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odora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
ALBION REED HODGDON, Editor-in-Chief 


^ 


ROLLA MILTON TRYON 
RADCLIFFE BARNES PIKE 
STEPHEN ALAN SPONGBERG ( Associate Editors 
GERALD JOSEPH GASTONY 
ALFRED LINN BOGLE 


Vol. 75 March, 1973 No. 801 
CONTENTS: 


Chemical, Cytological and Genetic Evidence for the Hybrid 
Origin of Aster blakei (Porter) House 


L. Michael Hill and O. M. Rogers .... sss 1 
Chromosome Numbers in Asters 
Paul Van Faasen and Fern Frank Sterk sss 26 


The Northern Limits of the Distributions of Hickories in 
New England 
Wayne E. Manning... 34 
Benthic Algae and Vascular Plants of the Lower Merrimack 
River and Adjacent Shoreline 
Arthur C. Mathieson and Richard A. Fralick .................... 52 
Some Morphological Aids in Distinguishing Nuphar micro- 
phyllum from Similar Aquatics 
Johonet C. Wicks 
(Continued on Inside Back Cover) 


The Nef England Botanical Club, Jne. 
Botanical Museum. Oxford St.. Cambridge. Mass. 02138 


RHODORA. — A quarterly journal of botany, devoted primarily to the 
flora of North America and floristically related areas. Price $10.00 
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 
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Scientific papers and notes, relating directly or indirectly to the 
plants of North America, will be considered by the editorial com- 
mittee for publication. Articles concerned with systematic botany 
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able. All manuscripts should be double-spaced throughout. Please 
conform to the style of recent issues of the journal. Illustrations 
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of publication. Extracted reprints, if ordered in advance, will be 
furnished at cost. 

Address manuscripts and proofs to Albion R. Hodgdon, 

Dept. of Botany, Nesmith Hall, 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 

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the next number of RHODORA, changes of address must be received 

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Second Class Postage Paid at Boston, Mass. 


MANUFACTURED BY 
THE LEXINGTON PRESS, INC. 
LEXINGTON, MASSACHUSETTS 


Rhodora 


JOURNAL OF THE 
NEW ENGLAND BOTANICAL CLUB 


Vol. 75 March, 1973 - No. 801 


CHEMICAL, CYTOLOGICAL AND 
GENETIC EVIDENCE 
FOR THE HYBRID ORIGIN OF 
ASTER BLAKEI (PORTER) HOUSE! 


L. MICHAEL HILL AND O. M. ROGERS 


The genus Aster is large and polymorphic, and many au- 
thors have cited the taxonomic complexity of this group 
(Anderson, 1929; Shinners, 1941; Rosendahl & Cronquist, 
1949). Fernald (1950) suggested that some of the varia- 
tion causing this complexity was due to hybridization, and 
this was anticipated by Wetmore & Delisle (1939), and 
verified by Avers (1953a), and Uttal (1962). The work of 
Avers on asters of the Heterophylli series demonstrated 
that many species could be crossed in cultivation, produc- 
ing hybrids (Avers, 1953a). However, she noted the ab- 
sence of such crossing in natural populations. She observed 
that the barriers to hybridization in nature were primarily 
ecological, relating to the absence of hybrid habitats (Av- 
ers, 1953b). 

A recent study by Pike (1970) presented morphological 
and geographical evidence indicating that Aster Blakei 
(Porter) House was of hybrid origin from A. acuminatus 
Michx, and A. nemoralis Ait. The morphological data fur- 


"Published with the approval of the Director of the New Hampshire 
Agricultural Experiment Station as Scientific Contribution No. 442. 

The work represents research done by the senior author in partial 
fulfillment of requirements for the Ph.D. degree. 


1 


2 Rhodora [Vol. 75 


ther indicated that introgression might be occurring in the 
direction of A. nemoralis. 

This paper considers further the origin of A. Blakei, uti- 
lizing cytological, genetic and chemical techniques. The 
chemical studies were conducted using thin-layer chroma- 
tography of phenolie compounds. These compounds were 
used as markers in detecting natural hybridization. The 
genetic techniques involved crosses and backcrosses of the 
putative parents, A. acuminatus and A. nemoralis. Self- 
pollinations also were made to synthesize an F. population. 
The cytological approach involved studies of mitotic chrom- 
osomes of the parental taxa and the F, hybrid, including 
specimens of A. Blakei cultivated in the greenhouse. Pollen 
stainability of the parents, F, hybrids, backcrosses and F. 
hybrids also were observed. 

The application of analytical chemistry to taxonomic 
studies has been increasingly successful as techniques have 
become more refined. The work of Turner & Alston (1959) 
and Alston & Turner (1962) on hybridization in Baptisia 
started the recent emphasis in chemotaxonomy. The species 
specific compounds of two putative parents in this genus 
both appeared in suspected hybrids. This has become a 
principle of phenolic biochemical systematics: the phenolic 
compounds found in hybrids represent a summation of the 
species-specific phenolic compounds found in the parents 
(Alston, 1965 and Harborne, 1968). Utilizing this princi- 
ple, other workers have confirmed hybridization in various 
genera. For examples, see Smith & Levin (1963) in Asple- 
nium, Jaworska & Nybom (1967) in Saxifraga, Olden & 
Nybom (1968) in Prunus, Fahselt & Owenby (1968) in 
Dicentra, and Walker (1969) in Petalostemon. The tech- 
nique has been useful in demonstrating introgression in 
Iris (Carter & Brehm, 1969) and in confirming an instance 
of intergeneric hybridization (Crang & Dean, 1971). 

Very few chemosystematic studies in Aster have been 
published to date. A preliminary survey of phenolics by 
Turner & Mabry (1964) on various members of the Aster- 
aceae cited only one species of Aster. The research of Abra- 


1973] Aster Blakei — Hill and Rogers 3 


hamson & Solbrig (1970) revealed no species-specific pheno- 
lies in the Heterophylli series and the authors advised 
against the use of phenolies in taxonomic considerations 
of this group. There have been no attempts to consider 
hybridization in Aster from a chemical point of view. 


SELECTION AND MORPHOLOGICAL 
ANALYSIS OF COLONIES 


Living material of A. acuminatus and A. nemoralis was 
provided by Dr. Pike from natural habitats in eastern 
Maine and brought to the greenhouse at the University of 
New Hampshire in the autumn of 1969. Specimens of A. 
Blakei were collected by the senior author near the south- 
ern shore of Lake Ossipee in New Hampshire during the 
same season. The specimens were potted in a mixture of 
peat and vermiculite and placed in a house maintained at 
24° C. When cold treatment was necessary, the plants were 
moved to a room maintained at 10°C. This treatment was 
applied from late November to mid-January of each year. 

Specimens subjected to morphological and chemical an- 
alysis were collected from three main areas. The first col- 
lection of twenty-five specimens came from the southern 
shore of Lake Ossipee in New Hampshire. A second col- 
lection of sixteen plants came from the Southern shore of 
Lake Winnisquam in New Hampshire. A final collection of 
fifty-four plants came from Great Wass Island in Washing- 
ton County, Maine. This latter collection consisted of a to- 
tal of fourteen specimens of A. nemoralis and A. acumina- 
tus in discrete colonies and forty specimens of A. nemoralis, 
A. Blakei and A. acuminatus collected at Ponds Point on 
the eastern tip of the island facing the Gulf of Maine. Aster 
acuminatus was collected from colonies in a wooded area 
of high elevation. Aster nemoralis was collected from col- 
onies in a small bog in the center of the island. These asters 
are clonal. Their stoloniferous habit is a characteristic of 
the species (Fernald, 1950). The clones at Lake Ossipee 
and Lake Winnisquam were well defined. One ramet was 
sampled from each clone at these locations. The clones were 


4 Rhodora [Vol. 75 


not well defined at Great Wass Island. Sampling at this 
location was based on morphological diversity throughout 
the area of the population. The plants collected were all 
scored utilizing the hybrid index described in detail by 
Pike (1970). 

The morphological analyses of specimens collected from 
these sites are summarized in Figure 1. Plants scoring 0-4 
were designated as A. nemoralis, 8-19 as A. Blakei and 25- 
30 as A. acuminatus (Pike, 1970). The population at Lake 
Winnisquam was essentially a variable population of A. 
Blakei (Figure 1a). The population at Lake Ossipee in- 
dexed as A. nemoralis or A. Blakei (Figure 1b). The pop- 
ulations at Great Wass Island contained the parental taxa 
in discrete colonies (Figure 1c), and all three taxa togeth- 
er in a local population at Ponds Point (Figure 1d). Fi- 
gure le summarizes the data in Figures 1c and 1d. 


0 1 2 3 4 5 6 7 8 9 w n 12 13 14 15 l6 7 18 19 20 21 22 23 24 25 26 27 28 29 30 
E 
5 
Oo 1 234 5 6 7 8 9 10 11 12 13 M 15 16 1 18 19 20 21 22 23 24 25 26 77 26 29 3 
D 
IT 
o ! 2 3 4 5 6 789 WN 12 13 14 15 16 V 18 19 20 21 2223 24 25 26 27 28 29 30 
c 
5 
o 12 3 45 6 7 8 9 10 1) 12 13 14 15 16 17 18 19 20 21 2223 24 25 26 27 28 29 30 
B 


| 
012 345 67 B 9 WW 121 M 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 
A 


Fig. 1. Morphological hybrid index of Asters at (a) Lake Winni- 
squam (b) Lake Ossipee and (e, d, e) Great Wass Island. 


1973] Aster Blakei — Hill and Rogers 5 


CHEMICAL DETECTION OF HYBRIDIZATION AND INTROGRESSION 


For chemical analyses, fresh basal leaves were shredded 
at the collection site into 10 ml. of methanol containing 1% 
1N HCl. The extracts were stored in the dark for 48 hours 
at room temperature and then concentrated to approxi- 
mately 1 ml. by evaporation with the aid of a hand dryer. 
Paper chromatography gave poor resolution in a compara- 
tive test with thin-layer chromatography. In contrast, thin- 
layer chromatography required only a small amount of ex- 
tract and gave excellent resolution. The aluminum-backed 
thin-layer plates were prepared by EM Reagents Division, 
Brinkman Instrument, Westbury, New York. 


Ten microliters of the sample were spotted on one cor- 
ner of a cellulose plate and chromatographed in the first 
dimension for 6.8 hours with n-butanol, glacial acetic acid 
and water (6:1:2). After drying overnight, the plate was 
chromatographed for 2-2 1/2 hours in the second dimension 
with 10% acetic acid containing 0.1% sodium acetate. Af- 
ter drying, the plate was viewed under visible and ultravio- 
let light. Colors of each spot under both conditions were 
recorded. These colors were again recorded after exposure 
to ammonia vapor. The plates were subsequently treated 
with spray reagents recommended by Block, et al., (1958) 
to distinguish among spots and to determine if these were 
phenolics. The sprays were: 1% alcoholic ferric chloride; 
1% aqueous basic lead acetate; 1% aqueous lead acetate; 
1% aqueous sodium carbonate; 1% alcoholic aluminum 
chloride and Benedicts reagent. Also, diazotized sulfanic 
acid was applied according to the specifications of Smith 
(1960). It was discovered that 1% aqueous lead acetate 
gave good distinctive colors under longwave ultraviolet 
light. Each spot was labeled with a number for identifica- 
tion on the basis of color of fluorescence, color reactions and 
position on the plate. The relative location of each spot to 
each other also was used as a criterion of identification. No 
attempts were made to determine the chemical structure of 
these compounds. 


o 


Rhodora [Vol. 75 


A premliminary survey of the phenolies present in the 
leaves of A. nemoralis, A. acuminatus and A. Blakei col- 
lected by Pike and the senior author revealed differences 
between the number of phenolics at various developmental 
stages (Hill, 1972). This source of variation was eliminated 
by studying the phenolics only at the time of full flowering. 

Chromatographs of the specimens collected from Lake 
Ossipee, Lake Winnisquam and Great Wass Island yielded 
numerous spots identified as phenolic compounds by the 
following criteria: (a) behavior in the developing solvents 
(Seikel, 1962) ; (b) reaction to sulfanilic acid; (c) colors 
under UV light before and after exposure to ammonia (AI- 


HOAc 


] «———— — ——— —1 
BAW 


Figure 2. Phenolic profile of Aster Blakei, A. nemoralis, and A. 
acuminatus. Spots 1, 2, 3, and 4 are specific to A. nemoralis; spots 


26, 27, and 28 are specific to A. acuminatus. The remaining spots are 
common to all three taxa. 


1973] Aster Blakei — Hill and Rogers 7 
5 

a 345 67 8 9 WN 12 13 14 W 16 17 18 19 20 21 22 23 24 25 26 27 28 
10 

5 

1 A 34 5 6 7 8 9 W-N 1213 4 1 16 y 18 9 202 22 23 24 25 26 27 28 


Figure 3. Frequency of occurrence of phenolies characteristic of 
(a) Aster nemoralis and (b) A. acuminatus in discrete colonies on 
Great Wass Island. Numbers refer to spots identified in Figure 2. 


ston, 1967) ; and (d) color reaction to phenolic-specific 
sprays (Block, et al., 1958). Twenty-eight spots were chos- 
en as diagnostic of these asters because of their high fre- 
quency of occurrence or because of their diagnostic value 
in identifying the parental taxa. 

The results of the chemical analysis of the Great Wass 
Island populations indicate that the parental taxa could be 
identified on a chemical basis alone. A. nemoralis, repre- 
sented in Figure 1c, contained four compounds which were 
species-specific. These were compounds numbered 1, 2, 3, 
and 4 shown in Figure 2. A. acuminatus, represented in 
Figure 1e, contained three compounds which were species- 
specific. These were compounds numbered 26, 27, and 28 in 
the phenolic profile in Figure 2. A histogram (Figure 3) of 
the frequency of occurrence of phenolies in the parental 
taxa in discrete colonies shows that twenty-one compounds 
were common to both species. The remaining seven pheno- 
lies clearly separate the parental species. 

A chemical analysis of the Ponds Point population on 
Great Wass Island revealed that the parental taxa main- 
tained a similar chemical integrity (Figures 4a, c). Aster 


8 Rhodora [Vol. 75 


Blakei contained a summation of the compounds specific to 
both parents but did not contain any new species-specific 
compounds (Figure 4b). The morphological and chemical 
evidence thus suggests that A. Blakei at Ponds Point orig- 
inated as a hybrid of A. acuminatus and A. nemoralis. It is 
clearly a chemical and morphological intermediate of the 
parental taxa. It is particularly significant that the three 
taxa could be identified on a chemical basis alone. 

The chemical analyses of A. Blakei at Lake Winnisquam 
and Lake Ossipee revealed similar phenolic profiles when 
compared to A. Blakei at Ponds Point. There were differ- 
ences in the frequency of occurrence of some compounds. 


10 


12.3 4 5 6 7 8 9 10 11 12 13 nM 15 16 V 18 19 20 21 22 23 24 25 26 27 28 
c 
10 
5 
12 3 4 5 6 7 8 9 O 1 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 
B 
10 
5 
12 3 4 5 6 7 8 9 111 12 13 4 15 1% 17 18 19 20 21 22 23 24 25 26 27 28 


Figure 4. Frequency of occurrence of phenolics characteristic of 
k I 

(a) Aster nemoralis (b) A. Blakei and (c) A. acuminatus at Ponds 

Point, Great Wass Island. Numbers refer to spots identified in Figure 

p. 


ú. 


1973] Aster Blakei — Hill and Rogers 9 


Table 1. Frequency (in %) of occurrence of compounds 
diagnostic of Aster nemoralis and A. acuminatus 
in relation to morphology in the population at 


Lake Ossipee, New Hampshire. 


Morphological 


Index Compound Number 

2 3 26 27 28 
1 50 100 100 
2 33 33 100 100 
3 67 67 100 
4 50 100 
8 75 25 75 
9 100 100 100 
11 100 50 100 
12 100 100 100 
13 100 100 100 
14 100 100 100 
15 100 100 100 
i 100 


The F, hybrids synthesized in the greenhouse demonstrated 
a high frequency of occurrence of A. accuminatus com- 
pounds in F, hybrids resulting from A. nemoralis female 
parents. The data on the asters collected from the Lakes and 
from the greenhouse are presented elsewhere (Hill, 1972). 

There was evidence for introgression at Lake Ossipee. 
Table 1 was constructed to show the relationship between 
chemical and morphological data using a method devised 
by Levin (1967). High frequencies of A. acuminatus com- 
pounds numbered 26 and 28 were found in plants which in- 
dexed as A. nemoralis. Specimens identified as A. Blakei 
contained only A. acuminatus phenolics. This suggests that 
introgression into A. nemoralis has been occurring at Lake 
Ossipee. A pictorialized scatter diagram was constructed 
according to the methods of Anderson (1949). The dia- 
gram was determined from the data on serrate vs. entire 


10 Rhodora 


NUMBER OF 


[Vol. 75 


PREPÉNT ie ARENT 


ë E ü 
ABSENT SPARSE ABUNDANT 


s. w' mw 


LEAVES 35-100 34-32 31-29 28-26 
SERRATE 3 
L 
E 
° E E ^35 
M 
A 
R 
c aa 
I 
' ' ana ** 
S 
ENTIRE " " 
o 
an 
[e] 1 2 3 
REVOLUTE FLAT 


LE 


Figure 5. Pictorialized scatter diagré 


AF MARGINS 


um of asters at Lake Ossipee. 


1973] Aster Blakei — Hill and Rogers 11 


leaf margins, revolute vs. flat leaf margins, zebra hairs, 
flower color and leaf number (Figure 5). Zebra hairs and 
serrate leaf margins, which are characteristics of A. acumi- 
natus, were present in many plants indexed as A. nemoral- 
is. This observation provides morphological evidence for 
the introgression of A. acuminatus into A. nemoralis at 
Lake Ossipee. Aster acuminatus was not present in this 
population. The habitat was a disturbed one which was lo- 
cated in an area of land development. There were woody 
habitats which could have supported A. acuminatus, but 
new roads cut off drainage and many of these areas were 
too moist and swampy to sustain A. acuminatus. It is sug- 
gested that the environmental disturbance removed this 
taxon, leaving A. Blakei to cross with A. nemoralis. The re- 
sult is the variable population found there today. 


GENETICS 


Specimens of A. acuminatus and A. nemoralis which were 
collected by Pike were crossed in the fall of 1970 and spring 
of 1971. The seeds from the fall 1970 crosses were germ- 
inated the following spring. These F, hybrids were then 
backcrossed to their parents. Crosses also were made be- 
tween various F, hybrids at that time. The resulting back- 
cross and recombinant progenies were grown during the 
spring of 1972. Crosses were performed in an insect-free 
house by rubbing the heads of two plants together when 
most of the flowers in a head were open and shedding pol- 


Table 2. Seed set of the cross Aster nemoralis X A. acu- 
minatus during two flowering seasons. 


Female Total Flowers Good Seed 
Season Parent Examined Seed Set (%) 
Fall, A. nemoralis 1180 205 17.4 
1970 A. acuminatus 1191 30 2.1 
Spring, A. nemoralis 2426 302 12.5 
1971 A. acuminatus 2297 26 1.1 


N 


5 


12 Rhodora [Vol. 


30 
25 


20 


o 12 34 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 2728 29 30 


Figure 6. Morphological hybrid index of the F, hybrids of the 
cross A. nemoralis X A. acuminatus. 


len. Seeds from the crosses were stored at room tempera- 
ture. Before germination they were subjected to a six to 
seven week treatment at 8°C. in Petri dishes containing 
moist filter paper. After this cold period, seeds were germ- 
inated at 24°C. in a peat-vermiculite mixture covered with 
ground sphagnum in pots enclosed in plastic bags. Ten days 
after germination the bags were removed. 


Crosses made between A. nemoralis and A. acuminatus in 
the fall of 1970 resulted in seed set that was high when A. 
nemoralis was the female parent (Table 2). The same re- 
sponse was obtained in the spring of 1970 using parents 
from a wider geographic source. The reason for this result 
probably rests in the presence of a maternal barrier to pol- 
lination or fertilization in A. acuminatus. 

The hybrid index of Pike (1970) was used to score 168 
F, hybrids. The results are represented in Figure 6. These 
specimens were morphologically intermediate and fell with- 
in the range assigned to Aster Blakei by Pike. Representa- 
tive specimens of A. Blakei collected from Lake Ossipee and 


1973] Aster Blakei — Hill and Rogers 13 


1 2419 


Figure 7. Aster Blakei from Lake Ossipee (right) and an F, hy- 
brid of the cross A. nemoralis X A. acuminatus (248, left). 


14 Rhodora [Vol. 75 


". 


Figure 8. Aster Blakei from Lake Ossipee (right) and an F. hy- 
brid of the cross. A. nemoralis X A. acuminatus (206, left). 


1973] Aster Blakei — Hill and Rogers 15 


F, hybrids from the cross A. nemoralis X A. acuminatus are 
compared in Figures 7 and 8. The morphological similari- 
ties are quite obvious. Many of the F, hybrids also resem- 
bled the specimens of A. Blakei collected from Lake Winnis- 
quam, Lake Ossipee and Great Wass Island. The F, hybrids 
were morphologically uniform for most characters. Coeffi- 
cients of variation calculated for the ten characters em- 
ployed in the hybrid index of Pike (1970) indicated the most 
variable characters to be internode length, the number of 
bracts subtending the peduncle and the number of heads 
(Hill, 1972). 

Backcrosses of the parents with the F, hybrid resulted in 
good seed set and seed germination (Table 3). The mater- 
nal barriers in A. acuminatus that existed for A. nemoralis 
pollen did not exist for pollen from the F,. The crosses 
made within the hybrid population also resulted in good 
seed set and seed germination, although germination was 


Table 3. Seed set and seed germination from crosses be- 
tween the parental taxa and their F, hybrid, 
Spring, 1971. 


Female Flowers Good # Seed Oo 
Parent Examined Seed Germ. Set (%) Germ. 
A. acuminatus 1267 345 158 27 46 
F, Hybrid 1213 476 310 39 65 
A. nemoralis 1594 428 211 27 51 
F, Hybrid 1306 225 111 17 49 


Table 4. Seed set and seed germination from the crosses 
made within the F, population, Spring, 1971. 


Type of Flowers Good # Seed % 
Cross Examined Seed Germ. Set (%) Germ. 
Intrasp. 784 303 124 41 41 


Sib 863 357 19 39 22 


16 Rhodora [Vol. 75 


01 2 3.4 5 6 7 8 9 10 11 12 13 14 15 16 V 18 19 2021 22 23 24 25 26 27 28 29 30 


aA a 


O ! 2 3 4 5 6 7 8 9 10) 1213 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 


B Fi HYBRID x A. NEMORALIS 


E 


3 4 7 B F 1011 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 
A MORALIS Fr HYBRID 
Figure 9. Morphological hybrid index of backcross and recom- 


binant progeny of Asters collected Summer, 1972. The female parent 
is represented first under each histogram. 


lower in progeny from sib-matings as compared with germ- 
ination of progeny from intraspecific crosses (Table 4). 
In the summer of 1972, 462 backcross and 110 recombin- 
ant progeny were collected and scored using the hybrid in- 
dex of Pike (1970). These are represented in Figures 9 and 
10. These data indicate that the range of variation as- 
signed to A. Blakei by Pike contains backcross and recom- 
binant progeny as well as hybrids. Chromatographs of six- 
ty of these specimens showed chemical evidence for intro- 
gression. However, the backcross or recombinant progeny 
could not be identified on the basis of chemistry alone. Thus, 


1973] Aster Blakei — Hill and Rogers 1T 


25 


20 


[n] 


901232245286 78 9 I D 13 M 1 16 1718 19 20 21 22 23 24 25 26 27 28 29 30 
B FL HYBRID x A. ACUMINATUS 


EET M 


DESC E ene ‘ini nmm 
A . ACUMINATUS x HYBRID 
ha AEA ER 


w 


Figure 10. Morphological hybrid index of backeross progeny of 
Asters collected Summer, 1972. The female parent is represented first 
under each histogram. 


chromatography might not be applicable in determining 
population structure within this aster complex, although 
this chemical technique has been successful in this regard 
in other genera, such as Coreopsis (Crawford, 1972) and 
Baptisia (McHale & Alston, 1964). Some of the backcross 
and recombinant progeny morphologically resembled some 
of the A. Blakei that the senior author has observed in his 
own collection as well as in Pike’s collection and the speci- 
mens in the Herbarium at the University of New Hamp- 
shire. 

The genetic results thus indicate that A. acuminatus and 
A. nemoralis can cross and produce a hybrid resembling A. 
Blakei. The hybrid can intercross with its parents and with 


18 Rhodora [Vol. 75 


Table 5. Mean pollen stainability (*«) of the backcross 
and recombinant progeny of asters collected in 
the summer of 1972. 


Female plants % 
Parent counted stained pollen 
A. nemoralis 52 81 
F, Hybrid 

A. acuminatus 71 79 
F, Hybrid 

A. nemoralis 

F, Hybrid 106 78 
A. acuminatus 

F, Hybrid 36 80 
F. Progeny 78 73 


itself. The complex appears capable of forming hybrid 
swarms which indeed have been observed by Pike (197 0) on 
some of the islands in the Bay of Fundy. He noted that 
some of these swarms show morphological skewness toward 
A. nemoralis. The data (Table 2) show that crossing be- 
tween the two parental taxa favors A. nemoralis as the 
female parent. If this crossing occurred in nature, there 
should be more seeds of the F, produced on the A. nemoral- 
is parent, and backcrosses would be more numerous in the 
wetter habitat of A. nemoralis. Progenies of the A. nemo- 
ralis backerosses would tend to be favored in preference to 
A. acuminatus backcrosses in moist habitats. On the other 
hand, backcrosses between A. acuminatus and the F, hy- 
brid were numerous and most were quite vigorous (Table 
3). Crosses between A. nemoralis and the F, hybrid pro- 
duced progeny which were numerous and vigorous when A. 
nemoralis was the female parent. A morphological skew 
was also noted toward A. nemoralis in the F. progeny (Fig- 
ure 9c). It thus appears that under greenhouse conditions, 
introgression could go either way, while in nature, the in- 


1973] Aster Blakei — Hill and Rogers 19 


trogression is in the direction of A. nemoralis. The deter- 
mining factor which would promote or inhibit introgression 
between A. acuminatus and A. nemoralis would thus be se- 
lection by the habitat. 


CYTOLOGY 


For root tip studies, 0.002 M 8-oxyquinoline was used as 
a pretreatment for 60-80 minutes at room temperature. The 
root tips were then stained and squashed in aceto-orcein ac- 
cording to the method of Huziwara (1957). Cover slips 
were smeared with Mayers albumin and dried over a flame 
to permit adhering of the preparation to the cover slip. Per- 
manent mounts were made according to the technique of 
McClintock (1929) with the following modifications; the 
slide and cover slip were separated in 10% acetic acid using 
the method of Celeraier (1956); the cover slip was then 
passed through changes of 1:1, 1:3, and 1:9 acetic alcohol 
and two changes of 95% ethanol. The slide and cover slip 
were then recombined in diaphane. For meiotic studies, 
flower buds were fixed in 1:3 acetic alcohol. Flowers were 
dissected in a small vial containing 70% alcohol. Each 
flower suitable for analysis was placed on a slide in a drop 
of aceto-carmine stain. Permanent slides were made as de- 
scribed above. Chromosomes were observed under oil at 
1125X. Photographs were taken with a Kodak camera 
mounted on a Spencer (A.O.) microscope with the prepara- 
tion under oil at 1455 X. 


The chromosome number of these taxa was determined to 
be 2n = 18. This was based on counts of mitotic and mei- 
otic chromosomes of asters collected from Lake Ossipee, 
Gould Pond in Milton, New Hampshire, Great Wass Island, 
North Lubec, Maine, and Campobello Island in New Bruns- 
wick, Canada. A total of forty-three specimens gave good 
counts. The chromosome numbers of A. Blakei and A. nem- 
oralis were new and were reported in the literature (Hill 
and Rogers, 1970). These specimens were not karyotyped. 
A representative plate of the mitotic chromosomes of A. 


20 Rhodora [Vol. 75 


Figure 11. Mitotic chromosomes of Aster nemoralis, Metaphase. 
(above) 

Figure 12. Meiotic chromosomes of Aster Blakei, Metaphase I. 
(below) The arrow indicates a loosely associated bivalent. 


1973] Aster Blakei — Hill and Rogers 2l 


nemoralis is shown in Figure 11. A plate of the meiotic 
chromosomes of A. Blakei shows nine bivalents, one of them 
loosely associated (Figure 12). The behavior of the meiotic 
chromosomes of all three taxa was compared. Both parental 
taxa formed nine bivalents at meiosis. No irregularities 
were observed. Pairing in A. Blakei was regular although 
loose associations were occasionally noted. Some bridges 
and lagging were observed in Anaphase I. Meiotic stages 
on 239 plates were observed. Of these, 93% showed nine 
bivalents, and the remaining few demonstrated the irregu- 
larities referred to above. Similar observations were noted 
in the F, population. 


