of floral parts. In some

462 Ећодога [Vol. 77

1005 STATION 2 —

STATION 3=—--.
STATION 67 ,

Biomass Grams X 10

MONTHS (1972)

cases vegetative turions without reproductive parts re-
mained intact and continued to grow.

A general decline in standing crop was apparent at
nearly all stations by late September, and although some
growth continued until December, no significant increase
in biomass was recorded at any station after October. The
process of floral abscission was not apparent at Station 6
until early October.

Horizontal transplants: Transplants were also initiated
between various stations during the spring. In all cases the
plants showed their maximum standing crop values at the
same time as the original populations from which they
were derived. For example, Zostera transplanted from
station 2 to 6 reached a maximum standing crop value in
mid-June, while the original material at Station 6 did not
peak until late September (Fig. 4). Zostera at Station 4
reached a maximum standing crop value in July.

1975] Zostera — Riggs & Fralick 463

FIG. 4

Se STATION 2
STATION 49———
STATION б<. :

A intertidal to subtidal (STA. 6) - ———.

20-4

Biomass Grams X 10
a
|

104

5

MONTHS

Transplants of Zostera plants (9.0 to 16.0 ст long) from
Station 2 to 8 died within a week. Differential responses
to transplanting were observed when smaller plants (2.0
to 8.0 em long) survived a transplant between the same
stations for more than 30 days.

DISCUSSION

Within the Great Bay Estuary system, Zostera first
initiated growth, flowered, and reached maximum biomass
at the stations nearest the coast. Stations further up in
the estuary initiated growth, flowered, and reached a maxi-
mum biomass some three months later than stations close
to the coast.

The initial growth of Zostera, in contrast to Setchell's
1929 findings, is neither restricted to a specific 5 degree

464 Ећодога [Vol. 77

temperature isotherm, nor to temperatures greater than
10° С. Although Zostera has been observed living beneath
winter sea ice (McRoy, 1969), to our knowledge its growth
has not previously been measured at temperatures below
10° С,

The initial period of maximum biomass did not appear
to be related to any specific temperature range. Thus,
plants at Stations 2 and 3 reached their maximum biomass
at 18° C, while it was attained at 25° C at Station 5 and
16° C at Station 6.

The flowering period is not restricted to a specific tem-
perature isotherm (15° to 20° C) as suggested by Setchell,
1929. Zostera flowered in the spring at Stations 2 and 3
when the temperatures were about 17.0° C. At Station 6
flowering did not take place until fall despite an increase
in temperature up to 28.0° C.

There was a correspondence between peak biomass and
high salinity at each site. Thus, Zostera plants at Stations
2 and 8 reached their first maximum biomass at salinities
of 26.0 о/оо and 24.0 о/оо respectively, while maximum bio-
mass at Station 5 was reached at a salinity of 25.0 0/00 —
40 days later than at Station 2. Zostera plants at Station 6
reached maximum biomass at a salinity of 27.0 o/oo, in
September, nearly 100 days later than at Stations 2 and 3.

The time of flowering was also correlated with salinity.
Thus, Zostera plants at Stations 2 and 3 initiated flowering
when the salinities were 26.0 and 24.0 o/oo. The plants at
Stations 5 and 6 also initiated flowers at the same salini-
ties, but approximately three weeks later than at Stations
2 and 3.

It appears that Zostera marina maintains its original
growth patterns even after being transplanted to new loca-
tions. Thus, transplants from Stations 2 and 4 to 6 fol-
lowed a pattern of growth and development similar to that
of the plants in their original colony. In no case did trans-
planted Zostera folow a sequence of growth and pheno-
logical development parallel to patterns in plants originat-
ing in the new location.

1975] Zostera — Riggs & Fralick 465

The horizontal transfer of Zostera colonies within the
estuary had no noticeable effects on their development.
Vertical transplants (intertidal to subtidal) frequently
showed a decrease in their rate of development. Thus,
when young seedlings (1 to 3.0 cm long) were transplanted
vertically their development was conspicuously slowed
(Fig. 4). In contrast, transplants from the subtidal to the
intertidal zone were consistently successful. Transplants
to brackish waters with salinities of 4.0 to 5.0 o/oo did
poorly. The greatest level of success in transplants to
brackish water occurred when plants 3 cm or less in length,
originating at stations close to the brackish water, were
transplanted with their rhizomes intact.

A distinct potential for vegetative propagation was ob-
vious throughout our transplant studies. In most cases
transplants of individual Zostera plants responded favor-
ably. The only exception occurred with transplants from
intertidal to subtidal locations.

A lack of clear correlation was evident between the onset
of the plants’ various phenological phases and temperature.
However, there was some indication that the onset of
various phenological stages may be, in part, related to
moderate salinity ranges of 24 to 27 o/oo. In all instances,
phenological development started in plants nearer the open
coast and progressed up the estuary. A tendency of Zostera
to form ecotypes between stations which differed in their
seasonal salinity patterns was obvious. However, further
studies to determine the effects of temperature related
salinity tolerances may be warranted. McMillan (1956)
has noted a tendency for some Graminae to adapt towards
different habitats. In addition Biebl and McRoy (1971)
have noted that Zostera may exhibit a physiological adap-
tation to a particular environment. This fact was obvious
in our investigations because Zostera could be collected in
distinctly subtidal habitats at depths as great as 6 meters.
Since there is no possibility of these plants ever being ex-
posed to the atmosphere, we consider them to be good
examples of adaptation to a subtidal existence.

466 Rhodora [Vol. 77

Our results indicate that Zostera marina may undergo
successful ecotypical adaptation toward a particular en-
vironment. This adaptation is probably more closely re-
lated to moderate salinities than specific temperatures.
We also have shown that transplants of Zostera between
habitats is feasible.

The mathematical formula employed in this investigation
proved to be an accurate method of assessing biomass for
Zostera without harvesting the plants.

LITERATURE CITED

BIEBL, R., & C. P. McRoy. 1971. Plasmic resistance and rate of
respiration and photosynthesis of Zostera marina at different
salinities and temperatures. Marine Biol. 8: 48-56.

JEPPS, M. W. 193i. Note on a marine Labyrinthula. Jour. Mar.
Biol. Assoc. U. K. 17: 833-838.

McRov, C. P. 1970. Standing stocks and other features of eel-
grasses (Zostera marina) populations on the coast of Alaska.
Jour. Fish. Res. Board Canada 27: 1811-1821.

RENN, C. E. 1934. Wasting disease of Zostera marina in Ameri-
can waters. Nature 134(3385): 416.

1935. A mycetozoan parasite of Zostera marina.

Nature 135: 544, 545.

1936a. The wasting disease of Zostera marina. Biol.

Bull. 70: 148-158.

1936b. Persistence of the eelgrass disease and para-
site on the American Atlantic coast. Nature 138: 507-508.

YouwG, E. L. 1937. Notes on the labyrinthulan parasite of the eel-
grass, Zostera marina. Bull Mt. Desert Island Biol. Lab. pp.
33-35.

1943. Studies on Labyrinthula, the etiologic agent of
the wasting disease of eelgrass. Am. Jour. Bot. 30: 586-593.

DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
AND JACKSON ESTUARINE LABORATORY

UNIVERSITY OF NEW HAMPSHIRE

DURHAM, NEW HAMPSHIRE 03824

OBSERVATIONS ON
BATRACHOSPERMUM (RHODOPHYTA)
IN SOUTHEASTERN WISCONSIN STREAMS

WILLIAM J. WOELKERLING

Knowledge of the genus Batrachospermum (Rhodophyta)
in Wisconsin stems mainly from the report of Prescott
(1951) who records four species from lotic environments
without reference to locality and with only scant ecological
data. Moreover, information on seasonal periodicity and
environmental conditions of the type obtained by Dillard
(1966) in North Carolina, Minckley and Tindall (1963) in
Kentucky, and Rider and Wagner (1972) in Pennsylvania
apparently is lacking for Wisconsin. (The last reference
includes a literature review of Batrachospermum ecology.)

The present investigation has been undertaken to gain
information on 1) the occurrence of Batrachospermum in
southeastern Wisconsin streams and 2) the environmental
conditions present at localities where this taxon grows.
During the course of the study 201 randomly selected
stream localities have been visited; Batrachospermum
plants oceurrec at 13 or 6.4% of these sites.

MATERIALS AND METHODS

At each station (Table 1) where Batrachospermum
plants were found, chemical and physical data on alkalinity,
carbon dioxide, hardness (calcium and total), nitrate ni-
trogen, orthophosphate, oxygen, pH, temperature, and
turbidity have been gathered using a Hach Water Analysis
Field Kit, model DR-EL, which employs microadaptations
from the “Standard Methods" handbook (American Public
Health Association 1965). In addition, some observations
on the type of substrate, relative current velocity, and
relative exposure to sunlight have been made.

At stations where Batrachospermum populations per-
sisted for extended periods, observations were made at
4-6 week intervals for 10-11 months to note any seasonal
changes.

467

[Vol. 77

Ећодога

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470 Ећодога [Vol. 77

Voucher specimens from all localities have been collected
and immediately preserved in FAA (10:7:2:1 95% etha-
nol: water :formalin:glacial acetic acid). Dried herbarium
specimens (bearing numbers prefaced by WJW) and per-
manent microscope slides using KARO as a mountant
(Woelkerling, 1970) as well as liquid preserved material
have been retained in the author’s personal collections,
currently housed at WIS. Species determinations have
been made primarily with the aid of the taxonomic key of
Israelson (1942); the papers of Kylin (1912), Prescott
(1951), Sirodot (1884), and Whitford and Schumacher
(1969) also have been consulted.

RESULTS AND DISCUSSION

Two taxa of Batrachospermum found during this study
have been identified to species. Batrachospermum borya-
num Sirodot, not recorded previously from Wisconsin,
occurred at seven localities (B, F, K, SC, SR, SS, T; ab-
breviations explained in Table 1), and with one exception
(locality B), it always grew mixed with other Batracho-
spermum taxa. Sexually mature plants ranged in size from
4-21 em with most plants averaging about 8-10 em tall.
Sexual plants have been encountered in all months except
February, March, and October; further study will probably
show that sexual] plants do occur throughout the year.

Batrachospermum moniliforme Roth has been found at
ten localities (BE, F, J, K, MC, N, SC, SR, SS, T), and
has been reported previously from Wisconsin (Prescott,
1951). Except for two stations (J, N), it grew mixed with
other Batrachospermum taxa. Sexually mature plants oc-
curred throughout the year and ranged in length from
3-25 cm with an average height of 6-9 ст.

In addition to the above two taxa, sterile (and thus spe-
cifically unidentifiable) plants of Batrachospermum have
been encountered at five localities (BE, M, Q, SC, T) from
January-March, May-July, and in December, They varied
in length from 3 cm to 15 cm. The size of some Wisconsin

1975] Batrachospermum — Woelkerling 471

plants (up to 25 ст) greatly exceeds the 6 ст maximum
recorded by Rider and Wagner (1972) and the 10 cm maxi-
mum found by Israelson (1942).

All Wisconsin populations of Batrachospermum observed
during this investigation grew at or near the headwaters
of spring-fed streams, thus agreeing with the findings of
Minckley and Tindall (1963). Rider and Wagner (1972)
also recorded their taxa from a spring-fed stream but
without mention of the headwater areas. Current veloci-
ties at Wisconsin localities never dipped below 10 cm/sec
and in most cases exceeded 25 cm/sec. In addition, all
Wisconsin localities but one (BE) contained rocky or
rocky-sandy bottoms and usually appeared free from heavy
siltation and high turbidity levels.

Except for seasonal fluctuations in temperature and
diurnal fluctuations in carbon dioxide and oxygen levels,
chemical ana physical conditions at any one locality tended
to remain within relatively narrow limits during the study
period. Conditions did vary considerably between localities,
however, and the taxa of Batrachospermum encountered
appear to tolerate a fairly wide range of environmental
conditions (Table 2).

Of particular note is the variation in carbon dioxide
levels. At no time did СО. levels exceed 24 ppm, and levels
as low as 1 ppm have been encountered. These values are
decidedly lower than those reported by Minckley and Tin-
dall (1963) for Batrachospermum sp. and by Rider and
Wagner (1972) for B. vagum, but they more or less agree
with the range in values measured by Rider and Wagner
(op. cit.) for B. moniliforme. Since both species found in
Wisconsin (B. moniliforme and B. boryanum) apparently
require free СО, for photosynthesis (Ruttner, 1960), data
from the present study strongly suggest that these taxa
can survive at very low concentrations of free CO., at
least for short periods of time.

At six of the thirteen localities (B, J, M, MC, N, Q),
Batrachospermum plants were encountered on only one
oecasion (Table 1), and data for three additional localities

412

Ећодога

TABLE 2

RANGE IN ENVIRONMENTAL CONDITIONS
UNDER WHICH BATRACHOSPERMUM OCCURRED

[Vol. 77

B. boryanum В. moniliformis B. spp.
Alkalinity 15-342 ppm 12-387 ppm 287-368 ppm
Carbon

Dioxide 2-23 ppm 1-23 ppm 2-20 ppm
Hardness,

Calcium 2-340 ppm 15-410 ppm 176-408 ppm
Hardness,

Total 22-442 ppm 29-500 ppm 352-420 ppm
Nitrate 1.0-4.8 ppm 0.2-16.5 ppm 2.0-10.0 ppm
Oxygen 8-20 ppm 5-21 ppm 5-17 ppm
pH 6.2-8.4 6.7-8.4 7.2-8.2
Ortho-

phosphate 0.007-4.0 ppm 0.02-4.9 ppm 0.02-8.1 ppm
Temperature 6°-18°C 1°-21°С 4°-22°С
Turbidity 0-7 J.U. 0-10 J.U. 0-15 J.U.
TABLE 3

RANGE IN SELECTED ENVIRONMENTAL
CONDITIONS AT SCUPPERNONG CK. (SC)
AND TICHIGAN CK. (T) DURING STUDY PERIOD

Environmental Factor 5С т
Alkalinity 275-310 ppm 300-360 ppm
Carbon Dioxide 2-23 3-18 ppm
Hardness, Ca 190-225 ppm 170-250 ppm
Hardness, Total 350-400 ppm 375-425 ppm
pH 7.5-8.4 7.7-8.3
Temperature 4-15°С 5-22°C

1975] Batrachospermum — Уое Кет та 473

(BE, SR, SS) are confined to three or four dates. Conse-
quently, information on seasonal changes is restricted to
limited observations at four localities (F, K, SC, T) from
either May or June, 1972 through April, 1973. Batracho-
spermum boryanum and В. moniliforme occurred in mixed
populations at all four sites; consequently, reference to
Batrachospermum in the ensuing discussion includes both
taxa.

The Fontana population (F) grew in very hard water
(Ca hardness = 250 ppm СаСО.; total hardness 410 ppm
CaCO.), swift flowing stream 1-2 m across whose tem-
tures ranged from 12°С in summer to 6°C in winter, Car-
bon dioxide levels varied from 2-23 ppm, pH from 7.8-8.4,
and alkalinity from 330-390 ppm. The stream bottom was
primary gravel. During most of the day the habitat was
exposed to full sunlight.

When first discovered in June, 1972, Batrachospermum
plants occurred in considerable numbers and reached
lengths of up to 18 ст. By July, 1972, however, most of
the plants had disappeared or were obviously moribund.
Several plants (preserved as WJW 3936) appearec heavily
caleified. The population disappeared entirely by August,
and new adult gametophytes did not become apparent until
December, 1972, when about a dozen plants up to 6 ст tall
were discovered. By January, 1973, Batrachospermum had
become the dominant alga in the stream with most plants
averaging 3-6 em in length. Throughout the remainder of
the study period (ending in April, 1973), Batrachosper-
mum maintained its dominance in the stream and plants
gradually increased in size to 12-15 cm on the average.

Similar seasonal fluctuation in population levels also oc-
eurred at Koshwego Springs (K), where certain chemical
and physica! conditions differed considerably from those
at Fontana. At Koshwego, the water was very soft (Ca
hardness = 15 ppm CaCO,; total hardness ~ 25-30 ppm
CaCO,), acid (pH varied from 6.2-7.0), and showed alka-
linity readings of 12-34 ppm. Stream bottom varied from
rocky to sandy to partially silty with Batrachospermum

474 Rhodora [Vol. 77

confined to rocky areas, At all times, the habitat was sub-
jected to deep shade. In other respects the two localities
appear more or less similar; at Koshwego, temperature
varied from 12°C in summer to 1.0°C in winter and CO,
levels fluctuated between 1 and 11 ppm.

In May, 1972, Batrachospermum plants up to & cm tall
dominated the stream vegetation, but by mid-July they had
become very moribund or had disappeared. Small (i.e.,
less than 2.5 cm tall) plants reappeared in considerable
numbers in November, 1972, and dominated the stream
vegetation throughout the winter. Noticeable increase in
size occurred between February (average size under 2.5
ст) and March, 1973 (average size 6 ст). Severe flooding
and silting of the stream occurred in late March and early
April, 1973, and the Batrachospermum population was al-
most entirely destroyed.

At the remaining two stations (SC, T) Batrachospermum
plants occurred throughout the year and formed the domi-
nant component of the algal vegetation during much of that
time. Both stations had environments (Table 3) similar to
that at Fontana except that one (SC) was largely shaded
throughout the day and the other (T) was exposed to full
sunlight during most of the day. In addition the latter (T)
had summer temperatures of 18-22°С or 5-10°C higher
than at the other hard water localities.

Immediately below the spring from which Scuppernong
Creek originates, Batrachospermum plants constituted the
dominant form of vegetation. Within 100 m, however,
angiosperm vegetation became dominant and the Batracho-
spermum population consisted only of scattered plants.
During winter months most plants encountered were 4-6 cm
long and during summer they were 8-10 cm long; one 15
ет tall plant was encountered.

The Tichigan Creek population of Batrachospermum
dominated the macroscopic vegetation throughout the year.
Summer plants generally did not exceed 10 cm in length;
winter plants all (i.e, December-April) were enormous in
size and reached lengths of up to 25 cm. The very large

19751 Batrachospermum — Woelkerling 415

size of these individuals as compared to the other winter
populations studied could not be accounted for on the basis
of the physical or chemical parameters examined during
the study.

The above observations suggest that different populations
of Batrachospermum (at least in Wisconsin streams) may
either produce mature gametophytes throughout the year
(SC, T) or show seasonal variation with an absence of ma-
ture plants from mid-summer to late fall (F, K). They also
suggest that maximum vegetative development can occur
in spring (F, K), summer (SC), or winter (T). Previous
American studies (Dillard, 1966, Minckley and Tindall,
1963, Rider and Wagner, 1972) all reported definite sea-
sonal fluctuations in Batrachospermum populations with a
disappearance of plants in summer and a reappearance in
fall. Yoshida (1959), however, makes mention of both
seascnal and year-round populations of Batrachospermum
in Japanese streams.

Various attempts have been made to account for seasonal
fiuctuation in population levels in terms of temperature
changes, changes in light intensity, and differences in cur-
rent velocity (see Dillard, 1966, Minckley and Tindall,
1963, Rider and Wagner, 1972, Yoshida, 1959). The re-
sults of the Wisconsin study, however, indicate that mature
plants and maximum vegetative development can occur
under both low and high light intensities and under both
summer and winter temperatures. Therefore, other fac-
tors, perhaps genetic, appear to be involved in determining
why different populations of the same species either persist
year round or show seasonal fluctuations. No relationship
to current velocity has been observed in this investigation.

The apparently consistent occurrence of Batrachosper-
mum in the headwater areas of spring-fed streams likewise
requires further investigation. Minckley and Tindall (1963)
suggest that the availability of unbound carbon dioxide
may be a controlling factor (their stream reportedly has
super-saturated СО, levels), but the relatively low CO,
levels found during this study again suggest that other

476 Ећодога [Vol. 77

factors may be involved, and additional study appears
warranted.

SUMMARY

The occurrence and some ecological aspects of Batracho-
spermum in southeastern Wisconsin streams have been
investigated. Batrachospermum boryanum (newly reported
for Wisconsin), B. moniliforme, and Batrachospermum sp.
occurred in 6.4% of the localities visited and were found
in both alkaline, hard water and acid, soft water environ-
ments. Depending upon the population, mature plants per-
sisted throughout the year or disappeared in summer and
fall, and they showed maximum vegetative development in
spring, or summer, or winter. The seasonal behavior does
not appear to be correlated entirely with changes in light
intensity or temperature. Likewise the apparent occurrence
of Batrachospermum near the headwaters of spring-fed
streams apparently cannot be explained solely on the basis
of greater availability of unbound carbon dioxide in these

habitats.

ACKNOWLEDGMENTS

Sincere thanks are due Mr. Robert Dietrich and Mr.
Warren Mueller for assistance in the gathering and pro-
cessing of data. This study was supported by grant No.
130376 from the Research Committee of the University of
Wisconsin Graduate School.

LITERATURE CITED

AMERICAN PUBLIC HEALTH ASSOCIATION, The American Water Works
Association, and the Water Pollution Control Federation. 1965.
Standard Methods for the Examination of Water and Waste-
water. ed. 12, 769 pp. New York.

DiLLARD, С. E. 1966. The seasonal periodicity of Batrachospermum
macrosporum and Audouinella violacea in Turkey Creek, North
Carolina. Jour. Elisha Mitchell Sci. Soc. 82: 204-207.

ISRAELSON, С. 1942. The freshwater Florideae of Sweden. Symb.
Bot. Upsal. 6: 1-135.

1975] Batrachospermum — Woelkerling 411

Кушм, Н. 1912. Uber die roten und blaue Farbstoffe der Algen.
Hoppe-Seylers Z. Physiol. Chem. 76: 897-425.

MiNCKLEY, W. L., & D. R. TINDALL. 1963. Ecology of Batracho-
spermum sp. in Doe Run, Meade County, Kentucky. Bull. Tor-
rey Bot. Club 90: 391-400.

Рвезсотт, 6. W. 1951. Algae of the Western Great Lakes Area.
946 pp. Cranbrook Inst. Sci.

River, D. E., & R. Н. Wagner. 1972. The relationship of light,
temperature, and current to the seasonal distribution of Batra-
chospermum (Rhodophyta). Jour. Phycol. 8: 323-331.

RuTTNER, Е. 1960. Carbon uptake in algae of Rhodophycean genus
Batrachospermum. Schweiz. Z. Hydrol, 22: 280-291.

Створот, 5. 1884. Les Batrachospermes. Organisation, functions,
dévelopement, classification. 299 pp. Paris.

WHITFORD, L. H., & G. J. SCHUMACHER. 1969. A Manual of the
Freshwater Algae in North Carolina. 313 pp. Raleigh, N. C.

WOoELKERLING, W. J. 1970. Acrochaetium botryocarpum (Harv.)
J. Ag. (Rhodophyta) in southern Australia. Br. Phycol. Jour.
5: 159-171.

Yosuipa, T. 1959. Life cycle of a species of Batrachospermum
found in Northern Kyushu, Japan. Jap. Jour. Bot. 17: 29-42.

DEPARTMENT OF BOTANY
UNIVERSITY OF WISCONSIN
MADISON, WISCONSIN 53706

A COUNTY CHECKLIST OF THE FERNS
AND FERN ALLIES OF KANSAS, NEBRASKA,
SOUTH DAKOTA, AND NORTH DAKOTA!

ALETA JO PETRIK-OTT

INTRODUCTION

To the present, there has been no comprehensive work
dealing exclusively with the ferns and fern allies of what
is considered to be the heart of the central plains and
prairies of the United States, i.e., Kansas, Nebraska, South
Dakota, and North Dakota.

Of the larger floras, Rydberg’s “Flora of the Rocky
Mountains and Adjacent Plains" (1917) extends eastward
to longitude 102? W in Nebraska, South Dakota, and North
Dakota. Rydberg's “Flora of the Prairies and Plains of
Central North America" (1932) includes the ferns and fern
allies of the aforementioned four states, but several taxa
reported to be in these states have never been verified,
many new taxa have been found since that time, and recent
research has resulted in many nomenclatural changes.
Fernald's account of the ferns and fern allies in *Gray's
Manual of Botany" (1950) covers only the area in Ne-
braska and Kansas east of longitude 96° W and thus
excludes the prairie and plains regions of the Dakotas,
Nebraska, and Kansas.

Smaller, individual state floristic treatments briefly cov-
ering the ferns and/or fern allies are represented by Keller-
man and Kellerman (1888), Saunders (1899), Bergman
(1918), Petersen (1923), Over (1932), Gates (1940),
Stevens (1950), and Winter, Winter, and Van Bruggen
(1959). Floristic regional and state studies exclusively
concerning ferns and/or fern allies are represented by
Wilson (1885), Cragin (1885, 1886), Bessey (1892), Reed
(1896), Fitzpatrick (1919, 1920a & b), Humtíeld (1951a

!Contribution Number 554, Virginia Institute of Marine Science,
Gloucester Point, Virginia 23062.

418

1975] Ferns — Petrik-Ott 479

& b), McGregor and Hartman (1956), McGregor (1960),
Brooks (1967, 1969), and Van Bruggen (1967).

Supporting specimens are not to be found for many
records given in the existing literature, and many of the
older records were based upon mis-identifications. The
records given here are based entirely upon specimens from
nine major herbaria in the region and my personal collec-
tions which are deposited in the University of Kansas
herbarium. The nine herbaria are (abbreviations after
Lanjouw and Stafleu, 1964) :

KANU = The University of Kansas, Lawrence, Kansas

кзс = Kansas State University, Manhattan, Kansas

KsTC = Herbarium of the Kansas State Teachers
College, Emporia, Kansas

MO — Missouri Botanical Garden, 2315 Tower
Grove Avenue, Saint Louis 10, Missouri

NDA == North Dakota State University and Experi-
ment Station, Fargo, North Dakota

NEB — University of Nebraska State Museum, Lin-
coln 8, Nebraska

spc == Department of Botany, South Dakota State
College, College Station, Brookings, South
Dakota

SDU = University Herbarium, University of South
Dakota, Vermillion, South Dakota

UMO = University of Missouri Herbarium, Colum-

bia, Missouri

The following list was compiled during the years 1968-
1970. Citations for each taxon are grouped alphabetically
by state and county. Only one specimen for each county
was chosen as representative. A map (Fig. 1) showing the
counties in each state is included. To conserve space, it was
considered sufficient to give only the collector's name and
his collection number for each representative county speci-
men. In certain cases, a collection number was not given
on the herbarium label ; therefore, the date of the collection,

480 Ећодога [Vol. 77

if indicated, was designated. It is realized that the county
distributions of certain taxa have probably been extended
since the termination of my study, but it has not come to
my attention that any new taxa have been added.

Because there has been much confusion concerning the
presence of certain taxa, which cannot be verified by exist-
ing specimens, a section on doubtful collections and ex-
cluded taxa has been appended to the checklist.

THE FERNS
FAMILY OPHIOGLOSSACEAE

Botrychium dissectum Spreng. var. dissectum Anleit.
Kennt. Gewachse 3:172. 1804.

Kansas: CHEROKEE CO.: №. L. McGregor 3870 (KANU).

Botrychium dissectum Spreng. var. obliquum (Muhl.)
Clute, Fern Bull. 10:76. 1902,

Kansas: ANDERSON Co.: L. K. Magrath 4988 (KANU). BOURBON
co.: S. Stephens 19492 (KANU). CHEROKEE CO.: Ё. L. McGregor
3860a (KANU). COFFEY CO.: Г. K. Magrath 4979 (KANU). DOUGLAS
co.: J. E. Bare 762 (KANU). FRANKLIN CO.: L. К. Magrath 5086
and A. Organ (KANU). JEFFERSON CO.: R. L. McGregor 4154
(KANU). LINN CO.: L. К. Magrath 4988 (KANU). MIAMI СО.; ЈУ. Н.
Horr and R. І. McGregor 3617 (KANU). WYANDOTTE Con: L. К.
Magrath 5056 (KANU).

Botrychium lunaria (L.) Swartz, Schrad. Journ. Bot. 1800
(2) :110, 1801.
North Dakota: MCHENRY С0.: О. A. Stevens 1530 (NDA).

Botrychium multifidum (S. G. Gmel.) Rupr. Beitr. Pflan-
zenk. Russ. Reich. 11:40. 1859,

Nebraska: FRANKLIN CO.: Е. M. Hussong 4689 (NEB).
South Dakota: CUSTER CO.: S. Stephens 35426 and R. Brooks
(KANU). PENNINGTON CO.: T. Van Bruggen 5042 (800).

Botrychium simplex E. Hitche. Amer. Journ. Sci. 6:103.
pl. 8. 1823.

South Dakota: LAWRENCE CO.: C. A. Taylor 7963 (SDC).

1975]

Ferns — Petrik-Ott

481

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Figure 1.
North Dakota.

County map of Kansas

482, Ећодога [Vol. 77

Botrychium virginianum (L.) Swartz, Schrad, Journ. Bot.
1800 (2) :111. 1801.

Kansas: ANDERSON C0.: R. L. McGregor 3211 (KANU). ATCHISON
co.: A. J. Petrik-Ott 752 (KANU). BOURBON CO.: A. J. Petrik-Ott
756 (KANU). BROWN CO.: R. L. McGregor 2886 (KANU). CHASE CO.:
B. B. Smyth, 19 Aug., 1902 (Ksc). CHAUTAUQUA CO.: P. H. Humfeld
842 (KANU). CHEROKEE CO.: A. J. Petrik-Ott 724 (KANU). COFFEY
co.: L. K. Magrath 3876 (KSTC). COWLEY co.: W. H. Horr and
R. L. McGregor, 23 July 1947 (KANU). CRAWFORD CO.: А. J. Petrik-
Ott 725 (KANU). DONIPHAN on: R. L. McGregor 12258 (KANU).
DOUGLAS on: R. L. McGregor 14217 (KANU). FRANKLIN CO.: R. L.
McGregor 5592 (KANU). GEARY CO.: А. S. Hitchcock, June 1896
(KSC). GREENWOOD CO.: S. Stephens 2914 (KANU). JACKSON CO.:
R. L. McGregor 2853 (KANU). JEFFERSON CO.: R. L. McGregor 880
(KANU). JOHNSON CO.: P. H. Humfeld 595 (KANU). LEAVENWORTH
co.: P. H. Humfeld 195 (KANU). LINN CO.: A. J. Petrik-Ott 763
(KANU). LYON co.: J. S. Wilson 2042 (KSTC). MARSHALL CO.: S.
Stephens 3291 (KANU). MIAMI CO.: 8. Stephens 2312 (KANU).
MONTGOMERY CO.: R. L. McGregor 12824 (KANU). NEMAHA CO0.:
L. K. Magrath 4506 (KANU). OSAGE CO.: S. Stephens 30602 (KANU).
POTTAWATOMIE C0.: S. Stephens 4710 (KANU). RILEY CO.: Miss
Thackrey, 24 May 1892 (KsC). SALINE CO.: J. Hancin 375 (KSC).
SHAWNEE CO.: L. D. Volle T15 (KANU). WABAUNSEE СО.: S. Stephens
4798 (KANU). WILSON CO.: L. С. Hulbert 3686 (KSC). WOODSON CO.:
R. L. McGregor 905 (KANU). WYANDOTTE CO.: A. J. Petrik-Ott 747
(КАМП).

Nebraska: BURT on: S. Stephens 32072 and №. Brooks (KANU).
cass CO.: A. J. Petrik-Ott 776 (KANU). CHERRY CO.: J. M. Bates
6043 (NEB). DAKOTA CO.: S. Stephen 32048 and R. Brooks (KANU).
DAWES on: Н. J. Webber 6103 (NEB). DOUGLAS CO.: W. Cleburne,
9 July 1897 (NEB). GAGE CO.: Н. L. Shantz, June 1904 (NEB). HALL
co.: J. M. Bates 3807 (NEB). HOLT CO.: R. Brooks 374 and S. Stephens
(KANU). LANCASTER CO.: Н. J. Webber 6102 (NEB). NEMAHA CO.:
5. Stephens 3660 (KANU). OTOE со.: Н. J. Webber, 1 June 1889
(MO). RICHARDSON CO.: A. J. Petrik-Ott 773 (KANU). SARPY CO.:
W. Cleburne, 3 Tune 1887 (NEB). SIOUX CO.: А. F. Woods 444 (NER).
THOMAS CO.: P. A. Rydberg 1467 (NEB). THURSTON CO.: S. Stephens
21066 (KANU).

North Dakota: RENSON CO.: Н. Е. Bergman 1890 (NDA). BOTTI-
NEAU CO0.: Н. Е. Rergman 2550 (NDA). cass co.: Н. F. Bergman
1637 (NDA). DUNN CO.: О. A. Stevens and D. Р. Moir, 14 Sept.
1956 (NDA). PEMRINA CO.: L. R. Waldron 1649 (NDA). RANSOM CO.:
S. Stephens 33426 and Р. Brooks (KANU). RICHLAND CO.: О. A.
Stevens 424 (NDA). STARK CO.: Mrs. M. Meissner, 20 Aug. 1939
(NDA).

1975] Ferns — Petrik-Ott 483

South Dakota: CLAY co: С. B. Smith, 22 Sept. 1966 (SDU). HARD-
ING CO.: S. S. Visher 7057 (SDU). LAWRENCE CO.: S. Stephens 7395
(KANU). LINCOLN C0.: N. Winter, Oct. 1954 (SDU). MARSHALL CO.:
T. Van Brugggen 4448 (SDU). ROBERTS CO.: R. Brooks 443 (KANU).

Ophioglossum engelmanni Prantl, Ber. Deutsch. Bot. Ges.
1:351. 1883.

Kansas: ALLEN CO.: L. K. Magrath 4977 (KANU). ANDERSON CO.:
R. Н. Thompson, 19 June 1949 (KANU). ATCHISON CO.: R. L. Me-
Gregor 887 (KANU). BOURBON CO.: S. Stephens 3978 (KANU). CHAU-
TAUQUA on: Р. L. McGregor 4186 (KANU). CHEROKEE CO.: R. L.
McGregor 15328 (KANU). COFFEY CO.: L. K. Magrath 3836 (KSTC).
COWLEY CO.: Rudy С. Koch 3468 (кѕс). DOUGLAS CO.: Р. H. Humfeld
1011 (KANU). ELK CO.: S. Stephens 10754 (KANU). JOHNSON CO.:
R. Н. Thompson, 22 April 1941 (KANU). LEAVENWORTH CO.: R. L.
McGregor 886 (KANU). MIAMI CO.: P. H. Humfeld 219 (KANU).
MONTGOMERY CO.: P. Н. Humfeld 107 (KANU). NEOSHO CO.: W. W.
Holland 2622 (KANU). WILSON cO.: R. L. McGregor, 30 June 1947
(KANU).

Ophioglossum vulgatum L, Sp. Pl. 2:1062. 1753.
Nebraska: CHERRY on: J. M. Bates 5679 (NEB).

FAMILY OSMUNDACEAE

Osmunda regalis L. var. spectabilis (Willd.) A. Gray, Man.
ed. 2:600. 1856.

Kansas: WILSON со.: L. L. Kiefer, 5 Oct. 1961 (KANU). WOODSON
cc.: W. H. Horr, 10 Јшу 1950 (KANU).

Nebraska: E. M. Hussong 6776 (ХЕВ).

FAMILY ADIANTACEAE

Adiantum capillus-veneris L, Sp. Pl. 2:1096. 1758.
South Dakota: FALL RIVER CO.: A. J. Petrik-Ott 784 (KANU).

Adiantum pedatum L. Sp. Pl. 2:1095. 1753.

Kansas: ALLEN CO.: R. L. MeGregor 968 (KANU). ATCHISON СО.:
Р. І. McGregor 2814 (KANU). BOURBON C0.: R. L. McGregor 979
(KANU). BROWN CO.: R. L. McGregor 904 (KANU). CHEROKEE CO.:
R. L. McGregor 983 (KANU). COFFEY CO.: R. L. McGregor 918
(KANU). CRAWFORD CO.: R. L. MeGroger 981 (KANU). DONIPHAN
co.: A. J. Petrik-Ott 753 (KANU). DOUGLAS CO.: R. L. McGregor
17356 (KANU). FRANKLIN on: R. L. McGregor 10043 (KANU).
JACKSON CO.: M. Reed, 1 Sept. 1889 (KSC). JEFFERSON CO.: R. L.

484 Rhodora [Vol. 77

McGregor 885 (KANU). JOHNSON CO.: P. Н. Humfeld 611 (KANU).
LEAVENWORTH CO.: №. L. McGregor 2792 (KANU). LINN CO.: A. J.
Petrik-Ott 762 (KANU). MIAMI CO.: R. L. McGregor 896 (KANU).
NEOSHO со.: M. D. Ewing, Summer 1891 (Ksc). OSAGE co.: R. L.
McGregor 891 (KANU). SHAWNEE CO.: L. D. Volle 131 (KANU).
WILSON CO.: Н. Danell, June 1890 (кѕс). Woopson co.: R. L. Mc-
Gregor 917 (KANU). WYANDOTTE СО.: S. Stephens 10863 (KANU).

Nebraska: cass on: S. Stephens 3743 (KANU). CUMING on: L.
Bruner, July 1880 (NEB). DOUGLAS со.: W. Cleburne, June 1878
(NEB). LANCASTER CO.: H. J. Webber 6128 (NEB). NEMAHA CO.:
J. M. Bates 5145 (NER). oTOE con: K. L. Johnson 1584 and D. E.
Dallas (KANU). PAWNEE СО.: J. E. Shue, May 1896 (NEB). RICHARD-
son CO.: A. J. Petrik-Ott 770 (KANU). ROCK со.: J. M. Bates,
13 Aug. 1900 (NEB). SARPY CO.: W. L. Tolstead and Н. C. Reynolds,
17 Мау 1942 (NEB).

Cheilanthes alabamensis (Buckley) Kunze, Linnaea 20:4.
1847.

Kansas: CHEROKEE СО.: R. L. McGregor 3865 (KANU).

Cheilanthes feei Moore, Index Fil. 38. 1857.

Kansas: BARBER co: R. L. McGregor 14950 (KANU). CHAUTAUQUA
co.: A. J. Petrik-Ott 745 (KANU). ELLSWORTH CO.: R. L. McGregor
1312 (KANU). FORD CO.: R. L. McGregor 10928 (KANU). HODGEMAN
co.: R. L. McGregor 10935 (KANU). LINCOLN co.: D. E. Lantz,
3 May 1898 (ach, OTTAWA CO.: A. J. Petrik-Ott 769 (KANU).
RUSSELL ou: L. C. Hulbert 4346 (KSC). STANTON CO.: R. L. Mc-
Gregor 16115 (KANU).

Nebraska: BANNER CO.: P. A. Rydberg 479 (NEB). SCOTTS BLUFF
со.: S. Stephens 5481 (KANU).

South Dakota: CUSTER CO.: S. Stephens 6167 (KANU). FALL RIVER
со.: S. Stephens 5797 (KANU). HARDING CO.: S. Stephens and R.
Brooks 425 (KANU). LAWRENCE CO.: S. Stephens 7475 (KANU).
MEADE C0.: Е. J. Palmer 37017 (мо). PENNINGTON CO.: S. Stephens
7134 (KANU).

Cheilanthes lanosa (Michx.) D. C. Eat. in Torr. Rep. U. S.
and Mex. Bound. Surv. 2:234. 1859.

Kansas: CHAUTAUQUA CO.: А. J. Petrik-Ott 745 (KANU). CHEROKEE
co.: R. L. MeGregor 11063 (KANU). ELK со.: R. L. McGregor 926
(KANU). MONTGOMERY Co: R. L. McGregor 946 (KANU). WILSON
co.: R. L. McGregor 925 (KANU). woopson Co: R. L. Мебтедот
914 (KANU).

Cheilanthes tomentosa Link, Hort. Berol. 2:42. 1833.
Kansas: CHEROKEE CO.: R. L. McGregor 3866 (KANU).

1975] Ferns — Petrik-Ott 485

Notholaena dealbata (Pursh) Kunze, Amer. Journ. Sci. II.
6:82. 1848.

Kansas: ALLEN CO.: R. L. McGregor 3885 (KANU). ANDERSON
co.: R. L. McGregor 3209 (KANU). BOURBON CO.: A. J. Petrik-Ott
759 (KANU). CHAUTAUQUA CO.: R. L. McGregor 951 (KANU). CHERO-
KEE co: R. L. McGregor 971 (KANU). COFFEY CO.: R. L. McGregor
910 (KANU). COWLEY CO.: R. G. Koch 1764 (KANU). CRAWFORD СО. :
R. L. McGregor 973 (KANU). DOUGLAS CO.: R. L. McGregor 874
(KANU). ELK C0.: R. L. McGregor 15880 (KANU). FRANKLIN CO.:
R. L. McGregor 3309 (KANU). GEARY СО.: C. C. Parry, April 1873
(MO). GREENWOOD CO.: R. L. McGregor 994 (KANU). JEFFERSON СО. :
P. Н. Humfeld 132 (KANU). JOHNSON on: R. L. McGregor 3435
(KANU). LABETTE CO.: R. L. McGregor 955 (KANU). LEAVENWORTH
co.: P. H. Humfeld 1019 (KANU). LINCOLN CO.: D. E. Lantz, 3 May
1898 (KSC). LINN CO.: M. Campbell, 28 June 1890 (KSC). MIAMI
co.: P. H. Humfeld 214 (KANU). MONTGOMERY CO.: A. J. Petrik-Ott
735 (KANU). MORRIS CO.: C. C. Parry, 29 April 1878 (мо). NEOSHO
co.: R. L. McGregor 957 (KANU). POTTAWATOMIE CO.: P. H. Hum-
feld 396 (KANU). RILEY со.: W. A. Kellerman 19 Aug. 1887 (KSC).
SHAWNEE CO.: B. B. Smyth 70 (KSC). WILSON co.: В. Г. Wagen-
(uecht 2231 (KANU). woopsoN Co.: R. L. McGregor 992 (KANU).
WYANDOTTE CO.: S. Stephens 10858 (KANU).

Nebraska: cass co.: Т. A. Williams 6778 (NEB).

Pellaea atropurpurea (L.) Link, Fil. Sp. Hort. Reg. Bot.
Berol. Cul. 59. 1841.

Kansas: ALLEN CO.: R. L. McGregor 3332. (KANU). ANDERSON CO.:
Р. L. McGregor 3209 (KANU). ATCHISON CO.: R. L. McGregor 2811
(KANU). BARBER CO.: Е. L. Richards 3716 (KANU). BARTON CO.:
S. Stephens 8380 (KANU). BOURBON CO.: A. J. Petrik-Ott 757 (KANU).
CHAUTAUQUA CO.: P. Н. Humfeld 855 (KANU). CHEROKEE CO.: Е. L.
Richards 3104 (KANU). CLAY co.: R. L. McGregor 5016 (KANU).
CLOUD en: S. V. Fraser 558 (Ksc). COFFEY CO.: R. Г. McGregor
913 (КАМ). COMANCHE CO.: R. L. McGregor 10944 (KANU). COWLEY
co.: S. Stephens 3020 (KANU). CRAWFORD CO.: C. L. Merritt, Sum-
mer 1941 (KSC). DONIPHAN o: F. Agrelius, Aug. 1913 (KSTC).
DOUGLAS CO.: В. L. McGregor 640 (KANU). ELK CO.: №. L. McGregor
15877 (KANU). ELLSWORTH co: R. L. McGregor 17326 (KANU).
FRANKLIN CO0.: R. L. McGregor 889 (KANU). GREENWOOD CO.: 8.
Stephens 9907 (KANU). JEFFERSON CO.: P. Н. Humfeld 133 (KANU).
JOHNSON cn: R. L. McGregor 3871 (KANU). KIOWA CO.: А. S.
Hitchcock, Aug. 1896 (KSC). LEAVENWORTH CO.: R. L. McGregor
2806 (KANU). LINCOLN CO.: P. H. Humfeld 965 (KANU). LYON CO.:
F. Agrelius. 10 April 1946 (KSTC). MARSHALL CO.: S. Stephens 4666
(KANU). MCPHERSON CO.: R. L. McGregor 13720 (KANU). MIAMI

486 Rhodora [Vol. 77

co.: Р. Н. Humfeld 160 (KANU). MONTGOMERY со.: A. J. Petrik-Ott
736 (KANU). NEOSHO СО.: W. W. Holland 872 (KANU). OSAGE CO.:
R. І. McGregor 3318 (KANU). OTTAWA CO.: A. J. Petrik-Ott 768
(KANU). RICE C0.: R. L. McGregor 2706 (KANU). RILEY со.: D. J.
Cashen, 24 Oct. 1920 (KSC). RUSSELL CO.: Р. L. McGregor 12676
(KANU). SALINE CO.: R. L. McGregor 2689 (KANU). SHAWNEE CO.:
P. H. Humfeld 145 (KANU). WABAUNSEE CO.: B. B. Smyth, 8 Aug.
1890 (UMO). WILSON CO.: B. L. Wagenknecht 2217 (KANU). WoOoD-
son C0.: R. L. McGregor 915 (KANU). WYANDOTTE CO.: S. Stephens
10860 (KANU).

Nebraska: BOX BUTTE CO.: M. P. Somer, Aug. 1909 (NEB). CASS
co.: H. J. Webber, 23 Мау 1887 (NEB). JEFFERSON CO.: А. F. Woods
and D. Saunders 2001 (NEB). NEMAHA CO.: J. M. Bates 6683 (NEB).