Pollen from 41 F,, 265 backcross, and 78 recombinant 
progeny were stained and scored. The pollen grains were 
stained with aniline blue in lactophenol, those staining dark 
blue being scored as fertile. The first 200 grains were scored 
for stainability under 10X magnification. The results for 
the backcross and recombinant progeny are listed in Table 
5. The mean pollen stainability was lower in these progeny 
than was the mean stainability in the F, hybrid (89-90% ) 
and parents (96-97%). The differences noted in Table 5 
were not statistically different. Although it is obvious that 
a certain degree of hybrid breakdown has occurred, it ap- 
pears that the F,, backcross and recombinant progeny syn- 
thesized in the greenhouse are fertile. 


CONCLUSIONS 


The chemical evidence indicated that specimens identified 
as A. Blake, were clearly intermediates of A. acuminatus 
and A. nemoralis. Identification of A. nemoralis, A. acu 
minatus and A. Blakei could be done on the basis of phenolic 
examination alone. This is also true of the F, hybrids of 
the cross A. nemoralis X A. acuminatus. Backcross or re- 
combinant progeny could be identified morphologically, but 
they could not be identified on the basis of chemistry alone. 

The genetic evidence indicated that the hybrid of A. 
acuminatus and A. nemoralis was attainable and was in- 


22 Rhodora [Vol. 75 


distinguishable from A. Blakei. The parental taxa can in- 
tercross with the F, hybrid and the F, hybrid can produce 
F. progeny. The production of a hybrid swarm in the 
greenhouse thus suggests the production of hybrid swarms 
in nature. The backcross and recombinant progeny were in- 
distinguishable from some specimens of A. Blake. 

The cytological evidence demonstrated that the chromo- 
some number for A. Blakei, A. acuminatus, and A. nemo- 
ralis was 2n. = 18. A. Blakei and the F, hybrid demon- 
strated regular meiosis most of the time. Pollen stainabili- 
ty with aniline blue in lactophenol was very high in both 
parental taxa, lower in the F, hybrid and even lower in the 
backcross and recombinant progeny. The evidence suggest- 
ed that A. Blakei was a fertile species hybrid. 

The data thus lead to a confirmation of Pike's (1970) hy- 
pothesis that A. Blakei is a hybrid of A. acwminatus and A. 
nemoralis. It is always difficult to assess the exact size of 
a species population in terms of the number, location, and 
size of individual sub- or local populations over a species 
range. Our sample sizes reflect only a small area that cov- 
ers the range of A. Blakei. Keeping this in mind, we wish 
to suggest that A. acuminatus and A. nemoralis are semi- 
or incipient species. They remain isolated by geography but 
cross within their overlapping ranges when the opportunity 
is presented. The major isolating barrier between the 
species would be the absence of hybrid or recombinational 
habitats when interspecific crossing occurs. 


ACKNOWLEDGMENTS 


Thanks are extended to Dr. Radcliffe B. Pike for his ef- 
forts in collecting the parental asters used in the genetic 
portion of this study, for his help in cultivating asters from 
seed and for his aid in locating the asters at Ponds Point, 
Great Wass Island. The hospitality of the summer residents 
near Ponds Point is also greatly appreciated. Thanks are 
also due to Dr. Albion R. Hodgdon for his help in locating 
the populations at Lake Ossipee and Lake Winnisquam. We 


1973] Aster Blakei — Hill and Rogers 23 


also thank Dr. L. C. Peirce, Professor and Chairman of 
Plant Science and Dr. Douglas Routley, Professor of Plant 
Science, both of the University of New Hampshire, for the 
critical reading of this manuscript. Voucher specimens for 
this work have been placed in the Herbarium of the Uni- 
versity of New Hampshire. 


LITERATURE CITED 


ABRAHAMSON, WARREN G., & OTTo T. SoLBRIG. 1970. Soil prefer- 
ences and variation in flavonoid pigments in species of Aster. 
Rhodora 72: 251-263. 

ALSTON, R. E. 1965. Flavonoid chemistry of Baptisia: A current 
reevaluation of chemical methods in the analysis of interspecific 
hybridization. Taxon 14: 268-274. 

1967. Biochemical systematies. In: T. Dobshansky, 

M. Hecht and W. C. Steere, eds., Evolutionary biology, Vol. 1. 

Appleton-Century Crosts, New York. pp. 197-305. 

, & B. L. Turner. 1962. New techniques in analysis 
of complex natural hybridization. Proc. Natl. Acad. Sci., U.S.A. 
48: 130-137. 

ANDERSON, EDGAR. 1929. Variation in Aster anomalous. Annals 
Missouri Bot. Gard. 15: 129-140. 

1949. Introgressive hybridization. John Wiley 
and Sons, Inc., New York. 109 pp. 

AVERS, CHARLOTTE, J. 1953a. Biosystematic studies in Aster. I. 
Crossing relationships in the Heterophylli. Amer. Jour. Bot. 40: 
669-675. 


1953b. Biosystematic studies in Aster. Il. 
Isolating mechanisms and some phylogenetic considerations. Evo- 
lution 7: 317-327. 

BLOCK, R. J., E. L. DURRUM, & G. ZwErG. 1958. A manual of paper 
chromatography and paper electrophoresis, Second edition. Aca- 
demic Press, Inc., New York. 710 pp. 

CARTER, LYNN C., & B. G. BREHM. 1969. Chemical and morphologi- 
cal analysis of introgressive hybridization between Iris tenax and 
Iris chrysophylla. Brittonia 21: 44-54. 

CELARIER, ROBERT P. 1956. Tertiary butyl alcohol dehydration of 
chromosome smears. Stain Technol, 31(4): 155-157. 

CRANG, RICHARD E., & H. L. DEAN. 1971. An intergeneric hybrid 
in the Sileneae (Caryophyllaceae) Bull. Torrey Bot. Club 98: 
214-217. 

CRAWFORD, DANIEL J. 1972. The morphology and flavonoid chemistry 
of synthetic infraspecific hybrids in Coreopsis mutica (Com- 
positae). Taxon 21: 27-38. 


24 Rhodora [Vol. 75 


FAHSELT, DIANE, & MARION OWNBEY. 1968. Chromatographic com- 
parison of Dicentra species and hybrids. Amer. Jour. Bot. 55: 
334-345. 

FERNALD, M. L. 1950. Grays Manual of Botany. Eighth edition. 
American Book Company, New York, 1632 pp. 

HARBORNE, J. B. 1968. The use of secondary chemical characters 
in the systematics of higher plants. In: J. G. Hawkes, ed., 
Chemotaxonomy and serotaxonomy. Academic Press, New York. 
pp. 173-191. 

HILL, L. MICHAEL. 1972. Chemical, cytological and genetic consid- 
erations of the possible hybrid origin of Aster Blakei (Porter) 
House. Doctoral Dissertation, The University of New Hamp- 
shire, Durham, New Hampshire. 84 pp. 

, & O. M. Rocers. 1970. Chromosome numbers of 
Aster Blakei and A. nemoralis. Rhodora 72: 437-438. 

Huzrwana, Y. 1957. Karyotype analysis in some genera of Com- 
positae. II. The karyotype of Japanese Aster species. Cytologia 
22: 96-112. 

JAWORSKA, H., & N. NvBOoM. 1967. A thin-layer chromatographic 
study of Saxafraga caesia, S. aizoides, and their putative hybrid. 
Hereditas 57: 159-177. 

LEVIN, D. 1967. An analysis of hybridization in Liatris. Brittonia 
19: 248-260. 

McOCuiNTOCK, B. 1929. A method for making aceto-carmin smears 
permanent. Stain Technol. 4: 53-56, 

MCHALE, JANICE, & R. E. ALSTON. 1964. Utilization of chemical pat- 
terns in the analysis of hybridization between Baptisia leucantha 
and B. sphaerocarpa, Evolution 18: 304-311. 

OLDEN, E. J., & N. NvBoM. 1968. On the origin of Prunus cerasus 
L. Hereditas 59: 327-345. 

PIKE, R. B. 1970. Evidence for the hybrid status of Aster Blakei 
(Porter) House. Rhodora 72: 401-436. 

ROSENDAHL, C. O., & ARTHUR CRONQUIST. 1949. The Asters of Min- 
nesota: a floristic study. Amer. Mid. Nat. 42: 502-512. 

SEIKEL, MARGARET K. 1962. Chromatographic methods of separation, 
isolation and identification of flavonoid compounds. In: T. A. 
Geissman, ed., The chemistry of flavonoid compounds. The Mac- 
Millan Company, New York. pp. 34069. 

SHINNERS, L. H. 1941. The genus Aster in Wisconsin. Amer. Midl. 
Nat. 26: 398-420. 

SMITH, DALE M., & D. A. Levin. 1963. A chromatographic study of 
reticulate evolution in the appalachian Asplenium complex. Amer. 
Amer. Jour. Bot. 50: 952-958. 

SMITH, I., ed. 1960. Chromatographic and electrophoretic tech- 
niques. Volume I, Interscience Publishers, New York. 617 pp. 


1973] Aster Blakei — Hill and Rogers 25 


TURNER, B. L., & R. E. ALSTON. 1959. Segregation and recombina- 
tion of chemical constituents in a hybrid swarm of Baptisia 
laevicaulis >< B. viridis and their taxonomic implications. Amer. 
Jour. Bot. 46: 678-686. 

,& T. J. MABRY. 1964. Partition chromatography as 
applied to taxonomic problems in the Asteraceae. Taxon 13: 11-14. 

UTTAL, L. J. 1962. Synthesis of Aster Herveyi. Rhodora 64: 113-117. 

WALKER, SALLY A. 1969. Cytological and chromatographic evidence 
for interspecific hybridization in Petalostemon. Annals Missouri 
Bot. Gard. 56: 261-267. 

WETMORE, RALPH, & ALBERT L. DELISLE. 1939. Studies in the gen- 
etics and cytology of two species in the genus Aster and their 
polymorphy in nature. Amer. Jour. Bot. 26: 1-12. 


DEPARTMENT OF BIOLOGY 
BRIDGEWATER COLLEGE 
BRIDGEWATER, VIRGINIA 22812 


DEPARTMENT OF PLANT SCIENCE 
NESMITH HALL 

THE UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, NEW HAMPSHIRE 03824 


CHROMOSOME NUMBERS IN ASTER 
PAUL VAN FAASEN AND FERN FRANK STERK 


Chromosome numbers are of ever-increasing importance 
to modern plant systematists. Although there have been 
numerous contributions made in this area over the past 
25 years, the chromosome numbers of most plants remain 
unknown. Even in some large genera which contain abun- 
dant and widespread species, many chromosome numbers 
are known from a single count, or from a few counts taken 
from a restricted portion of the range of a species. Con- 
sidering the value of the knowledge of chromosome numbers 
to evolutionary and taxonomic discussion (see Davis & 
Heywood 1963), it seems appropriate to expand this aspect 
of our knowledge whenever possible. 


As a necessary early step in a study of the variability 
and hybridizing capability in the Aster lateriflorus-simplex- 
pilosus complex, collections of these and other species of 
Aster were made, primarily in the northeastern United 
States, in 1970. Buds were collected in Carnoy’s 6:3:1 and 
maintained at ambient temperatures. Chromosome num- 
bers, using standard squash techniques, were determined 
in pollen mother cells undergoing meiosis, and, in a few 
cases, in root tip cells which were undergoing mitotic divi- 
sions. Chromosome counts from 32 populations of 17 dif- 
ferent species, along with locality data, are found in 
Table 1. These data include the first count reported for 
A. gracilis (n = 9); a count (n = 24) for A. concinnus, 
different from that previously reported for the species 
(2n — 46, Huziwara 1958) ; and a diploid count (n — 9) 
for A. undulatus, a species previously reported only as a 
tetraploid (» — 18, Avers 1953a, b). In addition, previous 
chromosome counts for those species included in this study 
are presented. Voucher specimens are deposited in the 
herbarium at Hope College. 


26 


1973] Chromosome Numbers — Van Faasen & Sterk 27 


Species 


Table 1. 


Chromosome 


Number 


Locality data 


Aster acuminatus 
Michx. 


Aster ciliolatus 
Lindl. 


Aster concinnus 
Willd. 


Aster cordifolius L. 


Aster gracilis 
Nutt. 


Aster junciformis 
Rydb. 


Aster lateriflorus 
(L.) Britt. 


io 


n —9 


“n= 9 


n 24 


n = 18 


n — 18 


i= 9 


an i} 


HH 


NEW HAMPSHIRE: CAR- 
ROLL CO. North Shore of 
Dan Hole Pond. Van Faa- 
sen 2712. 

MAINE: OXFORD CO. 2.6 
miles North of Bethel. Van 
Faasen 2725. 

VERMONT: ESSEX CO. 9.1 
miles west of North Strat- 
ford. Van Faasen 2730. 
MICHIGAN: EMMET co. 
1 mile South of Mackinaw 
City. Van Faasen 2787. 
PENNSYLVANIA:  MON- 
ROE CO. 0.5 mile North of 
Marshalls Creek. Van Faa- 
sen 2776. 

MAINE: OXFORD co. 2.6 
miles North of Bethel. Van 
Faasen 2724. 

NEW HAMPSHIRE: coos 
CO. 8.9 miles South of North 
Stratford. Van Faasen 
2728. 

NEW JERSEY: BURLING- 
TON CO. 1 mile North of 
Green Bank. Van Faasen 
2754. 

NEW HAMPSHIRE: coos 
CO. 0.5 mile North of Gor- 
ham. Van Faasen 2727. 
PENNSYLVANIA: MON- 
ROE CO. Delaware Water 
Gap. Van Faasen 2763. 


28 Rhodora [Vol. 75 


Chromosome 
Species Number Locality data 
»n- 16 NEW JERSEY: SUSSEX CO. 


7 miles South of Montague. 
Van Faasen 2780. 


Aster lowrieanus n—18 PENNSYLVANIA: MON- 


Porter ROE co. 5 miles North of 
Marshalls Creek. Van Faa- 

sen 2775. 
Aster macrophyllus n = 36 VERMONT: CALEDONIA 
L. co. 2.8 miles North of Gro- 


ton. Van Faasen 2746. 
n= 36 NEW HAMPSHIRE: GRAF- 

ToN co. 1 mile West of 

Warren. Van Faasen 2749. 


Aster patens n=10 NEW JERSEY: BURLING- 
Ait. TON co. Chatsworth. Van 
Faasen 2758. 
Aster pilosus ən — 48 MICHIGAN: OTTAWA CO. 
Willd. 3 miles South of Zeeland. 
Van Faasen 2674. 
Aster puniceus »-——8 MAINE: OXFORD CO. 0.5 
L. mile East of Bryant Pond. 


Van Faasen 2719. 

n-— 8 VERMONT: CALEDONIA CO. 
28 miles North of Groton. 
Van Faasen 2745. 


Aster sagittifolius n — 18 VERMONT: CALEDONIA CO. 
Wedem. 28 miles North of Groton. 
Van Faasen 2747. 
Aster simplex 24 — 32 MICHIGAN: ALLEGAN CO. 
Willd. New Richmond. Van Faa- 
sen 2685. 


n — 16 MAINE: CUMBERLAND CO. 
0.5 mile North of West 
Baldwin. Van Faasen 2717. 


1973] Chromosome Numbers — Van Faasen & Sterk 29 


Species 


Locality data 


Aster spectabilis 
Ait. 


Aster wmbellatus 
Mill. 


Aster undulatus 
L. 


Chromosome 

Number 
n = 16 
n = 16 
n — 16 
n = 82 
n = 82 
n = 36 
n= 9 
n=9 
n — 9 
n — 9 
n = 18 


MAINE: OXFORD CO. 2.6 
miles North of Bethel. Van 
Faasen 2723. 

NEW JERSEY: SUSSEX CO. 
0.5 mile North of Monta- 
gue. Van Faasen 2778. 
MICHIGAN: ALGER CO. 0.5 
mile South of Grand 
Marais. Van Faasen 2785. 
MAINE: YORK co. 1 mile 
west of Porter. Van Faasen 
2718. 

VERMONT: CALEDONIA CO. 
2.8 miles North of Groton. 
Van Faasen 2748. 


NEW JERSEY: BURLING- 
TON CO. 3 miles South of 
Chatsworth. Van Faasen 
2738; 


MAINE: YORK CO. 1 mile 
West of Porter, Van Faasen 
2714. 

MAINE: OXFORD co. 0.5 
mile East of Bryant Pond. 
Van Faasen 2720. 
VERMONT: ESSEX CO. 9.1 
miles West of North Strat- 
ford. Van Faasen 2736. 
MAINE: CUMBERLAND CO. 
0.5 miles North of West 
Baldwin. Van Faasen 2718. 
PENNSYLVANIA:  MON- 
ROE CO. 0.5 mile North of 
Marshalls Creek. Van Faa- 
sen 2777. 


N 
1 


I» 


30 Rhodora [ Vol. 


DISCUSSION 


Aster acuminatus Michx. Counts of » — 9 from three 
different populations confirm those for this species as pre- 
viously reported by Nelson (1966), Smith (1966), and Hill 
& Rogers (1970). 

Aster ciliolatus Lindl. This count (» — 36) is the same 
as reported for this species by Avers (1953a, b), Van 
Faasen (1963), and Lóve & Lóve (1964). 

Aster concinnus Willd. The chromosome count (n = 24) 
for this species, as determined in this study, places A. con- 
cinnus in that very diverse group of asters based on v — 8. 
Huziwara (1958) reported 2» — 46, an unusual count for 
Aster, for this species. He suggested that it was a “hypo 
hexaploid", presumably a product of aneuploid loss from 
2n — 48. 


Aster cordifolius L. The only previous tetraploid (» — 
18) counts for this species are by Van Faasen (1963) who 
also reported diploid plants. Avers (1953a, b, and 1954a, 
and 1957) reports only diploids for this species. 

Aster gracilis Nutt. This count, n = 9, is apparently 
the first count reported for this species and places the 
species in that large group of Aster species based on v = 9. 

Aster junciformis Rydb. A count of n = 16 confirms the 
previous count for this species made by Van Faasen (1963) 
but differs from the count n = 27 reported by Taylor 
(1967). 

Aster lateriflorus (L) Britt. Two tetraploid populations 
(n — 16) confirm that count by Van Faasen (1963) who 
also reported diploid and hexaploid populations of A. lateri- 
florus. Mulligan (1967) also reports n = 16 in this species. 


Aster lowrieanus Porter. A count of n = 18 confirms 
that count for this species made by Avers (1953a). Van 
Faasen's (1963) count » — 9 for a specimen tentatively 


identified as A. lowrieanus must be attributed to a variant 
form of A. cordifolius. 
Aster macrophyllus L. Counts of n = 36 confirm those 


1973] Chromosome Numbers — Van Faasen & Sterk 31 


reported by Huziwara (1941), Van Faasen (1963), Lóve 
& Lóve (1964), and Mulligan (1967). 


Aster patens Ait. Tetraploid counts of n = 10 were also 
reported by Huziwara (1941) and Avers (1954b). Jones 
(1968), however, reports a diploid count of n — 5. 


Aster pilosus Willd. Avers (1954b), Huziwara (1958) 
and Van Faasen (1963) also report hexaploid counts (2m 
— 48) for this species. Bostick (1965) reports the unusual 
count » — 12 for A. pilosus. 


Aster puniceus L. The count of n = 8 agrees with counts 
previously reported by Huziwara (1958), Van Faasen 
(1963) and Love & Love (1964, 1966). Chouksanova et al. 
report 2n — 36 for this species (1968). 


Aster sagittifolius Wedem. Tetraploid (n = 18) counts 
were also reported by Avers (1953a, b) and Van Faasen 
(1963), both of whom also reported diploid counts for this 
species. 

Aster simplex Willd. Counts of n = 16 and n = 32 con- 
firm previous counts by Avers (1954b), Huziwara (1958), 
and Van Faasen (1963). 


Aster spectabilis Ait. Solbrig et. al (1964) also reported 
n = 36 for this species. 


Aster wmbellatus Mill. Diploid (n = 9) counts for A. 
umbellatus were also reported by Huziwara (1962), Lóve 
& Lóve (1964, 1966), Jones (1966) and Van Faasen 
(1963). 

Aster undulatus L. Avers (1953a, b) also reported n = 
18 for this species but this is the first report of diploid 
plants (n = 9) for this species. 


ACKNOWLEDGEMENTS 


Fieldwork was suported, in part, by a Sigma Xi grant- 
in-aid to the senior author and laboratory work by the 
junior author was supported, in part, by a National Science 
Foundation Undergraduate Research Participation Grant. 


32 Rhodora [Vol. 75 


The authors are grateful for this support of their studies 
on the genus Aster. 


LITERATURE CITED 


Avers, C. J. 1953a. Biosystematic studies in Asters I. Crossing 

relationships in the Heterophylli. Amer. Jour. Bot. 40: 669-675. 
1958b. Biosystematic studies in Aster. II. Isolating 
mechanisms and some phylogenetic considerations. Evolution 7: 
317-827. 


1954a. Chromosome behavior in fertile triploid Aster 

hybrids. Genetics 39: 117-126. 

1954b. Documented chromosome numbers of plants. 
Madrofio 12: 210. 

1957. Fertile hybrids derived from a wide species 
cross in Aster. Evolution 11: 482-486. 

Bostick, P. E. 1965. Documented chromosome numbers of plants 
65: 2. Sida 2: 165-168. 

CHOUKSANOVA, N. A., et al. 1968. Data on karyology of the family 
Compositae Giseke. Citologija 10: 198-206. 

Davis, P. H., & V. H. HEYWOOD. 1963. Principles of angiosperm 
taxonomy. Van Nostrand, Princeton, NJ. 556 pp. 

HILL, L. M., & O. M. ROGERS. 1970. Chromosome numbers of Aster 
blakei and A. nemoralis. Rhodora 72: 437-438. 

HUZIWARA, Y. 1941. On the chromosomes of some species of Aster 
from foreign countries. Bot. Zool. 9: 75-76. 

1958. Karyotype analysis in some genera of Com- 

positae. V. The chromosomes of American Aster species. Jap. 

Jour. Genet. 33: 129-137. 

1962. Karyotype analysis in some genera of Com- 
positae IX. Chromosomes of European species of Aster. Bot. 
Mag. Tokyo 75: 143-149. 

JoNEs, S. B., JR. 1966. Compositae chromosome numbers. Bull. Tor- 
rey Bot. Club 93: 278-279. 

1968. Chromosome numbers in southeastern United 
States Compositae. Bull. Torrey Bot. Club 95: 393-395. 

Love, A. & D. Love. 1964. In IOPB Chromosome number reports 
I. Taxon 13: 100-110. 


& 1966. Cytotaxonomy of the alpine vascular 
plants of Mount Washington, Univ. Colorado Studies Ser. Biol. 
24: 1-74. 


MULLIGAN, G. A. 1967. In IOPB chromosome number reports XI. 
Taxon 16: 215-222. 


1973] Chromosome Numbers — Van Faasen & Sterk 33 


NELSON, A. D. 1966. In IOPB chromosome number reports VII. 
Taxon 15: 155-163. 

SMITH, E. B. 1966. In IOPB chromosome number reports VII. Tax- 
on 15: 155-163. 


SoLBRIG, O. T., et al. 1964. Chromosome numbers in Compositae V. 
Astereae II. Amer. Jour. Bot. 51: 513-519. 


TAYLOR, R. L. 1967. In IOPB chromosome number reports XIII. 
Taxon 16: 445-461. 


VAN FAASEN, P. 1963. Cytotaxonomic studies in Michigan asters. 
Mich. Bot. 2: 17-27. 


BIOLOGY DEPARTMENT 
HOPE COLLEGE 
HOLLAND, MICHIGAN 49423 


THE NORTHERN LIMITS OF THE DISTRIBUTIONS 
OF HICKORIES IN NEW ENGLAND 


WAYNE E. MANNING 


Five species of hickories are native to New England, 
mockernut hickory, Carya tomentosa (Poir.) Nutt., bitter- 
nut hickory, C. cordiformis (Wang.) K. Koch, shagbark 
hickory, C. ovata (Mill.) K. Koch, pignut hickory, C. glabra 
(Mill.) Sweet, and sweet pignut or red hickory, C. ovalis 
(Wang.) Sarg. The last species, at one time combined by 
Little (1953) with C. glabra, has recently been designated 
by him (Little, 1969) as C. glabra var. odorata (Marsh.) 
Little. 

The exact distribution in New England of these species 
is still not fully known. Except for some records in south- 
ern Quebec, the northern limits in New England represent 
the northeast limits of the species. The ranges given in 
various books and on the distribution maps are largely 
inaccurate and contradictory. This is due partly to lack of 
collecting, partly to misidentification and partly to guess- 
work where specimens are unknown. The greatest error 
is in the distribution of the mockernut. Some of the de- 
scriptions of the ranges are much condensed and indefinite, 
many incorrect; also maps on a smaller scale are usually 
highly generalized. The best maps in accuracy and detail 
not only for New England but also Canada and the rest 
of the United States are those of Little (1965, 1971). The 
greatest detail in range in New England is given by Blakes- 
lee and Jarvis (1911) and by Seymour (1969, 1970) ; these 
are not entirely correct or complete. Seymour gives the 
distribution by counties and towns and one must know the 
location of these to fully understand the distribution. 

This paper is an attempt to correct as far as possible 
the ranges of the hickories in New England, and thus show 
the northeast limits of the species. This will lay a founda- 
tion for future workers to make better maps and descrip- 
tions of ranges. Also clear are locations where future field 


54 


1973] Hickories — Manning 35 


work will be of value. For this study I have checked the 
identification of the hickories in the major herbaria of 
northern and central New England: University of Maine 
(MN, MAINE), U. of New Hampshire (NH, NHA), U. of 
Vermont (v, vT), U. of Massachusetts (MS, MASS), Dart- 
mouth College (H, HNH), Arnold Arboretum (A, A), Gray 
Herbarium (G, GH), New England Botanical Club (N, 
NEBC), Clark University (C, CUW), and Smith College, 
(S, SCHN). For each herbarium the first symbol is that 
which I will use in this paper, the second is the official 
one in Index Herbariorum (Lanjouw and Stafleu, 1964). 
Furthermore, I have recently collected rather extensively 
in Vermont, western New Hampshire, and northern Massa- 
chusetts; the field work has added over 50 towns to the 
lists of towns given by Seymour (1969). My specimens 
are deposited in various New England herbaria. The 
results of my study are given below in the lists of counties 
and towns under each species and on the maps. In the lists 
are given the symbols for the names of the herbaria where 
specimens are located. The symbol “wm” refers to my 
herbarium at Bucknell University, and “Z” refers to the 
observation by Dr. Robert Zimmerman, who did not col- 
lect specimens. For each species one map gives the detailed 
presently known distribution indicated for each town in 
Vermont, New Hampshire, Maine and Massachusetts where 
a specimen has been collected. A second map shows Little’s 
distribution with my corrections in the New England area; 
his distribution in eastern New York is included, in order 
to show where one might look for the species in adjacent 
areas in New England where no records are now known. 
With extremely few exceptions I have seen herbarium 
specimens for each record I have given. The maps and 
lists of towns are still incomplete as collections are incom- 
plete; there will undoubtedly be additions of towns within 
the ranges and some extensions of ranges with future 
collecting. Some of the towns I have given May represent 
planted trees as it is difficult to be sure of the situation 
from meager notes on herbarium specimens; even in the 


36 Rhodora [Vol. 75 


field there are often uncertainties. In Maine, Hyland and 
Steinmetz (1944) have definitely distinguished native from 
planted trees, removing pignut and bitternut from the list 
of native trees of Maine, and indicating in which counties 
the only native species, Carya ovata, is indigenous, and in 
which it is planted. 

It is well known that because of overlapping features 
and the tremendous variation hickories are very difficult 
to identify. Thus every species in New England may have 
hairy leaflets, hairy rachises, 7 leaflets. I have discussed 
the features used in identification, given suggestions for 
collecting specimens, and presented a detailed key to species 
using vegetative as well as reproductive features (Manning, 
1950). A less detailed and somewhat modified key to New 
England species is given below. A full distinction between 
C. glabra and C. ovalis is given in the key, despite the fact 
that the distribution is discussed later under C. glabra- 
ovalis complex. 

Under each species are given comments on distribution 
records. Changes from the maps of Little (1965, 1971) are 
indicated here as well as on the maps. Also given is a sug- 
gested description of the range in New England. 