South Dakota: CUSTER C0.: S. Stephens 6158 (KANU). FALL RIVER
со.: S. Stephens 5796 (KANU). LAWRENCE CO.: C. A. Taylor 7464
(spc). MEADE on: E. J. Palmer 37019 (мо). PENNINGTON CO.:
S. Stephens 7138 (KANU).

Pellaea glabella Mett. ex Kuhn, Linnaea 36:87. 1869.

Kansas: ALLEN CO.: R. L. McGregor 3331 (KANU). BOURBON CO.:
R. L. McGregor 3840 (KANU). CHASE CO.: F. Agrelius, 9 Aug. 1958
(KSTC). CHAUTAUQUA CO.: P. H. Humfeld 846 (KANU). COWLEY CO.:
S. Stephens 3020 (KANU). DOUGLAS CO.: R. L. MeGregor 649 (KANU).
FRANKLIN £0.: R. L. McGregor 892 (KANU). GEARY CO.: P. H. Hum-
feld 967 (KANU). GREENWOOD CO.: P. H. Humfeld T99 (KANU). JOHN-
SON C0.: R. L. McGregor 3433 (KANU). LABETTE CO.: P. M. Maus,
26 May 1927 (KSC). LEAVENWORTH CO.: P. H. Humfeld 187 (KANU).
LINCOLN CO.: D. E. Lantz, 3 May 1898 (Kach, LINN CO.: P. H. Hum-
feld 40 (KANU). LYON CO.: F. Agrelius, 10 May 1945 (KSTC). MAR-
SHALL CO0.: W. H. Horr 4616 (KANU). MONTGOMERY CO.: R. L. Mc-
Gregor 10839 (KANU). OTTAWA CO.: А. J. Petrik-Ott 767 (KANU).
POTTAWATOMIF CO.: P. H. Humfeld 395 (KANU). RILEY со.: W. А.
Kellerman, 28 Мау 1887 (кѕс). всСотт co.: F. Agrelius, 16 Aug.
1912 (KSTC). WABAUNSEE CO.: R. L. McGregor 12371 (KANU). WIL-
SON co.: P. H. Humfeld 120 (KANU). woopsoN co.: W. H. ‘Horr,
10 July 1930 (KANU).

North Dakota: ROWMAN CO.: О. A. Stevens and D. R. Moir 2304
(NDA). DUNN CO.: О. А. Stevens, 8 June 1938 (NDA). GOLDEN VALLEY
c0.: S. Stephens 23465 and R. Brooks (KANU). GRANT CO.: W. B. Bell
1395 (NDA). HETTINGER CO.: Mrs. M. Meissner, 10 Aug. 1938 (NDA).
MCKENZIE CO.: О. A. Stevens and D. R. Moir, 14 Sept. 1956 (NDA).
MORTON со: О. A. Stevens, 17 June 1945 (NDA). OLIVER CO.: О. A.
Stevens, T Aug. 1938 (NDA).

South Dakota: CLAY co.: W. H. Over 1680 (SDU). CUSTER CO.:
P. A. Rydberg 1191 (NEB). HARDING CO.: Over and Solem 12758
(SDU). LAWRENCE CO.: S. Stephens 7518 (KANU). PENNINGTON CO.:
S. Stephens 7350 (KANU).

1975] Ferns — Petrik-Ott 487

FAMILY ASPIDIACEAE

Athyrium filix-femina (L.) Roth, Tent. Fl. Germ. 3:65.
1799.

Nebraska: ADAMS CO.: J. M. Bates, 12 July 1913 (NEB). BOYD CO.:
J. M. Bates 1094 (NEB). BROWN CO.: F. Clements 2938 (NEB).

North Dakota: CAVALIER CO0.: O. A. Stevens 2504 (NDA). PEMBINA
CO.: R. Brooks 449 (KANU). RICHLAND CO.: O. A. Stevens 149 (NDA).

South Dakota: CUSTER co.: A. J. Petrik-Ott 792 (KANU). LAW-
RENCE CO.: Е. J. Palmer 31532 (MO). MINNEHAHA CO.: C. A. Taylor
7651 (SDC). PENNINGTON CO.: S. Stephens 7175 (КАКО).

Athyrium руспосагроп (Spreng.) Tidestr. Elys. Marianum
1:36. 1906.

Kansas: LEAVENWORTH CO.: J. Wilson 3729 (KANU). WYANDOTTE
co.: К. K. Mackenzie 307 (KSC).

Cystopteris bulbifera (L.) Bernh. Schrad. Neu. Journ. Bot.
А O 806.
South Dakota: ROBERTS CO.: A. J. Petrik-Ott 800 (KANU).

Cystopteris fragilis (L.) Bernh. Schrad. Neu. Journ. Bot.
1(2) :26. pl. 2. fig. 9. 1806.

Kansas: CRAWFORD CO.: W. W. Holland 963 (KANU). ELLSWORTH
со: А. L. McGregor 17324 (KANU). JOHNSON CO.: M. A. Carleton,
25 Aug. 1892 (KSC). MONTGOMERY CO.: F. Agrelius, 28 May 1930
(KSTC). RICE CO.: R. Г. McGregor 2704 (KANU). RILEY Co: H. F.
Bergman, 9 Sept. 1910 (NDA). SALINE CO.: R. L. McGregor 2688
(KANU). WILSON CO.: H. Willis 16 (KSTC). WOODSON Co.: R. Brooks
973 (KSTC).

Nebraska: ANTELOPE CO.: N. F. Petersen 1908 (NEB). BROWN CO.:
C. E. Bessey, 23 July 1887 (NEB). CASS Co.: S. Stephens 18922
(KANU). CHERRY C0.: W. L. Tolstead, 25 June 1937 (NEB). CUMING
со.: Г. Bruner, July 1880 (NEB). CUSTER CO.: Н. J. Webber 6121
(NEB). DAKOTA CO.: S. Stephens 21144 (KANU). DAWES CO.: 25.
Stephens 24606 and R. Brooks (KANU). DIXON CO.: F. Clements
2556 (NEB). DOUGLAS con: W. Cleburne, 28 June 1873 (NEB). HOLT
co.: J. М. Bates, 21 Aug. 1895 (NEB). JOHNSON со.: No collector
given, 189? (NER). KEYA РАНА CO.: W. Kiener 23692 (MO). KNOX
co.: W. T. Barker 2652 (KANU). LANCASTER CO.: T. Walton, 20
May 1886 (NEB). RICHARDSON CO.: J. M. Winter, July 1931 (8р0).
SAUNDERS CO.: 7. A. Williams, 24 June 1890 (NEB). SEWARD СО.:
Н. J. Webber 6116 (NEB). SIOUX co.: A. J. Petrik-Ott 783 (KANU).
THOMAS CO.: P. А. Rydberg 1479 (NEB). WEBSTER CO.: J. M. Bates,
17 April 1905 (NEB).

488 Ећодога [Vol. 77

North Dakota: BARNES CO.: H. F. Bergman 2292 (NDA). BENSON
co.: R. Brooks 456 (KANU). BILLINGS CO.: R. Brooks 419 (KANU).
DUNN со.: R. Brooks 395 (KANU). GOLDEN VALLEY СО.: S. Stephens
23453 and R. Brooks (KANU). GRANT CO.: О. A. Stevens, 10 July
1964 (NDA). MCKENZIE CO.: R. L. McGregor and J. E. Bare 1153
(KANU). MCLEAN CO.: №. Brooks 398 (KANU). MERCER CO.: R.
Brooks 391 (KANU). MORTON со.: R. Brooks 389 (KANU). PIERCE
co.: О. A. Stevens, 16 July 1939 (NDA). RANSOM СО.: S. Stephens
33423 and R. Brooks (KANU). SIOUX CO.: A. J. Petrik-Ott 797
(KANU). STARK CO.: C. H. Waldron, 23 June 1912 (NDA). STUTSMAN
со.: C. C. Schmidt, 5 July 1897 (NDA). WARD co.: R. Brooks 400
(KANU). WILLIAMS CO.: O. A. Stevens, 8 Aug. 1915 (NDA).

South Dakota: BRULE CO.: R. Brooks 461 (KANU). CLAY CO.:
W. Н. Over 11128 (SDU). CUSTER CO.: A. J. Petrik-Ott 789 (KANU).
DEWEY C0.: S. Stephens 83798 and R. Brooks (KANU). FALL RIVER
co.: S. Stephens 5795 (KANU). GRANT CO.: W. Н. Over 7033 (SDU).
GREGORY CO.: S. Stephens and R. Brooks 875 (KANU). HAAKON CO.:
S. Stephens and Р. Brooks 379 (KANU). HARDING C0.: S. Stephens
1922 (KANU). LAWRENCE on: A. J. Petrik-Ott 795 (KANU). MAR-
SHALL CO.: О. A. Stevens and D. Р. Moir, 30 May 1957 (NDA).
MEADE C0.: Е. J. Palmer 37069 (KANU). MINNEHAHA C0.: L. J.
Harms 2689 (KANU). PENNINGTON CO.: S. Stephens 7804 (KANU).
ROBERTS CO.: S. Stephens 21471 (KANU). STANLEY CO.: S. Stephens
and R. Brooks 377 (KANU). торр CO.: L. Stanley, 24 May 1966
(STU). UNION С0.: T. Van Bruggen 4917 (SDU). WASHABAUGH CO.:
J. E. Bare 1227 (KANU). ZIEBACH CO.: R. Brooks 380 and S. Stephens
(KANU).

Cystopteris protrusa (Weath.) Blasdell, Mem. Torrey Bot.
Club 21 (4) :41, 42. pl. 3. 1963.

Kansas: ANDERSON co: R. L. McGregor 3458 (KANU). ATCHISON
co.: Р. І. McGregor 2817 (KANU). BROWN CO.: R. L. McGregor
2937 (KANU). CHAUTAUQUA CO.: P. H. Humfeld 868 (KANU). CHERO-
KEE CO.: S. Stephens 4136 (KANU). DONIPHAN CO.: А. J. Petrik-Ott
755 (KANU). DOUGLAS С0.: P. H. Humfeld 383 (KANU). ELK CO.:
S. Stephens 10768 (KANU). FRANKLIN on: R. L. McGregor 10041
(KANU). GREENWOOD CO.: R. L. McGregor 3419 (KANU). JEFFERSON
co.: P. H. Humfeld 201 (KANU). JOHNSON CO.: R. L. McGregor
3431 (KANU). LEAVENWORTH CO.: A. J. Petrik-Ott 750 (KANU).
LINN со.: R. L. McGregor 3465 (KANU). MIAMI CO.: R. L. McGregor
2788 (KANU). MONTGOMERY C0.: L. С. Hulbert 3678 (KANU). NEOSHO
co.: W. W. Holland 943 (KANU). OSAGE CO.: R. L. McGregor 3316
(KANU). SHAWNEE on: A. J. Petrik-Ott 726 (KANU). WYANDOTTE
co.: A. J. Petrik-Ott 748 (KANU).

19751 Ferns — Petrik-Ott 489

Nebraska: ADAMS cn: J. M. Bates, 12 July 1913 (NEB). CASS CO.:
A. J. Petrik-Ott 777 (KANU). DOUGLAS CO: Н. J. Webber 6118
(NEB). LANCASTER СО.: E. B. Robinson, May 1895 (NEB). NEMAHA
со.: S. Stephens 3626 (KANU). OTOE CO.: Н. J. Webber 6115 (NEB).
RICHARDSON со.: А. J. Petrik-Ott 775 (KANU). SARPY CO.: W. Cle-
burne, 22 May 1888 (NEB). THURSTON CO.: S. Stephens 21067
(KANU).

Cystopteris X tennesseensis Shaver, Journ. Tennessee
Acad. Sci. 25 (2) :107. 1950.

Kansas: ALLEN CO.: S. Stephens 10688 (KANU). ANDERSON CO.:
R. L. McGregor 3457 (KANU). BOURBON CO.: А. J. Petrik-Ott 761
(KANU). CHASE CO.: F. Agrelius, 9 Aug. 1953 (KSTC). CHAUTAUQUA
CO.: A. J. Petrik-Ott 744 (KANU). CHEROKEE CO.: EJ Parmer
20889 (KANU). CLAY CO.: Г. Avery, 29 Sept. 1895 (KSC). COFFEY CO.:
J. E. Taylor, B. L. Taylor and L. K. Magrath 3820 (KANU). COWLEY
co.: W. H. Horr 3152 (KANU). DOUGLAS CO.: A. J. Petrik-Ott 727
(KANU). ELLSWORTH CO.: R. L. McGregor 2696 (KANU). FRANKLIN
co.: R. L. McGregor 9095 (KANU). GREENWOOD CO.: R. L. McGregor
17216 (KANU). JEFFERSON CO0.: R. L. McGregor 3452 (KANU).
JOHNSON en: R. L. McGregor 3876 (KANU). LEAVENWORTH CO.:
S. Stephens 19444 (KANU). LINN CO.: A. J. Petrik-Ott 766 (KANU).
MARSHALL CO.: S. Stephens 4665 (KANU). MIAMI CO.: P. H. Hum-
teld 212 (KANU). MONTGOMERY CO.: А. J. Petrik-Ott 734 (KANU).
NEOSHO CO.: S. Stephens 18763 (KANU). ОБАСЕ CO.: R. L. McGregor
2320 (KANU). RILEY CO.: J. B. S. Norton, 25 Aug. 1892 (M0). RUS-
SELL со: W. Н. Horr 3266 (KANU). SHAWNEE СО. : B. B. Smyth,
May 1897 (KSC). WABAUNSEE CO.: Р. І. McGregor 12372 (KANU).
WILSON Co.: Р. L. McGregor 3404 (KANU). WOODSON CO.: А. d.
Petrik-Ott 199 (KANU). WYANDOTTE on: R. L. McGregor 2418
(KANU).

Dryopteris carthusiana (УШ) H. P. Fuchs, Bull. Soc.
France 105:339. 1958.

Nebraska: BROWN со.: F. Clements 2939 (NEB). CASS CO.: E.
Stoner, 14 May 1892 (NER). CHERRY CO.: W. L. Tolstead 653 (NEB).
LANCASTER CO.: H. J. Webber 6122 (NEB). THOMAS CO.: P. vA.
Rydberg 1484 (NER).

North Dakota: cass co.: C. Waldron, 5 July 1909 (NDA). PEM-
BINA со.: Р. Brooks 457 (KANU). RANSOM CO.: R. A. Shunk, Aug.
1916 (NDA). RICHLAND CO.: О. A. Stevens 1313 (NDA).

Dryopteris cristata (L.) A. Gray, Man. ed. 1. 631. 1848.

Nebraska: HOOKER CO.: No collector given, 12 July 1895 (spc).
THOMAS со.: Р. A. Rydberg 1530 (NEB).

490 Ећодога [Vol. 77

North Dakota: PEMBINA co: Р. Brooks 453 (KANU). RANSOM CO.:
R. A. Shunk, Aug. 1916 (NDA).

Dryopteris filix-mas (L.) Schott, Gen. Fil. t. 9. 1834,

South Dakota: CUSTER CO.: S. Stephens 5896 (KANU). HARDING
со.: S. S. Visher 7067 (SDU). LAWRENCE CO.: A. J. Petrik-Ott 794
(KANU). MEADE CO.: P. A. Rydberg 1197 (SDC). PENNINGTON CO.:
C. A. Taylor 8072 (spc).

Dryopteris marginalis (L.) A. Gray, Man. ed. 1. 632. 1848.

Kansas: ANDERSON СО.: E. Hartman 585 (KSTC). CHAUTAUQUA
co.: A. J. Petrik-Ott 740 (KANU). CHEROKEE CO.: R. L. McGregor
984 (KANU). COFFEY CO.: R. І. McGregor 908 (KANU). ELK CO.:
R. L. McGregor 927 (KANU). GREENWOOD CO.: R. Brooks 469 and
S. Stephens (KANU). LEAVENWORTH CO.: R. L. McGregor 2791
(KANU). MONTGOMERY co: R. L. McGregor 3394 (KANU). SALINE
co.: J. Hancin 1718 (Ksc). wILsoN co.: B. L. Wagenknecht 2252
(KANU). WOODSON co.: A. J. Petrik-Ott 733 (KANU).

Gymnocarpium dryopteris (L.) Newm. Phytologist 4: app.
XXIV. 1851.

South Dakota: CUSTER CO.: S. Stephens 5995 (KANU). LAWRENCE
со.: E. J. Palmer 37587 (UMO).

Phegopteris hexagonoptera (Michx.) Fee, Gen. Fil. 243.
1850-52.

Kansas: CHEROKEE co: R. L. McGregor 3843 (KANU).

Polystichum acrostichoides (Michx.) Schott, Gen. Fil. pl. 9.
1834.

Kansas: CHAUTAUQUA CO.: A. J. Petrik-Ott 741 (KANU). CHEROKEE
CO.: S. Stephens 8589 (KANU). FRANKLIN CO.: S. Stephens 2890
(KANU). GREENWOOD CO.: W. H. Horr, 10 June 1939 (KANU). MONT-
GOMERY CO.: R. L. McGregor 949 (KANU). WILSON cn: R. L. Mc-
Gregor 922 (KANU). WOODSON CO.: R. L. McGregor 916 (KANU).

Polystichum munitum (Kaulf.) Presl, Tent, Pterid. 83. 1836.
South Dakota: PENNINGTON CO.: S. Stephens and R. Brooks 427
(KANU).

Thelypteris palustris Schott, Gen. Fil. t. 10. 1834.

Kansas: CHAUTAUQUA CO.: R. L. McGregor 2414 (KANU). DONI-
PHAN CO.: G. L. Clothier and H. N. Whitford, 31 July 1897 (xsc).
DOUGLAS CO.: R. L. McGregor 15065 (KANU). ELK со.: R. L. Mc-
Gregor 928 (KANU). ELLSWORTH CO.: O. A. Kolstad 1963 (KANU).
GREENWOOD C0.: R. Brooks 468 and S. Stephens (KANU). MONT-

1975] Ferns — Petrik-Ott 491

GOMERY co.: №. L. McGregor 3397 (KANU). POTTAWATOMIE CO.:
W. A. Kellerman, 29 Sept. 1888 (KSC). RILEY C0.: W. A. Kellerman,
29 Sept. 1888 (MO). SALINE CO.: J. Hancin 1761 (KSC). WASHINGTON
co.: T. С. Dodd, Jr. 25 (KSC). WILSON co: Р. Г. McGregor 921
(KANU). WwooDSON co.: R. L. McGregor 906 (KANU).

Nebraska: BLAINE CO.: R. Brooks 792 (KSTC). BROWN CO.: C. Е.
Bessey, 23 July 1887 (NER). BUFFALO CO.: Misses Smith and Lee
7704 (NEB). CHERRY CO.: 5. Stephens 8201 (KANU). DODGE CO.:
C. E. Bessey, 9 Aug. 1872 (NEB). DOUGLAS CO.: W. Cleburne, Aug.
1851 (NEB). FRANKLIN CO.: E. M. Hussong 6777 (NEB). HALL CO.:
BR. J. Lemaire 2239 (NEB). HOLT Co.: J. M. Bates, 31 July 1899
(NEB). JEFFERSON СО.: A. F. Woods and D. Saunders 2004 (NEB).
KEARNEY CC.: Dr. Hapeman, July 1892 (NEB). LOUP CO.: S. Stephens
6852 (KANU). NEMAHA CO.: J. M. Bates, 5 July 1910 (NEB). SHERI-
DAN on: F. Sandoz 378 (NEB). THOMAS CO.: Н. J. Webber, 12 July
if89 (KSC). WHEELER on: R. L. McGregor 19370 (KANU).

North Dakota: RANSOM со.: R. A. Shunk, 15 Aug. 1916 (NDA).

South Dakota: BENNETT CO.: W. Н. Over 15884 (SDU). PENNING-
TON co.: S. S. Visker 1614 (SDU). Topp CO.: T. Van Bruggen 5108
(SDU).

Woodsia obtusa (Spreng.) Torr. Geo. Rep. New York Nat,
Hist. Sur. 4:195 Albany. 1840.

Kansas: ALLEN CO.: P. Н. Humfeld 326 (KANU). ANDERSON CO.:
P. Н. Humfeld 291 (KANU). ATCHISON CO.: Н. W. Blocker 809 (KSC).
BARBER CO.: R. L. McGregor 14792 (KANU). BARTON CO.: S. Stephens
8379 (KANU). BOURBON CO.: A. J. Petrik-Ott 760 (KANU). CHAU-
TAUQUA С0.: P. Н. Humfeld 866 (KANU). CHEROKEE Со. А. J.
Petrik-Ott 723 (KANU). CLAY on: C. Weber 318 (KSC). CLOUD CO.:
S. V. Fraser 557 (KSC). COFFEY CO.: G. L. Clothier and Н. №. Whit-
ford, 31 Aug. 1897 (KSC). COWLEY Co: Palmer 21256 (KANU).
CRAWFORD CO.: R. L. McGregor 916 (KANU). DOUGLAS CO.: R. L.
McGregor 15074 (KANU). ELK CO.: S. Stephens 10767 (KANU). ELLS-
WORTH. со.: R: Г. McGregor 17325 (KANU). FRANKLIN CO.: J. E.
Bare 257 (KANU). GREENWOOD CO.: S. Stephens 15841 and R. Brooks
(KANU). JEFFERSON CO.: R. L. McGregor 883 (KANU). JOHNSON
co.: R. L. McGregor 897 (KANU). LABETTE CO.: S. Stephens 11044
(KANU). LEAVENWORTH CO.: A. J. Petrik-Ott 751 (KANU). LINCOLN
cc.: P. H. Humfeld 423 (KANU). LINN CO.: A. J. Petrik-Ott 765
(KANU). LYON CO.: F. Agrelius, Sept. 1897 (KSTC). MARSHALL CO.:
S. Stephens 4664 (KANU). MCPHERSON CO.: P. H. Humfeld 439
(KANU). MIAMI CO.: Р. Н. Humfeld 222 (KANU). MONTGOMERY CO.:
W. H. Horr E466 (KANU). NEMAHA CO0.: R. L. McGregor 908
(KANU). NEOSHO CO.: S. Stephens 18763 (KANU). OSAGE CO.: R. L:
McGregor 888 (KANU). OTTAWA CO.: S. Stephens 10913 (KANU).

492 Rhodora [Vol. 77

POTTAWATOMIE CO.: P. Н. Humfeld 393 (KANU). RICE CO.: E. L.
Richards 1625 (KANU). RILEY С0.: P. Н. Humfeld 406 (KANU).
RUSSELL C0.: P. Н. Humfeld 954 (KANU). SALINE CO.: P. Н. Hum-
feld 408 (KANU). SHAWNEE co.: L. D. Volle 324A (KANU). STAF-
FORD C0.: A. Н. Curtiss, 3 June 1905 (кѕс). SUMNER Co: D. Birk-
holz 2785 (KANU). WABAUNSEE СО.: S. Stephens 4826 (KANU).
WASHINGTON CO.: S. Stephens 10893 (KANU). WILSON on: R. L.
McGregor 962 (KANU). WOODSON CO.: A. J. Petrik-Ott 719 (KANU).
WYANDOTTE co.: P. H. Humfeld 193 (KANU).

Nebraska: DAKOTA CO.: S. Stephens 21145 (KANU). DOUGLAS CO.:
K. L. Johnson 1606 and D. E. Dallas (KANU). GAGE CO.: E. A. Fiala,
7 Sept. 1925 (NEB). HOOKER со.: W. Kiener 24000 (MO). JEFFERSON
со.: A. F. Woods and D. Saunders 2002 (NEB). LANCASTER CO.:
A. S. Hunter 6774 (NEB). PAWNEE CO.: C. Н. Barnard, 1892 (NEB).
SAUNDERS CO.: W. Cleburne, 26 July 1890 (NEB). THURSTON CO.:
S. Stephens 21119 (KANU).

Woodsia oregana D.C. Eat. Can. Nat. and Geol. N.S. 2:89.
1865.

Nebraska: BROWN CO.: S. Stephens 24459 and R. Brooks (KANU).
CHERRY CO.: W. L. Tolstead, 25 June 1937 (NEB). CUSTER CO.: A. J.
Petrik-Ott 786 (KANU). DAWES CO.: S. Stephens 5671 (KANU).
GREELEY on: R. Brooks 463 and S. Stephens (KANU). ROCK CO.:
S. Stephens 6916 (KANU). SCOTTS BLUFF CO0.: W. L. Tolstead, 28 Aug.
1941 (NEB). SHERIDAN CO.: S. Stephens 6188 (KANU). SIOUX CO.:
R. Brooks 475 and S. Stephens (KANU). THOMAS CO.: P. A. Rydberg
1479 (NEB).

North Dakota: BILLINGS co.: R. Brooks 418 (KANU). GOLDEN
VALLEY on: S. Stephens 23439 and R. Brooks (KANU). MCKENZIE
со.: R. Brooks 416 (KANU). MCLEAN co.: J. Lunell, 18 June 1916
(NDA). MORTON CO.: Р. Brooks 390 (KANU). SIOUX CO.: R. Brooks
385 (KANU).

South Dakota: CORSON CO.: S. Stephens and R. Brooks 383 (KANU).
CUSTER CO.: P. A. Rydberg 1199 (NEB). FALL RIVER CO.: А. Nelson
681 (SDU). HARDING CO.: S. Stephens 7889 (KANU). LYMAN CO.:
Р. Brooks 376 and S. Stephens (KANU). MINNEHAHA CO.: L. J.
Harms 2693 (KANU). PENNINGTON C0.: Р. L. McGregor 18786
(KANU). STANLEY CO.: R. Brooks 378 and S. Stephens (KANU).
WASHABAUGH CO.: J. E. Bare 1227 (KANU).

Woodsia scopulina D.C. Eat. Can. Nat. and Geol. N.S, 2:91.
1865.

South Dakota: CUSTER CO.: A. J. Petrik-Ott 785 (KANU). LAW-
RENCE Co: E. J. Palmer 37556-A (UMO). PENNINGTON CO.: S.
Stephens 7309 (KANU).

1975] Ferns — Petrik-Ott 493

FAMILY ASPLENIACEAE

Asplenium platyneuron (L.) Oakes ex D.C. Eat. Ferns N.
Amer. 1:24. 1878.

Kansas: ATCHISON CO.: Н. W. Blocker (KSC). BOURBON CO.: R. L.
McGregor 974 (KANU). CHAUTAUQUA CO.: A. J. Petrik-Ott 742
(KANU). CHEROKEE CO.: A. J. Petrik-Ott T22 (KANU). COFFEY CO.:
R. L. McGregor 909 (KANU). CRAWFORD C0.: R. L. McGregor 988
(KANU). DOUGLAS CO.: R. L. MeGregor 660 (KANU). ELK co.: R. L.
McGregor 929 (KANU). FRANKLIN CO.: R. L. McGregor 899 (KANU).
GREENWOOD CO.: R. L. MeGregor 17208 (KANU). JOHNSON CO.:
R. L. MeGreaor 898 (KANU). LABETTE CO.: R. L. McGregor 958
(KANU). LEAVENWORTH CO.: A. J. Petrik-Ott 749 (KANU). LINCOLN
co.: P. H. Humfe!d 960 (KANU). MONTGOMERY C0.: Р. І. McGregor
3396 (KANU). NEOSHO CO.: R. L. MeGregor 956 (KANU). WILSON
cn: R. L. MeGreaor 985 (KANU). WOODSON CO.: A. J. Petrik-Ott
731 (KANU).

Asplenium resiliens Kunze, Linnaea 18:331. 1844.

Kansas: BOURBON CO.: R. L. MeGregor 986 (KANU). CHAUTAUQUA
co.: R. L. McGregor 954 (KANU). CHEROKEE СО. : R. L. McGregor
11090 (KANU). ELK со.: R. L. McGregor 938 (KANU). GREENWOOD
en: R. L. McGregor 936 (KANU). LABETTE CO.: R. L. McGregor
969 (KANU). MONTGOMERY C0.: W. H. Horr E416 (KANU). WILSON
co.: R. L. McGregor 987 (KANU).

Asplenium septentrionale (L.) Hoffm. Deutsch. Fl, 2:12.
1795.

South Dakota: CUSTER CO.: S. Stephens and R. Brooks 429 (KANU).
PENNINGTON CO.: R. Brooks 477 and S. Stephens (KANU).

Asplenium trichomanes L. Sp. Pl. 2:1080. 1753.

Kansas: CHAUTAUQUA CO.: A. J. Petrik-Ott ТАЗ (KANU). GREEN-
woop on: R. L. McGregor 8416 (KANU). WILSON CO0.: F. Agrelius,
28 Мау 1930 (KSC). WOODSON со.: А. J. Petrik-Ott 730 (KANU).

South Dakota: PENNINGTON CO.: P. A. Rydberg 1193 (NEB).

Asplenium viride Huds. Fl. Angl. 385. 1762.
South Dakota: LAWRENCE CO.: S. Stephens апа R. Brooks 426
(KANU).

Camptosorus rhizophyllus (L.) Link, Hort. Berol. 2:69.
1833.

Kansas: ALLEN CO.: R. L. McGregor 3336 (KANU). ANDERSON CO.:
Р. L. McGregor 3458 (KANU). ATCHISON CO.: S. Stephens 10846
(KANU). BOURBON CO.: A. J. Petrik-Ott 758 (KANU). CHAUTAUQUA

494 Rhodora [Vol. 77

co.: Р. Н. Humfeld 843 (KANU). CHEROKEE CO.: R. L. McGregor
11163 (KANU). COFFEY СО.: R. L. McGregor 920 (KANU). COWLEY
co.: W. Н. Horr and R. L. McGregor, 23 July 1947 (KANU). CRAW-
FORD CO.: No collector given, по date (KSC). DONIPHAN CO.: F.
Agrelius, Aug. 1913 (KSTC). DOUGLAS en: R. L. McGregor 29
(KANU). ELK on: Р. L. McGregor 939 (KANU). FRANKLIN CO.:
S. Stephens 17463 (KANU). GREENWOOD Co.: W. H. Horr, 10 June
1989 (KANU). JOHNSON со.: Р. L. McGregor 2346 (KANU). LA-
BETTE CO.: R. L. McGregor 959 (KANU). LEAVENWORTH CO.: W. M.
Keith and B. L. Wagenknecht 2348 (KANU). LINN CO.: A. J. Petrik-
Ott 764 (KANU). MIAMI со.: P. Н. Humfeld 158 (KANU). MONT-
GOMERY CO.: A. J. Petrik-Ott 737 (KANU). NEOSHO cn: R. L. Me-
Gregor 960 (KANU). RILEY C0.: G. L. Clothier, 15 Sept. 1888 (Ksc).
WILSON CO.: Р. L. McGregor 2370 (KANU). WOODSON co.: R. L.
МеСтедот 919 (KANU). WYANDOTTE CO.: M. Reed, 26 Sept. 1889
(KSC).

FAMILY BLECHNACEAE

Matteuccia struthiopteris (L.) Tod. Syn. Pl. Acot. Vasc.
Sicilia 30. 1866.

North Dakota: BOTTINEAU oun: D. R. Moir, L. D. Potter and O. A.
Stevens 2042 (NDA). CASS со.: О. A. Stevens, 8 July 1922 (NDA).
GRIGGS CO.: No collector given, 11 June 1953 (NDA). MORTON CO.:
O. A. Stevens, 14 Aug. 1954 (NDA). PEMBINA CO.: R. Brooks 450
(KANU). RANSOM CO.: О. A. Stevens, 10 Sept. 1964 (NDA). RICH-
LAND CO.: W. J. Wanek 255 (NDA). ROLETTE CO.: D. T. Disrud 98
(NDA).

South Dakota: CUSTER Co.: S. Stephens 6105 (KANU). PENNING-
TON CO.: S. Stephens 7117 (KANU). ROBERTS CO.: A. J. Petrik-Ott
799 (KANU).

Onoclea sensibilis L. Sp. Pl. 2:1062. 1753.

Kansas: ATCHISON CO.: Н. W. Blocker 514 (KSC). CHAUTAUQUA
co.: R. L. Мебтедот 15046 (KANU). DONIPHAN CO.: G. L. Clothier
and H. N. Whitford, 31 July 1897 (кѕс). DOUGLAS со.: R. L. Me-
Gregor 704 (KANU). GREENWOOD CO.: S. Stephens 9908 (KANU).
JEFFERSON C0.: R. L. McGregor 4152 (KANU). JOHNSON со.: J. A.
Gummerwan, June 1888 (KSC). LEAVENWORTH CO.: P. Н. Humfeld
270 (KANU). MONTGOMERY C0.: W. H. Horr and R. L. McGregor
E421 (KANU). OTTAWA CO.: C. T. Rogerson, 4 June 1954 (кѕс).
SALINE CO.: Р. Н. Humfeld 448 (KANU). woopsoN co.: D. L. Marsh
549 (KANU).

Nebraska: BROWN C0.: S. Stephens 24433 апа R. Brooks (KANU).
BUFFALO CO.: Misses Smith and Lee, 1894 (NEB). CHERRY co.: S.

19751 Ferns — Petrik-Ott 495

Stephens 8251 (KANU). FRANKLIN CO.: Е. M. Низзопу 6775 (NEB).
HOLT on: F. Clements 2802 (NEB). JEFFERSON CO.: №. A. Price
7270 (NEB). LOUP CO.: S. Stephens 6832 (KANU). OTOE CO.: C. Dunn,
March 1904 (NEB). SAUNDERS CO.: No collector given, 16 July
1889 (SDC). SHERIDAN CO.: F. Sandoz 366 (NEB). THOMAS CO.:
P. A. Rydberg 1317 (NEB).

North Dakota: PEMBINA C0.: О. A. Stevens 1594 (NDA). RICHLAND
CO.: О. A. Stevens 833 (NDA).

South Dakota: BENNETT C0.: W. Н. Over 15886 (SDU). CUSTER
co.: S. Stephens 6016 (KANU). LAWRENCE CO.: А. C. McIntosh 558
(SDU). MINNEHAHA CO.: W. H. Over 11102 (SDU). PENNINGTON СО. :
A. J. Petrik-Ott 793 (KANU). STANLEY CO0.: H. E. Lee B154 (SDU).

FAMILY DENNSTAEDTIACEAE

Pteridium aquilinum (L.) Kuhn in v. d. Decken, Reisen in
Ost-Afrika 3(3) :11. 1879.

Kansas: CHEROKEE CO.: Ё. L. MeGregor 3863 (KANU).

North Dakota: PEMBINA CO.: R. Brooks 454 (KANU).

South Dakota: CUSTER CO.: P. A. Rydberg 1192 (NEB). LAWRENCE
со.: S. Stephens 7717 (КАКО).

FAMILY POLYPODIACEAE

Polypodium polypodioides (L.) Watt var. michauxianum
Weath., in Contr. Gray Herb. 124:31. 1939.

Kansas: CHAUTAUQUA CO.: R. L. McGregor 3379 (KANU).

Polypodium vulgare L. Sp. Pl. 2:1085. 1753.

South Dakota: CUSTER CO.: A. J. Petrik-Ott 790 (KANU). MINNE-
HAHA en: L. J. Harms 2688 (KANU). PENNINGTON CO.: S. Stephens
7227 (KANU). SHANNON CO.: S. S. Visher, 6 July 1923 (SDU).

FAMILY MARSILEACEAE

Marsilea mucronata A. Br. Amer. Journ. Sci. 3:55. fig. 2.
1847.

Kansas: BARBER C0.: R. L. McGregor 10701 (KANU). BARTON CO.:
S. Stephens and R. Brooks 851 (KANU). BUTLER CO.: P. H. Humfeld
798 (KANU). CHASE CO.: R. L. McGregor 3937 (KANU). CLARK CO.:
R. L. McGregor 4024 (KANU). CLAY CO.: S. Stephens 29475 (KANU).
CLOUD CO.: S. V. Fraser 560 (KSC). COFFEY CO.: J. F. True, 22 July
1938 (KSC). COWLEY CO.: S. Stephens 6583 (KANU). EDWARDS C0.:

496 Rhodora [Vol. 77

W. А. Kellerman, 21 Aug. 1884 (Kach, ELLIS CO.: R. L. McGregor
3286 апа W. Н. Horr (KANU). FINNEY CO.: R. L. McGregor 3991
(KANU). FoRD co.: R. L. MeGregor 3972 (KANU). GOVE CO.: A. S.
Hitchcock, July 1895 (KSC). GRAHAM CO.: G. L. Clothier, Aug.-Sept.
1898 (Ksc). GRAY co.: R. L. McGregor 3270 and W. Н. Нотт
(KANU). HARPER CO.: R. L. MeGregor 13744 (KANU). HARVEY CO.:
Р. L. McGregor 3504 (KANU). HASKELL CO.: R. L. McGregor 4001
(KANU). HODGEMAN CO.: R. L. MeGregor 3962 (KANU). KINGMAN
со.: Р. L. McGregor 7318 (KANU). KIOWA CO.: R. L. McGregor
4951 (KANU). LANE op: R. L. McGregor 3278 and W. H. Horr
(KANU). LYON CO.: J. S. Wilson 9749 (KSTC). MEADE CO.: R. L.
McGregor 3256 and W. H. Horr (KANU). MORTON CO.: E. L. Richards
3571 (KANU). OSBORNE CO.: J. Richardson and K. Robertson 856
(KANU). PAWNEE CO.: R. L. McGregor 3951 (KANU). POTTAWATOMIE
co.: О. A. Stevens, 18 June 1909 (Ksc). PRATT CO.: W. T. Barker
1809 (KANU). RENO CO.: E. W. Lathrop 2201 (KANU). REPUBLIC CO.:
J. B. S. Norton, 23 Sept. 1895 (KSC). RICE со.: R. L. MeGregor 3943
(KANY). RUSH on: A. S. Hitchcock, Aug. 1895 (KSC). SALINE CO.:
F. H. Humfeld 871 (KANU). ѕсотт on: A. S. Hitchcock, Aug. 1895
(KSC). SEDGWICK со.: R. L. McGregor 4047 (KANU). SEWARD CO.:
R. L. Мебтедот 4010 (KANU). SHERIDAN CO.: C. Weber 299 (KSC).
SHERMAN C0.: А. S. Hitchcock, June 1892 (KsC). STAFFORD CO.: I.
Ungar 629 (KANU). STANTON со.: F. Lagergren, 20 Тау 1951 (Ksc).
STEVENS CO.: R. L. Мебтедот 17015 (KANU). TREGO CO.: J. L.
Hutchison, 14 Aug. 1951 (KSC). WALLACE CO.: А. S. Hitchcock,
Aug. 1895 (Ksc). WASHINGTON on: R. L. McGrenor 1000 (KANU).
WICHITA CO.: Agrelius, Hall, Lovejoy, 1913 (KANU).

Nebraska: ANTELOPE CO.: N. F. Petersen, June 1909 (NEB). BOX
BUTTE on: W. L. Tolstead, 28 Aug. 1941 (NEB). CHASE CO.: W. L.
Tolstead, 4 Aug. 1941 (NEB). CLAY CO.: W. Kiener 22538 (NEB).
CUSTER CO.: J. M. Bates 2246 (NEB). DAWES CO.: J. M. Bates, 29
July 1890 (NER). DEUEL co.: E. M. Gilliard, 3 July 1893 (NEB).
DUNDY cn: W. Kiener 10513 (NEB). FILLMORE CO.: W. L. Tolstead,
21 July 1941 (NEB). FRANKLIN CO.: W. L. Tolstead, 5 Sept. 1941
(NFE). GOSPER CO.: W. Kiener 19386 (NEB). HALL CO.: R. Brooks
465 and S. Stephens (KANU). HAMILTON CO.: W. Kiener 17847
(NEB). HARLAN CO.: W. L. Tolstead, 5 Sept. 1941 (NEB). HAYES
co.: W. L. Tolstead, T Aug. 1941 (NEB). JEFFERSON CO.: E. F.
Lange, 3 Sept. 1892 (NER). KEARNEY CO.: P. A. Rydberg 6604
(МЕР). LANCASTER CO.: J. M. Bates, 4 Aug. 1898 (NEB). MERRICK
co.: 1. Mueller, 14 July 1938 (NEB). PERKINS CO.: W. L. Tolstead,
7 Aug. 1941 (NEB). PHELPS CO.: R. J. Lemaire 1499 (NEB). PIERCE
co.: N. F. Petersen, 2 July 1907 (NEB). SHERIDAN CO.: W. L. Tol-
stead, 25 Auc. 1941 (NER). WEBSTER СО.: J. M. Bates, no date
(NER). WHEELER CO.: R. Brooks 791 (KSTC).

1975] Ferns — Petrik-Ott 497

North Dakota: BARNES CO.: О. A. Stevens 1223 (NDA). BENSON
co.: J. Lunell, 10 Sept. 1905 (NDA). BOTTINEAU CO.: R. Smith, 20
Sept. 1912 (NDA). BOWMAN C0.: O. A. Stevens, 4 July 1949 (NDA).
BURLEIGH CO.: S. Stephens 33268 and R. Brooks (KANU). FOSTER
со: О. A. Stevens and W. A. Kluender, 21 Aug. 1935 (NDA). GRANT
со: W. B. Bell 1355 (NDA). KEDDER CO.: О. A. Stevens and D. R.
Moir, 22 Aug. 1956 (NDA). MCLEAN CO.: O. A. Stevens, 27 Aug.
1915 (NDA). MOUNTRAIL CO.: О. A. Stevens, 20 Aug. 1915 (NDA).
RAMSEY CO.: C. A. Geyer 71 (MO). SLOPE CO.: A. С. Fox, 26 July
1938 (NDA). WALSH со.: А. C. Foz, July 1938 (NEB). WARD CO.:
L. F. Lantenschlager 847 (NDA).

South Dakota: BROOKINGS co.: T. A. Williams, Sept. 1893 (мо).
BROWN en: R. L. McGregor 18733 (KANU). BUTTE CO.: A. Nelson
549 (SDU). CHARLES MIX CO.: J. Martin, 22 July 1965 (spu). CLARK
co.: Carter, Sept. 1896 (spc). CUSTER CO.: А. Nelson, 2 Sept. 1926
(SDU). DAVISON CO.: Dillman, 25 Aug. 1907 (spc). HANSON CO.:
Р. Rrooks 435 (KANU). HARDING CO.: S. Stephens and R. Brooks
493 (KANU). HUGHES со.: J. Martin, 24 July 1965 (SDU). HYDE CO.:
R. Brooks 460 (KANU). PERKINS CO.: S. S. Visher 7084 (SDU). SAN-
BORN co. W. Н. Over, 3 June 1921 (spU). STANLEY CO.: We.
Over 7082 (5007). WASHABAUGH CO.: S. S. Visher 7083 (50107).

Marsilea quadrifolia L. Sp. Pl. 2:1099, 1753.
Kansas: CHEROKEE on: О. A Kolstad and L. J. Harms 1581
(KANU). NEOSHO со.: W. W. Holland 1993 (KANU).

Pilularia americana A. Br. Monatsber. Akad. Berl. 1863:
435. 1864.

Kansas: HARVEY CO.: R. Brooks 467 and S. Stephens (KANU).
RENO cn: R. L. McGregor 3988 (KANU).
Nebraska: CHERRY cn: Р. L. MeGregor 20017 (KANU).

FAMILY AZOLLACEAE

Azola mexicana Presl, Abh. Bohm. Ges. Wiss. V. 3:150.
1845.

Kansas: BARBER CO0.: R. L. McGregor 14705 (KANU). BARTON СО.:
R. І. McGregor 5198 (KANU). COFFEY CO.: L. K. Magrath 3291-1
(KsTC). DOUGLAS CO.: №. L. MeGregor 4687 (KANU). JEFFERSON CO.:
R. L. McGregor 4166 (KANU). LINN CO.: O. A. Kolstad and L. J.
Harms 2690 (KANU). LYON со.: E. E. Garner 1110 (KsTC). NEOSHO
co.: W. W. Holland 732 (KANU).

Nebraska: FILLMORE CO.: W. Kiener 22552 (NEB). GARDEN CO.:
W. Kiener 23131 (NEB). KEITH C0.: W. Kiener 23066 (NEB). SEWARD
со.: W. L. Tolstead, 18 Aug. 1941 (NEB). THOMAS CO.: A. J. Petrik-
Он 781 (KANU).

498 Ећодога [Уо]. 77
THE FERN ALLIES

FAMILY EQUISETACEAE
Equisetum arvense L. Sp. Pl. 2:1061. 1753.

Kansas: ATCHISON C0.: R. L. McGregor 2807 (KANU). BROWN CO.:
Garner, Summer 1925 (Ksc). CLAY со.: Weber 409 (KSC). CLOUD
co.: S. V. Fraser 561 (KSC). DONIPHAN CO.: A. J. Petrik-Ott 754
(KANU). DOUGLAS CO.: №. L. McGregor 4234 (KANU). JACKSON
со.: R. L. McGregor 2844 (KANU). JEFFERSON CO.: R. L. McGregor
4153 (KANU). JOHNSON C0.: R. L. McGregor 3430 (KANU). LEAVEN-
WORTH CO.: R. L. McGregor 3074 (KANU). MARSHALL CO.: Stephens
4701 (KANU). POoTTAWATOMIE CO.: P. H. Humfeld 969 (KANU). RILEY
co.: F. C. Gates 15756 (KSC). SALINE on: J. Hancin 1255 (Kach,
SHAWNEE Co: B. P. Smyth 77 (KSC). WABAUNSEE CO.: T. M.
Barkley, 1 May 1963 (KANU). WASHINGTON CO.: T. C. Dodd, Jr.
145 (KSC). woopsoN со.: E. W. Lathrop 2190 (KANU). WYANDOTTE
со.: P. Н. Humfeld 1021 (KANU).