Key to New England species of Carya 


1. Buds completely yellow, scurfy, flattened ; leaflets 7-9 
(-11), usually narrow ; husk very thin, splitting slightly 
more than half-way; meat very bitter; bark tight. . 
eee tte e eens 2. Bitternut hickory, C. cordiformis 

1. Buds brown or gray, not scurfy, though sometimes with 
some yellow scales; meats usually sweet. ....-..--- 2 


2. Teeth (many of them) of leaflets with dense sub- 
terminal tufts of hairs; leaflets 5, rarely 7; over- 
wintering terminal buds rather slender, 13-22 mm. 
long, the outer dark brown bud-scales persistent; 
husk of fruit very thick, splitting to base; nuts 
white, angled, thin-shelled; bark of older trees 
shaggy. .......... 3. Shagbark hickory, C. ovata 


1973] Hickories — Manning 37 


2. Teeth glabrous or ciliate, without special tufts of 
hairs; nuts thick-shelled. ............. eee 3 


3. Leaflets 5-9, fascicled-hairy (tomentose) be- 
neath; branchlets stout; twigs and rachises of 
leaves often fascicled-hairy; overwintering ter- 
minal buds 10-15 mm. long, subglobose, the outer 
dark bud-scales finally deciduous, so buds gray 
silky-hairy ; husk medium thick, splitting to base; 
nuts brown, angled; bark tight, furrowed. .... 
UU EE 1. Mockernut hickory, C. tomentosa 


3. Leaflets 5-7, glabrous or rarely hairy beneath; 
twigs glabrous; rachises of leaves usually gla- 
brous or occasionally densely  puberulent; 
branchlets slender; overwintering terminal buds 
7-12 mm. long; husk thin; nuts whitish or light 
brown, smooth or slightly angled. ......... 4 
4. Fruit dark brown, shining, smooth; husk 

finally splitting only to middle or sometimes 
by one suture to near base; petiole not red; 
buds not yellow-dotted; lower leaflet surfaces 
inconspicuously yellow-dotted; terminal bud 
slender, pointed, the outer dark brown scales 
persistent or deciduous; bark tight. ....... 
e ee ales 4. Pignut hickory, C. glabra 


4. Fruit light brown, dull, warty; husk finally 
splitting to base, hence immature fruit 4- 
ridged to base; petiole often red; overwinter- 
ing terminal buds subglobose, essentially 
blunt, the outer dark bud-scales finally de- 
ciduous, the bud then gray, silky-hairy; 
lateral buds and lower leaflet surfaces con- 
spicuously yellow- and brown-dotted ; bark of 
older trees usually shaggy or loosely scaly. 

. 4. Sweet pignut, Red hickory, C. ovalis 


1. Carya tomentosa (Poir. Nutt., mockernut hickory 
As noted above, this species has had the greatest errors 
in published descriptions of ranges and especially in the 


38 Rhodora [Vol. 75 


maps. This is due in large part to the fact that the name 
Carya alba has been used by some botanists for mockernut, 
by others for shagbark. The name is now dropped as a 
*nomen confusum". Most maps showing clear ranges such 
as Little (1965), Hough (1907), Jaques (1941), and 
Harlow (1958) indicate the mockernut as occurring in New 
Hampshire and Vermont. Most older floras of Vermont 
(Burns and Otis, 1916, Dole, 1937), report the species in 
Vermont. However, the northernmost New England known 
records are in Middlesex and Hampshire (Mt. Warner, 
North Hadley) Counties, Massachusetts. All records men- 
tioned north of Massachusetts are based on incorrect 
identification. Strangely enough, the distribution given by 
Sargent (1933) was essentially correct, and more recent 
publications have confused the situation. Specimens are 
here reported from Cape Cod (Barnstable Co., Mass.) but 
are not shown by Little. 

Massachusetts: MIDDLESEX CO.: Arlington, N; Belmont, N; Concord, 
N; Framingham, N; Lincoln, N; Medford, N; Natick, N; Wakefield, N; 
Waltham, N. SUFFOLK CO.: Boston, N. NORFOLK CO.: Braintree, N; 
Brookline, N; Canton, N; Dover, N; Milton, N; Needham, N; Nor- 
wood, N; Wellesley, N. PLYMOUTH CO.: Marshfield, N. BRISTOL CO.: 
Dartmouth, N; Fall River, N; New Bedford, N; Swansea, N. BARN- 
STABLE CO.: Barnstable, wM; Bourne, N; Brewster, N; Dennis, N; 
Falmouth, N; Sandwich, wM; Wellfleet, N. DUKES CO.: Elizabeth 
Islands, N; Martha's Vineyard: Chilmark, G, N; Tisbury, N; West 
Tisbury, G. NANTUCKET CO.: Nantucket Id., G, N. WORCESTER CO.: 
Oxford, C. HAMPSHIRE CO.: Easthampton, WM; Hadley, G, S, WM; 
Northampton, WM; Southampton, G, WM. 

Maps 1 and 5 show the presently known distribution of 
the mockernut in New England. This may be stated verb- 
ally as: eastern Massachusetts (Middlesex Co. to Cape 
Cod, Martha's Vineyard, Nantucket) and Connecticut River 
Valley in Massachusetts (Hampshire and Hampden Co.), 
Rhode Island and Connecticut. It is absent in Maine, Ver- 
mont, New Hampshire, most of central and western Massa- 


chusetts. 


2. C. cordiformis (Wang.) K. Koch, bitternut hickory. 
Most maps and descriptions of ranges are essentially 


1973] Hickories — Manning 39 


' 
ri 


Yd -——— o 


/ Ma 
I = 
/ 
i PAU 
| 
e °° 


Map 1. Distribution of Carga, tomentosa, in central New England, 
limits indicated by heavy solid line. A dot designates one or more 
specimens collected in the town located there. 


correct for New England. A few of these report the species 
in Maine, central New Hampshire and central Vermont. 
No records are now known of these areas, and probably 
the maps are merely generalized. My studies show that 
the bitternut has a somewhat broader distribution in New 
Hampshire and Vermont than shown by Little. No speci- 
mens, however, have been located for the region next to 
Canada in northwestern Vermont. 

Vermont: CHITTENDEN CO.: Burlington, v; Charlotte, G, v; Rich- 
mond, H, N, v; South Burlington, V. ADDISON CO.: Middlebury, v; 
Panton, N, V. RUTLAND CO.: Brandon, N, S, v; Castleton, N, v; Ira, 
N, V; Poultney, H, N, NH, V; Rutland, G, H; Sudbury, N; Wallingford, 
G; Wells, N, V. BENNINSTON CO.: Arlington, N, v; Dorset, v; Pownal, 
v; Shaftsbury, N, V. CALEDONIA CO.: Peacham, H. ORANGE CO.: Brad- 
ford, H, N, NH, v; Newbury, H, N, NH, V; Thetford, H, N, V. WINDSOR 
co.: Hartland, G, H, N, NH, v; Norwich, NH; Springfield, N, v; 
Weathersfield, N, v., Windsor, WM. WINDHAM CO.: Brattleboro, 
N, V; Guilford, v, wM; Newfane, N, NH, V; Vernon, N, NH, V. 

New Hampshire: GRAFTON CO.: Hanover, H, N, v; Haverhill, H, N, 
NH. V; Lebanon, H, N; Orford, NH. SULLIVAN CO.: Charlestown, 


40 Rhodora [Vol. 75 


N, V; Claremont, WM; Plainfield, NH. CHESHIRE CO.: Alstead, N, V; 
Chesterfield, N, v; Hinsdale, H, N, NH, S, V; Keene, N, NH, V; Marl- 
boro, H, N, NH, V; Surry, NH; Swanzey, H, N, NH, V; Walpole, G, 
H, N, NH, V; Westmoreland, N, NH, V; Winchester, H, N, NH, V. 
STRAFFORD CO.: Durham, NH; Lee, NH. ROCKINGHAM CO.: Exeter, 
N; Kensington, NH; Newmarket, NH; South Hampton, NH. HILLS- 
BORO CO.: Hollis, NH. 

Massachusetts: ESSEX CO.: Andover, G, N; Ipswich, N. MIDDLESEX 
co.: Belmont, N; Cambridge, N; Medford, N; Winchester, N. SUFFOLK 
CO.: Revere, N. NORFOLK CO.: Bellingham, N; Brookline, G, N; Ded- 
ham, G, N; Milton, N; Norfolk, N, s; Quincy, N. BRISTOL CO.: Fall 
River, N; Swansea, N; Westport, N. PLYMOUTH C0.: Hingham, N. 
WORCESTER CO.: Boylston, MS; Leicester, N; Millbury, €, N; Paxton, €; 
Princeton, C, N; Spencer, WM; Worcester, C, N. FRANKLIN CO.: Buck- 
land, WM; Montague, N; Sunderland, N, v; Whately, N. HAMPSHIRE 
co.: Easthampton, s; Hadley, WM; Hatfield, wM; Huntington, WM; 
Northampton, WM; Westhampton, WM; Williamsburg, S. HAMPDEN 
co.: Chicopee, wM; Holyoke, s; Springfield, WM. BERKSHIRE CO.: 
Lanesboro, N; Sheffield, G, N; West Stockbridge, N. 

Maps 2 and 6 show the presently known distribution of 
the bitternut in New England. This may be expressed 
verbally as: southeastern New Hampshire (to southern 
Strafford Co.), western New Hampshire and eastern Ver- 
mont in the Connecticut River Valley (to Haverhill, N.H. 
and Peacham, Vt.), western Vermont (north to Burling- 
ton), Massachusetts (except for most of Plymouth Co., 
Cape Cod, Martha’s Vineyard, Nantucket, Berkshire Hills), 
Rhode Island(?) and Connecticut. It is absent in Maine, 
in central Vermont and central New Hampshire. Seymour 
(1969) reports the species from the towns of Cumberland 
and Tiverton in Rhode Island; however, R. L. Hauke, E. A. 
Palmatier and R. L. Champlin (personal communications) 
state that the tree does not seem to be native in the state, 
though it is escaping from cultivation in Kingston. 


3. C. ovata (Mill. K. Koch, shagbark hickory. 


Most maps and descriptions of ranges are essentially 
correct for this species. Some of these locate the shagbark 


Map 2. Distribution of Carya cordiformis in northern and central 
New England, limits indicated by heavy solid line. A dot designates 
one or more specimens collected in the town located there. 


41 


Hickories — Manning 


73] 


42 Rhodora [Vol. 75 


in northern and central parts of New Hampshire and Ver- 
mont or perhaps all of these states, but again these dis- 
tributions may merely be highly generalized. Seymour 
(1969) and Little (1965) record the tree on Cape Cod. I 
have seen the specimen involved and it is C. ovalis. De- 
spite the absence of the shagbark on Cape Cod, it occurs 
on Nantucket Island. There is a possibility that the trees 
there are planted, but Maria Owen (1888) reported the 
species as there a long time ago. Mackeever (1968) does 
not mention the shagbark on the island, though there is 
a specimen at the N. Y. Botanical Garden. I have seen only 
one specimen of C. ovata in Berkshire Co., but Hoffman 
(1922) reports the species as, “rich woods, open hillsides, 
common in the valley". My studies show that the shagbark 
occurs slightly further north in New Hampshire and Ver- 
mont than indicated by Little. The northernmost record 
for New Hampshire, Groton, in Grafton Co., is based on 
the observation of Dr. Robert Zimmerman when he was 
making an intensive study of the northern distribution of 
the oaks; he did not collect specimens of the hickory. 

As I have observed them, the trees are quite rare toward 
the north in the Connecticut River Valley, often repre- 
sented by one to few trees in one locality for a whole town; 
thus it is very easy to miss a record for a town. 

Vermont: CHITTENDEN CO.: Burlington, N, v; Charlotte, G; Col- 
chester, v; Milton, N; Shelburne, N; South Burlington, v; Williston, 
H. ADDISON CO.: Cornwall, v; Ferrisburg, G, N; Middlebury, v. RUT- 
LAND C0.: Brandon, H; Castleton, H, N, NH, V; Fair Haven, G; Ira, 
H, N, NH, V; Poultney, H, N, NH, v; Rutland, n; Sudbury, N; Wells, 
H, N, NH, V; West Haven, V. BENNINGTON CO.: Arlington, WM; 
Pownal, N, NH, V; Shaftsbury, WM; Sunderland, WM. WINDSOR CO.: 
Springfield, wM; Weathersfield, Z; Woodstock, H. WINDHAM CO.: 
Brattleboro, N; Guilford, wM; Newfane, H, NH, N, V; Townshend, N; 
Vernon, WM. 

New Hampshire: GRAFTON CO.: Groton, Z. SULLIVAN CO.: Charles- 
town, NH, WM. Claremont, WM. CHESHIRE CO.: Chesterfield, WM; 
Hinsdale, H, N, NH, S, V; Keene, H, N, NH, V; Marlboro, H, N, NH, V; 
Richmond, N, WM; Swanzey, H, N, NH, S, V; Westmoreland, H, N, 
NH, V; Winchester, G, N, NH, V. CARROLL CO.: Wolfeboro, NH. BEL- 
KNAP CO.: Alton, NH; Gilmanton, NH; Meredith, N; Sanbornton, N. 
STRAFFORD CO.: Dover, NH; Durham, NH; Farmington, NH; Lee, N; 


1973] Hickories — Manning 43 


Middleton, NH; Milton, NH; Rochester, N, NH. MERRIMACK CO.: 
Warner, N, NH. ROCKINGHAM CO.: Atkinson, N; Derry, N, NH; 
Fremont, N; Hampton Falls, N; Kensington, N; Nottingham, N; 
Stratham, N. HILLSBORO CO.: Amherst, N; Hancock, H, N, NH, V; 
Hillsboro, N; Mason, G, H, N; Merrimack, G, N. 

Maine: OXFORD CO.: Brownfield, MN; Paris, G, MN; Rumford, MN. 
CUMBERLAND CO.: Brunswick, MN; Cape Elizabeth, N, NH; Cumber- 
land, G, N, NH, S; North Yarmouth, N, NH; Westbrook, G, NH. YORK 
co.: Buxton, N; Eliot, N; Kittery, G, MN, N; Limington, N; North 
Berwick, G, N; Old Orchard, N; Saco, N; Sanford, MN; South Ber- 
wick, MN, N; Wells, N; York, G, MN, N. ANDROSCOGGIN CO.: Liver- 
more Falls, N; Turner, MN, N. SAGADAHOC CO.: Arrowsic, MN, NH; 
Bowdoinham, MN; West Bath, N; Woolwich, MN. SOMERSET CO.: 
planted?: Solon Village, MN; St. Albans, MN. KENNEBEC CO.: 
planted?: Augusta-Chelsea town line, MN; Clinton, N; Fayette, NH; 
Manchester, N; Winthrop, MN. 

Massachusetts: ESSEX CO.: Andover, G, N; Essex, N; Haverhill, N; 
Ipswich, G, MS; Methuen, N. SUFFOLK CO.: Boston, G, N; Revere, G, N. 
MIDDLESEX CO.: Ashby, N; Belmont, G; Cambridge, N; Concord, N; 
Groton, N; Malden, N; Medford, N; Pepperell, N; Tewksbury, N; 
Townsend, N; Wakefield, N; Woburn, N. NORFOLK CO.: Canton, N; 
Milton, N; Needham, N; Norfolk, s; Norwood, G; Wellesley, G. 
BRISTOL CO.: Dighton, N. PLYMOUTH CO.: Kingston, WM, NANTUCKET 
co.: Nantucket Id., N. WORCESTER CO.: Auburn, C; Barre, C, MS; 
Boylston, c; Charlton, c, N; Dudley, c; Hardwick, c, Ms; Harvard, 
c; Holden, c, N; Lancaster, c; Leicester, c; Leominster, N; Lunen- 
burg, N; Millville, c; New Braintree, c; Northboro, c; Northbridge, 
c; North Brookfield, c; Oakham, c; Oxford, c; Petersham, c; 
Princeton, c, N; Royalston, N; Rutland, c, N; Shrewsbury, C, N; 
Southbridge, c; Spencer, c, N; Sutton, c; Upton, c; Westboro, c, N; 
West Boylston, C; Worcester, C, MS. FRANKLIN C0.: Buckland, WM; 
Charlton, Ms, N; Deerfield, N, v; Gill, wM; Greenfield, wM; Montague, 
MS, N, V; New Salem, N; Northfield, wM; Orange, MS, WM; Shelburne, 
N; Sunderland, MS. HAMPSHIRE CO.: Amherst, MS, N; Belchertown, 
MS; Easthampton, wM; Enfield, Ms; Granby, Ms: Hadley, WM; 
Hatfield, Ms; Northampton, G, MS, N, S, v; South Hadley, s; Wil- 
liamsburg, S. HAMPDEN CO.: Granville, G, Ms; Holyoke, G, MS; 
Russell, N; Springfield, N. BERKSHIRE CO.: Sheffield, G, Ms. 

The records in Kennebec and Somerset counties, Maine 
are from the University of Maine herbarium, though 
Hyland ond Steinmetz (1944) state that the shagbark is 
locally planted in the following counties: s. Penobscot, 
Piscataquis, s. Somerset, c. Oxford, sc. Hancock, ec. Waldo, 


nw. Knox, e. Kennebec. 


44 Rhodora [Vol. 75 


Maps 3 and 7 show the presently known distribution of 
the shagbark in New England. This may be expressed 
verbally as: southwestern Maine (as far as Sagadahoc and 
southern Oxford counties) southeastern New Hampshire 
(as far as southeastern Grafton Co.), along the Connecticut 
River Valley in southwestern New Hampshire and south- 
eastern Vermont (as far as southern Sullivan Co., N.H. 
and Windsor Co., Vt.), western Vermont (as far north as 
Burlington), most of Massachusetts (except Cape Cod, 
Martha's Vineyard, and Berkshire Hills), Connecticut and 
Rhode Island. The shagbark is absent in most of central 
New Hampshire, central Vermont; it is found on Nan- 
tucket Island, Mass. 


4. C. glabra-ovalis complex, pignut hickories [C. glabra 
(Mill. Sweet, C. ovalis (Wang.) Sarg.] 

Carya glabra and C. ovalis have been treated by some 
authors (Sargent, 1933, Fernald, 1950, Gleason 1952, Sey- 
mour, 1969, 1970) as separate species, by others (Robin- 
son and Fernald, 1908, Little, 1953, 1969) as a single 
species, C. glabra. In any case there is tremendous varia- 
tion in bark, fruit shape, leaves. Thus Sargent, 1933, and 
especially Steyermark, 1963, not only recognize both species 
but also a number of varieties and forms under each of the 
two, based primarily on shape of the fruit; there are also 
pignuts with hairy leaves. At the present time I am 
recognizing two species, realizing that there seem to be 
intergradations (hybridization?) in certain features in 
some localities. I have not decided which varieties to 
recognize. The herbarium specimens of New England 
pignut hickories are extremely unsatisfactory: often only 
flowering material, or sterile early summer material] or 
poorly collected material without notes on bark. As already 
stated (Manning, 1950) one needs all possible features for 
separation of the species, typical leaves (early and late 
summer condition), true terminal buds in fall condition, 
mature fruit (preferably after fruit has fallen on the 
ground), and notes on bark. It is sometimes possible to 


1973] Hickories — Manning 45 


——————— —-* 


t 
jas» 
1 Dor 
I EAT a | 
Jet `~ 


^ 
$ 
1 


\ 


Map. 3. Distribution of Carya ovata in northern and central New 
England, limits indicated by heavy solid line. A dot designates one 
or more specimens collected in the town located there. The half dots 
in northern Maine designate towns where specimens have been 
collected but the trees were probably planted, not native. 


46 Rhodora [Vol. 75 


name well collected specimens in summer, sterile or fruit- 
ing, but in other cases it is nearly impossible. 


Considering the condition of the specimens available at 
the present time, for this study I am using the term Carya 
glabra-ovalis complex in order to give a general view of 
the distribution. Futhermore, in a great many cases the 
ranges seem to overlap, so this distribution may represent 
the distribution of each species. In the list of towns I have 
indicated by “g” and “o” the name given on the herbarium 
label, but this does not always have significance. I will 
use the term pignut for both. 


There is a wide variation in books with respect to the 
distribution of the pignuts in New England, whether the 
treatments represent one or both species. Many of the 
older books report pignuts as rather common in eastern 
and western Vermont. Most of these records are based on 
incorrect identification. Little’s (1965, 1971) maps are 
much more accurate reducing the localities to 2 in western 
and one in southeastern Vermont. However, I have travelled 
extensively through southeastern Vermont and have failed 
to find any pignuts there. I have found pignuts close by 
on 2 hills in southwestern New Hampshire and on one 
in northern Massachusetts (north of Bernardston) just 
2 miles south of the Vermont border. Little's maps do 
not show the pignuts found rather recently by Hodgdon 
in southeastern New Hampshire, nor those which occur on 
Martha's Vineyard and Nantucket Island. 


In western New Hampshire and all of Vermont, pignuts 
are located on only 4 isolated rather low hills, in general 
between 600 and 1000 feet elevation. The trees do not 
seem to occur on longer ridges or at higher elevation or 
in the valleys. On at least 2 of these hills, Meetinghouse 
Hill, Winchester, N.H. and Coon Hill near Lake Bomoseen, 
Rutland Co., Vt., the trees are very numerous near the top. 
On a third, Television Station Hill, Winchester, N.H., the 
few large trees near the top seem to be dying out. Very 
few hills of the numerous hills in the two states have been 


1973] Hickories — Manning š 47 


Map 4. Distribution of Carya glabra-ovalis complex in northern 
and central New England, limits indicated by heavy solid line. A dot 
designates one or more specimens collected in the town located there. 


48 Rhodora [Vol. 75 


Maps 5, 6. Distribution in New England and adjacent areas of 
Carya tomentosa and C. cordiformis respectively according to Little 
(1965, 1971), his limits indicated by a dotted line where they differ 
from mine, my corrections indicated by a heavy solid line. 


climbed in an attempt to locate hickories, and it is very 
possible that other hills with pignuts will be found. 


Vermont: RUTLAND CO.: Castleton, g, o: G, H, MS, N, NH, V, WM. 
BENNINGTON CO.: Pownal, g: G; o: H, MS, N, NH, V, WM. 


New Hampshire: CHESHIRE CO.: Winchester, g: N, WM; 0: N, V, 
WM. STRAFFORD CO.: Barrington, o: N, NH; Durham, 0: N, NH; 
Lee, o: N, NH. ROCKINGHAM CO.: Atkinson, o: N, NH; Derry, o: 
N, NH; Hampstead, g: NH; New Market, g: NH; Raymond, o: N, 
NH; Salem, g: NH; South Hampton, g: NH; Windham, g: NH. HILLS- 
BORO CO.: Pelham, g, 0: N, NH. 


Massachusetts: ESSEX C0.: Andover, g: N; Boxford, o: N; 
Ipswich, g: N; Lynn, g: WM; Manchester, g: N; North Andover, 
o: NH; Saugus, o: N. SUFFOLK CO.: Boston, g, o: G, N; Revere, 
g: N. MIDDLESEX CO.: Acton, o: WM; Arlington, o: N; Ashby, g: 


1973] Hickories — Manning 49 


Maps 7, 8. Distribution in New England and adjacent areas of 
Carya ovata and C. glabra-ovalis complex respectively according to 
Little (1965, 1971), his limits indicated by a dotted line where they 
differ from mine, my corrections indicated by a heavy solid line. 


WM; Belmont, g: N; Boxboro, g: N; Cambridge, g, o: N; Concord, 
g, o: N; Framingham, o: N; Lexington, g, o: G, N; Lincoln, g, o: 
N, WM; Medford, g, o: N; Melrose, g: N; Natick, g: N; Newton, 
o: N; Pepperell, o: NH; Reading, g: N; Stoneham, g: N; Wake- 
field, g: N; Waltham, o: N; Woburn, g, o: N. PLYMOUTH CO.: 
Hingham, o: N; Middleboro, g: N; Norwell, g: N. NORFOLK CO.: 
Canton, g, o: G, N; Cohasset, g, o: G, N; Medfield, o: N; Milton, 
g, 0: G, N, S; Needham, g: N; Norfolk, g, o: N, S; Norwood, g: G; 
Walpole, g: G; Wellesley, g: G; Weymouth, o: N. BARNSTABLE CO.: 
Barnstable, g, o: N, WM; Brewster, g, o: G, N; Chatham, o: N; 
Falmouth, o: N; Mashpee, g: WM; Sandwich, g: WM; Yarmouth, 
o: N. DUKES CO.: Martha’s Vineyard: Edgartown, o: N; West 
Tisbury, g: G, N. NANTUCKET CO.: Nantucket Id., o: N. BRISTOL 
co.: Easton, g, o: G; New Bedford, g: N; Swansea, o: N; Taunton, 
g: N; Westport, o: N. WORCESTER CO.: C. glabra: Bolton, C, N; 
Charlton, N; Hardwick, c; Lancaster, G; Leominster, c; Lunenburg, 


50 Rhodora [Vol. 75 


N; Mendon, c; Milford, c; Millbury, c; Sutton, c; Worcester, C, N. 
C. ovalis: Auburn, c; Barre, WM; Berlin, c; Blackstone, c; Boylston, 
c; Brookfield, c; Douglas, c; Holden, c; Harvard, C; Leicester, C; 
Lunenburg, N; New Braintree, C; Northboro, c; North Brookfield, C; 
Millville, c; Oakham, s, wM; Oxford, c; Paxton, c; Petersham, 
€, MS, WM; Shrewsbury, c; Southboro, C, N; Sterling, C, N; Sutton, 
c; Upton, €; Uxbridge, c; Webster, c; Westboro, C; West Boylston, 
c; West Brookfield, c; Worcester, G. FRANKLIN CO.: Bernardston, 
o: MS, N, V; Deerfield, g, o: G, N, V; Erving, g: N; Gill, o: N, V, 
WM; Greenfield, g, o: MS, N, S, V; Leverett, g: WM; Montague, o: 
N, V; Northfield, g: N, v; New Salem, o: N; Sunderland, g, o: 
MS, N, WM; Whately, g: N. HAMPSHIRE CO.: Amherst, o: WM; 
Belchertown, o: M, WM; Easthampton, o: S, WM; Granby, g: S; 
Hadley, g, o: S, WM; Hatfield, o: WM; Northampton, g. 0: A, G, 
MS, WM, V; South Hadley, o: WM; Westhampton, g: S, WM. HAMP- 
DEN CO.: Chicopee, g: N; Granville, g: G, MS, N; Holyoke, g. o: 
s, WM; Ludlow, g: MS; Southwick, g, o: N; Springfield, g, o: MS, 
N; Wales, o: N; Wilbraham, o: MS. BERKSHIRE CO.: Sheffield, g, o: 
G, H, N. 


Maps 4 and 8 show the presently known distribution of 
the pignuts in New England. This may be expressed ver- 
bally as southeastern New Hampshire, extreme southwestern 
New Hampshire, west central and extreme southwestern 
Vermont, most of Massachusetts (except Berkshire Hills), 
Rhode Island and Connecticut. It is not reported from 
Maine, eastern Vermont, nor most of New Hampshire and 
Vermont. 


LITERATURE CITED 


BEAN, R. C., C. H. KNOWLTON, A. F. HiLL. 1956. Eleventh report 
of the committee on plant distribution. Rhodora 58: 125-134. 
BLAKESLEE, A. F. and C. D. Jarvis. 1911. New England trees in 
winter. Storrs, Conn, 

Burns, C. P. and C. H. Oris. 1916. The trees of Vermont. Vt. 
Agr. Exp. Sta. Bull. 194, Burlington, Vt. 

DoLE, E. J., ed. 1937. The flora of Vermont, 3rd rev. ed. Burling- 
ton, Vt. 

FERNALD, M. L. 1950. Gray's Manual of botany, 8th ed. New York. 

GLEASON, H. A. 1952. The new Britton and Brown illustrated 
flora of the northeastern United States and adjacent Canada. 
New York. 


1973] Hickories — Manning 51 


HARLow, W. M. and E. S. HARRAR. 1958. Textbook of dendrology, 
4th ed. New York. 


HOFFMAN, R. 1922. Flora of Berkshire Co., Mass. Proc. Boston 
Soc. Nat'l Hist. 36: 171-382. 


HouaH, R. B. 1907. Handbook of the trees of northern states and 
Canada. New York. 


HYLAND, F. and F. H. STEINMETZ. 1944. Woody plants of Maine. 
Orono, Me. 


JAQUES, H. E. 1941. How to know the trees. Mt. Pleasant, Iowa. 

LANJOUW, J. and F. A. STAFLEU. 1964. Index Herbariorum, part 1. 
Herbaria of the world, 5th ed. Regnum Vegetabile 31. Utrecht, 
Netherlands. 

LITTLE, E. L., JR. 1953. Check list of native and naturalized trees 


of the United States. Agriculture Handbook no. 41. Washington, 
J.C. 


1965. Distribution range maps, in FOWELLS, 
H. A., Sylvies of forest trees of the United States. Agr. Hand- 
book no. 271. Washington, D.C. 

1969. Two varietal transfers in Carya (hick- 
ory). Phytologia 19: 186-190. 

1971. Atlas of United States trees. Vol. 1. 
Conifers and important hardwoods. U.S.D.A. Forest Service, 
Misc. Publ. no. 1146. 


MACKEEVER, F. C. 1968. Native and naturalized plants of Nan- 
tucket. Amherst, Mass. 

MANNING, W. E. 1950. A key to the hickories north of Virginia 
with notes on the two pignuts, Carya glabra and C. ovalis. 
Rhodora 52: 188-199. 

OwEN, M. L. 1888. A catalogue of plants growing without culti- 
vation in the county of Nantucket, Mass, Northampton, Mass. 

RoBINSON, B. L. and M. L. FERNALD. 1908. Gray’s new manual of 
botany. "7th ed. New York. 

SARGENT, C. S. 1933. Manual of the trees of North America. 2nd 
ed., rev. Boston. 

SEYMOUR, F. C. 1969. The flora of New England. Rutland, Vt. 

1970. The flora of Vermont. Burlington, Vt. 

STEYERMARK, J. A. 1963. Flora of Missouri. Ames, Iowa. 