Nebraska: BLAINE CO.: S. Stephens 24398 and Р. Brooks (KANU).
BOX BUTTE CO.: C. Н. Churchill, 5 July 1906 (NEB). воур со.: ЈУ. T.
Barker 3059 (KANU). BROWN со.: J. M. Bates, 25 June 1892 (NER).
BUFFALO CO.: S. Stephens 6767 (KANU). BURT CO.: S. Stephens 32067
and R. Brooke (КАМП). CASS con: S. Stephens 3768 (KANU). CEDAR
co.: R. Brooks 433 (KANU). CHERRY CO.: R. L. McGregor 19734
(KANU). CUMING Con: S. Stephens 36390 (KANU). DAWES CO.: S.
Stephens 5578 (KANU). DODGE CO.: L. Magrath 2615 (KSTC). DOUGLAS
co.: W. Cleburne, 23 May 1875 (NEB). GREELY CO.: R. L. McGregor
and J. E. Bare 430 (KANU). HALL CO.: R. J. Lemaire 2064 (NEB).
HOLT CO.: J. M. Bates, 22 June 1898 (NEB). HOOKER CO.: S. Stephens
17232 and R. Brooks (KANU). HOWARD on: J. E. Bare 390 and
R. L. McGregor (KANU). KEYA PAHA CO.: S. Stephens 6934 (KANU).
LANCASTER CO.: Pound, Clements, and Saunders, 2 May 1893 (NEB).
LINCOLN С0.: S. Stephens 15995 and R. Brooks (KANU). LOGAN
со.: S. Stephens 25068 and R. Brooks (KANU). NANCE CO.:
R. J. Lemaire 1971 (NFB). NEMAHA CO.: А. E. Watts, 5 May 1894
(SDU). OTOE CO.: Pound and Clements 4067 (NEB). RICHARDSON CO.:
S. Stephens 3563 (KANU). SARPY CO.: Pound and Saunders 4097
(NEB). SAUNDERS CO.: J. E. Weaver, May 1936 (NEB). SEWARD CO.:
H. J. Webber 6138 (NEB). SHERIDAN CO.: S. Stephens 6290 (KANU).
SHERMAN CO.: R. L. MeGregor 19801 (KANU). SIOUX CO.: S. Stephens
16364 апа R. Brooks (KANU). THOMAS CO.: P. A. Rydberg 1378
(NEB). THURSTON CO.: S. Stephens 21118 (KANU). WEBSTER CO.:
R. L. McGregor and J. E. Bare 792 (KANU).

North Dakota: BARNES co.: H. F. Bergman 373 (NDA). BENSON
со.: R. Brooks 457 (KANU). BILLINGS со.: О. A. Stevens, 21 June

1975] Ferns — Petrik-Ott 499

1961 (NDA). BoTTINEAU CO.: №. Brooks 402 (KANU). BURKE CO.:
R. Brooks 405 (KANU). cass co.: Н. Е. Bergman, 17 June 1909
(NDA). CAVALIER CO.: О. A. Stevens, 13 June 1956 (NDA). DIVIDE
co.: R. Brooks 407 (KANU). DUNN co.: R. Brooks 897 (KANU).
EDDY со.: О. A. Stevens and D. R. Moir, 10 Sept. 1956 (NDA).
GRAND FORKS CO.: V. Facey, 14 July 1960 (NDA). GRIGGS CO.: О. A.
Stevens, 30 July 1965 (NDA). LAMOURE CO.: J. F. Brenckle, 1905
(NDA). MCHENRY ou: О. A. Stevens, 16 June 1956 (NDA). MCKENZIE
co.: W. B. Bell 1038 (NDA). MOUNTRAIL co.: О. A. Stevens and
D. R. Moir, 13 Sept. 1956 (NDA). MORTON со.: О. A. Stevens, 25 July
1967 (NDA). PEMBINA CO.: R. Brooks 446 (KANU). RANSOM CO.:
W. B. Bell 290 (NDA). RICHLAND CO.: R. Brooks 445 (KANU). RO-
LETTE CO.: D. T. Disrud 1952 (NDA). SIOUX CO.: A. J. Petrik-Ott
795 (KANU). STARK C0.: О. A. Stevens, 17 Aug. 1954 (NDA). STEELE
CO.: О. A. Stevens, 7 July 1966 (NDA). WARD со.: L. F. Lauten-
schlager 425 (NDA). WILLIAMS CO.: R. Brooks 413 (KANU).

South Dakota: BROOKINGS on: D. Saunders, May 1897 (spc).
BUFFALO СО.: S. Stephens 32969 and R. Brooks (KANU). CLAY CO.:
W. H. Over 11012 (SDU). CODINGTON со.: J. R. Dugle 875 (spU).
CORSON CO.: S. Stephens and R. Brooks 381 (KANU). CUSTER CO.:
S. Stephens 6107 (KANU). DEUEL CO.: D. Saunders 8016 (am),
HAND CO.: J. E. Todd 16820 (SDU). HARDING CO.: S. Stephens and
R. Brooks 424 (KANU). LAWRENCE Di: S. Stephens 7578 (KANU).
LINCOLN co.: L. J. Harms 2604 (KANU). MARSHALL CO.: T. Van
Brugaen 446 (SDU). MINNEHAHA CO.: С. A. Taylor 9348 (spU).
MOODY CO.: C. A. Taylor and R. Lowell 10251 (spc). PENNINGTON
со.: S. Stephene 7216 (KANU). ROBERTS CO.: A. J. Petrik-Ott 798
(KANU). SANBORN CO.: S. S. Visher 4445 (SDU). UNION on: L. J.
Harms 2384 (KANU). YANKTON CO.: L. J. Harms 2832 (KANU).

Eauisetum X ferrissii Clute, Fern Bull. 12:22. 1904.

Kansas: ATCHISON co.: R. L. McGregor 2809 (KANU). BARTON
co.: Р. Г. McGregor 3944 (KANU): BROWN CO.: IF. Н. Horr, 10 July
1947 (KANU). CHAUTAUQUA CO.: B. Г. Wagenknecht- 1830 (KANU).
CHEROKEE CO.: E. L. Richards 3124 (KANU). CHEYENNE CO.: R. L.
McGregor 13582 (KANU). COWLEY CO.: W. T. Barker 3657 (KANU).
DONIPHAN CO.: B. L. Wagenknecht 3341 (KANU). DOUGLAS CO.:
E. B. Smith 412 (KANU). EDWARDS CO.: A. S. Hitchcock, 1 Sent.
1897 (KSC). ELLSWORTH CO.: R. L. McGregor 12898 (KANU). JACK-
SON co.: R. L. McGregor 2839 (KANU). JEFFERSON CO.: Hartman
1082 (KANU). LEAVENWORTH CO.: Hartman 1084 (KANU). LINCOLN
co.: P. H. Humfeld 957 (KANU). LINN CO.: R. L. McGregor 3467
(KANU). MEADE co.: R. L. McGregor 4016 (KANU). NEOSHO CO.:
W. W. Holland 2506 (KANU). POTTAWATOMIE CO.: S. Stephens 4767
(KANT). PRATT CO.: W. T. Barker 1574 (KANU). RENO CO.: R. L.

500 Ећодога [Vol. 77

McGregor 12476 (KANU). REPUBLIC СО.: D. К. Thomas, 1890 (KSC).
RILEY CO.: No collector given, 1 July 1889 (Mo). ROOKS CO.: Е.
Bartholomew, 25 June 1890 (NEB). SALINE CO.: P. Н. Humfeld 446
(KANU). SHAWNEE CO.: Hartman 1088 (KANU). TREGO СО.: M. Reed
and A. S. G., 8 July 1892 (KSC). WABAUNSEE CO.: S. L. Hunt 13
(anch, WASHINGTON CO.: W. H. Horr 4640 (KANU). WICHITA CO:
F. Agrelius, no date given (KsTC). WILSON со.: W. H. Haller,
16 June 1896 (Kach, woopsoN co.: S. Stephens 8383 (KANU).
WYANDOTTE CO.: Hartman 1090 (KANU).

Nebraska: BROWN C0.: S. Stephens 24439 and R. Brooks (KANU).
BUFFALO CO.: R. L. McGregor and J. E. Bare 1656 (KANU). CASS CO.:
S. Stephens 3771 (KANU). CEDAR CO.: S. Stephens 31978 and R.
Brooks (KANU). CHERRY CO.: S. Stephens 8167 (KANU). CUSTER
co.: J. M. Bates 2354 (NEB). DAKOTA CO.: S. Stephens 32054 and
Р. Brooks (KANU). DAWES CO.: Н. J. Webber 6134 (NEB). DIXON
co.: F. Clements 2543 (NEB). DODGE CO.: L. Magrath 2616 (KSTC).
DOUGLAS CO.: W. Cleburne, 8 July 1890 (NEB). DUNDY CO.: A. F.
Woods and D. Saunders 2000 (NER). HALL CO.: R. J. Lemaire 2456
(NEB). HOLT CO.: J. M. Bates, 2 Aug. 1892 (NEB). KEARNEY CO.:
P. A. Rydberg 475 (NEB). SARPY CO.: W. Cleburne, 10 July 1897
(NEB). SICUX CO.: W. L. Tolstead, 13 July 1940 (NEB). STANTON
CO.: R. Brooks 489 (KANU). THOMAS CO.: Н. J. Webber 6131 (NEB).
VALLEY СО.: S. Stephens 15620 and R. Brooks (KANU). WEBSTER СО.:
J. M. Bates, July 1903 (NEB).

North Dakota: BARNES co.: H. F. Bergman 248 (NDA). BENSON
co.: J. Luneli, 15 Aug. 1915 (NDA). CASS co.: О. A. Stevens 1083
(NDA). EMMONS CO.: R. Brooks 458 (KANU). GRIGGS CO.: О. A.
Stevens, 30 July 1965 (NDA). MCHENRY CO.: О. A. Stevens, 21 July
1966 (NDA). MCINTOSH CO.: R. Brooks 459 (KANU). MORTON CO.:
О. A. Stevens, 6 Aug. 1938 (NDA). PEMBINA CO.: R. Brooks 451
(KANU). RANSOM CO.: W. B. Bell, З July 1909 (NDA). RICHLAND
co.: R. Brooks 444 (KANU). WARD on: L. Е. Lautenschlager 658
(NDA). WILLIAMS CO.: О. A. Stevens, 14 Aug. 1941 (NDA).

South Dakota: JACKSON со.: W. Н. Over 7108 (SDU). MEADE CO.:
Snyder 307 (spc). PENNINGTON CO.: A. Nelson 514 (SDU). TURNER
co.: L. Messerli 532 (spu). vNION co.: L. J. Harms 2390 (KANU).
WASHABAUGH CO.: S. S. Visher 207099 (врт).

Enuisetum fluviatile L. Sn. Pl. 2:1062. 1758.

Nebraska: DouGLAs со.: E. Р. Walker, 25 May 1940 (NEB).
GARFIELD CO.: 27. M. Bates 4917 (NEB). HOLT со.: J. M. Bates,
27 June 1892 (NEB). SARPY CO.: W. L. Tolstead 41594 (NEB).

North Dakota: BENSON co.: J. Lunell, 30 June 1908 (NDA).
BOTTINEAU CO.: S. Stephens 28846 (KANU). MCHENRY CO.: J. Lu-
nell, 24 June 1908 (NDA). PEMBINA CO.: L. №. Waldron 1604 (NDA).
RICHLAND со.: О. A. Stevens, 11 July 1934 (NDA). ROLETTE CO.:

1975] Ferns — Petrik-Ott 501

D. T. Disrud and D. Disrud 347 (NDA). WARD CO.: L. F. Lauten-
schlager 783 (NDA).

South Dakota: GRANT CO.: D. Saunders and J. R. Towne, Aug.
1897 (spc).

Equisetum hyemale L. var. affine (Engelm.) A. A. Eat.
Fern Bull. 11:111. 1903.

Kansas: ALLEN C0.: B. Osborn 643R (KSC). ATCHISON CO.: R. L.
McGregor 2733 (KANU). CHEROKEE CO.: R. L. McGregor 2734
(KANU). CLAY CO.: C. Weber 316 (KSC). DONIPHAN CO.: S. Stephens
9349 (KANU). DOUGLAS on: P. H. Humfeld 153 (KANU). ELLSWORTH
со.: S. Stephens 17033 (KANU). FRANKLIN CO.: E. Hartman 644
(KANU). GREENWOOD CO.: R. L. McGregor 12339 (KANU). JACKSON
co.: R. L. McGregor 2843 (KANU). JEFFERSON CO.: P. H. Humfeld
138 (KANU). JOHNSON CO.: R. L. McGregor 3429 (KANU). LEAVEN-
WORTH co: P. H. Humfeld 196 (KANU). LINN CO.: S. Stephens
30999 (KANU). MARION CO.: M. H. Bartel, 14 Oct. 1960 (KSC).
MARSHALL Con: S. Stephens 3316 (KANU). REPUBLIC Con: G. Е.
Morley 742 (KANU). RILEY Co: F. C. Gates and M. Newcomb,
24 May 1928 (KANU). SUMNER СО.: G. L. Clothier and H. N. Whit-
ford, 23 Aug. 1897 (KSC). WASHINGTON CO.: W. H. Horr 4640
(KANU). WILSON on: H. Willis 7 (KSTC). WOODSON co.: E. W.
Lathrop 407 (KANU). WYANDOTTE CO.: A. J. Petrik-Ott 746 (KANU).

Nebraska: BUFFALO CO.: J. J. Thornbrer 54 (NEB). BURT CO.:
S. Stevens 21036 (KANU). CASS CO.: J. L. Morrison 935 (NEB).
CHERRY C0.: W. L. Tolstead, 24 Aug. 1941 (NEB). DAWES CO.: W. L.
Tolstead, 25 Aug. 1941 (NEB). DOUGLAS CO.: W. Cleburne, 8 Aug.
1902 (NEB). FRANKLIN C0.: J. M. Bates, 9 June 1891 (NEB). HARLAN
co.: Р. L. McGregor 19531 (KANU). NEMAHA CO.: S. Stephens 17706
(KANU). RICHARDSON CO.: A. J. Petrik-Ott 772 (KANU). SIOUX CO.:
A. J. Petrik-Ott 782 (KANU). THOMAS CO.: Р. A. Rydberg 1722
(NEB). THURSTON CO.: S. Stephens 21117 (KANU).

North Dakota: cass co.: O. A. Stevens 2834 (NDA). DICKEY CO.:
J. E. Bare 860 and R. L. McGregor (KANU). EDDY CO.: О. A. Stevens,
13 June 1960 (NDA). GRAND FORKS on: О. A. Stevens, 16 June 1953
(NDA). MCHENRY CO.: О. A. Stevens, 16 June 1956 (NDA). PEMBINA
co.: О. A. Stevens and D. R. Moir, 13 June 1958 (NDA). RANSOM
со.: S. Stephens 33487 and R. Brooks (KANU). RICHLAND CO.:
A. D. Stoesz, 2 June 1934 (NDA).

South Dakota: CUSTER CO.: S. Stephens 5993 (KANU). DEUEL CO.:
D. Saunders 8015 (SDC). HARDING CO.: S. S. Visher 7098 (SDU).
LAWRENCE CO.: S. Stephens 7424 (KANU). PENNINGTON CO.: 5.
Stephens 7217 (KANU). ROBERTS CO.: R. Brooks 440 and S. Stevens
(KANU). SHANNON co.: S. S. Visher, 27 June 1916 (spU). UNION
co.: R. Eslick, 8 July 1961 (SDU). YANKTON CO.: S. Stephens 21229
(KANU).

502 Ећодога [Vol. 77

Equisetum laevigatum A. Br. Amer. Journ. Sci. 46:87. 1844.

Kansas: ANDERSON CO.: B. Neill 285 (KSTC). BARBER CO.: W. Т.
Barker 799 (KANU). BARTON CO.: H. A. Stephens, 12 June 1958
(KSTC). BUTLER CO.: R. R. Weedon and L. K. Magrath 4204 (KANU).
CHEYENNE С0.: Н. A. Stephens, 14 June 1958 (KSTC). CLARK CO.:
R. L. McGregor 4025 (KANU). CLAY co.: Р. L. McGregor 2076
(KANU). COWLEY CO.: D. E. Dallas 578 (KANU). DECATUR CO.: W. H.
Horr 5046 (KANU). DICKINSON CO.: Hartman 1064 (KANU). DONI-
PEAN CO.: Hartman 1066 (KANU). DOUGLAS CO.: R. L. McGregor
3027 (KANU). ELLSWORTH CO.: R. L. McGregor 17107 (KANU).
FINNEY C0.: R. L. McGregor 4000 (KANU). GEARY CO.: Hartman
1072 (KANU). HAMILTON CO.: E. L. Richards 3012 (KANU). HARPER
co.: E. J. Palmer 21201 (UMO). HARVEY co: W. H. Horr and R. L.
McGregor 3724 (KANU). HODGEMAN CO.: R. L. McGregor 5188
(KANU). JEFFERSON CO.: G. Latham 880 (KANU). KINGMAN C0.:
k. L. McGregor 7306 (KANU). KIOWA CO.: R. L. McGregor and
W. H. Horr 3832 (KANU). LEAVENWORTH CO.: Hartman 1085 (KANU).
LOGAN CO.: S. Stephens 31297 and R. Brooks (KANU). MARION CO.:
P. H. Hwmfeld 790 (KANU). MARSHALL CO.: S. Stephens 3337
(KANU). MCPHERSON C0.: L. J. Harms 1054 (KANU). MEADE CO.:
W. Н. Horr and R. L. McGregor 4053 (KANU). MORTON CO.: E. L.
Richards 2499 (KANU). OTTAWA CO.: W. T. Barker 2545 (KANU).
POTTAWATOMIE CO.: Hartman 1086 (KANU). PRATT CO.: W. T.
Barker 1611 (KANU). RENO со.: E. Hartman 889 (KANU). REPUBLIC
co.: G. E. Morley 190 (KANU). RICE со.: R. L. McGregor 12563
(KANU). RILEY CO.: P. Zavos 29 (KSTC). SALINE CO.: Hartman
1974 (KANU). всотт CO.: R. L. McGregor 18007 (KANU). SEDGWICK
co.: M. A. Carleton, 27 May 1889 (KSC). SHAWNEE CO.: Hartman
1078 (KANU). SHERMAN CO.: О. A. Kolstad and L. J. Harms 1822
(KANU). STAFFORD CO.: I. Ungar 511 (KANU). SUMNER Co: J. W.
Johnson 75 (KSTC). TREGO CO.: S. Stephens 31211 and R. Brooks
(KANU). WALLACE СО.: R. L. McGregor 18579 (KANU). WICHITA
co.: Agrelius and Agrelius, 22 Aug. 1912 (KANU). WOODSON CO.:
W. H. Horr, 10 July 1930 (KANU).

Nebraska: BOYD co.: R. L. McGregor 19402 (KANU). BOX BUTTE
co.: C. H. Churchill, 5 July 1906 (NEB). BROWN co.: J. M. Bates,
June 1893 (NEB). BURT CO.: S. Stephens 82066 and R. Brooks
(KANU). CASS CO.: S. Stephens 32146 and R. Brooks (KANU). CEDAR
co.: R. Brooks 434 (KANU). CHASE oo: J. E. Bare 1403 and R. L.
McGregor (KANU). CHERRY CO.: S. Stephens 8225 (KANU). CUSTER
co.: A. J. Petrik-Ott T80 (KANU). DAKOTA CO.: S. Stephens 32035
and А. Brooks (KANU). DAWES co: S. Stephens 5565 (KANU).
PAWSON CO.: R. L. McGregor and J. E. Bare 1619 (KANU). DIXON
CO.: F. Clements 2542 (NEB). DODGE CO.: W. Kiener 29692 (NEB).
DOUGLAS CO.: W. Kiener 24571 (NEB). DUNDY CO.: S. Stephens and

1975] Ferns — Petrik-Ott 503

R. Brooks 358 (KANU). GARDEN CO.: S. Stephens and R. Brooks 359
(KANU). GRANT CO.: R. L. McGregor 19682 (KANU). HALL CO.:
R. J. Lemaire 1559 (NEB). HOOKER CO.: Р. A. Rydberg 1801 (NEB).
KEARNEY CC.: S. Stephens 6722 (KANU). KEITH CO.: S. Stephens
and R. Brooks 361 (KANU). KEYA РАНА CO.: S. Stephens 34319 and
R. Brooks (KANU). KNOX со.: S. Stephens and R. Brooks 373
(KANU). LANCASTER CO.: W. Kiener 29597 (NEB). LINCOLN CO.:
S. Stephens 15947 and R. Brooks (KANU). LOUP CO.: S. Stephens
6901 (KANU). MADISON со.: W. Kiener 29715 (MO). MCPHERSON
со.: S. Stephens 24986 and R. Brooks (KANU). NANCE CO.: B.
Osborn 1168R (мо). NEMAHA CO.: J. W. Gehling, 18 April 1935
(SDU). oTOE CO.: В. Osborn 815R (MO). PHELPS CO.: S. Stephens
24268 апа R. Brooks (KANU). RICHARDSON CO.: S. Stephens 3596
(KANU). SARPY CO.: W. Cleburne, 21 May 1879 (NEB). SAUNDERS
со.: T. B. Croat 2119 (KANU). SCOTT BLUFF CO.: S. Stephens and
R. Brooks 363 (KANU). SHERIDAN CO.: S. Stephens and R. Brooks
430 (KANU). SHERMAN со.: W. T. Barker 2889 (KANU). SIOUX CO.:
S. Stephens 16397 and R. Brooks (KANU). THOMAS со: R. L. Me-
(Gregor 19651 (KANU). WEBSTER on: R. L. McGregor 18642 (KANU).
WHEELER CO.: W. T. Barker 2955 (KANU).

North Dakota: BARNES co.: O. A. Stevens 1218 (NDA). BENSON
С2.: J. Lunell, 3 July 1910 (NDA). BILLINGS con: Р. Brooks 420
(KANU). BOTTINFAU CO.: O. A. Stevens, 15 July 1966 (NDA). BUR-
LEIGH CO.: S. Stephens 33250 and R. Brooks (KANU). CASS con: Lee
1205 (NDA). DICKEY co.: J. E. Bare 859 and R. L. McGregor
(KANU). DUNN со.: R. Brooks 396 (KANU). EDDY COo.: О. А.
tevens, 18 July 1966 (NDA). EMMONS CO.: О. A. Stevens 825 (NDA).
GOLDEN VALLEY CO.: О. A. Stevens, 11 July 1962 (NDA). GRAND FORKS
co.: J. D. Walp, 19 June 1936 (NDA). GREGORY со.: S. Stephens
24071 and R. Brooks (KANU). HETTINGER CO.: О. A. Stevens and
W. А. Kluender 137 (NDA). LAMOURE co: J. E. Bare 922 and R. L.
McGregor (KANU). MCHENRY CO.: J. E. Bare 1039 and R. L. Ме-
Gregor (КАМП). MCKENZIE C0.: О. A. Stevens 1625 (NDA). MCLEAN
со.: V. Rudd, 11 July 1958 (NDA). MERCER cn: R. Brooks 393
(KANU). MORTON CO.: S. Stephens 33614 and R. Brooks (KANU).
MOUNTRAIL CO.: J. E. Bare 1128 and №. L. McGregor (KANU). RAN-
SOM CO.: О. A. Stevens, 7 July 1959 (NDA). RICHLAND CO.: О. A.
Stevens 98 (NDA). SARGENT CO.: Н. L. Bulley, 10 June 1891 (NDA).
SHERIDAN CO.: 0. А. Stevens, 9 July 1959 (NDA). "SIOUX со.: Р.
Brooks 388 (KANU). SLOPE CO.: R. Brooks 422 (KANU). STARK CO.:
О. A. Stevens, 17 Aug. 1954 (NDA). STEELE CO.: О. A. Stevens, 7 July
1966 (NDA). STUTSMAN CO.: Н. Е. Bergman 89 (NDA). WARD CO.:
HR. Brooks 401 (KANU). WILLIAMS CO.: О. A. Stevens, 20 June 1945
(NDA).

504 Rhodora [Vol. 77

South Dakota: BEADLE CO.: C. A. Treadwell 104 (NEB). BROOKINGS
со: W. T. Barker 2697 (KANU). BUTTE CO.: W. Н. Over 17433
(SDU). CAMPBELL CO.: Г. A. Hanna 128 (Mo). CLAY co.: У. Н.
Over 7104 (SDU). CODINGTON co.: J. E. Dugle 109 (SDU). CORSON
со.: S. Stephens and Р. Brooks 382 (KANU). CUSTER CO.: S. Stephens
6064 (KANU). FALL RIVER CO.: S. Stephens 5790 (KANU). HAMLIN
co.: R. Brooks 486 and S. Stephens (KANU). HARDING CO.: S.
Stephens 7841 (KANU). HUTCHISON CO.: V. L. Harms 454 (KANU).
LAWRENCE CO.: S. Stephens 7401 (KANU). LINCOLN CO.: L. Mes-
serli 321 (SDU). МЕАРЕ CO.: S. Stephens 8097 (KANU). MINNEHAHA
co.: C. A. Taylor 9349 (spc). моору CO.: J. Н. Wilde and C. A.
Taylor 9287 (SDC). PENNINGTON CO.: S. Stephens 7118 (KANU).
PERKINS CO.: S. Stephens 8029 (KANU). ROBERTS CO.: R. Brooks 441
and S. Stephens (KANU). SULLY CO.: S. Stephens 33077 and К.
Brooks (KANU). TODD CO.: Г. Stanley 199 (sbU). TRIPP CO.: S.
Stephens 34119 and R. Brooks (KANU). UNION C0.: R. Eslick, 17
June 1962 (SDU). WASHABAUGH CO.: S. S. Visher 7101 (sbU). YANK-
TON on: L. J. Harms 2533 (KANU).

Equisetum palustre L. Sp. Pl. 2:1061. 1753.

North Dakota: RANSOM со.: R. Brooks 1554 and S. Stephens 33495
(KANU).

Equisetum pratense Ehrh. Hannov. Mag. 22:138. 1784.
North Dakota: PEMBINA C0.: №. Brooks 448 (KANU).

South Dakota: LAWRENCE CO.: S. Stephens 7544 (KANU). PEN-
NINGTON со.: R. Brooks 481 and S. Stephens (KANU). ROBERTS CO.:
R. Brooks 488 (KANU).

Equisetum scirpoides Michx. Fl. Bor. Amer. 2:281. 1803.

South Dakota: LAWRENCE CO.: G. N. Jones and F. F. Jones 14862
(MO).

Equisetum sylvaticum L. Sp. Pl. 2:1061. 1753.

North Dakota: CAVALIER C0.: О. A. Stevens and D. Е. Moir 1767
(NDA). PEMBINA CO.: R. Brooks 447 (KANU).

South Dakota: CUSTER CO.: A. J. Petrik-Ott 791 (KANU). LAW-
RENCE cn: Е. J. Palmer 37347 (UMO). PENNINGTON СО.: C. A.
Taylor 8112 (spc).

FAMILY ISOETACEAE

Isoetes butleri Engelm. Bot. Gaz. 3:1. 1878.
Kansas: CHEROKEE CO.: A. S. Hitchcock 1068 (KSC).

1975] Ferns — Petrik-Ott 505

Isoetes melanopoda Gay et Durieu, Bull. Soc. Bot. France
11:102. 1864.

Kansas: SALINE co.: J. С. Hancin 2447 (KANU). WILSON CO.:
R. L. McGregor 10267 (KANU).

Nebraska: CLAY CO.: W. L. Tolstead, b July 1941 (NEB). FILL-
MORE со.: ЈУ. L. Tolstead, 11 July 1941 (NEB). HAMILTON CO.:
W. Kiener 17851 (NEB). HARLAN CO.: W. Kiener 11850 (NEB).
KEARNEY cn: W. L. Tolstead, 23 July 1941 (NEB).

South Dakota: MELLETTE on: W. H. Over 15878 (500).

FAMILY LYCOPODIACEAE

Lycopodium obscurum L. Sp. Pl. 2:1102. 1753.
South Dakota: LAWRENCE CO.: P. A. Rydberg 1185 (NEB).

FAMILY SELAGINELLACEAE

Selaginella engelmanii Hieron. Hedwigia 39 (5) :294. 1900.

Nebraska: HOLT со.: W. T. Barker 2970 (KANU).

North Dakota: BILLINGS on: R. Brooks 417 (KANU). BOTTINEAU
co.: O. A. Stevens, 25 July 1962 (NDA). BURKE CO.: R. Brooks 406
(KANU). DUNN CO.: R. Brooks 394 (KANU). EDDY CO.: О. A. Stevens
2110 (NDA). GOLDEN VALLEY CO.: H. F. Bergman 1158 (NDA).
GRANT CO.: Bell 632 (NDA). MCHENRY Co: L. R. Haldron 2837
(NDA). MCKENZIE on: R. Brooks 415 (KANU). MOUNTRAIL CO.:
О. A. Stevens and D. R. Moir, 12 July 1960 (NDA). SIOUX CO.: О. A.
Stevens, 5 Aug. 1965 (NDA). SLOPE CO.: R. Brooks 421 (KANU).
STARK cn: H. F. Bergman, 21 June 1910 (NDA). WARD CO.: J. Lunell,
1 July 1909 (NDA). WILLIAMS co.: R. Brooks 412 (KANU).

South Dakota: CUSTER con: P. A. Rydberg 1184 (NEB). HARDING
co.: Over and Solem 11358 (SDU). PENNINGTON CO.: S. Stephens
7310 (KANU). PERKINS CO.: S. Stephens 8106 (KANU).

Selaginella rupestris (L.) Spring., Flora 21 (12) :149, 182.
1838.

Kansas: CHAUTAUQUA CO.: A. J. Petrik-Ott T39 (KANU). DOUGLAS
co.: R. L. McGregor 4663 (KANU). ELK CO.: R. L. McGregor 966
(KANU). FRANKLIN CO.: R. L. McGregor 1003 (KANU). GREENWOOD
co.: R. L. McGregor 3410 (KANU). JOHNSON CO.: R. L. McGregor
1004 (KANU). LEAVENWORTH on: R. L. McGregor 999 (KANU).
MONTGOMERY con: R. L. McGregor 996 (KANU). NEOSHO CO.: R. L.
McGregor 967 (KANU). WILSON CO.: R. L. McGregor 995 (KANU).
woopson co: A. J. Petrik-Ott 720 (KANU).

506 Ећодога [Vol. 77

Nebraska: ANTELOPE Co: N. Е. Petersen, T Aug. 1907 (NEB).
BLAINE CO.: S. Stephens 24376 and R. Brooks (KANU). BOX BUTTE
co.: C. Н. Churchill, about 1907 (NEB). BROWN co.: W. Kiener 29757
(NEB). CHERRY CO.: J. M. Bates, June 1891 (NEB). HOLT Co.: J. М.
Winter, 28 June 1931 (SDU). KEYA PAHA CO.: W. Kiener 23695
(MO). LOUP co.: S. Stephens 6858 (KANU).

North Dakota: PEMBINA co: R. Brooks 455 (KANU).

South Dakota: CODINGTON со.: W. H. Over 15474 (SDU). CUSTER
CO.: A. J. Petrik-Ott 788 (KANU). HANSON CO.: L. J. Harms 2759
(KANU). MINNEHAHA CO.: L. J. Harms 2659 (KANU). PENNINGTON
CO.: S. Stephens T104 (KANU). TURNER on: J. Н. Martin 594 (spU).

DOUBTFUL COLLECTIONS AND EXCLUDED TAXA

Adiantum pedatum: This species was reported by Bessey
(1892) for the Black Hills of South Dakota, but discounted
by himself in his 1898 and 1900 publications. Rydberg
(1917) again reports the presence of Adiantum pedatum
in the Black Hills, although he states that the section in his
manual concerning ferns was contributed by Miss Margaret
Slosson. Miss Slosson, at the time of writing the fern sec-
tion, was situated at the New York Botanical Garden. This
leaves the possibility that there is a specimen, at the New
York Botanical Garden, which I have not seen.

McIntosh (1949) reports the presence of Adiantum peda-
tum in his work on the Black Hills, but his information is
based upon Rydberg (1917), as is Winter, Winter, and
Van Bruggen’s (1959). Van Bruggen (1967), however,
states that there is a specimen of Adiantum pedatum that
was collected in Lawrence County, on the north slope of
Spearfish Mountain by F. L. Bennett in 1941. Brooks
(1969) states that this particular station has never been
relocated.

I have not seen the Bennett specimen or any other speci-
mens of Adiantum pedatum from the Black Hills and,
therefore, in my work, exclude it from this region.

Asplenium trichomanes: There is a specimen of Asplen-
ium trichomanes, located in the sp: herbarium and orig-
inally misidentified as Cystopteris sp., from Devil’s Lake,
Ramsey Co., North Dakota, collected by W. H. Over, 22

1975] Ferns — Petrik-Ott 507

June 1908. The herbarium label accompanying this speci-
men is quite confusing. The locality of collection written
on the label is Devil’s Lake, but the printed heading on the
label reads, “Flora of South Dakota, State Geol. and Biol.
Survey Collections, Plants of Clay Co., S.D.". There is no
place in South Dakota known as Devil’s Lake and therefore,
someone later wrote in ‘North Dakota’ on the label. Devil’s
Lake is located in Ramsey County, North Dakota, but this
record still remains questionable, due to the confusing label
information and the lack of a suitable habitat in the area
of Devil’s Lake to support this species.

Athyrium filix-femina: KANSAS: OSAGE on: C. Н.
Stokely, May 1895 (Kach This specimen was collected in
the town of Burlingame, Kansas. I would, however, be
hesitant to accept this as a valid record since there is no
suitable habitat near Burlingame where this fern might be
found. Furthermore, it is commonly found in cultivation
and C. H. Stokely is known to have collected many cultivars.

Cryptogramma acrostichoides: NEBRASKA: FRANKLIN
CO.: E. M. Hussong, 1893 (NEB). The collection locality
for this specimen is given as Franklin, Nebraska and no
habitat information is given. It consists of only a few
fertile segments. Cryptogramma acrostichoides is normally
found in rocky crevices, on rocky slopes or slides. Such
habitats are not to be found near the town of Franklin nor
in the county. The closest collection site for this fern is in
the Rocky Mountains of Colorado where it is common at
altitudes of 7,000 to 12,000 feet. It would be difficult to
accept this as a valid record for Franklin County, Nebraska.

Dryopteris goldiana: KANSAS: LEAVENWORTH CO.: J.
Wilson, 1871 (KANU). This specimen was collected in the
town of Leavenworth, Kansas. Mr. Wilson is known to
have cultivated ferns and several of his cultivated speci-
mens are on deposit in the KANU herbarium, This speci-
men does not bear the abbreviation ‘cult.’ on the label
although many of Wilson’s other specimens do. McGregor
and Hartman (1956) note: “It... seems as though the

508 Ећодога [Vol. 77

labels were prepared by someone other than Wilson, for
the same handwriting appears on other labels in the her-
barium. It seems certain that the individual writing the
labels merely omitted the ‘cult.’ inadvertently ..., and
thus introduced errors in our records.” Wilson gives no
habitat data or exact locality for his specimen. It is un-
likely that there is any natural area near Leavenworth or
anywhere in Kansas that would meet the habitat require-
ments of Dryopteris goldiana.

Equisetum variegatum: This species has been erroneously
reported many times as occurring in Kansas, Nebraska,
South and North Dakota. I saw several specimens identi-
fied to this species, but they were all depauperate forms of
Equisetum laevigatum. On the basis of the specimens I
observed, this species is probably best excluded from the
list of Equisetums for the four-state area.

Matteuccia struthiopteris: Cragin (1885) reports Mat-
teuccia struthiopteris from Lawrence, in Douglas County,
Kansas upon the word of Carruth. It is unlikely that it
grows any place in Kansas in nature. I have seen it doing
quite well in several gardens and greenhouses in Lawrence.

Ophioglossum vulgatum: Gates (1940) reports Ophio-
glossum vulgatum from Douglas County, Kansas upon the
basis of a specimen at the University of Kansas. The KANU
herbarium has no such specimen. Clausen (1938) cites a
specimen collected in Crawford County, Kansas on June
15, 1929 by F. A. Riedel and states that the specimen is on
deposit in the herbarium of the New York Botanical Gar-
den. Since I have not seen the specimen mentioned by
Clausen (1938), or any duplicates, to check its validity, I
am hesitant to include Ophioglossum vulgatum for Kansas
until further evidence is secured.

Osmunda claytoniana: This species has been erroneously
reported from Nebraska by Rydberg (1932).

Polystichum acrostichoides: NEBRASKA: No county
given, L. Bruner, no date given (NEB). No locality, other

1975] Ferns — Petrik-Ott 509

than Nebraska, or date of collection is given for this speci-
men. It is probably best considered as a doubtful record.

Thelypteris noveboracensis: KANSAS: POTTAWATOMIE
co.: G. Shimp, 30 April, no year given (KSC). This speci-
men was collected near the town of St. George, Kansas.
The habitat requirements of Thelypteris noveboracensis
are not likely to be met in Kansas. Furthermore, Miss
Shimp is known to have collected flowering plants from
near St. George that are garden plants and its seems possi-
ble that this specimen came from a similar habitat.

NEBRASKA : DOUGLAS C0.: W. Cleburne, 6 Oct. 1893 (NEB).
This specimen was collected in the city of Omaha, Nebraska.
In the NEB herbarium is placed, also, a specimen collected
on September 20, 1894, by W. Cleburne in a garden in
Omaha. It is likely that the 1893 specimen was also col-
lected in a garden and this information not mentioned on

the label.

ACKNOWLEDGEMENTS

This checklist has been made possible through the kind
cooperation of the curators at the following herbaria:
KANU. KSC, KSTC, MO, NEB, NDA, SDC, SDU and UMO, who
loaned me innumerable specimens for examination. The
accumulation of the data presented here has also been
greatly enhanced by financial assistance from the National
Science Foundation Graduate Traineeship Program, The
University of Kansas Committee on Systematics and Evo-
lutionary Biology (GB-446X, Dr. George W. Byers, Prin-
cipal Investigator) and the Sigma Xi Committee on Grants-
in-Aid of Research. Special thanks are due my husband,
Dr. Franklyn D. Ott, for his encouragement throughout the
writing of this paper, Mr. and Mrs. Leo J. Petrik, Dr.
Ronald L. McGregor, Mr. Ronald Weedon, Mr. Wayne
Robuck, Mr. Kermit Johnson, and Miss Mary Wiegand who
assisted me with field collecting, Mrs. Martha W. Rhodes
for assistance in proof-reading the manuscript, and Mrs.
Ann Tyska who typed the final manuscript.

510 Ећодога [Vol. 77

LITERATURE CITED

BERGMAN, Н. F. 1918. Flora of North Dakota. Bienn. Rep. North
Dakota Agric. Coll. Surv. 6: 151-372.

BEssEY, C. E. 1892. Ferns of the Black Hills. Amer. Nat. 26: 252-
253.

1898. The southern maidenhair fern in the Black Hills
of South Dakota. Bot. Gaz. 26: 211.
1900. One thousand miles for a fern. Asa Gray Bull.
8: 2-6.
BRooks, R. 1967. Ferns in Kansas. Kansas School Nat. 13: 1-15.
1969. Тће ferns of the Black Hills. Trans. Kansas
Acad. Sei. 72: 109-136.

CLAUSEN, R. T. 1938. A monograph of the Ophioglossaceae. Mem.
Torrey Bot. Club 19: 5-171.

CRAGIN, F. W. 1885. Notes on the ferns of Kansas. Bull. Wash-
burn Lab. Nat. Hist. 1: 58-60.

1886. Further notes on Kansas ferns. Bull. Wash-
burn Lab. Nat. Hist. 1: 175-176.

FERNALD, M. L. 1950. Gray’s Manual of Botany. 8th Ed. American
Book Со. New York. 1xiv + 1632 pp.

FITZPATRICK, T. J. 1919. The fern flora of Nebraska. Proc. Iowa
Acad. Sci. 26: 311-326.

—————,. 1920a. The fern flora of Nebraska — I. Amer. Fern
Jour. 10: 5-15.

1920b. The fern flora of Nebraska — II. Amer. Fern
Jour. 10: 33-44.

GATES, К. C. 1940. Annotated List of the Plants of Kansas: Ferns
and Flowering Plants (Flora of Kansas). Department of Botany,
Kansas State College Contribution No. 391. 266 pp.

HUMFELD, P. Н. 1951a. А checklist of Kansas pteridophytes. Amer.
Fern Jour. 41: 79-85.

19515. А checklist of Kansas pteridophytes. Amer.
Fern Jour. 41: 79-85.

KELLERMAN, W. A. & W. A. KELLERMAN. 1888. Analytical Flora
of Kansas. Published by the Authors, Manhattan, Kansas. 197
рр.

LANJouw, J. & Е. А. STAFLEU (Eds.). 1964. Index herbariorum.
Part I-— The Herbaria of the World. 5th Ed. Regnum Veg. 31.

MCGREGOR, К. L. 1960. Ferns and allies in Kansas. Amer. Fern
Jour. 50: 62-66.

‚ & E. L. HARTMAN. 1956. Notes of Kansas ferns.
Amer. Fern Jour. 46: 84-87.

McINTOSH, А. С. 1949. А botanical survey of the Black Hills of

South Dakota. Black Hills Eng. 28: 3-74.

1975] Ferns — Petrik-Ott 511

Омек, W. Н. 1932. Fiora of South Dakota. 'The University of
South Dakota, Vermillion, S. D. 161 pp.

PETERSEN, N. F. 1923. Flora of Nebraska. A List of the Ferns,
Conifers, and Flowering Plants of the State with Keys for their
Determination. 3rd Ed. Published by the Author, Plainview,
Nebraska. 220 pp.

REED, М. 1896. Ferns of Wyandotte County. Trans. Kansas Acad.
Sei. 14: 150-51.

RYDBERG, P. А. 1917. Flora of the Rocky Mountains and Adjacent
Plains. Published by the Author, New York. xii == Ор:

1932. Flora of the Prairies and Plains of Central
North America. New York Botanical Garden, New York. 969
pp.

Saunpers, D. А. 1899. Ferns and flowering plants of South Dakota.
South Dakota Agric. Exp. Sta. Bull. No. 64. pp. 101-227.

Stevens, O. A. 1950. Handbook of North Dakota Plants. North
Dakota Institute for Regional Studies, Fargo, М. D. 824 pp.

VAN BRUGGEN, T. 1967. The pteridophytes of South Dakota. Proc.
South Dakota Acad. Sci. 46: 126-144.

WILSON, J. 1885. Annotated list of ferns found in Kansas. Bull.
Washburn Lab. Nat. Hist. 1: 56-58.

Winter, J. M., C. WINTER, & T. VAN BRUGGEN. 1959. А Checklist
of the Vascular Plants of South Dakota. Department of Botany,
State University of South Dakota, Vermillion, South Dakota.

176 pp.

DEPARTMENT OF PLANT SCIENCES
TEXAS A & M UNIVERSITY
COLLEGE STATION, TEXAS 77848

SEASONAL OCCURRENCE AND ECOLOGY
OF MARINE ALGAE IN A
NEW HAMPSHIRE TIDAL RAPID'

NORMAN B. REYNOLDS? AND
ARTHUR C. MATHIESON

Water motion is a major factor determining the growth
and abundance of benthic plants in the ocean since it en-
hances their metabolism (Conover, 1968) and assists in
the propagation of plants and their dispersal to new areas.
Several workers (see Lewis, 1964) have discussed the
effects of wave action on the species composition and dis-
tribution of seaweeds. Exposed coastal sites typically have
more productive and diverse algal populations than shel-
tered locations, presumably because of differential water
motion. Sheltered embayments with strong tidal currents
are also rich, productive habitats (Lewis, 1968; Schwenke,
1971). To date, few detailed studies have been conducted
of tidal rapid communities except for the comprehensive
biological-hydrographical studies at Lough Ine, Ireland
(Kitching and Ebling, 1967). In the present paper we
describe the seasonal occurrence and ecology of seaweeds
at the Dover Point, New Hampshire tidal rapid.

The tidal rapid at Dover Point, New Hampshire, has
several unique features which distinguish it from other
tidal rapids. Foremost, it is located in the middle of the
Great Bay Estuary System, rather than near the open
coast (Fig. 1). Secondly, its substrate is stable even
though it is partially composed of small pebbles and cob-
bles. Thirdly, it has one of the fastest currents on the east
coast of North America (Anon., 1969).

1Published with the approval of the New Hampshire Agricultural
Experiment Station as Scientific Contribution Number 673.

?Present address: State University of New York, Cortland, N.Y.

512

1975] Marine Algae — Reynolds & Mathieson 513
3
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Point
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Mass. Уб

Wi 55" 6 70 45. |

Fig. 1. The New Hampshire Coast апа the Great Bay Estuary
System.

514 Ећодога [Vol. 77

MATERIALS AND METHODS

Monthly collections of seaweeds were made during low
tides from September, 1967, to September, 1969. Seasonal
collections of sublittoral plants were made (by SCUBA)
during 1968. Diving was only possible for about 20 min-
utes at dead low tide; it was further restricted by the
severe winter weather. No drift specimens are reported.
All specimens were processed immediately after collection.
Herbarium voucher specimens (a total of 1,264) were
deposited in the Algal Herbarium of the University of
New Hampshire (NHA). The specimens were identified
according to Taylor (1957), with the exception of Fucus
(Powell, 1957a, 1957b, 1963), Porphyra (Conway, 1964a,
1964b), and Laminaria (Wilce, 1965). The revised nomen-
clature of Parke and Dixon (1968) was applied whenever
possible. The longevity of the plants was designated ac-
cording to Feldman’s (1951) terminology.