BOTANICAL LABORATORY 
BUCKNELL UNIVERSITY 
LEWISBURG, PENNSYLVANIA 17837 


BENTHIC ALGAE AND VASCULAR PLANTS 
OF THE LOWER MERRIMACK RIVER AND 
ADJACENT SHORELINE 


ARTHUR C. MATHIESON AND RICHARD A. FRALICK! 


Introduction 


The present investigation was initiated as a part of a 
broad ecological study of the Merrimack River Estuary in 
Massachusetts (Miller, et al., 1971). The project was con- 
ducted by Normandeau Associates for the U. S. Army 
Corps of Engineers in order to predict the biological effects 
of a possible fresh water diversion on the ecology of the 
Estuary. Jerome, et al. (1965) have evaluated the marine 
resources of the Estuary, but they have conducted only 
limited botanical studies. In the present account we sum- 
marize the composition and distribution of seaweeds and 
vascular plants from the Merrimack River Estuary. A 
comparison with the algal species diversity in the Hampton- 
Seabrook and the Great Bay Estuary Systems of New 
Hampshire is also given. 

Collections and observations of intertidal algae and vas- 
cular plants were made at 19 stations in the Merrimack 
River Estuary during the summer and fall of 1971 
(Fig. 1). Vascular plants were studied at 15 stations 
(Table I), while algae were studied at 13 locations 
(Table II). Representative specimens of plants from each 
site were collected and have been deposited as voucher 
specimens in the Herbarium of the University of New 
Hampshire (NHA). The type and quantity of substrate 
available for benthic plants at each site were noted. The 
nomenclature of the Second Revised British Checklist 
(Parke and Dixon, 1968) was applied for most taxa of 
seaweeds, while the Eighth Edition of Gray's Manual 
(Fernald, 1950) was employed for vascular plants. 


‘Published with the approval of the Director of the New Hamp- 
shire Agricultural Experiment Station as Scientific Contribution 
Number 635. 


52 


1973] Merrimack Plants — Mathieson & Fralick 53 


The breakwater at station 1 provides the maximum 
amount of stable substrate for benthie organisms. The 
intertidal areas at the remaining stations are composed 
primarily of scattered rock outcrops, boulders, pebbles and 
junk, interspersed with sand or mud. The largest amount 
of solid rock is usually evident in the upper intertidal zone 
and the substrate tends to grade into sand-mud in the 
lower shore. From the mouth of the river to station 11 
there is a reduction in the amount of rock (particularly 
large outcrops) and a progressive increase in the deposi- 
tion of mud on the shore. A buildup of extensive peat-like 
material is evident in the upper intertidal zone at many 
stations throughout the estuary, where the roots of Spar- 
tina spp. stabilize muddy surfaces and allow colonization 
of seaweeds, vascular plants and invertebrates. 


Miller, et al. (1971) summarized the salinity distribu- 
tion in the Merrimack River Estuary. A typical gradient 
is evident, with maximum salinities of 30 o/oo or higher 
on the open coast at Salisbury Beach and fresh water up- 
stream near stations 18 and 19. Brackish waters are found 
between the two extremes. Salinity encroachments at open 
coastal levels occur from 2 to about 5 miles upstream, 
depending upon the season. At high tide the limits of salt 
intrusion vary from 4.3 to 10.9 miles from the mouth; they 
range from 3.1 to 6.8 miles at low tide. Gross fluctuations 
of temperature and salinity are evident (both daily and 
seasonally) at any site, particularly in the intermediate 
brackish water areas. 


Vascular Plants 


Thirty-one taxa of vascular plants were found in the 
marshy habitats of the Merrimack River Estuary 
(Table I). A fairly uniform distribution of salt-marsh 
plants was apparent from stations 4 to 12. Species con- 
sistently present included Solidago sempervirens, Spartina 
alterniflora, S. patens, Salicornia europaea, Atriplex patula 
and Limonium carolinianum. Station 13 was characterized 


54 Rhodora [Vol. 75 


by a diminished salt-marsh flora. New associations were 
present at stations 16 and 17 in the form of Scirpus validus, 
Scrirpus maritimus, Acorus calamus and Zizania aquatica. 
These four species are essentially brackish to freshwater 
inhabitants and represent a marked change in association 
away from some of the more persistent halophytes such as 
S. alterniflora, S. patens, Salicornia europaea and Solidago 
sempervirens. A detailed description of species composi- 
tion at each station investigated is presented in Table I. 


Intertidal Algae 


A total of 31 taxa of benthic algae was collected 
(Table II). Twelve Chlorophyceae (green algae), eleven 
Phaeophyceae (brown algae) and five Rhodophyceae (red 
algae) were identified. A detailed evaluation of the Cyan- 
ophyceae (blue-green algae), Xantophyceae (yellow-green 
algae) and Bacillariophyceae (diatoms) was beyond the 


FIGURE 1 


Intertidal Collecting Stations In the Merrimack River Estuary 
1. Open ocean side of the breakwater at Salisbury Beach 
2. Estuarine side of the breakwater at Salisbury Beach 
3. Breakwater near Badgers Rocks 
4. Western bank of Plum Island River underneath the bridge con- 
necting Plum Island 
5. Black Rock Point 
6. Morrill Creek 
7. Rocks just upriver from Coffin Point 
8. Waterfront at Newburyport just west of the power generating 
station 
9. Twin Rocks, on the Salisbury side of the river 
10. North End Boat Club 
11. Rock promontory on Ram Island 
12. In the main channel on a rocky promontory of Carr Island 
13. North side of the river across from Eagle Island 
14. Rock promontory at Salisbury Point 
15. On the north shore just upriver of the factories 
16. On the north shore just upriver of the Seahorse Marina 
17. On the south shore just down river of the Artichoke River 
18. On the south shore between the Artichoke and Indian Rivers 
19. South shore just upriver of the Groveland Bridge 


Merrimack Plants — Mathieson & Fralick 55 


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60 Rhodora [Vol. 75 


scope of the present investigation, although some data was 
collected. For example, the colonial diatom, Amphipleura 
rutilans, was a conspicuous component at stations 2, 5, 6 
and 9. Vaucheria sp. and various blue-green algae such as 
Lyngbya, Oscillatoria and Merismopedia formed a con- 
spicuous mat amongst Spartina roots at stations 5, 6, 9, 
10, 11, 13, 15 and 17. 


Details of species composition and distribution of sea- 
weeds are summarized in Table II. The maximum number 
of species was found at stations 1, 2 and 5; beyond sta- 
tion 5 there was a rapid and progressive reduction in 
species numbers. Red algae appeared to be least tolerant 
of reduced salinities. Three of the five species dropped out 
at station 2, and no red algae were found beyond station 7. 
Brown algae showed a wider distribution than red algae, 
but even so they were not found upstream of station 9, and 
their largest number of species was found at station 1. 
Green algae were the most cosmopolitan of the three major 
groups, with Enteromorpha erecta extending to the low 
salinities of station 15. The yellow-green alga, Vaucheria 
sp. and the green alga, E. erecta, were the most wide- 
spread of all seaweeds. Several blue-green algae are prob- 
ably equally tolerant to reduced salinities, but a lack of 
specific identifications precluded a precise evaluation. 


An inspection of Table II indicates that most seaweeds 
exhibit a cosmopolitan distribution, for they occur both 
on the open coast and within the estuary. Vaucheria sp., 
Enteromorpha groenlandica, Monostroma oxyspermum, 
Fucus vesiculosus var. spiralis, and Polysiphonia fibrillosa 
are considered to be truly estuarine, for they were never 
found on the open coast. Spongomorpha arcta, Porphyra 
leucosticta, and Ptilota serrata appear to be coastal forms, 
since they were not found within the mouth of the river. 


Discussion 


The variety and abundance of rock are major factors 
restricting growth and distribution of algae in the Merri- 


1973] Merrimack Plants — Mathieson & Fralick 61 


mack River Estuary. The breakwater at stations 1 and 2 
provided maximum stability and surface area for the 
growth of seaweeds, and the highest species numbers and 
biomass of algae were evident at these two sites. The 
reduced species and biomass (based on a qualitative assess- 
ment of all species) upstream of station 5 can be attributed, 
at least in part, to unsuitable substrate. Most rocks upriver 
of this station were mud covered, and it is obvious that 
films of mud and silt will inhibit the attachment and growth 
of many algal species. In addition, small cobbles and 
pebbles, characteristic of upriver stations, are unsuitable 
as substrate for many larger plants, because of their 
instability. Only crustose algae such as Hildenbrandia 
prototypus and Pseudendoclonium marinum were found on 
such rocks. Vaucheria sp., Enteromorpha spp., and various 
blue-green algae were the only forms collected on the 
muddy surfaces stabilized by the roots of Spartina alterni- 
flora and Spartina patens. The Spartina spp. play an im- 
portant role in the formation of substrate suitable for 
algal colonization. 

In contrast, the rocky substrate at stations 1 and 2 was 
not suitable for attachment and colonization of estuarine 
vascular plants, and progressive increases in numbers of 
estuarine vascular plants were observed in relation to a 
decrease in the amount of rocky substrate upstream. 
Therefore, an increase in biomass and species numbers of 
vascular plants upriver can be attributed to suitable sub- 
strate, including small rocks and fibrous peat. Maximum 
species numbers occurred at stations 5, 7 and 11. Beyond 
station 11 the reduction in species number (but not bio- 
mass) probably resulted more from sub-optimal hydro- 
graphic factors than from suitability of substrate. Scirpus 
validus, Scirpus maritimus, Acorus calamus, and Zizania 
aquatica accounted for nearly all the plant biomass at 
many of the latter stations. 

Spatial and temporal variations of hydrographic factors 
in the Merrimack River Estuary, particularly the low up- 
stream salinities, restrict the longitudinal distribution of 


62 Rhodora [Vol. 75 


many species (Mathieson and Burns, 1971; Mathieson, 
et al., in press). Algal species having limited tolerances 
to temperature and salinity changes would not be expected 
to migrate upstream for any distance. As suggested earlier, 
Spongomorpha arcta, Porphyra leucosticta, and Ptilota 
serrata have a distinctly coastal distribution, and they did 
not extend inland of station 1. Other species exhibited 
gradations of tolerances to temperature and salinity fluctu- 
ations within the estuary. The most tolerant ones exhibited 
the widest distributions (e.g., Enteromorpha erecta and 
Vaucheria sp.), while the less tolerant ones had limited 
estuarine distributions (e.g., Elachista fucicola and Peta- 
lonia fascia). For the algae the most conspicuous reduc- 
tion in species diversity occurred between stations 5 and 6; 
it was probably caused by the greater fluctuations of tem- 
perature and salinity and the limited amount of solid sub- 
strate. 

Pollution is an important limiting factor in algal distri- 
bution and abundance (North, et al., 1964; Patrick, 1964). 
A comparison of the species composition of seaweeds from 
the Merrimack River Estuary with that of the Hampton- 
Seabrook Estuary (Mathieson and Fralick, 1972) and 
the Great Bay Estuary Systems (Mathieson, Reynolds, and 
Hehre, in press) of New Hampshire indicates a paucity 
of total species and species numbers per station in the 
Merrimack. A total of 118 taxa of seaweeds was collected 
from the Hampton-Seabrook Estuary and the adjacent 
open coast, while over 150 species were found within the 
vicinity of the Great Bay Estuary System. The low algal 
species diversity (only 31 taxa) from the Merrimack River 
Estuary is probably also due to the extreme domestic and 
industrial pollution of this interstate river. Jerome et al. 
(1965) and Miller, et al. (1971) indicate that the Merri- 
mack is one of the most polluted rivers in New England. 

The concept of species diversity has been applied exten- 
sively in evaluating eutrophication of freshwater habitats. 
In general, a decrease in species diversity is a typical 
response to an increase in either domestic and/or industrial 


1973] Merrimack Plants — Mathieson & Fralick 63 


pollution (Patrick, 1964). Under polluted conditions, a 
few tolerant species tend to dominate in large numbers 
and high biomass. The abundance of many Ulotrichalean 
green algae such as, Enteromorpha spp., Ulva lactuca, and 
Monostroma sp. typifies a polluted estuarine habitat (Cot- 
ton, 1910; Fritsch, 1935). These species are not only 
tolerant of extremes in pollution, but of gross fluctuations 
in hydrographic factors. 

In summary the paucity of the algal flora may be attrib- 
uted to the following: (1) limited solid substrate for 
benthic species; (2) probably to the high degree of pollu- 
tion of the Merrimack River Estuary. The abundance of 
certain algae definitely seems to be related to the pollution 


factor. 


The authors wish to thank Dr. Donald Normandeau, of 
Normandeau Associates, who provided a boat, outboard 
motor and other support to complete the investigation. Dr. 
B. Miller, formerly with the same company, is also acknowl- 
edged for his assistance with the collection of specimens. 
We also express our appreciation to Dr. A. Hodgdon for 
his identification of several vascular plants. 


REFERENCES CITED 


CorTON, A. D. 1910. On the growth of Ulva latissima in water 
polluted by sewage. Bull. Misc. Inform. Roy Bot. Gard. Kew, 
pp. 15-19. 

FERNALD, M. L. 1960. Gray's Manual of Botany, 8th Edition, 
American Book Company, New York, 1632 pp. 

FRITSCH, F. E. 1935. The structure and reproduction of the 
algae. Vol. I, Cambridge Univ. Press, 791 pp. 

JEROME, W. C., A. P. CHESMORE, C. O. ANDERSON, JR., and F. GRICE, 
1965. A study of the marine resources of the Merrimack River 
Estuary. Monograph Ser. No. 1, Div. of Mar. Fish., Dept. of 
Natl. Res,, The Comm. of Mass., 90 pp. 

MATHIESON, A. C. and R. L. Burns. 1971. Ecological studies of 
economic red algae I. Photosynthesis and respiration of Chondrus 
crispus Stackhouse and Gigartina stellata (Stackhouse) Batters. 
J. Exp. Mar. Biol. Ecol. 7: 197-206. 

, and R. A. FRALICK. 1972. Investigations of New 

England marine algae. V. The algal vegetation of the Hampton- 


64 Rhodora [Vol. 75 


Seabrook Estuary and the open coast near Hampton, New Hamp- 

shire. Rhodora 74: 406-435. 

, N. B. REYNOLDS and E. HEHRE. Ibid. II. Distri- 
bution of benthonic marine algae in the Great Bay Estuary 
System. Nova Hedwigia (in press). 

MILLER, B., D. NoRMANDEAU, G. PIEHLER, P. HALL, A. MATHIESON, 
R. FRALICK, D. TURGEON, P. MAHONEY and W. OWEN. 1971. 
Ecological study Merrimack River Estuary — Massachusetts, 
Presented to the U. S. Army, Corps of Engineers by Normand- 
eau Associates, Inc. and Vast Inc., 342 pp. 

NoRTH, W. J., K. A. CLENDENNING, L. G. JONES, J. B. LACKEY, D. L. 
LEIGHTON, M. NEUSHUL, M. C. SARGEANT and H. L. SCOTTEN, 
1964. Investigation of the effects of discharged wastes on kelp, 
State Water Quality Control Bd., Publ. 26, 124 pp. 

PARKE, M. and P. S. Drxon. 1968. Check-list of marine algae — 
second revision. J. Mar. Biol. Assoc., U.K. 48: 783-832. 

PATRICK, R. 1964. A discussion of natural and abnormal diatom 
communities. In D. F. JACKSON (Ed.) Algae & Man, Plenum 
Press, pp. 185-204. 


DEPARTMENT OF BOTANY AND THE 
JACKSON ESTUARINE LABORATORY OF THE 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, NEW HAMPSHIRE 03284 


SOME MORPHOLOGICAL AIDS IN DISTINGUISHING 
NUPHAR MICROPHYLLUM 
FROM SIMILAR AQUATICS 


JOHONET C. WICKS 


This study is an outgrowth of my recent discovery that a 
specimen of Nuphar microphyllum (Pers.) Fern in the Uni- 
versity of New Hampshire Herbarium had been erroneous- 
ly labelled Nymphoides cordata (Ell.) Fern because of the 
apparently identical macroscopic appearance of the small 
floating leaves despite the presence of filmy submersed foli- 
age characteristic of Nuphar. This specimen, collected in 
Adams Pond in Pittsfield, N. H., constitutes a first record 
for Merrimack County and, more significantly, represents 
the most southern collection point in the state known at 
the present time. Nuphar microphyllum occurs generally 
in Maine and in Massachusetts, and to a more limited ex- 
tent in Vermont and Connecticut, but has been reported in 
New Hampshire only in Coós and Carroll counties accord- 
ing to F. C. Seymour (1969). Research at the Gray Her- 
barium yielded two specimens from the same locality in 
Hanover in Grafton County in the early 1900’s; and, more 
recently, C. Barre Hellquist has added another Grafton 
County station at Lily Pond in Livermore. Perhaps New 
Hampshire ponds have not been as thoroughly botanized as 
have those in Maine, Vermont and Massachusetts. Future 
investigation may disclose the fact that N. microphyllum 
occurs more frequently, if not much more abundantly, in 
the state than has heretofore been suspected. 

The pH and alkalinity of ponds sometimes affect the dis- 
tribution of species. Professor Hellquist kindly furnished 
readings taken by him at some ponds where Nuphar micro- 
phyllum occurs in Maine, Vermont and in Coés, Carroll and 
Grafton counties in New Hampshire. In New Hampshire 
the pH ranges from 6.0 to 6.7 whereas in Maine and Ver- 
mont, pH values are generally in the 7 range up to 7.9. 
The alkalinity readings for New Hampshire run from 2.5 


65 


66 Rhodora [Vol. 75 


ppm to 9.0 ppm of carbonates and bicarbonates but are 
considerably higher, 9.0 ppm to 60.0 ppm, in Vermont and 
Maine. The above limited data seem to indicate that N. 
microphyllum is adapted to a pH close to neutral and tol- 
erates a wide range of alkalinity. Perhaps the more acidic 
waters of New Hampshire ponds and the lower concentra- 
tion of carbonates and bicarbonates do not favor growth of 
this species. 

Seymour lists not only Coós and Carroll counties but also 
Cheshire County for Nuphar X rubrodiscum Morong, a 
fertile hybrid between N. variegatum Engelm. and N. 
microphyllum. In the light of the Grafton County records 
of N. microphyllum cited above and the Merrimack County 
record reported in this paper, it would seem plausible that 
hybridization could have occurred in these counties al- 
though no hybrids have as yet been reported. Examination 
of the Cheshire County specimens, one each in the Gray 
and New England Botanical Club herbaria from Gilmore 
Pond in Jaffrey in 1897, revealed them to truly be N. X 
rubrodiscum according to both the author and A. R. Hodg- 
don. Although it has not yet been reported, one would ex- 
pect to find N. microphyllum, one of the parents, in the 
county and possibly in the same pond. When and if it is 
discovered, it will constitute the southernmost record for 
the state. However, as the two Jaffrey specimens were 
collected prior to 1900, it is conceivable that the species 
could have since disappeared. 

It is sometimes difficult to differentiate the hybrid from 
either parent as the range of parental characteristics is a 
broad one in regard to such morphological features as the 
size of the leaf, the diameter of the petiole and the size of 
the flower. Figure 1A, a comparison of the minimum and 
maximum lengths of the floating leaves in the three species 
as indicated in current manual descriptions, shows a con- 
siderable overlap which makes identification troublesome, 
if not impossible, on the basis of leaf size alone. It is then 
advisable to examine the diameter of the petiole as shown 
in figure 1B. The petioles of Nuphar microphyllum leaves 


67 


Nuphar microphyllum — Wicks 


1973] 


‘OZIS [en3o* ITL suorsuəutIp [IV ““pydnN JO saemoy 
Jo Supr wnwtxew pue winuTUIU oAnveredun, ') -«mjdnw Jo 
se[orjed jo srojeurIp aAnjw1eduro) 'g -"4nudnw Jo S9AU9[ SULO ay} 
jo sqySue[ umturxeur pue urnunuru əaeaedwog v :[ əng 


g J | 
|] | UNEIA N 8 x 

: ul» (ys ¢ 

[- -j umosrpo4qna — "NI 3 wo 09" 
Lr mW. OF 1 

[H wunjjâqydomi u "NJ ga md Ot 


T 


V ~ 

WNJEBIIBA N up oc 

r up LO 1 

PUE DEN " 
w"nasipo4qna N a up Ot 

: up s 

' un[p£udog2rur.— "NJ ur» 90r 


68 Rhodora [Vol. 75 


are markedly filiform. Careful measurement of petioles of 
all three species in the New England Botanica] Club Her- 
barium indicated a significant overlapping of petiole diam- 
eters. A specimen with a leaf and petiole within the size 
range of both N. microphyllum and N. X rubrodiscum 
would still be puzzling. The final recourse then is to the 
flower or fruit. Figure 1C illustrates the comparative 
widths of the flowers of the three species when laid open, 
following dimensions given in Gray’s Manual. For the 
first time, there is no overlap between N. microphyllum 
and N. X rubrodiscum. Furthermore, the petals and sta- 
mens of the flower of N. microphyllum are promptly decid- 
uous whereas, in the other two species, they persist at the 
base of the young fruit. I examined the specimens of all 
three species in the New England Botanical Club Herba- 
rium and, with the approval of A. R. Hodgdon, reannotated 
four of them. Two were transferred from N. microphyl- 
lum to N. X rubrodiscum and two were changed from N. 
variegatum, one each to N. microphyllum and N. X vru- 
brodiscum. 

The genus Nuphar is characterized by dimorphic leaves : 
filmy, undulate submersed leaves and the well known firmer 
floating leaves. In his descriptions of the three species of 
Nuphar, Fernald (1950) mentions only N. microphyllum 
as possessing filmy submersed leaves. Fassett (1966) fails 
to note the presence of this foliage in any of the three spe- 
cies. It is of interest that Otto Brunfels in 1530 in his il- 
lustrated herbal, Herbarium Vivae Eicones, included a 
drawing of Nuphar luteum, a European species, depicting 
both types of foliage. 

The thin submersed leaves tend to disappear in Nuphar 
variegatum as the plant grows larger and older whereas 


they generally seem to persist in N. microphyllum (figure 
2A) and to a lesser degree in the hybrid, N. X rubrodis- 


cum. Perhaps their persistence in N. microphyllum ac- 
counts for their being mentioned by Fernald. These obser- 
vations are based on a study of the New England Botanical 
Club specimens: of 23 specimens of N. microphyllum with 


1973] Nuphar microphyllum — Wicks 69 


Figure 2: A. Nuphar microphyllum habit X 14. B, C, D. Nuphar 
variegatum, seedling stages X !4. 


70 Rhodora [Vol. 75 


basal parts, 19 have filmy submersed foliage; of 11 speci- 
mens of N. X rubrodiscum, T have filmy leaves; of eight 
folders of N. variegatum containing a great many speci- 
mens, only 3 sheets have this foliage. 

The seedlings of all three species have membranaceous 
foliage with filiform petioles (figure 2B, C, D). It would 
require careful observation to distinguish between the 
three species in the seedling stage. Determination of spe- 
cies could possibly be made on the basis of seedling size if 
the seeds of each species are of different sizes which has 
yet to be established. The size of seedlings, however, could 
be confounded by variability in age. One way to solve this 
problem would be to germinate seeds of each species and 
to observe differences in size of seedlings with age as a 
control factor. However this procedure, while of value 
under controlled conditions, would not shed much light on 
relative age and size in the field. Perhaps the location of 
a seedling could indicate its species although in mixed col- 
onies this criterion would be inadequate. While examining 
specimens of Nymphoides cordata in the New England 
Botanical Club Herbarium, I found a sheet of Nuphar 
seedlings, possibly those of N. microphyllum, labelled as 
seedlings of Nymphoides. 

Nuphar microphyllum, as shown in figure 2A, appears to 
be a composite of a Nuphar seedling with attached floating 
leaves of Nymphoides cordata. The close resemblance be- 
tween the floating leaves of N. microphyllum and those of 
Nymphoides in regard to both size and shape is startling 
and can lead to a confusion of the two species especially 
when underwater foliage is not present and when Nym- 
phoides lacks the characteristic cluster of thick roots on 
the petiole just below the surface of the water. Figure 3 
shows the similarity in shape between the floating leaves 
of the two species as well as the variability within each 
species. These are reduced leaf tracings from herbarium 
specimens. The shape and width of the sinus show re- 
markable variation. Some specimens of N. microphyllum 
exhibit a wide V-shaped sinus while others have a very 


1973] Nuphar microphyllum — Wicks 71 


UU 
JG 
UU 


Nymphoides cordata 


Be 


Sie 


Figure 3. Similarity and variability in leaf shape of the floating 
leaves of Nuphar microphyllum and Nymphoides cordata. 


12 Rhodora 


KD 


Figure 4: A. Nymphoides cordata, habit X 1⁄2. 


C. Young plant X !$. 


[Vol. 75 


B. Seedling X !$. 


1973] Nuphar microphyllum — Wicks ia 


narrow sinus with almost parallel sides. The shape of the 
sinus of the floating leaves is similar to that of the sub- 
mersed foliage on the same plant. 

Nymphoides cordata also possesses dimorphic leaves. In 
addition to the floating cordate leaves, the plant has smaller 
rhombic submersed leaves characteristic of the seedling 
stage and of the young plant (figure 4A). The illustration 
of the seedling (figure 4B) depicts the generation of a new 
plant from the rhizome whereas the drawing of the young 
plant (figure 4C) seems to indicate that a new individual 
can also arise by vegetative propagation from the detach- 
ment, submergence and subsequent rooting of a leaf and 
petiole with its group of thickened roots. The underwater 
leaves are absent in many, but not all, of the specimens 
studied. Their frequent absence in more mature plants 
may account for the fact that neither Fernald nor Fassett 
mentions these submersed leaves in their descriptions of 
the species. Drawings of Nymphoides in both Fassett’s 
book and in that of N. Hotchkiss (1967) have no basal 
parts, only the floating leaf and the short, spur-like thick- 
ened cluster of roots attached to the petiole at which point 
the flowers arise. 

In conclusion, it is sometimes difficult to distinguish 
Nuphar microphyllum from other similar aquatic plants. 
When the entire plant including the floating leaves, filmy 
submersed leaves, flower or fruit are present, it cannot be 
mistaken for Nymphoides cordata; if only the floating leaves 
and submersed foliage are present, it could be confused 
with N. X rubrodiscum; when only the floating leaves are 
evident, it could be taken for either N. X rubrodiscum or 
Nymphoides. This confusion points out the desirability of 
collecting as complete specimens as possible. 

The illustrations were prepared by the author from speci- 
mens in the University of New Hampshire Herbarium 
(NHA) from localities in Maine and New Hampshire. 

I greatly appreciate the privilege of using the New Eng- 
land Botanical Club and Gray herbaria without which this 
study would have been impossible. 


74 Rhodora [Vol. 75 
LITERATURE CITED 


Fassert, N. C. 1966. A Manual of Aquatic Plants. Madison: 
The University of Wisconsin Press. 

FERNALD, M. L. 1950. Gray’s Manual of Botany, Eighth Edition. 
New York: American Book Company. 

Horcukiss, N. 1967. Underwater and Floating-Leaved Plants of 
the United States and Canada. Washington, D.C.: Resource 
Publication 44. 

PEAsE, A. S. 1964. A Flora of Northern New Hampshire. Cam- 
bridge: New England Botanical Club, Inc. 

SEYMOUR, F. C. 1969. The Flora of New England. Rutland: The 
Charles E. Tuttle Company. 


DEPARTMENT OF BOTANY 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, N.H. 03824 


A RE-EVALUATION OF POTAMOGETON 
FIBRILLOSUS FERN. (POTAMOGETONACEAE)? 


ROBERT R. HAYNES AND JAMES L. REVEAL 


In 1932, Fernald proposed Potamogeton fibrillosus based 
on specimens collected in Wyoming, Idaho, Oregon and 
Washington. He noted that the new species closely resem- 
bled P. foliosus Raf. in its foliage, peduncles and small dor- 
sally keeled fruits. It was said to differ from P. foliosus by 
its stipules which disintegrate into *rope-like" fibers and 
are thus open or convolute, and by having fruits with a less 
developed keel and more nearly median beak. In his closing 
comment on the species, Fernald (1932; p. 52) stated his 
reservations in assigning specific rank to this new entity: 
“The plant will doubtless be found to have a broader range, 
when it may prove to be a marked geographic variety of the 
wide-spread P. foliosus." 

Evidence from our field work (Haynes as part of his doc- 
toral studies and Reveal as part of his review of Potamoge- 
ton for the Intermountain Flora), from examination of most 
of the cited specimens noted by Fernald (1932) and from 
a review of more recently gathered material from other lo- 
calities substantiates his closing remarks. Several collec- 
tions, especially from Yellowstone National Park of Wy- 
oming and from southeastern Oregon, are distinct from P. 
foliosus. In general, these specimens differ from P. foliosus 
not only in having the stipules disintegrating into fibres and 
fruits with poorly developed keels, but also in having cylin- 
dric, interrupted inflorescences and leaves with basal glands 
— the last two characters not mentioned by Fernald, but 
present even on the holotype. 

Other collections from the Yellowstone region are not so 
distinct. One such collection (Haynes 3849) has fruits, in- 
cluding the keel, which are identical with those of Potamo- 


"Contribution from the Botany Department, The Ohio State Univer- 
Sity (Paper No. 828). The work of the junior author was supported 
by National Science Foundation Grant GB-22645. 


15 


76 Rhodora [Vol. 75 


geton foliosus, but the inflorescence is capitate and the stip- 
ules only rarely disintegrate into fibers. However, basal 
glands are present. Flowering specimens from Albany Co., 
Wyoming, made by C. L. Porter (3473) are similar to P. 
fibrillosus in having basal glands and short, clavate pedun- 
cles. However, in this collection, the stipules only rarely 
disintegrate into fibers. Flowering specimens obtained by 
Maguire (21578) in northern Utah commonly produce fi- 
bers when the stipules decay, but basal glands are rarely 
found. Unpublished data of Haynes indicate, for other 
species of Potamogeton, that the degree of connation of the 
stipules is not a reliable taxonomie character and can not 
be used successfully in distinguishing species. 

Hitchcock (1969) regarded Potamogeton fibrillosus as a 
distinct species, but conceded its close relationship with P. 
foliosus. Porter (1963) reduced P. fibrillosus to synonymy, 
regarding it only as a local form of P. foliosus produced by 
the rather warm-water conditions of “geyser formations 
and hot springs" in the Yellowstone area. Porter stated 
(1963; p. 9) "The warm water hastens disintegration of 
the sheaths by bacterial action." This is not the case, how- 
ever, in northern Utah or southeastern Oregon. Here the 
water is cool, usually fresh (although slow-moving or in- 
frequently standing), and the stipules still break down. As 
we know of sites where both entities can be found in close 
proximity, and at the same degree of development, it seems 
unlikely that mere bacterial activity (or some other type ot 
mechanical breakdown) could account for the fibrous stip- 
ules. Nevertheless, because this entity, in most situations, 
can be distinguished quite easily from P. foliosus, we think 
it should be taxonomically recognized. However, since many 
intermediate forms can be detected, we can not justify the 
specific rank and therefore propose: Potamogeton foliosus 
Raf. var. fibrillosus (Fern.) Haynes & Reveal, stat. & comb. 

nov., based on P. fibrillosus Fern., Mem. Amer. Acad. 

Arts 17:51. 1932. — In warm spring, margin of Harney 

Valley, at “P” Ranch, Harney Co., Oregon, 22 Jun 1901, 

Cusick 2598. Holotype, GH! Isotypes, F, K, MO, ORE, us! 


1973] Potamogeton — Haynes & Reveal 77 


From var. foliosus, the var. fibrillosus may be distin- 
guished by its poorly developed keel, cylindric and inter- 
rupted inflorescence, fibrous stipules and basal glands pres- 
ent at the base of most leaves. 

Distribution. — In waters, often warm, of shallow lakes, 
springs, streams and rivers, from southwestern Washing- 
ton southward through southeastern Oregon, hence east- 
ward to southeastern Wyoming. Fruiting from mid-June 


to mid-September. 

Representative Specimens. — Oregon: HARNEY CO.: warm springs 
near Burns, 17 May 1927, Henderson 8867 (GH); in irrigating ditch 
at Frenchglen, 15 Jul 1927, Thompson 12068 (NY, WTU). MALHEUR 
co.: Otis Creek, 20 Jun 1896, Leiberg 2340 (GH, US). Utah: CACHE 
co.: slow stream in meadow, 3 mi ne of Logan, 21 May 1939, Maguire 
16685 (MO, NY, UTC); slow-moving stream in meadow, 2 mi nw of 
Logan, 23 Jun 1942, Maguire 21578 (F, GH, MO, NY, US, UTC). Wash- 
ington: PIERCE Co.: In outlet to Lake Spanaway, near Tacoma, 4 Aug 
1933, Thompson 9657 (GH, WTU). UNKNOWN LOCATION: Washington 
Territory, 1883, Brandegee 1127 (GH). Wyoming: ALBANY C0.: N. 
Fork of Poole Creek, 5 Aug 1944, C. L. Porter 3474 (NY, RM, US). YEL- 
LOWSTONE NATIONAL PARK [or Park Co.]: Firehole River, 24 J ul 1906, 
Jepson 2540 (GH); Midway Geyser Basin, Firehole River, in warm, 
swift-flowing water, 30 Aug 1971, Haynes 3850 (GH, MICH, MO, NY, 
os); Upper Geyser Basin, 1 Sep 1878, Richardson s.n. (GH); Upper 
Nez Perce Creek, in swift flowing water, 30 Aug 1971, Haynes 3849 
(MICH, OS). 


LITERATURE CITED 

FERNALD, M. L. 1932. The linear-leaved North American species of 
Potamogeton, section Axillares. Mem. Amer. Acad. Arts 17: 
1-183. 

Hitcucock, C. L. 1969. “Potamogetonaceae.” Im: Hitchcock, C. L. 
et al. “Vascular plants of the Pacific Northwest.” Univ. Wash. 
Publ. Biol. 17 (1): 158-175. 

Porter, C. L. 1963. “13. Najadaceae.” In: A flora of Wyoming. 
Part II. Wyoming Agric. Exp. Sta. Bull. 404: 6-12. 


COLLEGE OF BIOLOGICAL SCIENCES 
THE OHIO STATE UNIVERSITY 
COLUMBUS, OHIO 43210 

and 

DEPARTMENT OF BOTANY 
UNIVERSITY OF MARYLAND 
COLLEGE PARK, MARYLAND 20742 


THE AUDOUINELLA COMPLEX (RHODOPHYTA) 
IN THE WESTERN SARGASSO SEA 


WILLIAM J. WOELKERLING 


The macroscopic vegetation of the western Sargasso Sea 
is known mainly from four accounts (Harvey 1852, Boerge- 
sen 1914, Winge 1923, Parr 1939) which deal almost exclu- 
sively with the genus Sargassum. Except for two recent 
papers (Carpenter 1970, Woelkerling 1972b), only iso- 
lated records of other macroscopic or epibiotic algae from 
this region have appeared in the literature (Collins 1917, 
Conover and Sieburth 1964, Farlow 1914, Hentschell 1921, 
Prat 1935). These reports leave a number of points to be 
clarified including identification to species in most cases. 
To date only one reference (Conover and Sieburth 1964, 
p. 150) to an unidentified audouinelloid alga in this region 
has been published. 


The present study was initiated after an analysis of 
several samples of Sargassum from the western Sargasso 
Sea revealed the presence of numerous epiphytic audoui- 
nelloid plants. This paper incorporates data from six 
cruises to the Sargasso Sea by vessels of the Woods Hole 
Oceanographic Institution and includes a morphotaxonomic 
account of the audouinelloid algae found to date. 


MATERIALS AND METHODS 


Samples of Sargassum were gathered with the aid of a 
long-handled dip net, immediately preserved in 1:10 forma- 
lin in sea water, and brought back to shore for subsequent 
study. Vouchers of all collections have been prepared in 
the form of permanent microscopic slides (Woelkerling 
1970) or as liquid preserved material in 10:1 70% ethanol 
in glycerine. Herbarium numbers designated WJW are 
those in the author’s personal collections; other herbarium 
abbreviations follow Lanjouw and Stafleu (1964). Line 


78 


1973] Audouinella — Woelkerling 79 


drawings have been made with the aid of a Leitz drawing 
head microscope attachment. 

In all cases morphological data is based on plants found 
in the Sargasso Sea and represents, wherever possible, the 
results of analyses of populations rather than of isolated 
plants. 

The taxonomic proposals of Woelkerling (1971) have 
been adopted during this study, including the use of older 
generic names in cases of possible synonomy in order to 
avoid making new and unnecessary nomenclatural com- 
binations. In species descriptions, the designation L/D 
refers to the ratio of cell length to cell diameter. 


GENERAL OBSERVATIONS 


Of the six species found to date, Colaconema infestans 
(Howe et Hoyt) comb. nov. occurs in nearly all collections, 
and Audouinella daviesii (Dillwyn) Woelk., A. hallandica 
(Kylin) comb. nov., and A. thuretii (Bornet) Woelk. ap- 
pear fairly frequently as well. Audouinella microscopica 
(Naegeli) Woelk. and Calaconema scundata (Lyngbye) 
comb. nov., in contrast, have been encountered on only 
several occasions. 

Monosporangia occur in all species and tetrasporangia 
also have been found in A. thwretii; sex organs, however, 
are apparently wanting. The formation of reproductive 
bodies is in noteworthy contrast to the situation in Sargas- 
sum fluitans (Boergesen) Boergesen and S. natans (L) J. 
Meyen, the two most conspicuous algae of the Sargasso 
Sea, which apparently never become fertile (see, however, 
Parr 1939, p. 49). 

The audouinelloid algae probably represent a permanent 
component of the Sargasso Sea flora rather than being a 
temporary invading element (see Woelkerling 1972b) 
since they frequently epiphytize Sargassum fluitans and 
S. natans, which are apparently endemic to this region. 
However, all species found to date in the Sargasso Sea are 
also reported from Bermuda, the Caribbean, or the North 


80 Rhodora [Vol. 75 


American mainland (Boergeson 1915, 1924; Collins and 
Hervey 1917; Taylor 1960). It is likely that these land 
masses act as distribution centers; in fact several collec- 
tions examined during the present study have been attached 
to hosts (e.g. the grass Spartina) from inshore areas. 

In general, the morphotaxonomic conclusions reached by 
Woelkerling (1971) are supported by this investigation. 
Differences between species in which the prostrate system 
is dominant and those in which the erect system is domi- 
nant have become more apparent to the author as a result 
of the current investigation, and it appears that at least 
two distinct habits occur among species with multicllular 
prostrate systems: 1) a stoloniferous habit in which the 
prostrate filaments are more or less widely creeping and 
give rise to erect filaments at irregular intervals, and 2) 
a more or less caespitose habit in which the prostrate fila- 
ments are generally pseudoparenchymatously united into a 
disc or funiform mass which, in turn, gives rise to a 
number of erect filaments in close proximity to one an- 
other. In stoloniferous forms, the prostrate system usually 
predominates while in caespitose forms, the erect system 
usually predominates. 


KEY TO SPECIES 


l. Prostrate system normally absent; plants attached to substrate by 
a single basal cell which may rarely divide to form severa] acces- 
sory cells. 

2. Cells generally 5-104 long, more or less barrel-shaped, com- 
monly isodiametric or broader than long ........... ee 
— S eener eehnae Audouinella microscopica. 

2. Cells generally 10-304 long, more or less cylindrical, L/D 
usually more than 2 sssr Audouinella hallandica. 

1. Prostrate system present, filamentous or pseudoparenchymatous. 
3. Plants more or less stoloniform; prostrate system exceeding 

erect system in length; erect filaments rarely over 904 long 
and commonly unbranched .................. Colaconema infestans. 

3. Plants more or less caespitose; prostrate system not exceeding 
erect system in length; erect filaments commonly over 2004 
long and branched. 


1973] Audouinella — Woelkerling 81 


4. Sporangia, in part at least, grouped in clusters of 3-8 in 
axils of lateral branches ................... Audouinella, daviesii. 

4. Sporangia usually solitary or in pairs, not clustered in 
axils of lateral branches. 

5. Chromoplasts distinctly stellate; prostrate system at 
first a parenchyma-like group of cells; sporangia com- 
monly on short (3-5 celled) lateral branches... eene 
p e ee dert S E EPOR 5 S as pa Colaconema secundata. 


Chromoplasts parietal lobate; prostrate system fila- 
mentous in young stages; sporangia situated on longer 
lateral branches ............................. Audouinella thuretii. 
Audouinella daviesii (Naegeli) Woelkerling 1971: 28, Figs. 
TUE 
Acrochaetium daviesii (Dillwyn) Naegeli 1861: 405, 
Figs. 26-27. Boergesen 1924: 25, Fig. 8. 1827: 25, 
Fig. 15. Taylor 1960: 307. 
Callithamnion daviesii (Dillwyn) Lyngbye 1819: 129 
(only as to binomial). 
Ceramium daviesii (Dillwyn) C. Agardh 1817: XXVII. 
Chantransia daviesii (Dillwyn) Thuret in Le Jolis 1863: 
106. Kylin 1907: 117, Fig. 27. Rosenvinge 1909: 104, 
Fig. 34. 
Conferva daviesii Dillwyn 1809: 73, Suppl. pl. F. 
Rhodochorton daviesii (Dillwyn) Drew 1928: 172. Na- 
kamura 1944: 106, Fig. 5. 
Trentepohlia daviesii (Dillwyn) Areschoug 1847: 338. 
Note: Further synonomy is given by Woelkerling (1971, 
p. 28). 


Plants epiphytic or epizoic, caespitose, up to 5 mm. tall; 
original spore non-persistent. Prostrate system consisting 
of branched filaments more or less forming a pseudoparen- 
chymatous disc. Erect filaments freely and irregularly 
branched, commonly attenuate and ending in multicellular 
hair-like prolongations. Cells of main axes and laterals 
cylindrical, 8-15, wide and (15-) 20-60, long, L/D (1.75-) 
2-4(-5) ; cells sometimes tapering to 2-81 wide and 20-80, 
long (L/D up to 35) near the apices; each cell containing 
a parietal lobate chromoplast with one pyrenoid. 


or 


82 Rhodora [Vol. 75 


Monosporangia ovoid, 8-13, wide and 13-17 (-22) long, 
in clusters of 3-8 on branched stalks or singly or in pairs 
on 1-2 celled stalks, situated on the lowermost cells of 
laterals or sometimes more scattered. 

Other reproductive structures not observed in material 
examined. | 

Type Locality. — Bantry Bay, Ireland (Hutchens) ; lo- 

cality for H. Davies collection not given by Dillwyn 
(1809). 

Holotype. — NMW. 

Distribution. — Nearly cosmopolitan. 

Hosts. — Hydroids, Sargassum, and Spartina | ( Angio- 

spermae) fragment in Sargasso Sea; a wide variety 
of algae, marine angiosperms, and invertebrates else- 
where. 

Specimens examined. — Sargasso Sea: 26° 57'N-72° 58'W, 26.iv. 
1970, Moore (ww 2648); 28°N-70°W, 4.iii.1970, Volkmann (WJW 
2429) ; 31°N-69° 29'W, 3.iii.1970, Volkman (wJw 2380), (wJw 2364) ; 
32? 09'N-64? 58'W, 16.V.1970, Woelkerling (WIW 2667); 34°N-70°W, 
7.iii.1970, Volkmann (WJWw 2396), (wJw 2409); 37?N-70^W, 12.v. 
1970, Woelkerling (wyw 2621); 38° 22’N-70°58'W, 12.x.1970, Volk- 
mann (WIW 2888); 39° OT'N-10? 35'W, 16.viii.1970, Moore (WJW 
2930). Ireland: Bantry Bay, prior to 1809, Hutchins (NMW, Dillwyn 
Collection, type). 

Populations of A. daviesii examined during this study 
agree in all essentials with the type material and with 
plants described in the accounts of Rosenvinge (1909) and 
Woelkerling (1971). Multicellular hair-like prolongations 
with poorly developed plastids often devoid of pyrenoids 
occur very frequently in Sargasso Sea plants. 

Clustered monosporangia are common in most collections 
(2621 represents the only exception), but tetrasporangial 
and sexual individuals (see Woelkerling 1971 for accounts 
of these stages) have not been encountered. 


Audouinella hallandica (Kylin) comb. nov. Figs. 1-4. 
Acrochaetium hallandicum (Kylin) Hamel 1927: 20, 
Figs. 19-21; 1928: 114, Figs. 19-21. 


1973] Audouinella — Woelkerling 83 


Figs. 1-4. Audouinella hallandica (Kylin) comb. nov. Fig. 1. 
Monosporangial plant. Figs. 2-4. Variation in shape of basal cell. 
Note chromoplasts (Fig. 3). 


84 Rhodora [Vol. 75 


Chantransia hallandica Kylin 1906: 123, Fig. 8. Rosen- 
vinge 1909: 93, Figs. 21-23. 

Chromastrum hallandicum (Kylin) Papenfuss 1945: 321. 

Kylinia hallandica (Kylin) Kylin 1944: 13, 15, Fig. 7. 

Rhodochorton hallandicum (Kylin) Rosenvinge 1935: 7. 

Acrochaetium dufourii (Collins) Boergesen 1915: 19. 
Hoyt 1920: 470, Fig. 26. Collins Jn Collins, Holden, 
and Setchell 1909: 1594 (Nom. Nud.). Collins and 
Hervey 1917: 96. Taylor 1960: 305. 

Chantransia dufourii Collins 1911: 187. 

Kylinia dufourii (Collins) Kylin 1944: 13. 

Acrochaetium, sargassi Boergeson 1915: 17, Figs. 7-10. 
Taylor 1925: 129. 1928: 134, pl. 22, Figs. 1-5. 1960: 
306. 

Chantransia sargassi (Boergeson) DeToni 1924: 45. 

Kylinia sargassi (Boergesen) Kylin 1944: 13. 


Plants epiphytic, up to 1 mm. tall, original spore per- 
sisting as a unicellular base the same size or somewhat 
larger than other cells and giving rise to 1-2 main axes. 
Erect filaments moderately and irregularly branched, occa- 
sionally tapering towards the tips; unicellular hairs not 
observed. Cells cylindrical, 4-7, wide and 10-30, long (L/D 
2-5), each containing a parietal lobate chromoplast and one 
pyrenoid. 

Monosporangia ovoid, 6-8, wide and 8-11 (-15), long, 
singly or in pairs, sessile or stalked, scattered over the 
erect filaments adaxially and occasionally abaxially as well. 

Other reproductive structures not observed. 

Type Locality. — Hogardsgrund, Halland, Sweden. 

Holotype. — Apparently not designated by Kylin. Ma- 

terial on three prepared slides in LD dated 13.vii.1904 
has been chosen as lectotype. 

Distribution. — Sargasso Sea; Europe, Atlantic Coast 

of North America. 

Hosts. — Sargassum natans, Sargassum sp. and hy- 

droids in Sargasso Sea; elsewhere on a variety of 
algae. 


1973] Audouinella — Woelkerling 85 


Specimens examined. — Sargasso Sea: 31°N-69° 29'W, 3.iii.1970, 
Volkmann (WJW 2385) ; 34° N-70° W, 10.1.1970, Volkmann (wJw 2210), 
7.iii.1970, Volkmann (wsw 2401); 36°N-70° 36'W, 9.xii.1970, Moore 
(wJyw 2944). Sweden: Hogardsgrund, Halland Coast, 13.vi.1904, 
Kylin (Lp, lectotype). United States: Beaufort, North Carolina, 
13.vii.1908, Hoyt (FH, Co-type of Acrochaetium  dufourii Collins). 
Virgin Islands: St. Thomas harbour, 26.xii.1905, Boergesen (C, type 
of Acrochaetium sargassi Boergesen). 


Specimens of Audowinella hallandica from the Sargasso 
Sea compare favorably with lectotype material from the 
Halland Coast of Sweden. Neither the lectotype nor any 
Sargasso Sea collections contain sexual plants; these, how- 
ever, have been described from Denmark (Rosenvinge 
1909, p. 93, Figs. 21-22), France (Hamel 1927, p. 20, Figs. 
19-20), and the Virgin Islands (Boergesen 1915, p. 17, 
Figs. 7-10 — as Acrochaetium sargassi) . 


The taxa originally described as Acrochaetium sargassi 
Boergesen and Chantransia dufourii Collins are here con- 
sidered conspecific with Audouinella hallandica after criti- 
eal comparisons of type collection material. Plants in all 
three type collections show virtually the same range of cell 
and spore dimensions and also show good morphological 
agreement in other respects. Taylor (1960, p. 302) has 
attempted to distinguish Acrochaetium sargassi from A. 
dufourii on the basis of slight differences in cell width and 
basal cell size, but neither of these characters has proven 
reliable as a result of this study. Boergesen (1915, p. 19) 
also expressed some doubt about the taxonomic differences 
between the two taxa. 


The morphotaxonomic relationships of A«wdowinella hal- 
landica to other audouinelloid algae appear to be very 
complex and involve at least 15 other taxa (including 
Chantransia parvula, regarded by Rosenvinge (1909) and 
Hamel (1927) as conspecific with Audowinella hallandica). 
Pending critical studies of the types and other collections 
of all taxa involved, the relationships of A. hallandica to 
other species in the complex necessarily remain uncertain. 


86 Rhodora [Vol. 75 


Audouinella microscopica (Naegeli) Woelkerling 1971: 33, 
Figs. 10, 283A; 1972a: 85, Figs. 1-14. 

Acrochaetium | microscopicum | (Naegeli in Kuetzing) 
Naegeli 1861: 407, Figs. 24-25. 

Callithamnion microscopicum Naegeli in Kuetzing 1849: 
640. 

Chantransia microscopica (Naegeli in Kuetzing) Batters 
in Schiffner 1916: 136, Figs. 13-18. 

Chromastrum microscopicum (Naegeli in Kuetzing) 
Papenfuss 1945: 322. 

Kylinia microscopica (Naegeli in Kuetzing) Kylin 1944: 
13. Papenfuss 1947: 437. 

Rhodochorton microscopicum (Naegeli in Kuetzing) 
Drew 1928: 151, 163. 

Acrochaetium catenulatum Howe 1914: 84, pl. 31, Figs. 
12-18. 

Chantransia catenulata (Howe) DeToni 1924: 44. 

Kylinia catenulata (Howe) Kylin 1944: 13. 

Rhodochorton catenulatum (Howe) Nakamura 1941: 
273, 280, Fig. 1. 

Acrochaetium collopodum (Rosenvinge) Hamel 1927: 
81. 

Chantransia collopoda (Rosenvinge) Rosenvinge 1909: 
81. 

Chromastrum collopodum (Rosenvinge) Papenfuss 1945: 
320. 

Kylinia collopoda (Rosenvinge) Kylin 1944: 18, 15, Fig. 
6. 

Acrochaetium compactum Jao 1936: 241, pl. 10, Figs. 
6-14. 

Chromastrum compactum (Jao) Papenfuss 1945: 321. 

Kylinia compacta Papenfuss 1947: 436. 

Acrochaetium crassipes (Boergesen) Boergesen 1915: 
20, Figs. 11-13. Boergesen 1927: 12, Fig. 5. Collins 
and Hervey 1917: 96. Howe 1918: 511. Taylor 1941: 
75. 

Chantransia crassipes Boergesen 1909: 1, Fig. 1. Tay- 
lor 1928: 134, pl. 28, Fig. 16. 


1973] Audouinella — Woelkerling 87 


Chromastrum crassipes (Boergesen) Papenfuss 1945: 


$21. 
Kylinia crassipes (Boergesen) Kylin 1944: 13. Taylor 
1960: 300. 


Acrochaetium microfilum Jao 1936: 240, pl. 10, Figs. 
1-5. (Non A. microfilum Levring 1945: 12, Fig. 4. = 
A. Levringii Papenfuss 1947: 436). 

Acrochaetium | moniliforme (Rosenvinge) Boergesen 


1915: 22. 

Chantransia moniliformis Rosenvinge 1909: 99, Figs. 
28-29. 

Chromastrum moniliforme (Rosenvinge) Papenfuss 
1945: 322. 


Kylinia moniliformis (Rosenvinge) Kylin 1944: 13. 
Rhodochorton moniliforme (Rosenvinge) Drew 1928: 
151, 164. 


Plants epiphytic or epizoic, up to 75, tall; original spore 
persisting as a unicellular base slightly smaller to slightly 
larger than other cells. Filaments of erect system 1-3, 
commonly acurate, simple or with a few secundly to irregu- 
larly arranged laterals. Cells barrel shaped to cylindrical, 
5-9, wide and 5-10, long (L/D .75-2) ; each cell contain- 
ing a parietal irregularly lobate chromoplast with one 
pyrenoid. Unicellular hairs up to 75, long occur. 

Monosporangia ovoid, 4-7» wide and 5-8, long, terminal 
or lateral, single or rarely in pairs, sessile or stalked, 
adaxially seriate or occasionally more scattered. 

Other reproductive structures not observed. 

Type Locality. — Torquay, England. 

Holotype. — L, No. 940285 . . . 306. 

Distribution. — Nearly cosmopolitan. 

Hosts. — Dictyota, Sphacelaria, and hydroids in Sar- 

gasso Sea; a wide variety of algae elsewhere. 

Specimens examined. — Sargasso Sea: 32^ 09’N-60° 58'W, 16.v. 
1970, Woelkerling (ww 3232); 34^N-70^W, 10.1.1970, Volkmann 


(WJW 2215), 7.iii.1970, Volkmann (Wsw 2408). England: Torquay, 
1845, Naegeli (L 940285 . . . 306, type). 


88 Rhodora [Vol. 75 


Two of the three Sargasso Sea collections (WJW 2215, 
wJw 2408) contain only several plants each of A. micro- 
scopica. The other collection (3232) contains numerous 
plants on a fragment of Dictyota collected about 14 km. off 
Bermuda; it seems likely that the host had drifted out 
from shore. The apparent rarity of A. microscopica in the 
western Sargasso Sea suggests that it may not be a perma- 
nent component of the flora. 

Woelkerling (1972a) has recently presented a detailed 
account of this species including descriptions of sexual 
stages and has reduced to synonomy of a number of allied 
taxa cited as references in this account. 


Audouinella thuretii (Bornet) Woelkerling 1971: 36, Figs. 
12, 24. 

Acrochaetium thuretii (Bornet) Collins et Hervey 1917: 
98. Taylor 1960: 310. 

Chantransia thuretii Bornet. Collins 1900: 49 (Nom. 
Nud.) 

Chantransia thuretii (Bornet) Kylin 1907: 119, Fig. 28. 

Rhodochorton thuretii Drew 1928: 171. 

Chantransia corymbifera Thuret in LeJolis 1863: 107 
(in part; see Papenfuss 1945: 313 under Acrochae- 
tium bornetii. 

Chantrancia efflorescens var thuretii Bornet 1904: XVI, 
pl. 1. Collins 1906: 196. 


Plants epiphytic, caespitose, up to 5 mm. tall; original 
spore nonpersistent. Prostrate system consisting of 
branched filaments more or less forming a pseudoparen- 
chymatous disc. Erect filaments freely and irregularly 
branched and sometimes tapering towards the tips; uni- 
cellular hairs not present. Cells of main axes and laterals 
8-12 (-16) » wide and 20-80, long (L/D (2-)3-5(-8)), some- 
times tapering to 4-7, wide near the apices; each cell con- 
taining a parietal lobate chromoplast and one pyrenoid. 

Monosporangia ovoid, 8-12, wide and 16-24, long, soli- 
tary or occasionally in pairs, sessile or sometimes stalked, 


1973] Audouinella — Woelkerling 89 


generally situated adaxially on the lower portions of lateral 
branches but occasionally more scattered and/or terminal. 
Tetrasporangia ovoid to globose, 16-24, wide and 20-30, 
long, solitary or occasionally in pairs, sessile or occasionally 
stalked, scattered over laterals and main axes. 

Other reproductive structures not observed. 

Type Locality. — Cherbourg, France. 

Holotype. — PC. 

Distribution. — Nearly cosmopolitan. 

Hosts. — Sargassum natans and Sargassum sp. in Sar- 

gasso Sea; elsewhere on a variety of algae and marine 
angiosperms. 

Specimens examined. — Sargasso Sea: 28°N-70°W, 4.ii1.19770, Volk- 
mann (WJw 2424); 31°N-69° 29'W, 3.iii.1970, Volkmann (WJW 
2379); 33? 58.5'N-69^ 56.5'W, 15.v.1970, Woelkerling (WIW 2592) ; 
34°N-70° W, 10.1.1970, Volkmann (WJW 2203); 36°N-70° 36'W, 9.xii. 
1970, Moore (WJw 2934), (waw 2943) ; 37° N-70° W, 12.v.1970, Woelk- 
erling (WIW 3221); 38° 22’N-70° 58'W, 12.x.1970, Volkmann (wJW 


2889). 
Specimens of Audowinella thuretii agree in general with 


descriptions of Rosenvinge (1909) and Woelkerling (1971). 
Sexual plants are described in detail by Kylin (1907). 

The relationship of A. thwretii to other audouinelloid 
algae requires further investigation. Two similar species 
— Acrochaetium avrainvillae Boergesen (see Boergesen 
1915, p. 48, Figs. 47-49) and A. Nemalionis (DeNotaris) 
Bornet (see Collins and Hervey 1917, p. 98; Taylor 1960, 
p. 314) — occur in the Sargasso Sea region (Virgin Is. and 
Bermuda, respectively), but until the types and other col- 
lections of all three taxa can be compared, they are best 
maintained as distinct species. The major difference be- 
tween A. Nemalionis and the Sargasso Sea specimens of 
Audouinella thuretit is apparently the lack of a funiform 
prostrate system in the latter, and this difference is of 
doubious taxonomic value (Woelkerling 1971). 


Colaconema infestans (Howe et Hoyt) comb. nov. Figs. 5-6. 
Acrochaetium infestans Howe et Hoyt 1916: 116, pl. 14. 
Howe 1918: 511. Hoyt 1920: 473, pl. CXVIII. 