General weather conditions (rainfall, air temperatures
and occurrence of ice) were recorded during field observa-

N
ж
P d
Dover Point d "
Piscataqua River e
Slow x
Current

"n,

Fast
Current

Little Bay > Ж

М

Fig. 2. Dover Point and surrounding areas.

200 Yards

Newington

1975] Marine Algae — Reynolds & Mathieson 515

tions. Biweekly records of surface water temperatures,
salinities and dissolved oxygen concentrations were moni-
tored at dead low tide in the high and low current areas at
Dover Point (Fig. 2). Salinities were recorded in the field
with a set of hydrometers (G. M. Mfg. Co., New York) ;
all of the values were corrected to 15°C. Temperatures
were recorded with a common immersible thermometer.
Oxygen concentrations were determined by use of a modi-
fied Winkler method (Hach Chem. Co., Ames, Iowa).
Diurnal variations of temperature, salinity, oxygen con-
centration and current speed were recorded on five separate
occasions. Salinities and temperatures were recorded at
multiple depths with an Electrodeless Induction Sali-
nometer (G. M. Mfg. Co., New York). The current speed
was recorded with a Little Captain boat speedometer
(Swift Instrument Company of Boston, Massachusetts),
which was modified with a six-foot well tube. The accuracy
of the instruments is + !4 knots.

DESCRIPTION OF AREA AND ENVIRONMENTAL FACTORS

Dover Point is located at latitude 47°07'05” N. and longi-
tude 7094950" W. in Dover, New Hampshire (Fig. 1).
It is 5 miles northwest of Portsmouth, New Hampshire,
and 5 miles east southeast of Durham, New Hampshire, at
the junction of the Piscataqua River and Little Bay. All of
the waters entering and leaving Little Bay, Great Bay and
their five tributaries pass through the constricted channel
at Dover Point, which is 470 yards wide and has, a maxi-
mum depth of 34 feet.

The rock outcroppings at Dover Point are of the Eliot
formation (Novotny, 1968). A variety of substrate types
are present; they grade from boulders to cobbles, pebbles,
sand and mud. The rocks are composed of mica schist with
garnet crystals, phyllite, pegmatite-quarts, feldspar and
metasiltstone. Smaller rocks and pebbles are stabilized by
extensive mussel populations, which allow them to support
relatively large plants. Most of the collections were ob-

[Vol. 77

Rhodora

516

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1975] Marine Algae — Reynolds & Mathieson 517

tained in the areas of high currents under the bridge
(Fig. 2). The locations to the northeast and southwest of
the bridge are mud flats which have reduced currents. The
substrate in the high current area grades from rock near
shore into mud at approximately 20 feet below mean low
water.

The tides at Dover Point are semi-diurnal, and they
occur 87 minutes later than those of the adjacent open
coast (Anon., 1965). The tidal amplitude is 6.8 feet. A
maximum tidal current of 5.5 knots occurs 2.5 to 5 hours
after low tide (Fig. 3). During ebb tide the cuzrent is
concentrated on the Newington side of the channel; it is
accompanied by a slight back eddy on the Dover Point side.
Organisms in the lower intertidal and subtidal zones are
exposed to strong currents at least 50% of the time (flood
tide), while organisms in the upper intertida] zone are
rarely exposed to strong currents. Subtidal organisms
beyond the second piling (Fig. 2) are exposed to a nearly
continuous current of high intensity.

The annual range of water temperatures was —2.0? to
19.0°C (Fig. 4). Diurnal temperature fluctuations at Dover
Point depend on the two water masses involved —i.e.,
Great Bay and the Atlantic Ocean. Little or no diurnal
variation (0-0.5°C) occurred during the winter. The larg-
est diurnal variation (5°C) occurred during the late spring
and summer. The annual range of salinity at Dover Point
was 7.5 о/оо at spring runoff to 31.0 0/oo in the late sum-
mer (Fig. 4). It usually varied from 23-29 o/oo. The
greatest diurnal fluctuation of salinity was recorded during
March, 1969 (i.e., 10 о/оо). In general there was a decrease
in salinity from low tide until one hour after the predicted
low. Thereafter it rose until high tide at which time it
decreased for two to three hours. The dissolved oxygen
concentration varied seasonally, with peak values occurring
in the spring (12-14 ppm) and minimal values in the late
summer (6-8 ppm). No obvious differences in temperature,
salinity and oxygen values were found between the high
and low current areas.

518

(996) 4jui|og

Rhodora

— ———

'AM JJ A SOND: Ј

|

FM

high current area

low

SONDJ

10 –
5
0
0
0

| 1
(о чу
СЧ e

(2) амәј (udd)ueDAxQ

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'F'NAMW JIJIAISIO 2

1969

1968

Fig. 4. Seasonal temperature, salinity and oxygen variations.

1975] Marine Algae — Reynolds & Mathieson 519

SPECIES COMPOSITION

The algal flora at Dover Point shows a high diversity of
species for an estuarine habitat. Eighty-three taxa of sea-
weeds were recorded from the tidal rapid site at Dover
Point, including 39 Rhodophyceae, 24 Chlorophyceae and
20 Phaeophyceae. Table I summarizes the number and
kind of seaweeds encountered at Dover Point as well as at
adjacent coastal and estuarine locations. Seventy-one of
the taxa found at Dover Point were also found on the ad-
jacent open coast (Mathieson & Fralick, 1972; Maithieson,
Hehre & Reynolds, in press). In contrast, only 59 taxa
from Dover Point were found in adjacent estuarine sites
with calm waters (Mathieson, Reynolds & Hehre, in press).
The red and brown algae at Dover Point showed more af-
finities to the open coast than the green algae (Table I).

SEASONAL OCCURRENCE

Tables II-IV summarize the monthly occurrence and
longevity of each species based on three years of monthly
collections at Dover Point. The maximum number of spe-
cies of Rhydophyceae were found in July (Table V). Peak
numbers of Chlorophyceae were evident in May, while the
Phaeophyceae showed peaks in April and May (Table V).
Table V also records the total number of seaweeds collected
per month; the largest number of species occurred during
May to July. The low numbers recorded in January and
September resulted from a lack of subtidal collections.

Several taxa were rare at Dover Point. Bryopsis plu-
тоза, Enteromorpha compressa, Monostroma leptodermum,
Ascophyllum nodosum forma scorpioides, Fucus distichus
spp. evanenscens, Myrionema strangulans, Sphacelaria cir-
rosa, Ceramium strictum, Dermatolithon pustulatum, Glo-
iostphonia capillaris, Melobesia lejolisii and Rhodochorton
purpureum were only found once during the entire three
year period. Of particular interest was the sporadic oc-
currence of Gloiosiphonia capillaris, for it was only seen
during a two week period and it was represented by 10
plants within a 10 ft? area. A comparison of Tables П-ТУ

520 Rhodora [Vol. 77

shows that six of the above species are annuals, which
might be expected to have a sporadic distribution. Other
seaweeds such as Callithamnion baileyi, Dasya pedicellata,
Lomentaria orcadensis, Polysiphonia harveyi, P. lanosa
and Laminaria longicruris were collected twice during the
36 month study. Monostroma pulchrum, Gracilaria foli-
ifera, Porphyra miniata and Chordaria flagelliformis were
collected three times. Laminaria longicruris, Gracilaria
foliifera and Polysiphonia lanosa are perennials, while all
the others are annuals (Tables III and IV).

Forty-two of the 83 taxa collected at Dover Point were
considered to be perennials (Table VI). Most of the green
algae were annuals (79%). A larger portion of the brown
(65%) and red algae (62%) were perennials. Two dis-
tinct types of annuals are present at Dover Point: seasonal
and aseasonal. Seasonal annuals occur during a certain
season and subsequently disappear. Monostroma grevillei,
Spongomorpha arcta, Bangia fuscopurpurea, and Urospora
penicilliformis are examples of late winter and/or spring
annuals. Chorda tomentosa, Bryopsis plumosa, Antitham-
nion cruciatum, Callithamnion baileyi, Ceramium strictum
and Dasya pedicellata are summer annuals. Aseasonal an-
nuals, such as Ectocarpus confervoides, Petalonic. fascia,
Scytospihon lomentaria, Enteromorpha intestinalis and
Ulothrix flacca occur all year long, although they may have
peaks in the spring and summer. Aseasonal annuals are
represented by several generations of plants.

Two major types of perennial algae are also evident at
Dover Point. Ascophyllum nodosum, Fucus vesiculosus,
Laminaria spp., Pseudendoclonium marinum, Ahnfeltia
plicata, Chondrus crispus, Gigartina stellata, Phyllophora
membranifolia and Rhodymenia palmata are typical of the
most common type where the whole plant is perennial.
Other species such as Elachista fucicola, Pilaiella littoralis,
Cladophora sericeae and Phycodrys rubens perenniate by
a portion of the thallus. Transitional forms are also evi-
dent between the two types. Knight and Parke (1931)
designate plants as pseudperennials if a small persistent
portion of the thallus can regenerate the intact plant.

1975] Marine Algae — Reynolds & Mathieson 521

DISCUSSION

The algal flora at Dover Point is “open coastal” in char-
acter for it has a very productive and diverse flora, as well
as a large number of species in common with the open
coast. It should be emphasized that adjacent, estuarine
areas, that lack currents but have similar substrate and
hydrographic conditions support a less diverse and reduced
vegetation. Lewis (1964) reports a similar biological char-
acterization of tidal rapids in Scotland and Ireland. In
addition he records the “dying out” of more delicate algae
in favor of more robust forms with increased water flow.

Kitching and Ebling (1967) state that the major in-
fluence of tidal currents is exerted on the sublittoral zone.
They also suggest that tidal currents are analogous to wave
action, for they prevent the deposition of sediment, reduce
local extremes of temperature and oxygen, and exert strong
mechanical pull. Moore (1966) states that tidal rapids
support open coastal invertebrates, even though the loca-
tions are essentially sheltered. The sublittoral zone at
Dover Point has more “open coastal” species than the
littoral zone.

Pronounced seasonal fluctuations of algal species were
recorded at Dover Point, with the largest number of spe-
cies occurring in July and the lowest in the winter. The
wide range of hydrographic conditions, particularly tem-
perature, probably causes the seasonal differences in its
flora. Williams (1948, 1949) and Coleman and Mathieson
(1975) have also recorded a wide range of annuals in areas
with pronounced temperature fluctuations. A combination
of boreal (e.g., Dumontia incrassata, Porphyra umbilicalis)
and warm temperate annuals (e.g., Callithamnion baileyi,
Dasya pedicellata) are present in the Great Bay Estuary
System, because of the wide range of temperatures ( Math-
ieson, Reynolds and Hehre, in press). The seasonal occur-
rence of organisms at Dover Point is very similar to that
at the adjacent open coast near Portsmouth, New Hamp-
shire (Mathieson, Hehre & Reynolds, in press), except that
a larger number of spring annuals is evident during April
on the open coast.

[Уо]. 77

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[Vol. 77

Rhodora

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ЯУЯОАНаОЯУНа AO ALIAHONO'I АМУ ЯОМЯНЧПООО TVNOSVUS TII ATaVL

Marine Algae — Reynolds & Mathieson 529

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[Vol. 77

Rhodora

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1975] Marine Algae — Reynolds & Mathieson 3531

TABLE V.

NUMBERS OF TAXA OF RHODOPHYCEAE, PHAEOPHYCEAE,
AND CHLOROPHYCEAE COLLECTED AT DOVER POINT
DURING VARIOUS MONTHS, 1967-1969

J ЕМ А MJ JA SO ND

Rhodophyceae 19 19 21 19 25 24 27 22 15 21 20 21
Phaeophyceae 10 13 18 16 15 .14.19^ HL. 1f 319 11 12
Chlorophyceae Sut. 19 18 15:19^139- 1L :12 .14 113512

Total 38* 43 46 48 55 50 53 44 38* 48 42 45

*No subtidal collections were made

TABLE VI.

NUMBERS OF PERENNIAL RHODOPHYCEAE,
PHAEOPHYCEAE AND CHLOROPHYCEAE AT DOVER POINT,

1967-1969
Number of taxa ` Total % of total % of

potentially Number taxa which perennial

perennial of taxa were perennial taxa/class
Rhodophyceae 24 39 29 62
Phaeophyceae 13 20 16 65
Chiorophyceae 5(7) 24 6(8) 21 (29)
Grand Total 42 (44) 83 61 (64) 49(52)

ACKNOWLEDGEMENTS

We would like to thank Dr. A. Hodgdon for his critical
review of the manuscript. In addition we express our grati-
tude to Drs. William Flahive and Richard Burns for as-
sistance in the collection of field data.

LITERATURE CITED

ANON. 1965. Tide tables, high and low water prediction, east
coast of North and South America, including Greenland, 1966.
U. S. Dept. of Commerce, Coast and Geodetic Survey. 289 pp.
Washington, D.C.
1969. Tidal current tables, Atlantic coast, North
America, 1969. U. S. Dept. of Commerce, Coast and Geodetic
Survey. 200 pp. Washington, D.C.

582 Rhodora [Vol. 77

COLEMAN, D. С., & А. С. MATHIESON. 1975. Investigations of New
England marine algae VII: Seasonal occurrence and reproduc-
tion of marine algae near Cape Cod, Massachusetts. Rhodora
77: 76-104. |

CONOVER, J. T. 1968. The importance of natural diffusion gradients
and transport of substances related to benthic marine plant
metabolism. Bot. Mar. 40: 1-9.

Conway, Е. 1964а. Autecological studies of the genus Porphyra:
I. The species found in Britain. Brit. Phycol. Bull. 2: 342-346.

1964b. Autecological studies of the genus Porphyra:
It. Porphyra umbilicalis (L.) Jour. Ag. Brit. Phycol. Bull. 2:
549-363.

FELDMANN, J. 1951. Ecology of marine algae. Рр. 313-334 i:
С. M. SMITH (ed.) Manual of Phycology. Ronald Press, New
York.

KITCHING, J. A., & Е. J. EBLING. 1967. Ecological studies at Lough
Ine. Adv. Ecol. 4: 197-291.

KNIGHT, M., & M. W. PARKE. 1931. Manx algae. Mem. Liverpool
Marine Biol. Comm. 30: 1-147.

Lewis, J. R. 1964. The ecology of rocky shores. xii 323 pp.
English Univ. Press Ltd., London.

1968. Water movements and their role in rocky shore
ecology. Ecology. Sarsia 34: 13-36.

MATHIESON, А. С., & К. A. FRALICK. 1972. Investigations of New
England marine algae V. The algal vegetation of the Hampton-
Seabrook Estuary and the adjacent open coast near Hampton,
New Hampshire. Rhodora 74: 406-435.

, E. HEHRE, & N. B. REYNOLDS. Investigations of New
England marine algae I. A floristic and descriptive ecological
study of the marine algae of Jaffrey Point, New Hampshire.
Nova Hedwigia, in press.

, N. REYNOLDS, & E. HEHRE. Га П. The species com-
position, distribution and zonation of seaweeds in the Great
Bay Estuary System and the adjacent open coast of New
Hampshire. Nova Hedwigia, in press.

МоокЕ, Н. В. 1966. Marine Ecology. xi + 493 pp. John Wiley &
Sons, New York.

Novotny, К. М. 1968. Geologic Map of the Seacoast Region, New
Hampshire Bedrock, N.H. Dept. Resources and Econ. Develop-
ment.

PARKE, M. & P. S. Drxon. 1968. Checklist of British marine
algae — second revision. Jour. Mar. Biol. Ass. U.K. 48: 783-832.

PowELL, H. T. 1957a. Studies in the genus Fucus L. I. Fucus
distichus L. emend. Powell. Jour. Mar. Biol. Ass. U.K. 36: 407-
432.

1975] Marine Algae — Reynolds & Mathieson 533

1957b. Studies in the genus Fucus L. II. Distribu-
tion and ecology of forms of Fucus distichus L. emend. Powell
in Britain and Ireland. Jour. Mar. Biol. Ass. U.K. 36: 668-798.

1963. Speciation in the genus Fucus L. and related
genera. Pp. 63-77 in: J. P. HARDING & N. TEBBLE, eds. Specia-
tion in the sea. System. Ass. Publ. No. 5.

SCHWENKE, Н. 1971. Water movements II. Plants. Pp. 1091-1121
in: О. KINNE, ed. Marine Ecology. Vol. I, Pt. 2.

TAYLOR, W. R. 1957. Marine Algae of the Northeast coast of
North America. viii + 509 pp. Univ. of Michigan Press, Ann
Arbor.

WiLCE, В. T. 1965. Studies in the genus Laminaria. ПІ. А re-
vision of the North Atlantie species of the Simplices Section of
Laminaria. Botanica Gothberg. 3: 247-256.

WILLIAMS, L. G. 1948. Seasonal alternation of marine floras at
Cape Lookout, North Carolina. Am. Jour. Bot. 35: 682-695.
1949. Marine algal ecology at Cape Lookout, North

Carolina. Bull. Furman Univ. 31: 1-21.

DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
AND JACKSON ESTUARINE LABORATORY
UNIVERSITY OF NEW HAMPSHIRE

DURHAM, NEW HAMPSHIRE 03824

BOOK REVIEW:
THE FLORA OF ESSEX COUNTY,
MASSACHUSETTS!

JAMES P. POOLE

In 1880 a Flora of Essex County, Massachusetts was
published by John Robinson who had been in charge of the
herbarium of the Peabody Museum in Salem, Massachusetts
for a number of years, eventually becoming Director of
the Museum. His flora consisted of a list of the species
then growing or reported to be growing in the County,
often with the names of the collectors but with only scant
information concerning the habit or habitat or the localities
where the species were found. The genera were listed
under families that were referred to as “orders” in the
summary, following the fifth edition of Gray’s Manual of
Botany. That publication, in turn, followed DeCandolle as
modified by Bentham and Hooker’s Genera Plantarum. In
addition to the vascular plants, listed as Exogens, the flora
2180 included mosses, liverworts, lichens, and algae; the
fungi were covered by only a brief reference.

In 1951 Stuart Harris, then Professor of Biology at Bos-
ton University, acceded to a request to write а modern
flora of the County for publication by the Peabody Museum.
From that date on he devoted what time he could spare
from his teaching duties and other varied interests to in-
tensive study in the field and in some of the herbaria in
which early collections from Essex County had been de-
posited. After many years he completed the Catalog of the
Flora and some of the accompanying sections, but unfortu-
nately did not live to complete a preface or write some of
the introductory sections he had planned. These portions
have been supplied by others. Dorothy Eastman Snyder,

iThe Flora of Essex County, Massachusetts by Stuart Kimball
Harris. Peabody Museum, Salem, Massachusetts. 1975. vii + 269 pp.
Map of the County on the inside cover. Sketch of Sweet Bay flower
on title page. $12.50

534

1975] Book Review 535

Consultant in Natural History at Peabody Museum, with
Stuart Harris, authored the preface and also served as
Editor. Other sections included: a Foreword by Richard
Jefferson Eaton, former Curator of the herbarium of the
New England Botanical Club, who also contributed a brief
biographical sketch of the author; one on Geology in Rela-
tion to the Flora by Sarah Fraser Robbins, Director of
Education at the Museum; another by Robert Lincoln
Goodale, M.D., entitled History of Botanical Collecting,
with biographical information about some of the collectors
of Robinson’s time and earlier; and a short section on
Climate by Hurd C. Willett, Professor Emeritus of the
Department of Meteorology, M.I.T. An End Map was
drawn by Charles F. Allen II, Assistant at the Museum.
The sections contributed by Dr. Harris included: a Catalog
of Species; a Statistical Summary of genera, species, vari-
eties, forms, and hybrids listed by families with each group
of taxa designated as Native or Introduced; a list of Spe-
cies Exclusae; and a list of collectors named in the Cata-
log with information as to the period and area of activity
of each of them. A list of references is also included.

In the Catalog the arrangement of the families and the
nomenclature follow Gray’s Manual, 8th edition, 1950, with
few exceptions, but the specific epithets are in lower case
as recommended in the International Code of Botanical
Nomenclature, 1972. Under the genera the species are
listed with both Latin and common names, with a brief
statement as to habitat and frequency, followed by the
names of towns or cities where collected, the names of the
collectors, with collectors’ numbers and date of collection
when available. For the varieties, forms and hybrids, only
the locality where collected, the collectors, collectors’ num-
bers, and dates are given, except that information as to
habitat and frequency is included for hybrid species. Pre-
ceding the Catalog is a list of towns and cities numbered
in the order quoted in the Catalog. These numbers are
spotted on the End Map.

586 Ећодога [Vol. 77

In the text there are а few misstatements апа typo-
graphic errors, and in the Catalog there are a number of
errors, mostly typographic. In the list of references and
also in the section listing the publications by the author
there are numerous mistakes, mostly in the year or the
volume number of the publication cited. In addition there
are some discrepancies which should be recorded. A few
of the species included in the Species Exclusae are also
listed in the Catalog. It seems probable that Dr. Harris
may have decided that some of the questionable species
should be included in the Catalog but failed to find the time
later to delete the names from the Species Exclusae. Again,
if the Statistical Summary is checked against the Catalog,
it will be discovered that there are instances in which the
two do not agree. Any corrections in the Statistical Sum-
mary must affect the Total Summary by Minor Groups on
page 23. Thus it is impossible to make any exact compari-
son in totals with Robinson’s Flora. In Harris’s Flora the
figure given for the Grand Total Taxa (1767) is the same
number as that given for the total species. It is evident
that it fails to include the totals for varieties, forms, and
hybrids.

These discrepancies and errors are difficult to explain
but, knowing the competence of Stuart Harris and his
scholarly standards, it seems most probable that the incon-
sistencies and most of the other errors would have been
remedied if he could have lived to make that final check
that is so essential before any manuscript goes to the
printer.

Fortunately, none of these difficulties or errors are of
enough significance to detract from the interest or value of
this publication. The catalog of species constitutes an
authoritative checklist of the flora of the County and, as
emphasized in the Foreword (p. vii), Essex County is note-
worthy from a phytogeographical point of view. It is an
area where Canadian zone species overlap traces of a
southern flora, where the northern Habenaria blephari-
glottis grows in the shelter of an indigenous colony of

1975] Book Review SA

Magnolia virginiana, Moosewood (Acer pensylvanicum)
occurs rather abundantly within a few rods of the sea-cliffs
at Manchester, and Hobble-bush (Viburnum alnifolium)
not far away in the same town, both of the last two species
characteristic of more northern upland woods.

Harris’s list serves very effectively to up-date Robinson’s
Flora by adding numerous species that were either not
present or had not been reported in 1880, and it gives much
additional information not included by Robinson. It fur-
nishes extensive coverage for an area that has lacked ade-
quate treatment in a single volume for nearly a century.
It also stands as a testimonial to the competence of Stuart
Harris as a taxonomist, and the sections contributed by
others serve as a fitting memorial to a highly respected
botanist whose life was all too short.