90 , Rhodora [Vol. 75 


Figs. 5-6. Colaconema infestans (Howe et Hoyt) comb. nov. 
Habit of monosporangial plant (Shaded portion represents host). 


1973] Audouinella — Woelkerling 91 


Chantransia infestans (Howe et Hoyt) DeToni 1924: 64. 

Chromastrum infestans (Howe et Hoyt) Papenfuss 
1945: 324. 

Kylinia infestans (Howe et Hoyt) Papenfuss 1947: 438. 
Taylor 1960: 301. 

Rhodochorton infestans (Howe et Hoyt) Drew 1928: 
151, 187. Nakamura 1944: 118, Fig. 13. 

Rhodochorton membranaceum | auct. non. (Magnus) 
Hauck: Collins and Hervey 1917: 148. 


Plants partly endozoic, more or less stoloniform, up 90, 
tall, exclusive of hairs. Prostrate system consisting of 
branched, stoloniferous filaments creeping just beneath the 
surface of the host, sometimes becoming very conjested and 
appearing pseudoparenchymatous; cells cylindrical to ir- 
regular in shape, 4-6 (-10) » wide and 5-40, long (L/D 1-8). 
Erect filaments arising more or less perpendicularly from 
the prostrate system, simple or sparingly and irregularly 
branched, occasionally bearing terminal hairs up to 125, 
long. Cells cylindrical, 3-7, wide and 6-30, long (L/D 2-5), 
each bearing a single parietal lobate chromoplast with one 
pyrenoid. 
Monosporangia ovoid, 4-6(-8), wide and 6-10(-15),4 
long, sessile or stalked, borne singly or in pairs, scattered 
over the erect filaments. 
Other reproductive structures not observed. 
Type locality. — A reef about 37km. off of Beaufort, 
North Carolina. 

Holotype. — NY. 

Distribution. — Sargasso Sea; Bermuda, Japan, North 
Carolina. 

Hosts. — Hydroids. 


Specimens examined. — Sargasso Sea: 26? 50’N-71° 48'W, bii. 
1970, Volkmann (WJW 2420); 28°N-70°W, 4.iii.1970, Volkmann 
(wzw 3432); 31°N-69° 29W, 3.iii.1970, Volkmann (wsw 2389) ; 
34°N-70°W, 7.iii.1970, Volkmann (wsw 2403), 30.vi.1970, Volk- 
mann (WJw 2725), 6.vii.1970, Volkmann (wsw 2749); 35° 54’-70° 
30'W, 13.viii.1970, Moore (wyw 2902); 36°N-70° 96'W, 9.xii.1970, 
Moore (wyw 2949), (wzw 2938); 36° 28'N-70? 29"W, 15.viii.1970, 


92 Rhodora [Vol. 75 


Moore (Ww 2907); 37° 30'N-70° W, 8.vii.1970, Volkmann | (WJW 
2706); 38° 22'N-70° 58'W, 12.x.1970, Volkmann (WIW 2887); 39° 
O7'N-70° 35'W, 16.viii.1970, Moore (wsw 2917), (wzw 2929); 39° 
30'N-71°W, 6.x.1970, Volkmann (WJW 2810). United States: Beau- 
fort, N. Carolina, 11.viii.1914, Radcliffe (NY, type). 

Specimens of Colaconema infestans from the Sargasso 
Sea agree in general with the type material on a prepared 
slide in NY. This species occurs very commonly in the Sar- 
gasso Sea and has been found in nearly every collection of 
Sargassum bearing the hydroid hosts. Erect filaments 
rarely exceed 10 cells in length, and laterals normally do 
not exceed four cells in length. 

The relationships of C. infestans to a number of other 
audouinelloid algae of similar morphology requires clari- 
fication. Acrochaetium effusum Levring 1953, p. 479, Figs. 
13F-G and Rhodochorton penetrale Drew 1928, p. 187, pl. 
44, Figs. 57-58, pl. 45, Figs. 59-60 do not appear to differ 
significantly from Colaconema infestans and a comparison 
of the types will probably show the three taxa to be con- 
specific. Chantransia endozoica Darbishire 1899, p. 13, pl. 1 
appears to have larger cells, and further study is needed 
to determine whether intermediate forms occur. Pending 
the results of such investigation, C. endozoica and Cola- 
conema infestans are maintained as distinct species. 

The relationships of Colaconema. infestans to a number 
of adouinelloid taxa growing on plant hosts also requires 
clarification, especially since host specificity does not ap- 
pear to be a reliable criterion of specific distinction (Woel- 
kerling 1971). Endophytic taxa of similar morphology 
include Acrochaetium antillarum Taylor 1942, p. 78, pl. 2, 
Figs. 3-4, A. endophyticum Batters 1896, p. 386, Chan- 
transia emergens Rosenvinge 1909, p. 128, Fig. 55, and 
Colaconema porphyrae (Drew) Woelkerling (see Drew 
1928, p. 188, pl. 46, Figs. 70-75; Woelkerling 1971, p. 50, 
Figs. 20, 27B), and critical studies on the types and other 
collections of these taxa may show some or all to be con- 
specific. 

Further investigations are also needed to determine 


1973] 


Audouinella — Woelkerling 93 


^ ! 
e 
iP | 
» | 
»Z ii 
zs 3 
a KP iy 
OG | 
We W/ £ 
A y 
tps y 
Zo lint ay 
fo P y 
Lx (8) 
l 
c» jel/ 
y 
V/ 
AO 
PET i 
Vati 8 
| Y J 
r^ Í by, ear 
i Zam 
fE 
HAZ 
OSS 


Figs. 7-8. 


Colaconema secundata (Lyngbye) comb. nov. Fig. 7. 
Chromoplasts in vegetative cells. Fig. 8. Habit of fairly smal] mono- 
sporangial plant. 


94 Rhodora [Vol. 75 


whether or not Colaconema bonnemaisoniae Batters and 
related taxa (see Woelkerling 1971, p. 42) possibly repre- 
sent prostrate system stages of C. infestans. 

Specimens of Collins and Hervey (1917, p. 148) from 
Bermuda referred to Rhodochorton membranaceum (Mag- 
nus) Hauck and distributed in the PBA, Vol. XLIV, No. 
2194 (Collins, Holden, and Setchell 1917) have been ex- 
amined and found to be plants of Colaconema infestans. 
Howe (1918, p. 511) reached the same conclusion. 


Colaconema secundata (Lyngbye) comb. nov. Figs. 7-8. 
Acrochaetium. secundatum (Lyngbye) Naegeli 1861: 
405. 
Callithamnion secundatum (Lyngbye) C. Ag. 1828: 187. 
Ceramium secundatum (Lyngbye) C. Agardh 1824: 132. 
Chantransia secundata (Lyngbye) Thuret Jn Le Jolis 


1863: 106. 
Chromastrum secundatum (Lyngbye) Papenfuss 1945: 
323. 


Kylinia secundata (Lyngbye) Papenfuss 1947: 437. 

Callithamnion daviesii var. secundatum Lyngbye 1819: 
129. pl. 41, Fig. B4-6. 

Acrochaetium luxurians (J. Agardh) Naegeli 1861: 405. 

Callithamnion luxurians J. Agardh 1851: 14. 

Chantransia luxurians (J. Agardh) Kylin 1907: 117, 
Fig. 26. 

Acrochaetium virgatulum (Harvey) Bornet 1904: XXII. 
Boergesen 1927: 14, Figs. 7-8. Chapman 1963: 56. 
Hoyt 1920: 473, Figs. 29-30. Taylor 1941: 75. 

Callithamnion virgatulam Harvey In Hooker 1833: 349. 

Chantransia virgatula (Harvey) Thuret In LeJolis 1863: 
106. Rosenvinge 1909: 109, Figs. 37-41. 

Chromastrum virgatulum (Harvey) Papenfuss 1945: 
$28. 

Kylinia virgatula (Harvey) Papenfuss 1947: 437. 

Rhodochorton virgatulum (Harvey) Rosenvinge 1935: 
7. 


1973] Audouinella — Woelkerling 95 


Trentepohlia virgatula (Harvey) Farlow 1881: 109. 


Plants epiphytie or epizoic, more or less virgate, up to 
2mm. tall; original spore non-persistent. Prostrate system 
at first a parenchyma-like group of cells, later forming a 
small, more or less circular pseudoparenchymatous disc of 
one to several cell layers. Erect filaments nearly simple to 
moderately branched, commonly bearing short laterals 
giving plant a virgate appearance; terminal and pseudo- 
lateral hairs common. Cells of erect filaments cylindrical 
(6-) 8-12, wide and (10-) 20-70, long (L/D 1-7), each 
with a distinctly stellate chromoplast and one pyrenoid. 

Monosporangia ovoid, (6-) 9-13, wide and (10-) 16-24, 
long, sessile or stalked, solitary, in pairs or occasionally in 
threes, commonly crowded laterally or terminally on the 
shorter laterals or occasionally more scattered. 

Other reproductive structures not observed. 

Type locality. — Kvivig, Faeroes Islands (on “Conferva 

rupestris”). 

Holotype. — C. 

Distribution. — Sargasso Sea; Atlantic Shores of North 

America, Canary Islands, Europe. 
Hosts. — Sargassum fluitans, S. natans, and hydroids in 
Sargasso Sea; a wide variety of algae elsewhere. 

Specimens examined. — Sargasso Sea: 28°N-70°W, 4.iii.1970, Volk- 
mann (WJW 2425); 31?N-69? 29W, 3.iii.1970, Volkmann (WJW 
2367); 39? 30’N-71°W, 6.x.1970, Volkmann (wJw 2865). England: 
Torquay, prior to 1833, Griffiths (TCD, type of Acrochaetium virgatu- 
lum (Harvey) Bornet). Faeroes Islands: Kvivig, 19.vi.1817, ? (C, 
Herb. Lyngbye, type). Sweden: Kattegat Channel, no date, ? (LD 
35117, type of Callithamnion luxurians J. Agardh). 

The few specimens of Colaconema secundata from the 
Sargasso Sea agree well with type material from the 
Faeroes Islands. Although an extremely variable species, 
C. secundata can be distinguished from other audouinelloid 
algae by the following combination of characters (in addi- 
tion to cell and spore dimensions): 1) Spore germinating 
to form a distinctive parenchymatous group of cells which 
may later proliferate (see Kylin 1907, p. 115, Fig. 24; 


96 Rhodora [Vol. 75 


Rosenvinge 1909, Figs. 37-41) ; 2) Cells with a distinctly 
stellate chromoplast with a centrally located pyrenoid, and 
in many cases 3) sporangia densely crowded on short 
lateral branches and 4) numerous terminal hairs, often 
terminating 1-2 celled branchlets. Tetrasporangia have 
not been observed but are reported by Hehre and Mathieson 
(1970), Kylin (1907, 1944), and Rosenvinge (1909) among 
others. Sexual stages remain unknown. 

Unicellular hairs occur very commonly in C. secundata 
and often terminate 1-2 celled lateral branchlets, thus ap- 
pearing stalked (Fig. 8). 

In agreement with Hamel (1927, 1928) and Rosenvinge 
(1909), the taxa originally described as Callithamnion 
luxurians J. Agardh (1851, p. 14) and C. virgatula Harvey 
In Hooker (1833, p. 349) are considered conspecific with 
Colaconema secundata. The type collections of all three 
have been examined during this study and found to agree 
in all essential features. Since the specific epithet “secun- 
data", first used by C. Agardh (1824, p. 132), predates the 
specific epithet *virgatula" (Harvey 1833) used by Hamel 
(1927, 1928) and Rosenvinge (1909) by nine years, it has 
nomenclatural priority. 

Some authors (e.g. Kylin 1944, Taylor 1957) have main- 
tained C. secundata and Callithamnion (— Acrochaetium, 
Kylinia) virgatula as distinct species on the bases of dif- 
ferences in the number of layers in the prostrate system 
or on slight differences in height, branching, or cell size, 
but as noted by Rosenvinge (1909), and as observed in 
New England collections (Woelkerling, unpublished data), 
considerable variation occurs in all cases, and species 
limits between the two taxa cannot be drawn reliably. 

The taxonomy and relationships of Colaconema secundata 
to other taxa are quite involved and will be dealt with at 
a later date. 


SUMMARY 


The Awdowuinella, complex is represented in the Western 


1973] Audouinella — Woelkerling 97 


Sargasso Sea by four species of Audouinella and two species 
of Colaconema, all newly recorded for this region. These 
species apparently constitute a permanent component of 
the Sargasso Sea flora, reproducing asexually by mono- 
spores and/or tetraspores. Species with a stoloniferous 
habit tend to produce more extensive prostrate systems 
than erect systems, whereas caespitose species usually pos- 
sess better developed erect systems than prostrate systems. 
Critical comparisons of type and other collections indicate 
that Acrochaetium dufourti (Collins) Boergesen and A. 
sargassi Boergesen are conspecific with A«wdowinella hal- 
landica (Kylin) comb. nov., and Acrochaetium luxurians 
(J. Agardh) Naegeli and A. virgatula (Harvey) Bornet 
are conspecific with Colaconema secundata (Lyngbye) 
comb. nov. Detailed descriptions of Sargasso Sea collec- 
tions together with a taxononmic key are provided. 


ACKNOWLEDGMENTS 


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 to the directors of 
various herbaria (C, L, LD, NMW,NY, TCD) for the loan of 
critical specimens. Miss Susan Heller kindly assisted in 
the preparation of drawings. 

This study was supported in part by grant GB-13250 from 
the National Science Foundation to the Systematics-Ecology 
Program, Marine Biological Laboratory, Woods Hole, 
Massachusetts. 


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1906. Acrochaetium and Chantransia in North 

America. Rhodora 8: 189-196. 

1911. Notes on algae. X. Rhodora 13: 184-187. 
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„ and Hervey, A. B. 1917. The algae of Bermuda. 

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., HOLDEN, I., and SETCHELL, W. A. 1909. Phycotheca 

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1917. Phycotheca 


Boreali Americana. XLIV. 

CONOVER, J. and SrEBURTH, J. 1964. Effect of Sargassum distribu- 
tion on its epibiota and antibacterial activity. Botanica mar. 
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DARBISHIRE, O. V. 1899. Chantransia endozoica Darbish., eine neue 
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1973] Audouinella — Woelkerling 99 


DE TowNi, J. B. 1924. “Sylloge Algarum omnium hucusqua Cogni- 
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DiLLWYN, L. W. 1802-1809. “British Confervae." (London.) [For 
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139-224, Plates 37-48. 

FARLOW, W. G. 1881. The marine algae of New England. Rep. 
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. 1914. The vegetation of the Sargasso Sea. Proc. 
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HAMEL, G. 1927. “Recherches sur les genres Acrochaetium Naeg. 
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1928. Sur les genres Acrochaetium Naeg. et Rhodo- 
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Harvey, W. H. 1833. Cryptogamia algae. Div. II. Confervoideae. 
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E . 1852. Nereis Boreali-Americana. Part I. Mela- 
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HENTSCHELL, E. 1921. Ueber den bewuchs auf den treibenden 
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Hower, M. A. 1914. The marine algae of Peru. Mem. Torrey bot. 
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1918. Algae, pp. 489-540. In Britton, N. L. Flora 
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„ and Hoyt, W. D. 1916. Notes on some marine algae 
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Hoyt, W. D. 1920. Marine algae of Beaufort, N.C. and adajacent 
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Jao, C. C. 1936. New Rhodophyceae from Woods Hole. Bull. Tor- 
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KUETZING, F. T. 1849. “Species Algarum." (Leipzig.) 

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(Uppsala.) 


100 Rhodora [Vol. 75 


1907. “Studien ueber die Algenflora der Schwedischen 
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1947. Further contributions toward an under- 
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1973] Audouinella — Woelkerling 101 


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1941. Notes on the marine algae of Texas. Pap. 

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1942. Caribbean marine algae of the Allan Han- 

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bot., Suppl. Ser. Suppl. 1. 91 pp. 
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1972b. Some algal invaders on the north- 
western fringes of the Sargasso Sea. Rhodora 74: 295-298. 


DEPARTMENT OF BOTANY 
UNIVERSITY OF WISCONSIN 
MADISON, WISCONSIN 53706 


SALT CONCENTRATIONS IN GROUND WATERS 
BENEATH RHIZOPHORA MANGLE AND 
AVICENNIA GERMINANS 


LoRING MORROW 
AND NORTON H. NICKERSON' 


The purpose of this investigation was to determine 
whether salinity of ground water beneath Rhizophora 
mangle L., (red mangrove), and Avicennia germinans 
(L.)L. (black mangrove), was correlated with the dis- 
tribution of these species. It was expected that there might 
be a mixture of fresh ground water and tidal salt water 
and that the levels of resulting salinity might be a deciding 
factor in their distribution. Moog (1963) stated on the 
basis of his preliminary studies that A. germinans was 
regulated by this salt variation. He found that in salt 
pools where there was a high concentration of salt (higher 
than sea water) the growth of A. germinans was dwarfed 
and the trees grew poorly, growing only approximately 
2 em. per year. Individuals of the marsh area, he reported, 
thrived, growing 8 to 30 cm. per year. Moog ascribed this 
difference to the fact that they were in contact with fresh 
water daily at high tide when the underlying fresh-water 
table was raised due to tidal pressure. 

The method of testing was to drill a hole near the base 
of the individual of a particular species with a three-foot- 
long hand auger. Next a small hand pump with a two-foot- 
long hose was put into the hole and a sample of water was 
taken. This sample was kept in a sealed glass test tube 
and later that day tested for conductivity with a Beckman 
conductivity bridge, Model RC-19, which gives its results 


"This work was supported by grants from the Hurdle Hill Founda- 
tion, the Arnold Bernhard Foundation and the Faculty Research Fund 
of Tufts University. Thanks are gratefully extended to Dr. R. A. 
Howard, Director, Arnold Arboretum, Harvard University, for per- 
mission to utilize his as yet unpublished updated Bahamian plant 
nomenclature. 


102 


1973] Rhizophora — Morrow & Nickerson 103 


in micromhos per centimenter (umho/cm). Conductivity is 
the reciprocal of resistivity and is expressed in mho/cm 
(mho = 1/ohm, mho = 1/10" ohms). This technique does 
not indicate salinity directly, but does give conductivities 
which can be compared to the conductivities of sea water 
and of fresh water. A higher conductivity would mean 
higher concentration of salts, of which, in sea water, sodium 
chloride would be predominant. 

The samples were taken during January, 1971, on five 
cays (Coakley, Bowe, Ben Rolle, Hummingbird and Cul- 
mer’s) of the Jewfish Chain, which runs westward at 23.5°N 
from George Town, Great Exuma, in the Bahamas. Hum- 
mingbird Cay and the other islands are discussed further in 
a study now in preparation (Nickerson et al, 1973). 

Several samples of open sea water were tested; they dis- 
played a conductivity of 62.7k umho/cm. In contrast, fresh 
water from a well on Hummingbird Cay, site of the Hum- 
mingbird Cay Foundation's Biological Laboratory, displayed 
a conductivity of 1.03 »mho/cm. These values mean re- 
sistances of approximately 16 ohms for 1 cm of seawater and 
1000 ohms for 1 em of fresh water. They also mean for sea- 
water, about 37,000 ppm NaCl at 259C. and for the well 
sample, about 450 ppm NaCl at 25°C. 

Very few Laguncularia racemosa (L.) Gaertn. (white 
mangrove) were tested because the soil was usually too dry 
beneath them to obtain a sample. The one that could be 
tested had a conductivity of 62.9k. This species is not dealt 
with further in this study. 

Fifteen samples were taken from areas in which Rhizo- 
phora mangle grew exclusively. The average conductivity 
was 63.2k, approximately that of sea water. 

Sixteen samples were taken from areas where Avicennia 
germinans occurred exclusively. They had a much higher 
average conductivity (72.1k), with the highest individual 
sample at 88.8k. In areas where both A. germinans and R. 
mangle were found in approximately the same numbers, the 
average of eight samples was 67.7k, approximately midway 


104 Rhodora [Vol. 75 


between the other two averages. These data are summarized 
in Table I. 

The areas where A. germinans was found were usually 
depressions, often partly filled with sand, where the salt 
ocean water flowed in at high tide and stayed even after the 
tide went out. This condition would form a salt pool, which 
the results show, because the water apparently evaporated 
and left the salt behind. Our results were quite consistent. 
There were only two black mangrove areas with salinity 
equal to sea water, and we were unable to determine the 
shape of the underlying rocks in these areas with any de- 
gree of reliability. These results are interpreted to mean 
that A. germinans grows where it does because of a higher 
concentration of salt in the ambient water than that found 
in sea water. Whether A. germinans needs the higher con- 
centration of salt or whether it just tolerates it is another 
question to which these data do not speak. 

Moog (1963) noted that the osmotic pressure in A. germi- 
nans is 21.5 atm. and in R. mangle it is 27 atm. The energy 
required to remove only water from sea water should be less 
with increased internal osmotic pressure and thus it would 
presumably be easier for R. mangle to obtain fresh water 
from sea water. A. germinans, however, secretes salt (NaCl) 
in large quantities on its leaves: hence it must absorb salt 
water from its environment. It is also possible that A. ger- 
minans grows in salt pools because of the lack of competition 
from other plants. There are solid stands of A. germinans 
in inland salt-poo] areas on several islands of the Jewfish 
chain. 

Rhizophora mangle is always found in either open sea 
water or in tidal flat areas that are covered with large 
quantities of new sea water each day at high tide. The con- 
ductivity of water samples obtained beneath these R. mangle 
colonies at the time of testing was always close to that of 
sea water. Davis (1943) reported that R. mangle will live at 
least six years in a greenhouse with only fresh water, which 
indicates that it can perhaps adapt to fresh water. Against 
this evidence is a color infra-red aerial photograph of the 


1973] Rhizophora — Morrow & Nickerson 105 


Marco Island area of western Florida, which shows a pro- 
gressive color change, interpreted as slow dying of man- 
grove swamps from which the sea has been excluded by 
diking (Nickerson, 1971, Plate 4.) 

Our evidence indicates that R. mangle survives only in 
areas where salinity approximates that of sea water, and 
where sea water is free to exchange with the ocean at each 
tide. It indicates that A. germinans thrives in areas where 
salinities consistently exceed that of sea water, and where 
there is not a free exchange of root-bathing waters with the 
ocean at each tide. 


TABLE I 
TABLE OF CONDUCTIVITIES 
FOR EACH WATER SAMPLE 
(in thousands of »mhos/cm) 


Rhizophora & 
Rhizophora Avicennia Avicennia Laguncularia 


61.7 82.1 62.8 62.9 
62.4 88.8 69.9 
57.7 11.5 68.8 
64.6 80.5 12.8 
65.6 70.2 Tei 
62.6 12.3 63.1 
63.5 70.2 67.3 
55.5 65.5 64.9 
55.7 63.9 av 67.7 
58.7 69.2 
65.0 71.9 
69.9 78.6 
65.5 68.3 
70.5 74.4 
69.2 65.2 

av. 63.2 61.0 


av. 72.1 


106 Rhodora [Vol. 75 


LITERATURE CITED 

Davis, J. H., JR. 1943. The Natural features of South Florida. De- 
partment of Conservation Geological Bulletin 25: 1-31. 

Moos, A. O. D. 1963. A preliminary investigation of the significance 
of salinity in the zonation of species in salt-marsh and mangrove 
associations. South African Journal of Science 59(3): 81-86. 

NICKERSON, N. H. 1971. Ecology and Everyman. Tufts Review, 
Winter 1970-71. Medford, Mass. pp. 20-25. 

NICKERSON, N. H., SEMPLE, J. C., HOWARD, R. A., PHIPPEN, W. and 
TSCHUNKO, A. 1973. Ecology and vegetation of Hummingbird 
Cay, Jewfish Island Chain, Great Exuma, Bahamas. (in prepa- 
ration. 

DEPARTMENT OF BIOLOGY 

TUFTS UNIVERSITY 

MEDFORD, MASSACHUSETTS 02155 


VARIATION OF SEED SIZE IN 
ATRIPLEX PATULA VAR. HASTATA (L.) Gray 


FRANK R. DRYSDALE 


Ungar (Rhodora 73:548-551, 1971) demonstrated seed 
dimorphism in Atriplex patula var. hastata (L.) Gray col- 
lected from saline marshes surrounding Lincoln, Nebraska. 
One kind of seed is small (1.0-1.7 mm) having a hard black 
testa and the other is usually larger (1.3-2.6 mm) having a 
soft yellowish-brown testa. Ungar (1971) shows a bimodal 
distribution of this condition. He has informed me that in- 
dividuals of A. patula var. hastata do not yield equal num- 
bers of each seed type, but he did not know the relative 
yields of each type. 

In order to investigate this matter further, six lots of 
four to fourteen individuals each of A. patula var. hastata 
were harvested from an Alexander Township, Athens Coun- 
ty, Ohio salt pan during October, 1971. Seeds were separated 
from fruits by chaffing. Small black seeds and larger brown 
seeds from each lot were counted (Table 1) and placed in 
separate envelopes. Samples from each category in each 
lot were placed on a piece of filter paper. With a dissecting 


1973] Atriplex — Drysdale 107 


Table 1. Seed samples of Atriplex patula var. hastata. 


Lot 
1 2 3 4 5 6 
Number of plants 4 7 6 7 8 14 
Total small seeds 255 961 87 1039 815 3 773 
Sample size (sm. seed) 37 24 10 23 21 21 
Total large seeds 14 T2 ` 298 56 63 25 
sample size (lg. seed) 14 22 AS 9 18 16 


Table 2. Size characteristics for seed of Atriplex patula 
var. hastata. 


Lot 
1 2 3 4 5 6 
Seed size (mm) Number Sampled 

1.0 1 1 2 
1.1 5 4 3 2 3 
1.2 11 5 4 13 7 9 
1.3 8 9 4 4 i 5 
1.4 9 4 2 2 4 3 
1.5 6 3 2 2 
1.6 3 2 4 2 5 3 
1.7 d T 2 3 4 
1.8 4 9 2 2 3 
1.9 1 2 1 5 1 
2.0 1 3 1 2 1 1 
2.1 4 1 1 1 
2 2 2 2 
2.6 1 


microscope and a calibrated ocular micrometer the diameter 
of each seed in each sample was measured (Table 2). By 
multiplying the reciprocal of the proportion of seeds sam- 
pled from each envelope times the number of seeds in each 
diameter size class an estimate of the absolute number of 
seeds in each size class was made (Table 3). 


108 Rhodora [Vol. 75 


Table 3. Estimated number of seed in each size class in 
each lot sampled of Atriplex patula var. hastata. 


Lot 

Seed Total 
Size (mm) 1 2 3 4 5 6 Seeds 
1.0 40 45 74 159 
1.1 34 160 136 78 111 529 
1.2 76 201 a5 589 272 335 1508 
1.3 55 93061 a5 182 272 185 1090 
1.4 62 160 17 91 156 76 562 
1.5 30 7 43 3 79 
1.6 3 43 9 12 18 5 87 
1.7 3 2 12 11 6 31 
1.8 4 30 2 12 7 b 54 
1.9 1 7 6 18 2 33 
2.0 1 10 2 12 4 2 30 
2.1 13 2 4 2 21 
2.2 7 4 11 
2.6 2 2 


Figure 1 presents the estimated number of seeds in each 
size class in all lots. The log values for the estimated num- 
ber of seeds in each size class in each lot are plotted in 
Figure 2. 

These data indicate that rather than presenting equal 
numbers of seeds in each major size category, the large and 
the small of Ungar, there is a log normal distribution of 
seeds over each size class with considerably fewer seeds in 
the larger classes. They do not explain the presence of 
seemingly different colors of seeds; though pigmentation of 
the testa may be a product of pigment density and surface 
area with larger seeds having a lighter coloration due to 
more diffuse pigmentation. The data indicate that in the 
population seed size is log normally continuous, as opposed 
to bimodal. Also, the attendant observation of seeds of both 
colorations and major categories (large and small) on the 


1973] Atriplex — Drysdale 109 


15 - 
14 F 
13 - 
12 ge 
z we 
c 
z 
wo 
mii 
[9] 
^-^ 
er 
a 
m 
m 
o = 
^ 8 
> 7r 
c 
3 
E 
i < 
ael . 
e 
5k 
ALT 
3i- 
2T 
Te regia M 
m ta 
. erba oo | 


pi L 1 i L 1 1 L L 1 1 L + + ) a 
1.0 1.1 2 13 1.4 1.5 1.6 1.7 1.8 1.9 2.0 2.1 2.2 2,3 2.4 2.5 2.6 
SIZE (mm) 


Figure 1. Estimated number of seeds in each size class for all 
lots of Atriplex patula var. hastata. 


110 Rhodora [Vol. 75 
CUT 
- 
L m 
500L 
L 
s. 2° 
5- 5. 
L 2* 4 
2 “42, 
z 4- 5 
c 
6- 
z100 
> [ ae 
m F 6 5° 18 6- 
m 
; QU 
O 50 
m a rd 5s 2° 
3* 3e 
Mi = 1- le 2° 
m 
m 
o 
wn - 5. 
3s 3° 
— Ae ge . 2 
m 4* 5* 4 4 
e 10 = 3. 2* 
M [ 2* 5* 2e 2° 
Q F ó» 4+ 
^ 5r 6» oe 
H = 1. 5» 5. 3s 
" és 1] Ve 
L 36 6* 3*6 36 3* 
1 l L i L 1! 1 l 1 Low lq L 1 1 1 4 J 
10 n1 L2 1.3 1.4 1.5 1.6 1.7 1.8 1.9 2.0 2.1 2.2 2.3 2.4 2.5 2.6 
SIZE (mm) 
Figure 2. Log values for estimated numbers of seed in each size 


class for each of six lots of Atriplex patula var. hastata. 


same plant indicates that seed morphism may be develop- 


mental. 


DEPARTMENT OF BOTANY 
OHIO UNIVERSITY 
ATHENS, OHIO 45701 


FLORAL DIMORPHISM 
IN 
RACHICALLIS AMERICANA (JACQ.) HITCH. 
(SALTWATER-BUSH) 


NORTON H. NICKERSON! 
and 
JOSEPH W. TRIPP 


Hummingbird Cay is one of ten large oolitic limestone 
islands in the Jewfish Chain which runs westerly at 23.5°N 
from the George Town area of Great Exuma in the Baha- 
mas. The island’s ecology and flora are more fully described 
elsewhere (Nickerson et al., in preparation). One member 
of the rocky-shore seaside plant communities is Rachicallis 
americana (Jacq.) Hitch. (Rubiaceae), listed by Britton and 
Millspaugh (1920) as a monotypic genus with 4 stamens 
and a half-superior, 2-celled ovary with slightly two-lobed, 
thick styles. Observation of many individual plants during 
early July of 1968 and 1969, the wet season when they are 
near their flowering peaks, showed that some bushes pro- 
duced flowers with only bifurcate stigmas clearly visible, 
while others showed only 4 anthers. Since no mention was 
made by Britton and Millspaugh of this seeming dioecism, 
a careful study was made. 

Many flowers on each of several different bushes were ex- 
amined in the field in 1968. In 1969, branches laden with 
flowers in all stages of anthesis were collected from 32 ran- 
domly selected plants in five different locations. Ten flowers 
from each of these branches were carefully examined in the 
laboratory with a binocular microscope. Each flower was 
dissected by slitting the tube and flattening it slightly to 
observe the filaments, anthers, stigmas and style. The sta- 
men-style form of each flower and each branch was noted 


"This research was supported by grants from the Hurdle Hill 
Foundation to the Cape Cod Museum of Natural History, Brewster, 
Mass. and to Tufts University, and from the Arnold Bernhard 
Foundation to Earlham College, Richmond, Indiana. 


111 


112 Rhodora [Vol. 75 


and recorded. Pollen samples from flowers of each branch 
were compared with those of each other branch with a com- 
pound microscope. Dried specimens of representative 
branches were prepared and are deposited in the Herbarium 
of Tufts University, Medford, Mass. In 1970, 25 of these 32 
plants were checked again; each exhibited the same sexual 
form as they had the previous year. 

Foliage and branching habits of all the plants appeared to 
encompass the same amounts of variation. Flowers were 
dimorphic. Those in which the stigmas were clearly visible 
were perfect and had anthers on very short filaments well 
down the tube. Pollen was present. Those in which anthers 
were visible had longer filaments, occasionally with insertion 
higher up the tube, but not with anthers exserted. Stigmas 
and styles in such flowers were abortive, regardless of floral 
age. Ovaries were essentially equal in size in all flowers at 
this stage. All the flowers on any one plant were found to 
be identical with regard to form of stigmas, styles and fila- 
ments. The data are summarized in Table I. 

Flowers of the two types occurred on separate plants. 
Both types of plants occurred at all but one of the sites 
sampled. Pollen was abundantly produced by both floral 
types and appeared alike in size and color under the com- 
pound microscope. Fruits were produced only on perfect- 
flowered individuals. 

Perfect-flowered plants and male plants of Rachicallis 
americana occur in approximately equal numbers in this 
part of the Bahamas. There was no obvious correlation be- 
tween geographic and/or ecological situation and ex- 
pressed floral form, as Vuilleumier (1967) had noted might 
be present. Three plants of each type were known to be con- 
sistent in the sexual form of their flowers for the three sea- 
sons of 1968, 1969 and 1970. This case of floral dimorphism 
seems to be a further documentation of Vuilleumier's (1967) 
stand that many cases of floral dimorphism have been un- 
reported by taxonomists. 

The description of Rachicallis americana (Jacq.) Hitch. 
should be amended as follows: Plants of two types, bearing 


1973] Rachicallis — Nickerson & Tripp 113 


TABLE I 
Floral Dimorphism in Rachicallis americana, (Jacq.) Hitch. 
Sexual Form of Individual Plants 


Location of Plant Perfect Flowers Male Flowers 
Great Exuma, So. Dock 
(southeast-facing exposure) 6 3 


Tommy Young Cay 
(west-facing exposure) 9 3 


Culmer's Cay 
(south-facing exposure) 3 


t 


Hummingbird Cay 


South Coast 4 5 
Northeast Coast 0 3 
Total number of plants, 
each sexual expression : 16 16 


dimorphic flowers either exclusively perfect with 4 stamens 
on very short filaments below the prominent bifurcate stig- 
ma, or exclusively with 4 anthers visible at the mouth of 
the tube and above the short aborted style and stigma. 
Fruits formed only on perfect-flowered plants. 


LITERATURE CITED 


BRITTON, N. L. & C. F. MinrsPAUGH. 1920. (1962 reprint). The 
Bahama Flora. Hafner Publishing Co., Inc., N.Y., N.Y. 695 pp. 

NICKERSON, N. H., SEMPLE, J. C., HowaRD, R. A., PHIPPEN, W., & 
TSCHUNKO, A. 1973. Ecology and Vegetation of Hummingbird 
Cay, Jewfish Island Chain, Great Exuma, Bahamas. (in prep- 
aration) 

VUILLEUMIER, B. S. 1967. The origin and evolutionary development 
of heterostyly in the Angiosperms. Evol. 21: 210-226. 


DEPARTMENT OF BIOLOGY 
TUFTS UNIVERSITY 
MEDFORD, MASSACHUSETTS 02155 


UNIVERSITY OF MASSACHUSETTS 
AMHERST, MASSACHUSETTS 01002 


BOOK REVIEW 


AQUATIC AND WETLAND PLANTS OF THE SOUTHWESTERN 
UNITED STATES! 


This large and extensively illustrated treatment of the 
aquatic and wetland flora of Oklahoma, Texas, New Mexico, 
and Arizona, published by the Environmental Protection 
Agency, is timely, for it emphasizes the importance of basic 
information about plants for water conservation and water 
quality studies and programs. The Corrells interpret the 
aquatic and wetland environment broadly so that it includes 
many habitats ecologically peripheral to the rivers, streams, 
lakes, marshes and swamps of the region. This scope is im- 
portant because it stresses the totality of wet areas of a 
watershed — the wet medows, bogs, and seepages that act 
as sponges to hold precipitation, the river and stream banks 
and floodplains that are important in erosional and deposi- 
tional processes, as well as the lakes and rivers. The botany 
of the whole area involved in water quality and water use 
is thus included. The basic environmental message of this 
book — the integrity of the whole watershed and the basic 
role of plant life in its maintenance — should be appreciated 
by persons in charge of watershed protection and utiliza- 
tion. 

The book is obviously important in another way, being a 
manual of a significant portion of the flora of the south- 
western United States. The aquatic and wetland flora in- 
cludes about 2100 species, representing 567 genera and 129 
families. Most of the species grow in wet soil or in season- 
ally wet places. The Cyperaceae, Gramineae and Compositae 
are the dominant families, with a total of 590 species, rath- 
er few of them strictly aquatic. The wide range of plants 
brought into the treatment is also indicated, for example, 


‘Donovan S. Correll and Helen B. Correll. Aquatic and Wetland 
Plants of the Southwestern United States, i-xv, pp. 1-777, figs. 1-789. 
Environmental Protection Agency. 1972. (Price $7.75, Superintendent 
of Documents, United States Government Printing Office, Washington, 
D.C. 20402, Stock number 5501-0177). 


114 


1973] Book Review — Tryon 115 


by the presence of Dodecatheon, Primula and Trifolium 
(alpine springs and wet meadows), of Marsilea and Elan- 
tine (vernal pools), of Populus and Salix (borders of wa- 
tercourses), and of Amaranthus (periodically inundated 
areas along rivers). The excellent illustrations of about 800 
species, the keys, and the descriptions all facilitate accur- 
ate identification. The manual will have a wide use as a 
source of information about the species and for determin- 
ations. A printing error may be noted, the names on two 
figures have been reversed: Fig. 570 should be named My- 
riophyllum pinnatum, and Fig. 571 should be Myriophyllum 
heterophyllum. 

This sizable volume adds to the impressive scope of works 
by Dr. Donovan Correll including books on the orchids of 
North America, on the potato and its wild relatives, on the 
ferns and fern allies of Louisiana, of Texas, and of Chihua- 
hua, Mexico, as well as the most recent large publication on 
the flora of Texas (with M. C. Johnston). Dr. Helen Cor- 
rell has contributed to the successful completion of these 
and has assumed a major role in the research and prepara- 
tion of this publication on aquatic and wetland plants. This 
book by the Corrells is an important source of information 
on plants of the southwestern United States and will be 
especially useful for environmental studies related to water 
resources. 


ROLLA TRYON 
GRAY HERBARIUM AND DEPARTMENT OF BIOLOGY, 
HARVARD UNIVERSITY 


A NEW ALASKAN SPECIES OF 
ERIGERON L. (COMPOSITAE)' 


STEPHEN A. SPONGBERG 


During a study of variation patterns and the systematics 
of monocephalous species of Erigeron indigenous in arctic 
and cordilleran North America (Spongberg, 1971), a new 
species closely allied to the polymorphic, arctic-alpine spe- 
cles, E. grandiflorus W. J. Hooker, was discovered among 
herbarium collections from Alaska. It is a pleasure to name 
the new taxon in honor of Professor Eric Hultén, foremost 
authority on the Alaskan flora. 


Erigeron Hultenii Spongberg, sp. nov. Figure 1. 

Herba parva monocephala, e caudice perenni gracili de- 
cumbentique exoriens. Folia basalia pauca rosulata, 12-30 
mm. longa, 2-6 mm. lata, lanceolata ad oblanceolata; folia 
caulina 3-5, foliis basilaribus similia, autem sessilia et supe- 
riora parum minora. Jnvolucrum e phyllariis uniserialibus 
ca. 30 constatum; pagina abaxialis phyllarii trichomatibus 
multiserialibus unicellularibus dimorphis subtiliter hirsuta. 
Capitula gynomonoecia ; radii flosculi exteriores femine ligu- 
lati 20-45, limbo corollae 5-9 mm. longo, 1-2 mm. lato; disci 
flosculi multi, corolla 2.5-3 mm. alta ad anthesin infundi- 
buliformi; rami stylares 0.65-0.75 mm. longi. Achenia 
ovoidea hirsuta quadri-costata, costa omne filum vasculare 
adscendens in pariete achenii superjecta.  Holotypus: 
Alaska: Campbell Creek Valley, 11 miles north of Anchor- 
age, August 7, 1965, L. Strutz, s.n. (S). 


Small, herbaceous perennial from a slender caudex, 2-4 
mm. in diameter; flowering stem(s) monocephalous, 3-6.5 
cm. high, pubescent with uniseriate trichomes. Basal leaves 
few, 2-5, lanceolate to oblanceolate, 1.2-3 cm. long, 2-6 mm. 


'I wish to thank Dr. Elizabeth A. Shaw of the Gray Herbarium, 
Harvard University, for preparing the Latin description, and Miss 
Marion Seiler of the University of North Carolina, Chapel Hill, for 
her careful and precise skill in preparing the illustration. 


116 


1973] New Erigeron — Spongberg 117 


es 


Se ab deste ee 


ERAS Ta ye Ses 


PPE 
PIT DESEE 


Figure 1. A-F, Erigeron Hultenii: A, habit, X 1 1/3; B, head 
with involucre and ligulate florets, as pressed, X 11/3; C, phyllary, 
abaxial surface, ca. X 5; D, ligulate floret, ca. X 5; E, detail of basal 
portion of ligulate floret, ca. X 10; F, disc floret, ca. X 10. 


118 Rhodora [Vol. 75 


wide, with an apiculate apex, short, basally dilated petioles, 
and slightly thickened, often red-tinged and ciliate entire 
margins ; blades with a prominent midvein and 2 less prom- 
inent lateral veins, sparsely pubescent, moreso along the 
veins; cauline leaves 3-5, alternate, resembling the basal 
leaves but sessile, reduced in size only slightly along ths 
stem. Involucre shallowly hemispherical, comprised of ca. 
30 phyllaries in 1 series; phyllaries with prominent mid- 
veins, linear, 5-7 mm. long, 0.7-1.0 mm. wide, with recurved, 
acuminate apices; abaxial surfaces sparsely pubescent with 
glandular multiseriate and nonglandular uniseriate tri- 
chomes, the adaxial surface glabrous. Ligulate florets 20- 
45, the corolla drying pink to lavender, the limb linear, 5-9 
mm. long, 1-2 mm. wide, often trifurcate at the blunt apex; 
tubular portion of the corolla ca. 1.5 mm. long, the style 
projecting well out of the orifice. Disc florets numerous, 
forming a disc 11-14 mm. in diameter ; corollas pale yellow, 
funnel-form at anthesis, 2.5-3 mm. high, the 5 lobes 0.65- 
0.7 mm. high; anthers free at anthesis, the thecae 0.8-0.85 
mm. long with a short appendage, 0.2-0.25 mm. long ; pollen 
98% normal, 19-24 » in diameter; style branches 0.65-0.75 
mm. long, the tips deltoid, collecting hairs not extending to 
the apices. Achenes ovoid, 4-ribbed (the ribs corresponding 
to 4 vascular traces); surface finely hirsute with short 
bristles; carpopodia 0.08-0.1 mm. high, 0.18-0.24 mm. 
broad; pappus single, comprised of 14-20 lucid, barbellate 
bristles up to 3.5 mm. long; pappus of the ligulate florets 
slightly exceeding the involucre, pappus of the disc florets 
equaling or slightly exceeding the disc corollas. 


On the basis of its morphology, Erigeron Hultenii ap- 
pears to be transitional between E. peregrinus (Pursh) 
Greene and species of the monocephalous complex, which 
centers around F. grandiflorus and includes, among other 
species, E. simplex Greene, E. Muirii A. Gray, E. hyper- 
boreus Greene, and E. wniflorus L. ssp. eriocephalus (Vahl) 
Cronquist. Erigeron peregrinus is widespread in cordilleran 
North America, and Cronquist (1947, 1968) has considered 


1973] New Erigeron — Spongberg 119 


it to be the least derived species of sect. Erigeron. In its 
four-bundled achene vascularization, its linear or strap- 
shaped ligulate corollas, and its cauline leaves that remain 
relatively unreduced in size at progressively higher nodes, 
E. Hultenii resembles E. peregrinus. In its monocephalous 
habit, its reduced stem height and internode distance, its 
disc floret and carpopodium morphology, and its reduced 
number of pappus bristles, E. Hultenii is aligned with spe- 
cies of the monocephalous complex. 

Judging from correlations between pollen size, pollen 
abortion percentages, chromosome number, and breeding 
behavior data obtained in studies of other monocephalous 
Erigeron species (Spongberg, 1971), E. Hultenii is prob- 
ably a sexual diploid or tetraploid (2n = 18 or 36). Its 
close morphological resemblance to some plants of E. grand- 
iflorus from Alaska, a taxon comprised of apomictic tri- 
ploids (2n = 27) in the southern, Rocky Mountain part of 
its range (and, on the basis of pollen data, apomictic and 
triploid in Alaska as well), suggests that E. Hultenii may 
have been involved in the synthesis of certain apomictic 
races of E. grandiflorus. Although known only from the 
type locality (narrow endemism is frequent in Erigeron), 
this hypothesis assumes E. Hultenii had a once wider dis- 
tribution. 


LITERATURE CITED 


CRONQUIST, A. 1947. Revision of the North American species of 
Erigeron, north of Mexico. Brittonia 6: 121-302. 

1968. The evolution and classification of flowering 
plants, x + 396 pp. Boston. 

SPONGBERG, S. A. 1971. A systematic and evolutionary study of 
North American arctic and alpine monocephalous species of Erige- 
ron (Compositae). Unpubl. Ph.D. Dissertation, Univ. North Caro- 
lina, Chapel Hill. 


ARNOLD ARBORETUM 
HARVARD UNIVERSITY 
CAMBRIDGE, MASSACHUSETTS 02138 


TRIBAL CLASSIFICATION OF TRIOSTEUM 
(CAPRIFOLIACEAE) 


WALTER H. LEWIS and PAUL R. FANTZ 


Traditionally the small genus Triostewm of disjunct 
Asian and eastern North American distribution has been 
included with Viburnum in the tribe Viburneae (Fritsch, 
Bot. Centralbl 50: 137-139, 1891; Wagenitz, Engler's 
Syllabus der Pflanzenfamilien, 473-475, 1964). This classi- 
fication has been questioned in recent years as research 
from several disciplines has extended our understanding 
of the Caprifoliaceae, research to which we are now able 
to add data from cytology. 

Wilkinson (Amer. J. Bot. 36: 481-489, 1949) concluded 
that Triosteum differs too greatly from Viburnum by floral 
anatomy and morphology to be included in the same tribe. 
She found the former generalized and unspecialized com- 
pared with the unusual and complex structure of the flower 
of Viburnum. The irregular corolla and conspicuous sepal 
lobes of Triostewum reminded Wilkinson of some members 
of the Lonicereae and Linnaeeae, although she believed that 
the genus might be segregated as a distinct tribe. Metcalf 
& Chalk (Anatomy of the Dicotyledons 2: 152-158, 1950) 
noted that cork usually arises in the pericycle region of 
the stem of Triostewm, which is also characteristic of the 
Linnaeeae, Lonicereae and Diervilleae, whereas its origin 
is superficial in species of Viburnum. Adding further ques- 
tion to the alignment of Triostewm, Erdtman (Pollen 
Morphology and Plant Taxonomy, Angiosperms, 97-99, 
1966) noted that its pollen was essentially similar to that 
of the Linnaeeae-Lonicereae-Diervilleae group and he in- 
cluded Triosteum in the Lonicereae. Moreover, Bohm & 
Glennie (Canad. J. Bot. 49: 1799-1807, 1971) found the 
presence of only flavone glycosides in Triosteum, which 
supports its removal from the Viburneae and relocation 
closer to Lonicera. In fact, Hutchinson (The Genera of 
Flowering Plants 2: 81-89, 1967) suggested that Triosteum 
is probably little more than a reduced Lonicera, with 


120 


1973] Triosteum — Lewis & Fantz 121 


annual stems from a woody perennial rhizome, and the 
connate leaf-bases similar to those of the more advanced 
species of Lonicera. On this basis, however, Hutchinson 
separated the genus as the monotypic tribe Triosteae. 

Members of the family differ in chromosome size, base 
number and ploidy. Sambucus and Viburnum have chromo- 
somes much larger than all other genera varying in length 
from 4-5 » (Sax & Kribs, J. Arnold Arb. il: 147-153, 
1930) with x — 8 or more commonly 9 for Viburnwm 
(Egolf, J. Arnold Arb. 43: 132-172, 1962) and x — 18 and 
19 for Sambucus (Ourecky, Amer. J. Bot. 57: 239-244, 
1970). The remaining species studied have chromosome 
sizes averaging only 1-2 ». Of these, members of the Lin- 
naeeae typically have z = 8 primarily at the polyploid 
level, or less frequently x — 9 (Symphoricarpus) and then 
commonly polyploid. The Diervilleae characteristically have 
r — 9 and like Symphoricarpus are commonly polyploid. 
The Lonicereae also predominantly have x — 9, but largely 
at the diploid level. 

To these results we add the chromosome number of 
Triosteum: 2n = 18 (x = 9) based on T. perfoliatum L. 
from Missouri (Boone Co. Columbia, Dunn s.m. (MO); 
Dallas Co., Bennett Springs State Park, Kastler s.n. (MO) ; 
Franklin Co., Gray Summit Arboretum, Fantz 1386 (M0); 
and Maries Co., ca. 2 miles NE of Vienna on Rt. 2, Fantz 
1128 (MO)). All counts are from root squashes, one plant 
from each population. 

As the chromosomes of Triosteum approximate the size 
of those found for members of the Linnaeeae-Lonicereae- 
Diervilleae group in agreement with data from gross mor- 
phology, anatomy, palynology and chemistry, the exclusion 
of the genus from the Viburneae is clear. Having a base 
number of 9 and ploidy level of 2x, similar to members of 
the Lonicereae, we propose the inclusion of Triostewm in 
this tribe. We find no evidence for its separation as a 
unique tribe. 


DEPARTMENT OF BIOLOGY, WASHINGTON UNIVERSITY AND 
THE MISSOURI BOTANICAL GARDEN 
ST. LOUIS, MISSOURI 63110 


CHROMOSOME PAIRING IN OBLIGATELY 
APOGAMOUS FERNS; PELLAEA ATROPURPUREA 
AND PELLAEA GLABELLA VAR. GLABELLA 


SYLVIA JANE RIGBY 


Obligate apogamy is a type of asexual reproduction, which 
is found in several genera of ferns, including Pellaea, the 
cliff brakes. Among the species of this genus which show 
obligate apogamy are two which can be found in north- 
eastern North America, namely Pellaea atropurpurea (L.) 
Link and P. glabella Mett. ex Kuhn. As part of a more 
general investigation of these two species (Rigby 1968) 
cytological studies were made on their sporangia during 
sporogenesis. 

In obligately apogamous ferns the sporophyte arises 
directly from the gametophyte without fertilization, while 
viable spores are still produced by meiosis as in sexually 
reproducing ferns. Therefore, a compensating mechanism 
is necessary to keep the chromosome number from being 
repeatedly reduced by meiosis from one generation to the 
next. Such a mechanism was observed by Steil (1919) and 
described in detail by Manton (1950). In the final mitotic 
division before meiosis the chromosomes of the spore 
mother cells divide, but the cells themselves do not, so that 
restitution nuclei are formed. Thus, instead of the spo- 
rangium containing sixteen spore mother cells each with 
the same chromosome number as the sporophyte plant, 
the sporangium contains only eight spore mother cells, 
each with double the sporophytic chromosome number. 
These spore mother cells can then go through a regular 
meiosis, producing thirty-two viable spores, each with the 
same number of chromosomes as the sporophyte. 

This type of compensating mechanism does not occur in 
all the sporangia of the apogamous plant. Of the four types 
of sporangia found by Manton (1950) in varying propor- 
tions in the apogamous ferns she investigated, the two 
which seem to be most useful for cytogenetic studies are: 


122 


1973] Apogamous Ferns — Rigby 123 


Type One— There is no premeiotic doubling of the 
chromosome number. Sixteen spore mother cells are formed 
and meiosis begins, but many of the chromosomes do not 
pair. This lack of pairing and irregular disjunction leads 
to spore abortion. 

Type Two — The premeiotic doubling of the chromosome 
number gives eight spore mother cells, in each of which 
meiosis can take place with regular pairing between the 
newly doubled chromosomes. Thirty-two viable spores are 
produced. 

Type Two sporangia, with their regular bivalents, can 
be used to determine the chromosome number of the species. 
In the Type One sporangia the chromosomes may not con- 
sist of two homologous sets, for pairing is irregular. Since 
many apogamous ferns are triploid or of a higher ploidy 
level and may owe their origin to hybridization, Manton 
(1950) suggests that the degree of homology between the 
genomes of the original parents of the cross may be deduced 
from examination of the pairing patterns in Type One 
sporangia. 

Pellaea atropurpurea is an apogamous triploid, and the 
most common form of P. glabella var. glabella (the eastern 
variety of this species) is an apogamous tetraploid. The 
chromosome numbers of these two taxa have been estab- 
lished as 87 for P. atropurpurea and 116 for P. glabella 
var. glabella (Manton 1950; Tryon & Britton 1958). 
During the present study Type One sporangia of the two 
taxa were examined to ascertain, as far as possible, the 
relative proportions of univalents, bivalents and multi- 
valents present in each. It was thought that these pairing 
patterns might give some indication as to the origin of 
these taxa, that is, whether they had arisen as autopoly- 
ploids or had begun as hybrids between two or more species. 


MATERIALS AND METHODS 


The sources of material used for studying the meiotic 
chromosomes were as follows: 


124 Rhodora [Vol. 75 


Pellaea atropurpurea (L.) Link 


(i) Spore-grown specimen. Source of spores, collection 
by A. Monette, Campbell’s Bay, Quebec (DAO 
26361). 


(ii) Spore-grown specimen. Source of spores, collection 
by T. M. C. Taylor, Fairmount Hotsprings, British 
Columbia (UBC 37640). 


P. glabella Mett. ex Kuhn var. glabella 

(i) Spore-grown specimen. Source of spores, collection 
by J. B. Clark, Laclede Co., Missouri (0AC 37706). 

(ii) Material collected in the field, by S. J. Rigby, Rock- 
wood, Ontario. 


Fertile pinnae were fixed in 3 parts absolute ethanol: 
1 part glacial acetic acid (previously saturated with ferric 
acetate) for 3 to 8 days at 4°C and then transferred to 
70% ethanol and stored at 4°C until examined. Sporangial 
squashes were made using the propionic-iron-haematoxylin 
technique described by Lu (1967), with the modification 
that Solution A was diluted with about an equal volume 
of 45% acetic acid, since otherwise there was a tendency 
for the iron to precipitate when the two solutions were 
mixed. Slides were made permanent by ringing the cover 
slip with Hoyer’s mounting medium (formula given in 
Alexopoulos & Beneke 1952, p. 2). The chromosomes were 
observed and photographed under interference contrast 
(Zeiss-Nomarski) with an oil immersion objective (Pla- 
napo, N.A. 1.3). Because the chromosomes in the Type One 
spore mother cells do not separate readily at metaphase, it 
was hoped that the three dimensional effect of the inter- 
ference contrast might make it easier to distinguish be- 
tween two or more overlapping chromosomal associations. 
In eases where the cells could not be flattened into a single 
focal plane, photographs were taken of the same cell in 
several focal planes. Tracings from these photographs 
were compared with the original slides and used to make 
the explanatory diagrams. 


1973] 


Apogamous Ferns — Rigby 


125 


C d 

Figure 1. Spore mother cell from Type One sporangium of 
Pellaea atropurpurea (Campbell’s Bay material) at meiosis. X 2000. 
a, b, c: Cell photographed in three different focal planes. d: Ex- 


planatory diagram, Interpretation: Univalents (black) 20, Bivalents 
(white) 20, Trivalents (stippled) 9. 


126 Rhodora [Vol. 75 


Figure 2. Spore mother cell from Type One sporangium of Pel- 
laea glabella var. glabella (Laclede Co. material) at meiosis. 2000. 
(Cell ruptured by pressure.) a, b, c: Cell photographed in three 
different focal planes. d: Explanatory diagram, Interpretation: 
Univalents (black) 15, Bivalents (white) 36, Trivalents (stippled) 
7, Quadrivalents (hatched) 2. 


127 


Apogamous Ferns — Rigby 


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128 Rhodora [Vol. 75 


OBSERVATIONS 


The chromosomes of the Type One spore mother cells tend 
to remain clumped together at meiosis and do not spread 
out well when squashed. Nine Type One cells were found 
that spread enough to be suitable for analysis, three from 
P. atropurpurea and six from P. glabella var. glabella. 
Photographs and explanatory diagrams of one cell from 
each of these taxa are shown in Figures 1 and 2, and the 
chromosomal association frequencies are summarized in 
Table 1. Even with the limited amount of material studied, 
it appears that in both taxa at least two thirds of the 
chromosomes are involved in some kind of association. 
Bivalents are more common than the higher order multi- 
valents, but the relative frequencies of bivalents and uni- 
valents in the two samples of P. glabella var glabella differ: 
in the Rockwood material the univalents outnumber the 
bivalents, while in the material from Laclede County the 
proportions are reversed. In the Campbell's Bay material 
of P. atronurpurea the freauencies of univalents, bivalents 
and trivalents can be related to the basic chromosome 
number of 29, tvpical of the genus, but the material from 
Fairmount Hotsprings shows more than 29 associations. 


DISCUSSION 


Although the sample of Type One spore mother cells was 
not large enough for a meaningful statistical analysis, even 
within this small sample certain trends are indicated. In 
both species the proportions of different types of chromo- 
somal associations show some variation from one population 
to another. In P. atropurpurea the material from Campbell's 
Bay suggests three sets of chromosomes, of which two sets 
possess a high degree of homology to each other, as shown 
by the formation of bivalents, while the third set has a 
lesser degree of homology to the other two, so that only 
about a quarter of its chromosomes go into the formation 
of trivalents, with the remainder being present as uni- 
valents. This type of chromosomal association might 


1973] Apogamous Ferns — Rigby 129 


indicate that P. atropurpurea arose from a cross between 
two distinct, though related species, one of which con- 
tributed two sets of chromosomes to the hybrid while the 
other contributed only one set. However, the cell from the 
Fairmount Hotsprings material, with its 31 bivalents, does 
not fit in with this pattern but rather suggests a more 
complex type of relationship. 