JESUP HERBARIUM
DEPARTMENT OF BIOLOGICAL SCIENCES
DARTMOUTH COLLEGE, HANOVER, N.H. 03755

CAREX GEYERI: REVISITED
PAUL E. ROTHROCK

In this journal over 35 years ago, R. T. Clausen and
Н. A. Wahl (1939) first reported the occurrence of Carex
geyeri Boott (a typically Rocky Mountain sedge ranging
from British Columbia and Alberta to Colorado, Utah, and
northern California) in the Eastern United States. This
site:

By limestone outcroppings, in dry deciduous
woods on bluffs and slopes on west side of Spring
Creek near west boundary of grounds of State
Penitentiary, Rock, north of Lemont, Centre Co.
[Pennsylvania ]

apparently still represents the only known locality for this
species east of Colorado.

With the assistance of Dr. Wahl, I was able to relocate
Carex geyeri in May, 1975. The area has remained undis-
turbed and, except for sapling growth, mostly unchanged.
Clumps of C. geyeri were observed in 3 distinct areas
along a quarter mile stretch of the bluff and have been
seen at various times in similar habitats slightly down-
stream (Wahl, personal communication). These clumps
showed a full range of vigor: fully half were setting fruit,
others had aborted fruit (perhaps due to lack of proper
pollination?), while several clumps were only persisting
vegetatively.

The bicentric distribution of this sedge may have arisen
either through introduction by man in recent years or by
some more natural means. Hermann (1970) favors the
first hypothesis, but Clausen and Wahl made definite note
of the natural undisturbed aspect of the habitat. Further-
more, it should be noted that the nearest major travel
routes, i.e., railroads, highways, or waterways, are at least
several miles away thereby making introduction by man
unlikely. Since other East-West disjuncts are known (see

538

1975] Carex Geyeri — Rothrock 539

Wood, 1970, for a full discussion) and considering the
relatively sizable area these plants cover, the localized, dis-
tinctive nature of the habitat, and the period of time over
which C. geyeri has been known for this site (first dis-
covered in 1932), I believe that the occurrence of this sedge
in central Pennsylvania represents a relict disjunct of
phytogeographical importance.

LITERATURE CITED

CLAUSEN, R. T., & Н. A. WAHL. 1959. Plants of central Pennsyl-
vania. Rhodora 41: 28-34.

HERMANN, Е. J. 1970. Manual of the Carices of the Rocky Moun-
tains and Colorado Basin. Agriculture Handbook No. 3874.
U. S. D. A. Washington, D. C. 397 pp.

Woop, C. E., Jr. 1970. Some floristic relationships between the
Southern Appalachians and Western North America. Pp. 331-404
in: P. С. Нот Ер., The distributional history of the biota of
the South Appalachians. Part II: Flora. Va. Polytech. Inst. &
State Univ., Blacksburg, Va.

202 BUCKHOUT LAB
PENNSYLVANIA STATE UNIVERSITY
UNIVERSITY PARK, PENNA. 16802

INTERNATIONAL REGISTER OF COMPUTER
PROJECTS IN SYSTEMATICS

SPONSORED BY
THE INTERNATIONAL ASSOCIATION FOR PLANT TAXONOMY
AND THE SOCIETY OF SYSTEMATIC ZOOLOGY

CALL FOR INFORMATION ON PROJECTS, PROGRAMS
AND DATA FILES

The above two international associations are the prime
sponsors of an International Register of Computer Projects
In Systematics. For the purpose of the Register, system-
aties includes taxonomy, biosystematics, evolution, and
biogeography of all biological taxa. The Register also
welcomes information about nonbiological data files of use
to systematics (e.g., the long range weather data tapes of
the U.S. Weather Bureau). For the present, our project
is a Register, which hopefully can direct people to the
source of information desired. Depending on demand, it
could be extended into a repository and clearing house for
computerized files of systematic value.

As in the first such Register (see Taxon 19:63-76[1970] )
we welcome systematic information on computerized data
files about living organisms, preserved organisms, experi-
mental data, literature files, etc. We also welcome informa-
tion on well-written and documented computer program
packages (other than basic statistics) that are of value
for systematic research and/or teaching.

If you or a colleague use computers in systematics (or
definitely plan to), please write to the Chairman of the
Register, and request as many copies of the Register
Questionnaire as you have separate projects or program
packages. You will be helping systematics in general by
avoiding duplication of effort and by contributing to our
attempts to minimize the incompatibility of computerized
systematic data, or programs generated on different proj-
ects. You will be helping yourself because not only might

540

1975] Computer Projects — Crovello 541

you discover that someone else has already written the
program, or computerized the data that you want, but also
the data and programs you have created may be useful to
others, thus enhancing their value.

The Register will be computerized and available for
customized search requests by September 1976. As demand
warrants it, published summaries will also appear. This
Register will be compatible with a similar Register for all
of biology that Crovello is organizing for the American
Institute of Biological Sciences.

Please address all suggestions, requests for information,
and for Register Questionnaires, to:

THEODORE J. CROVELLO, CHAIRMAN
INTERNATIONAL REGISTER
DEPARTMENT OF BIOLOGY
UNIVERSITY OF NOTRE DAME

NOTRE DAME, INDIANA 46556 U.S.A.

Volume 77, No. 812, including pages 441-552, was issued Dec. 31, 1975

100004

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by
ALFRED LINN BOGLE, Editor-in-Chief

ROLLA MILTON TRYON
STEPHEN ALAN SPONGBERG А
Associate Editors
GERALD JOSEPH GASTONY
RICHARD EDWIN WEAVER

VOLUME 77

1975

The New Lugland Botanical Club, Sue.

х

Botanical Museum, Oxford Street, Cambridge, Mass. 02188

INDEX TO VOLUME 77

Entries include: authors; title words such as geographic area; and
new records, systematic revisions, or ecological studies by taxa. Lists
or tables of taxa and maps are also classified spearately under “Lists”
and “Maps”. Note that only one page number per article is given
for each taxon entered. New scientific names and combinations are

printed in bold face type.

Alsophila Brooksii X Nephelea
portoricensis 451, A. dryopter-
oides X М. portoricensis 442,
А. hotteana X Nephelea sp.
452

Amorpha, revision of North
American 337, Key to Species
340, californica 345, californica
var, californica, var. napensis
348, apiculata 350, herbacea
352, key to varieties of herba-
cea 357, var. floridana 358, A.
crenulata 360, georgiana 362,
key to varieties of A. georgi-
ana, var. georgiana, var. con-
fusa 366, A. canescens 367, nana
373, glabra 377, paniculata 384,
nitens 386, laevigata 388, roe-
meriana 391, ouachitensis sp.
nov. 394, fruticosa 397

Antithamnion antillarum 26

Ascophyllum nodosum 16

Asterocytis ramosa 22

Ayensu, Edward S., Report on
Endangered and Threatened
Plant Species of the United
States 355

Barker, William Т.,
fread, C.)

Baskin, Jerry M. and Carol C.,
Geographical Distribution of
the Cedar Glade Endemic Viola
Egglestonii 427-429

Batrachospermum, Observations
in Southeastern Wisconsin
Streams 467

Boerhavia spicata 423

(see God-

ogle, A. Linn, Editor's Note to
Contributors 163-164
Butomus umbellatus 160

Cantino, Philip D., Boerhavia
spicata (Nyctaginaceae), The
North American Species, In

Northwestern Argentina, 4283-
426

Cape Cod Algae, map 77, Sea-
sonal occurrence and reproduc-
tion, figures 78, 79, tables 86-
96, Distribution, tables 86-102,
Longevity, table 97-99, Annuals
and Perennials, tables 100-105

Carex geyeri 537

Ceramium fastigiatum, 28, gra-
cillimum 27

Chimaphila maculata, in
Hampshire 436

Chondria 31

Chordaria flagelliformis 13

Chromosome Counts of Composi-
tae From The United States,
Mexico and Guatemala 171-195

Cladophora dalmatica, laetevirens,
6, socialis 7

Coleman, Douglas C. and Arthur
C. Mathieson, Investigations of
New England Marine Algae
VII: Seasonal Occurrence and
Reproduction of Marine Algae
Near Cape Cod, Massachusetts,
76-104

Colpomenia sinuosa 18

Compositae, Chromosome Counts
of figure 174-175, Table 182-
191

New

545

546 Rhodora

Computer projects in systematics,
International register of 539

Conant, David S., Hybrids in
American Cyatheaceae 441-455

Crotalaria sagittalis, Facultative
Dwarfism, Notes On the Le-
guminosae II. 141

Crouania attenuata 28

Crovello, "Theodore J., Inter-
national Register of Computer
Projects in Systematics 539-40

Crow, Garrett E., Book Review:
Winter Keys to Woody Plants
of Maine 438-439

Cyatheaceae, Hybrids in Ameri-
сап 441

Cyathea arborea 448, irregularis,
jamaicensis 453

Dasya rigidula 30

Davidse, Gerrit (See Spellman,
D.L.)

De Plantis Toxicariis E. Mundo
Novo ‘Tropicale Commenta-
tiones XII, Notes on Biody-
namic Piperaceous Plants 165-
170

De Wolf, Gordon P. Jr., The
Flora of Concord: A Review
334

Dictyota sp. 11

Diplochaete solitaria 4

Dwyer, John D. (See Spellman,
D.L.)

Ectocarpus elachistaeformis 8

Elachistea lubrica 12

Endangered and Threatened Spe-
cies of the United States, A
Report on 335

Enteromorpha sp. 4

Ericaceace, Bog, Apparent Eco-
typic Differences In The Water
Relations Of Some Northern 53

Erythrocladia recondita; subin-
tegra 23

[Vol. 77

Erythrotrichia carnea 23

Feldmannia irregularis 8

Ferns and Fern Allies, A. County
Checklist of in Kansas, Ne-
braska, South Dakota and
North Dakota 478

Fosliella farinosa, 25; lejolisii,
25, 26

Fralick, Richard A. (See Riggs,
S.A.)

Fucus vesiculosus 16

Geranium sibiricum 162

Giffordia conifera 9; G. littoralis
10; G. mitchelliae 9; G. san-
driana 10

Godfread, Carolyn, Butomaceae:
A New Family Record for
North Dakota, 160-161

Griffithsia radicans 29

Handlos, Wayne L., The Taxo-
nomy of Tripogandra (Comme-
linaceae) 213-333

Harriman, Neil A., Geranium
sibiricum L. (Geraniaceae) in
Wisconsin 162

Herposiphonia secunda, 32; te-
nella 32

Heterosiphonia wurdemanni 30

Hypoglossum tenuifolium 31

Index Holmensis Project, Appeal
for Support 336

Instructions to Contributors 551

Isthmoplea sphaerophora 15

Jania adherens 24; capillacea, 25

Keil, David J., Pectis humifusa
New to the Flora of the United
States 145-146

Keil, David J. and Tod F. Stuessy,
Chromosome Counts of Com-
positae from the United States,
Mexico, and Guatemala 171-
195

1975]

Kent Island, А Vegetational Sur-
vey of the Vascular Plants of
196

Kiger, Robert W., Papaver in
North America North of Mex-
ico 410-422

Kral, Robert, Rudbeckia auricu-
lata (Perdue) Kral, A Species
Distinct From R. fulgida Ait.
44-52

Lactuca muralis, in Maine 434;
in British Columbia 435

Leguminosae, Notes on the 141

Lichens, Maritime and Marine 147

LISTS (TABLES)

Batrachospermum, localities,
abbreviations and sampling
dates 468-469, range in envi-
ronmental conditions for 472,
range in selected environ-
mental conditions for Scup-
pernong Ck. and Тісһіғап
Ck. 472

Belize Monocotyledoneae 113-
139

Botanical Collectors in Belize

110-112
Cape Cod Marine Algae Dis-
tribution, Longevity, Domi-

nant Annuals and Perennials,
Seasonal Occurrence, Repro-

ductive Periodicities 86-103
Chromosome Counts of Com-
positae from the United
States, Mexico, and Guate-
mala 182-191
Frequency of Kent Island

Shrubs and Herbs 201
Herbaceous and Woody Species
Cover, Kent Island 207-208
Marine Algae from Nahant,
Massachusetts 152-155, 430-
483

Index to Volume 77 547

Marine Lichens from Nahant,
Massachusetts 147-148

Seaweed, composition at Dover
Point, New Hampshire 522-
525, monthly occurrence and

longevity of 527-529, num-
bers of taxa from Dover
Point 530

Soil analysis Data for Kent
Island Transects 204

Zostera marina, Characteristics
of eight stations within the
Great Bay Estuary 457

Lophosiphonia cristata, 33

Magnolias, The Deciduous of
West Florida 64, acuminata 70,
Ashei 66, pyramidata 71, tripe-
tala, 72

MAPS
Amorpha canescens, Distribu-
tion of 371, Amorpha spp.,

in Southeastern U.S. 395, in
Southwestern U.S. and Mex-
ico 349, fruticosa, in U.S.
404, glabra, schwerinii and
paniculata, in U.S. 380, nana,
in U.S. 376, nitens, laevigata,
roemeriana, ouachitensis in
U.S. 389

Belize, Administrative / Political
Districts 106

Kansas, Nebraska, South Da-
kota and North Dakota,
County maps 481

Marine Algae Stations on Cape
Cod 77

New Hampshire Coast and the
Great Bay Estuary System,
map 513, Dover Point and
surrounding areas 514

Vegetation Map, Kent Island,
New Brunswick 198

548 Rhodora

Viola egglestonii, geographical
distribution 428

Marchand, Peter J., Apparent
Ecotypic Differences іп the
Water Relations of Some
Northern Bog Ericaceae 53-63

Marine Algae of Nahant, Massa-
chusetts 149-158

Mathieson, Arthur C. (see Cole-
man, D.C.), (see Reynolds,
Norman B.)

McCain, John W., A Vegetational
Survey of the Vascular Plants
of the Kent Island Group,
Grand Manan, New Brunswick
196-209

Miller, Ronald F., The Deciduous
Magnolias of West Florida,
64-75

Monocotyledonae of Belize 105;
list of, 113-139

Monostroma pulchrum 5

Nahant Algae 149, recent addi-
tions to 430

Nephelea balanocarpa Х Nephe-
lea woodwardioides 452

Notes On the Leguminosae П.
Facultative Dwarfism In Cro-
talaria sagittalis 141-144

Padina sp. 12

Papaver, in North America north
of Mexico 410, Papaver sp. 411,
key to species 412, somniferum
413, rhoeas, dubium 414, cali-
fornicum, hybridum 415, orien-
tale 416, alpinum, alboroseum,
walpolei 417, mecconnellii 418,
lapponicum 419, nudicaule 420

Pectis humifusa, new record for
U.S. 145

Peperomia emarginella, glabella
var. melanostigma 165; serpens
166

[Vol. 77

Percursaria percursa 5

Petalonia fascia 14

Petrik-Ott, Aleta Jo, A County
Checklist of the Ferns and
Fern Allies of Kansas, Ne-
braska, South Dakota, and
North Dakota 478-511

Phragmites communis 159

Piperaceae (see Schultes, К. К.)

Piper Allenii, auritum, Bartlin-
gianum 166; dactylostigmum,
P. erythroxyloides sp. nov. 167;
hispidum, P. cf. interitum 169;
P. sp. 170

Plant Collecting іп Belize, а
Historieal Introduction 105

Polysiphonia delicatula, lanosa,
34; poko 35

Poole, James P., Book Review:
The Flora of Essex County,
Massachusetts 533-536

Poole, James P. Chimaphila
maculata (L.) Pursh in New
Hampshire 436-437

Poole, James P., Lactuca muralis
(L.) Gaertn. In Maine 434

Pothomorphe umbellata 170

Pseudotetraspora marina 3

Punctaria latifolia, plantaginea
14

Pylaiella littoralis 10

Reviews: Flora of Concord 334,
Flora of Essex County 533-
536, Winter Keys to Woody
Plants of Maine 438-439

Reynolds, Norman B. and Arthur
C. Mathieson, Seasonal Occur-
rence and Ecology of Marine
Algae in a New Hampshire
Tidal Rapid 512-532

Riggs, Stanley A. and Richard
A. Fralick, Zostera marina L.,
Its Growth and Distribution in
The Great Bay Estuary, New
Hampshire 456-466

1975]

Rothrock, Раш Е., Carex geyeri:
Revisited 537-538

Rudbeckia auriculata, a Species
Distinct From R. fulgida 44,
stat. nov. 49; R. fulgida var.
fulgida, R. fulgida var. um-
brosa, Plate 45, Habit sketches
46

Sargassum, Conspectus of Taxa
in the western Sargasso Sea
(key) 17, fluitans 18, hystrix
19, natans 19, “A”, 18, 20; “B”,
13.20, 16,01; "D^, 17, 21

Saxifraga Aizoon, S. cernua 41

Saxifrages on Mount Washing-
ton 41

Schultes, Richard Evans, Пе
Plantis Toxicariis E Mundo
Novo Tropicale Commenta-
tiones XII. Notes on Biody-
namic Piperaceous Plants 165-
170

Scytosiphon lomentaria 15

Shuey, Allen С. A Red-petioled
Form of Thalia geniculata
L. from Central Florida 210-
2152

Spellman, David L., John Р.
Dwyer and Gerrit Davidse., A
List of the Monocotyledoneae
of Belize Including a Historical
Introduction to Plant Collect-
ing in Belize 105-140

Spermothamnion investiens 29

Sphacelaria fucigera 11

~

Spongomorpha arcta 7

Stalter, Richard, Phragmites
communis in South Carolina
159

Statement of Ownership, inside
back cover, December, 1975
Steele, Frederic L., Saxifrages on

Mount Washington 41-43
Stuessy, Tod F. (see Keil, David
J.)

Index to Volume 77 549

Thalia geniculata, L. f. rheu-
moides Shuey, forma nova, A
Red-Petioled Form From Cen-
tral Florida 210-212

Tralau, Hans, Appeal for Sup-
port for the Index Holmensis
Project 336

Tripogandra (Commelinaceae)
the Taxonomy of 213-333;
Historical Review 214; Morph-
ology 218; Cytology, Breeding
Systems and Pollinators 231;
Measurements 284; Systematic
Account 235; Key to the Spe-
cles of 237; T. amplexans sp.
nov. 242, amplexicaulis 245, an-
gustifolia 249, brasiliensis, sp.
nov. 253, disegra 254, diuretica
comb. nov. 259, encolea 262,
glandulosa 265, grandiflora
268, guerrerensis 272, kru-
seana 275, montana 276, multi-
flora, neglecta sp. nov. 287,
palmeri 288, purpurascens 291,
purpurascens subsp. purpuras-
cens 293, purpurascens subsp.
australis 297, saxicola 298, ser-
rulata 301, warmingiana 311

Vegetational Survey of The Vas-
cular Plants of the Kent Island

Group, Grand Manan, New
Brunswick 196-209
Viola egglestonii, geographical

distribution of 427

Watson, Thomas Jr. (see Wither-
spoon, John T.)

Webber, E. E., Maritime and
Marine Lichens from Nahant
147-148

Webber, E. E., Phycological Stud-
ies from the Marine Science
Institute, Nahant, Massachu-
setts I: Introduction And Pre-
liminary Tabulation of Species
at Nahant 149-158

550

Webber, Е. E., Recent Additions
to the Marine Algal Flora of
Nahant, Massachusetts 430-433

Wilbur, Robert L., A Revision of
The North American Genus
Amorpha (Leguminosae-Psora-
leae) 337-409

Windler, Donald R., Notes on the
Leguminosae II: Facultative
Dwarfism in Crotalaria sagit-
talis L. 141-144

Witherspoon, John T. and
Thomas Watson Jr., Lactuca
muralis in British Columbia

485

Woelkerling, William J., Obser-
vations On Batrachospermum
(Rhodophyta) in Southeastern
Wisconsin Streams 467-477

Rhodora

[Vol. 77

Woelkerling, William J., On the
Epibiotic and Pelagic Chloro-
phyceae, Phaeophyceae, and
Rhodophyceae of the Western
Sargasso Sea 1-40

Wrangelia argus 30

Xylem Water Potential of Some
Bog Ericaceae Figure 57, Diur-
nal Trend of Water Potential,
Figure 59

Zostera marina, growth and dis-
tribution in New Hampshire
456, hydrographic factors 459,
seasonal growth 460, standing
crop 461, horizontal transplants
462

II ШШШ

1753 00343
1975] E

INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA

Manuscripts should be submitted in duplicate and should
be double-spaced or preferably triple-spaced (not on cor-
rasable bond), and a list of legends for figures and maps
provided on a separate page. Footnotes should be used
sparingly, as they are usually not necessary. Do not indi-
eate the style of tvpe through the use of capitals or under-
scoring, particularly in the citations of specimens, except
that the names of species and genera may be underlined to
indicate italies in discussions. Specimen citations should
be selected critically especially for common species of broad
distribution. Systematic revisions and similar papers should
be prepared in the format of “The Systematics and Ecology
of Poison-Ivy and the Poison-Oaks," W. T. Gillis, Rhodora
73: 161-237, 370-443. 1971, particularly with reference to
the indentation of keys and synonyms, Papers of a floristic
nature should follow, as far as possible, the format of
“Contribution to the Fungus Flora of Northeastern North
America. У.” Н. E. Bigelow & M. Е. Barr, Rhodora 71: 177-
203. 1969. For bibliographic citations, a recommended list
of standard journal abbreviations is given by L. Schwarten
& Н. У. Rickett, Bull. Torrey Bot. Club 85: 277-300. 1958.

RHODORA December, 1975 Vol. 77. No. 812
CONTENTS

Hybrids in American Cyatheaceae.
о Д е WEE E 441

Zostera marina L., Its Growth and Distribution in the Great Вау
Estuary, New Hampshire.
Stanley A. Riggs, Jr., and Richard А. Fralick .......................... 456

Observations on Batrachospermum (Rhodophyta) in Southeastern
Wisconsin Streams.
МОИ JL WOollbrlig E раны наанаа 467

A County Checklist of the Ferns and Fern Allies of Kansas,
Nebraska, South Dakota, and North Dakota.
Aleta Јо 21 udi: ——XÁ—Ó 478

Seasonal Occurrence and Ecology of Marine Algae in a New
Hampshire Tidal Rapid.

Norman B. Reynolds and Arthur C. Mathieson ........................ 512
Book Review: The Flora of Essex County, Massachusetts.

Juda P. Ромо LLLI ланын a 534
Carex geyeri: Revisited.

К ТТ: АЖ Rogo D m ТЬ: EE 538
International Register of Computer Projects in Systematics.

Tid J. Ороо арсена аанын 540
Так го Voluts TT, Steng нан нан 545
Instructions fo СОВО О qure cute aie inn ann es mn 551

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