In P. glabella var. glabella several probable quadriva- 
lents were seen in the Type One spore mother cells from 
both populations, although their total proportion was low 
(about 10%). The presence of these quadrivalents sug- 
gests at least some homology among the four sets of 
chromosomes which originally contributed to the chromo- 
some complement of this tetraploid. If quadrivalents, tri- 
valents and bivalents are all cosidered, then on the average 
about 73% of the chromosomes in each cell are involved 
in some kind of chromosomal association, which again 
would suggest a fairly high degree of homology among 
these chromosome sets. This seems to indicate that P. 
glabella. var. glabella originated either as an autopolyploid 
or from a cross between two closely related taxa. Because 
the species complex P. glabella includes two morphologi- 
cally distinguishable, sexually reproducing diploid entities, 
namely var. occidentalis (E. Nelson) Butters, found in 
northwestern North America, and a diploid form of var. 
glabella, which so far has been reported only from Missouri 
(Wagner et al. 1965), possible ancestors for the tetraploid 
form of var. glabella can be found within the species com- 
plex. 

While the patterns of choromosomal association observed 
in this study may suggest certain degrees of homology be- 
tween the chromosome sets within each polyploid, it should 
be borne in mind that the degree of homology among the 
original sets of chromosomes of each polyploid may not be 
the only factor affecting chromosome pairing. Neither 
P. atropurpurea nor P. glabella var. glabella is a newly 
formed hybrid (or autopolyploid) but rather each has 
been reproducing asexually for many generations. During 


130 Rhodora [Vol. 75 


this time changes may have taken place in the chromosome 
structure and in the genetic factors which govern chromo- 
some pairing, so that the chromosomal associations seen at 
meiosis in the Type One spore mother cells are no longer 
fully indicative of the degree of homology between the 
chromosome sets involved. An indication that changes have 
taken place since these polyploids were first formed is the 
variation in chromosomal association frequencies between 
populations within each taxon. In fact, as Wagner (1963, 
1970) points out, even in newly formed hybrids pairing 
may be influenced by genetic factors and therefore may 
not be a precise indicator of the degree of chromosome 
homology. The cytological observations made in this study 
cannot therefore be regarded as conclusive proof of the 
amount of homology between the various chromosome sets 
of the polyploids, but could serve as useful supporting 
evidence when used in conjunction with other types of 
observation, such as morphological characteristies. 


SUMMARY 


In obligately apogamous ferns the chromosome number 
is kept constant from generation to generation by a pre- 
meiotic doubling of the chromosome complement in some 
(but not all) of the spore mother cells, which then go 
through meiosis to produce viable spores. Spore mother 
cells in which such premeiotie doubling does not take place 
have irregular chromosome pairing at meiosis. 

Pellaea atropurpurea is an apogamous triploid, while the 
most common form of P. glabella var. glabella is an apo- 
gamous tetraploid. In spore mother cells of these two poly- 
ploids — in which the chromosome number had not doubled 
when the chromosomes were observed at meiosis — unival- 
ents, bivalents and multivalents were all seen to be present. 
The patterns of chromosomal association observed could 
be interpreted as indicating homologies between the 
chromosome sets of the original parents of these polyploids, 
or they may be, at least in part, due to changes in chromo- 
some structure and in the genetic factors governing 


1973] Apogamous Ferns — Rigby 131 


pairing that have taken place through many generations 
of asexual reproduction. 


ACKNOWLEDGEMENTS 


The author wishes to express her appreciation to Dr. 
D. M. Britton and Dr. Alice Tryon for their assistance 
and advice in this study. Financial assistance from the 
National Research Council of Canada, the Ontario Graduate 
Fellowship Program and the Ontario Department of Uni 
versity Affairs is gratefully acknowledged. 


LITERATURE CITED 


ALEXOPOULOS, C. J., & E. S. BENEKE. 1952. Laboratory manual 
for introductory mycology. Burgess Publishing Co., Minneapolis. 

Lu, B. C. 1967. The course of meiosis and centriole behavior 
during the ascus development of the ascomycete Gelasinospora 
calospora. Chromosoma 22: 210-226. 

MANTON, I. 1950. Problems of cytology and evolution in the Pteri- 
dophyta. Cambridge Univ. Press, Cambridge. 

RIGBY, S. J. 1968. An investigation of Pellaea glabella Mett. ex 
Kuhn and Pellaea atropurpurea (L.) Link and their relation- 
ships. Master’s thesis. University of Guelph. 

STEIL, W. N. 1919. A study of apogamy in Nephrodium hirtipes 
Hk. Ann. Bot. 33: 109-132. 

Tryon, A. F. & D. M. BRITTON. 1958. Cytotaxonomic studies on 
the fern genus Pellaea. Evolution 12: 137-145. 

WAGNER, W. H. JR. 1963. Biosystematics and taxonomic categories 
in lower vascular plants. Reg. Veg. 27: 63-71. 

1970. Evolution of Dryopteris in relation to the 

Appalachians. pp. 147-192, in P. C. Holt (ed.), The distribu- 

tional history of the biota of the southern Appalachians, Part II: 

Flora. Research Division Monograph 2, Virginia Polytechnic 

Institute and State University. 

, D. R. FARRAR, & K. L. CHEN. 1965. A new 

sexual form of Pellaea glabella from Missouri. Am. Fern Jour. 

55: 171-178. 


DEPARTMENT OF BOTANY AND GENETICS 
UNIVERSITY OF GUELPH 
GUELPH, ONTARIO, CANADA 


THE PAST AND PRESENT GEOGRAPHICAL 
DISTRIBUTION OF PETALOSTEMON FOLIOSUS 
AND NOTES ON ITS ECOLOGY 


JERRY M. BASKIN AND CAROL C. BASKIN 


Petalostemon foliosus Gray, a member of the Legumi- 
nosae, is a rare plant species that now appears to be 
restricted to a few localities in central Tennessee and 
northern Alabama. However, at one time it also occurred 
in at least four counties (LaSalle, Kankakee, Kane and 
Will) in northeastern Illinois. Herbarium specimens from 
these Illinois counties have been cited by Gambile (1953) 
and (or) Wemple (1965), and specimens are still present 
in various herbaria (F, GH, ISC, US, NY and ILL). Among 
the collections in these herbaria the latest one is dated 
12 September 1912 (L. M. Umbach 5715, in the Iowa State 
University Herbarium, collected from river flats, Romeo, 
Will County, Illinois). This may possibly represent the last 
collection of P. foliosus in Illinois. Jones (1963) and 
Wemple (1965, 1970) state that the species probably is 
now extinct in Illinois. 


Although the species has persisted until the present in 
several localities in central Tennessee and in at least two 
locations in northern Alabama, it has not been collected 
widely and, therefore, very little is known about its dis- 
tribution in central Tennessee and northern Alabama. In 
a recent revision of the genus, Wemple (1965) cites speci- 
mens from Davidson, Franklin and Rutherford Counties in 
middle Tennessee and from Knox County in east Tennes- 
see. All of the specimens that he examined from middle 
Tennessee were collected since about 1915, from a cedar 
glade area across from Stones River Park, 2 miles north- 
west of Murfreesboro, Rutherford County, Tennessee. 
Sharp et al. (1960) list the species as occurring in David- 
son and Rutherford Counties in central Tennessee and 
Mahler (1970) lists it from the same two counties and 


132 


1973] Petalostemon — Baskin & Baskin 133 


from Knox County in east Tennessee. Baskin and Caudle 
(1967) first reported the species from Alabama, in Frank- 
lin and Morgan Counties. 

While making botanical observations in central Tennes- 
see, we have found the species in several counties where it 
previously has not been reported. These counties include: 
Wilson County, Cedars of Lebanon State Forest, 6 August 
1971 (Baskin & Baskin 1177); Maury County, 2.7 miles 
east of Columbia on Co. Rd. 50, 23 August 1966 (Baskin 
& Caudle 522) ; Marshall County, 1 mile east of U. S. 431 
on St. Rd. 99, 18 August 1970 (Baskin & Baskin 869). In 
addition, there is a specimen from Williamson County, 
Tennessee in the Vanderbilt University herbarium (VDB) 
from along Sneed Road west of Hillsboro Avenue toward 
the Harpeth River (Kral 29001). Actually, this popula- 
tion was discovered earlier by Dr. Elsie Quarterman of 
Vanderbilt University, but apparently she did not place 
a specimen in the herbarium at Vanderbilt. 

During August 1971, we visited the previously known 
locations of P. foliosus in Franklin and Morgan Counties 
Alabama, and Marshall, Maury, Williamson and Wilson 
Counties Tennessee to see if the species was still present 
in these locations. In all the locations visited we found 
at least a few plants of the species. From our recent obser- 
vations on living populations of the species and from avail- 
able specimens and literature, we have constructed a map 
of the known past and present geographical distribution 
of the species (Fig. 1). Although we do not know of 
specific locations for P. foliosus in Rutherford County, 
Tennessee, at least two collections were made there in the 
1960's. One, L. E. Franklin & J. D. Freeman 2481, on 
26 July 1962 (VDB), and the other, D. Isley & D. Wemple 
9420, on 11 September 1964 (Isc). Although there have 
not been any recent colleetions of P. foliosus in Davidson 
and Franklin Counties, Tennessee, there is no reason to 
believe that the species is not present there because seem- 
ingly suitable cedar glade habitats are available, and the 
species does occur in nearby counties. 


134 Rhodora [Vol. 75 


9 e Ohio 
Ind. 


s ° 
T enn. o; 
o O 


Miss. Ala. Ga. 


o Living populations, I97l 


* Herbarium records 


Fig.l. Past and present geographical distribution of Petalos- 
temon foliosus. 


1973] Petalostemon — Baskin & Baskin 135 


The occurrence of P. foliosus on the “U. T. Farm" in 
Knox County, Tennessee is represented on the distribution 
map; it probably was transplanted to that site and no 
longer survives there. The only specimen that seems to have 
been collected from this location is 20 September 1938, 
Lahner (TENN). Sharp et al. (1960) do not cite this loca- 
tion. 

Gleason (1952) and Gleason and Cronquist (1963) also 
give the species as occurring in Ohio, but its occurrence 
there is doubtful. None of the other literature that we 
examined refers to any specimens collected in Ohio, and 
Weishaupt (1968) does not include the species in her 
Vascular Plants of Ohio. 

The habitat and distribution of P. foliosus have been 
described in various taxonomic and floristic works as fol- 
lows: river banks, Interior Low Plateau, Tennessee to 
Illinois (Small, 1933) ; rocky hills, glades and river banks, 
northern Illinois to Tennessee (Fernald, 1950); river 
banks, Illinois, Ohio and Tennessee (Gleason, 1952; Glea- 
son and Cronquist, 1963); river banks and gravelly soil, 
rare or probably now extinct in Illinois (Jones, 1963) and 
cedar glades, barrens, Tennessee, northern Illinois and 
Alabama (Mahler, 1970). 

In central Tennessee and northern Alabama P. foliosus 
always is associated closely with cedar glades. On the 
typical Lebanon limestone glades (for description see 
Quarterman, 1950), P. foliosus usually occurs at the edges 
of open glades in the transition zone between open glades 
and glade thickets or woods, a habitat that is shaded during 
at least part of the day. In this situation P. foliosus often 
is associated with Astragalus tennesseensis Gray, Ascle- 
pias verticillata L. and Sporobolus vaginiflorus (Torr.) 
Wood. In one glade in Marshall County, Tennessee that 
is more prairie-like than typical Lebanon limestone glades, 
P. foliosus occurs in an opening that receives full daylight 
throughout the growing season, as well as near a shrub 
thicket composed of Symphoricarpos orbiculatus Moench, 
Rhus aromatica Ait. and Forestiera ligustrina (Michx.) 


136 Rhodora [Vol. 75 


Poir. The open, prairie-like portions of this glade, where 
P. foliosus grows, are dominated by Panicum capillare L. 
Here a layer of mineral soil (30-45 em deep) overlies solid 
bedrock. Other plants on this glade with prairie affinities 
include Cacalia tuberosa Nutt., Helenium autumnale L. 
and Desmanthus illinoensis (Michx.) MacM. Petalostemon 
foliosus grows in a somewhat similar open, prairie-like 
situation in Morgan County, Alabama and Williamson 
County, Tennessee. In Morgan County, Alabama its most 
frequent associates are Cassia fasciculata Michx., Rud- 
beckia triloba L. and R. hirta L., and in the glade in Wil- 
liamson County, Tennessee Astragalus tennesseensis and 
Delphinium virescens Nutt. are frequent associates. 

Since apparently very little is known about the ecological 
life cycle of P. foliosus, we believe that it is not inappro- 
priate to report here our observations on the phenology of 
this species in the field. In March new buds located at 
or slightly below the soil surface at the base of the previous 
year’s growth (shoot) begin growth. By early June the 
new shoot is 15-20 cm tall. Growth of the shoot in height 
continues until July by which time plants are typically 
40-65 cm tall. An individual plant usually bears one shoot 
with many inflorescences, but some plants may have several 
shoots with many inflorescences each. Flower buds ap- 
parently are formed in late June or early July and by 
mid-July large flower buds are visible. Flowering begins 
in late July or early August and continues through August. 
The peak of flowering is approximately mid-August. The 
only insect that we have observed visiting the flowers of 
this species is a bumblebee (Bombus sp.). By early Octo- 
ber seeds are ripe and by mid-October the shoots are dead. 
However, after the shoots die they remain erect and hold 
the fruiting heads into the winter. Seeds are not shed 
immediately upon ripening but are dispersed from the 
plants from late fall to early spring. Our observations 
indicate that seed set is good. 

Germination in the field occurs in April and by late May 
the seedlings have several leaves. Numerous seedlings of 


1973] Petalostemon — Baskin & Baskin 137 


P. foliosus were present in the Morgan County, Alabama 
population on 6 April 1972. These seedlings were in the 
cotyledon or first leaf stage. As in many members of the 
Leguminosae (Crocker and Barton, 1953), seed coats of 
most of the freshly-matured seeds of P. foliosus are im- 
permeable to water, but will germinate to nearly 100% 
when mechanically scarified. However, in nature the seed 
coat must be softened by natural agents before the seeds 
can germinate. All the seeds produced in a particular seed 
crop do not soften by the following spring, and it takes 
several years for the coats of all seeds in a particular crop 
to soften. We planted 400 seeds on greenhouse potting soil 
in flats on 11 October 1969 and placed them on a bench in 
a non-heated room of a greenhouse at the University of 
Kentucky. Eleven of the seeds germinated during the fall 
of 1969, indicating that only a very small percentage of the 
freshly-matured seeds had permeable or soft coats. After 
overwintering in the greenhouse, 21 additional seeds germ- 
inated in the spring of 1970, and to date after three over- 
winterings) only 89 of the seeds have germinated. During 
the spring of 1970 most of the germination in the non-heat- 
ed room of the greenhouse occurred between 8 April and 6 
May when the average daily maximum and minimum tem- 
peratures in the greenhouse were 23.7 and 11.3?C, respec- 
tively. During the spring of 1971 most of the germination 
occurred bewteen 1 April and 20 April when the aver- 
age daily maximum and minimum temperatures in the 
greenhouse were 23.6 and 8.3°C, respectively. 

It appears that very few of the seedlings survive to ma- 
turity as relatively very few juvenile plants can be found 
in the field in the summer and fall. Our observations indi- 
cate that seedlings are drought intolerant and that many 
of them are killed by summer drought. We observed dead 
seedlings during a short dry period in late May of 1971 
and others that were badly wilted. Since the species does 
not reproduce vegetatively and since very few seedlings 
survive, perennation is a most important factor in main- 
taining populations of this species. Further evidence that 


138 Rhodora [Vol. 75 


suggests that seedling survival is poor is that the num- 
ber of plants in the small populations of this species does 
not seem to be increasing. For example, in the populations 
of P. foliosus in Franklin County, Alabama and in Maury 
County, Tennessee where the populations consist of less 
than 25 plants each, there was no noticeable change in 
number of plants between 1966 when we first discovered 
the populations and 1971 when we last observed them. 

Petalostemon foliosus is a rare and *endangered" species. 
Although one of the populations (Morgan County, Ala- 
bama) consists of hundreds of individual plants and an- 
other one (Williamson County, Tennessee) has several 
dozen plants in it, all the other populations that we have 
seen have less than 25 plants. As we have mentioned in à 
previous article on Astragalus tennesseensis (Baskin, 
Baskin and Quarterman, 1972), »nother rare cedar 
glade endemic, the glade habitats in middle Tennessee 
rapidly are being destroyed or disturbed. Some of the 
glade habitats that have been destroyed once supported 
populations of P. foliosus. For example, until 1967 there 
was a sizeable population of P. foliosus on a glade 2 miles 
northwest of Murfreesboro in Rutherford County, Tenn. 
In 1967 this area was bulldozed in preparation for con- 
struction of a box factory and, of course, the entire popu- 
lation was destroyed. 

Both the Morgan and Franklin County populations of 
P. foliosus are in a precarious position as is true for the 
one population of Astragalus tennesseensis that we know 
of in Alabama. The large population, in Morgan County, 
is located mostly along side a narrow, gravel road and if 
the road is widened to any extent most, if not all, of this 
population will be destroyed. The population of P. foliosus 
in Franklin County, Alabama is located on the right-of-way 
of County Road 79 and should this two lane road be 
widened this small population (17 plants in 1971) will 
undoubtedly be destroyed. 

The past and present geographical distribution of P. 
foliosus and A. tennesseensis are very similar. Both species 


1973] Petalostemon — Baskin & Baskin 139 


once had two centers of distribution; one in middle Ten- 
nessee and northern Alabama and another in northern 
Illinois. Both apparently have become extinct in Illinois 
and now are restricted to middle Tennessee and northern 
Alabama. Undoubtedly habitat destruction in the present 
geographical range of these two species will continue with 
obvious consequences. It seems reasonable to predict, then, 
that the two species will become extremely rare and maybe 
even extinct in this century. 

Travel funds for this project were provided by a grant 
from the University of Kentucky Research Committee. 
This support is gratefully acknowledged. 


LITERATURE CITED 


BASKIN, J. M. and C. CAUDLE. 1967. Petalostemon foliosus in Ala- 
bama. Rhodora 69: 383-384. 

BASKIN, C. C., J. M. BASKIN and E. QUARTERMAN. 1972. Observa- 
tions on the ecology of Astragalus tennesseensis. Amer. Midl. 
Natur. 88: 167-182. 

Crocker, W. and L. V. BARTON. 1953. Physiology of seeds. Chron- 
ica Botanica Company, Waltham, Mass. 267 pp. 

FERNALD, M. L. 1950. 8th Ed. Gray’s manual of botany. Amer. 
Book. Co., New York. 1632 pp. 

GAMBILE, W. G., JR. 1953. The Leguminosae of Illinois. Illinois 
Biological Monographs. Vol. XXII, No. 4. The University of 
Illinois Press, Urbana. 117 pp. 

GLEASON, H. A. 1952. The new Britton and Brown illustrated flora 
of the northeastern United States and adjacent Canada. The 
New York Botanical Garden. Lancaster Press, Inc., Lancaster, 
Penn. Vol. III. 589 pp. 

and A. CRoNQUIsT. 1963. Manual of the vascular 
plants of northeastern United States and adjacent Canada. D. 
Van Nostrand Comp., Inc., Princeton, N. J. 810 pp. 

Jones, G. N. 1963. Flora of Illinois. 3rd Ed., The American Mid- 
land Naturalist Monograph No. 7. The University of Notre 
Dame Press, Notre Dame, Indiana. 401 pp. 

MAHLER, W. F. 1970. Manual of the legumes of Tennessee. Jour. 
Tenn. Acad. Sci. 45: 65-96. 

QUARTERMAN, E. 1950. Major plant communities of Tennessee 
cedar glades. Ecology 31: 234-254. 

SHARP, A. J., R. E. SHANKs, H. L. SHERMAN and D. H. Norris. 
1960. A preliminary checklist of dicots of Tennessee. ms. 


140 Rhodora [Vol. 75 


SMALL, J. K. 1933. Manual of the southeastern flora. University 
of N. C. Press, Chapel Hill. 1554 pp. 

WEISHAUPT, C. G. 1968. Vascular plants of Ohio (Revised Edi- 
tion). Wm. C. Brown Book Co., Dubuque, Iowa. 280 pp. 
WEMPLE, D. K. 1965. Revision of the genus Petalostemon (Legu- 
minosae). Ph.D. Thesis, Iowa State University (Libr. Congr. 
Card No. Mic. 66-3015). 321 pp. Univ. Microfilms. Ann Arbor, 

Mich. 


1970. Revision of the genus Petalostemon (Legu- 
minosae). Iowa St. Jour. Sci. 45: 1-102. 


DEPARTMENT OF BOTANY 
UNIVERSITY OF KENTUCKY 
LEXINGTON, KENTUCKY 40506 


BOOK REVIEW 


FLORA OF THE GALAPAGOS ISLANDS! 


This splendid volume was published in 1971. The first 
copies to reach the Galápagos Islands were brought there 
in June of that year by Dr. William A. Weber. In the in- 
teresting review that later appeared under his name in 
Science (1972) he explained that he was able to test the 
book on location with the first study tour conducted there 
for university credit. 

Quite by chance the present reviewer was privileged to 
make a similar test of the book in June and July of that 
same year. With my wife, I arrived at Baltra Island in the 
Galápagos on June 22, 1971 to take the two weeks' cruise 
on the “Golden Cachalot?” under Lindblad auspices. The 
first day out the owner, Mr. David Balfour, learning that 
I was a botanist, placed an unused copy of the book now 
under review in my hands whereupon the excursion be- 
came not only a most pleasant experience in natural history 
but an education in Galápagos botany. The copy of the 


'"WiGGINS, IRA L. and DUNCAN M. PORTER. 1971. Stanford Univer- 
sity Press, Stanford, California. XVII, 998 pp. -- plates. 
$37.50. 


1973] Book Review — Hodgdon 141 


Galápagos Flora I used may have been one brought by Dr. 
Weber, but if it were not, it must have reached there about 
the same time. I can lay claim to being the first "tourist" 
to have made extended use of the Flora in understanding 
the plants of the Galápagos. 

The Flora of the Galápagos Islands became a focus of 
attention during many relaxed moments on that memorable 
cruise among the isJands. Specimens were brought back 
from journeys ashore by enthusiastic fellow tourists and 
the very helpful staff were much interested in increasing 
their considerable stores of knowledge of the plant life of 
the islands. Mr. Balfour in particular supported my en- 
deavors by supplying a smail plant press and, from time to 
time, bringing back some exceptionally interesting speci- 
mens to put in it. 

It should be emphasized that the islands have been set 
aside as a National Park by Ecuador. Collecting there- 
fore had to be limited to fragments of plants or to few 
specimens in the case of abundant species. For ready iden- 
tifieation more complete specimens would have been more 
suitable. Surprisingly, late June and early July, 1971, 
proved to be somewhat cool and moist with occasional 
showers even at sea level. Therefore, the drying of plants 
was helped greatly by putting the press in a compartment 
above the engine room. 

In view of the fact that I had made no plans to carry on 
any botanical work at all and that I had no knowledge, on 
reaching the Galápagos Islands, that a Flora of it had 
even been thought of, it is interesting to record that some 
120 species of vascular plants were collected and most of 
these were identified during the cruise. More than 40 of 
the 120 proved to be endemics which is a higher propor- 
tion than the 32% endemism recorded by Wiggins and 
Porter for the entire vascular flora. Probably at least 100 
other species were identified but not collected during the 
cruise. This speaks well for the usefulness of the book to 
a botanist who had no previous acquaintance with this or 
floras of similar tropical areas. Since the known Galápagos 


142 Rhodora [Vol. 75 


flora consists of 642 species it was possible to attain some 
familiarity with nearly one third of the species of the 
islands. Had this book not been available with its work- 
able keys, abundant and clear illustrations, detailed maps 
and range data, good descriptions as well as the helpful 
discussions of the plants of the several vegetational zones, 
hardly a beginning could have been made. 

It was pointed out by Wiggins and Porter in their intro- 
duction on pp. 29 and 30 that much more field work needs 
to be carried on in the higher parts of the islands compris- 
ing the SCALESIA, MICONIA and FERN-SEDGE zones. They 
conclude (p. 30) with this statement, “There undoubtedly 
are many niches that have not yet been seen or recorded: 
Isabella especially, owing to its great area, is only minimal- 
ly known." Weber (1972, p. 290) makes this assertion after 
having used the book in the islands during his visit in 
June 1971, “The strongest impression I have from the ex- 
perience is that the book is still a preliminary treatment. 
The islands will not be adequately known until there are 
resident botanists studying the flora and until the Darwin 
Research Station is equipped for the handling, drying and 
proper storage of botanical materials." At the Darwin Sta- 
tion on Santa Cruz Island on July 3, I talked with Daniel 
Weber who had assembled a neat collection of about 1000 
specimens. He had collected in difficult parts of the islands 
and told me of many new stations to add to those in the 
Flora. Specifically he stated that he had found Pernettya 
Howellii in two localities on Isabella Island, whereas Wig- 
gins and Porter reported this endemic species only from 
Santa Cruz Island. My own limited experience convinces 
me also that the Flora of the Galápagos Islands for all 
its excellence and thoroughness, is but a beginning. On 
Santa Maria in a tidal pool near a much visited lagoon 
hardly off the beaten track I found Ruppia maritima in 
abundance. Daniel Weber did not know of this station 
and Wiggins and Porter state of the species (p. 928) 
* .. known only from Isabella and Genovesa, but to be 
expected elsewhere." 


1973] Book Review — Hodgdon 143 


Very recently Johnson and Raven (1973) have made 
extensive use of information extracted from the Flora 
about the numbers of species, percentage of endemism, 
etc., in relation to area of the different islands and in rela- 
tion to the vegetational zones where endemics and other 
species are found. They point out that almost all other 
islands which have been studied in detail have a high de- 
gree of endemism in the moist uplands, whereas in the 
Galápagos the degree of endemism in the moist uplands is 
low. They fail to make entirely clear what they mean by 
moist uplands for, according to Wiggins and Porter, the 
heaviest rainfall in the Galápagos Islands falls in the alti- 
tudinal zone dominated by Scalesia forest between 180 and 
about 400 m elevation on southern slopes and 300 to 500 
m on northern slopes. The considerable areas above this 
zone receive less rainfall than the Scalesia forest. Johnson 
and Raven (loc. cit. p. 895) state that Colinvaux in a paper 
in press *has shown that the period from about 10,000 to 
30,000 years ago (radiocarbon dates) was very much drier 
than the present. In the light of his findings we propose 
that the great majority of the nonendemic plants in the 
moist uplands have reached the Galápagos Archipelago 
within the past 10,000 years. Moist uplands in their pres- 
ent form seem to have existed in the Archipelago only dur- 
ing this period of time." If by moist uplands Johnson and 
Raven include the Scalesia forests which presumably they 
do, they should advance some explanation for the evolution 
of a modified rain forest situation dominated by several 
species of the endemic genus Scalesia, the endemic Psidium 
galapageium and another endemic, Pisonia floribunda, the 
two last named of which grew to a size and in numbers 
sufficient to be shipped from the Galapagos to the main- 
land as lumber in earlier times. Presumably this type of 
forest or at least the species which comprise it must have 
been in the process of evolution ostensibly under reason- 
ably moist conditions for more than 10,000 years. It seems 
a bit early to make sweeping generalizations about the 
time of arrival of the floras in the several zones and to 


144 Rhodora [Vol. 75 


extrapolate from the meager data yet available. Above the 
Scalesia zone and intermingling with the fern-sedge vege- 
tation of the highest parts of the islands there is a domi- 
nance of Miconia Robinsoniana and above this, amongst 
the low and dense t&n-sedge type of vegetation on several 
of the larger islands, the very beautiful tree fern, Cyathea 
Weatherbyana is found. The former was named in honor 
of Benjamin Lincoln Robinson, the first editor of Rhodora 
who prepared the first Galápagos flora while the latter 
honors the distinguished specialist on ferns, Charles A. 
Weatherby. It was particularly interesting to me to ob- 
serve these species growing side by side with the familiar 
Lycopodium clavatum and Pteridium aquilinum nearby. 
The presence of the endemic Miconia and Cyathea would 
seem to depend on reasonably moist conditions having ob- 
tained for some time in the areas where they are not grow- 
ing or in places not far away. It is also entirely probable 
that the amount of speciation that has taken place in a 
flora may not be a very good indication of the age of the 
flora. This is not to say that the upper levels of these 
islands may not have received most of their species in re- 
cent times. Further exploration in these neglected parts 
of the islands is urged by Wiggins and Porter. The per- 
centage of endemism may still prove to be low when these 
areas are carefully studied. What the Galápagos really 
need most is plenty of study and not too many guesses. 
Whatever may be our eventual understanding of the 
phytogeography of these extraordinary islands, the Wig- 
gins and Porter Flora stands as a magnificent base for 
future work. It is a sumptuous work, attractive to peruse, 
with its 16 pages including 96 individual photographs in 
color of scenic features, vegetation and individual plants. 
There is an introduction or preliminary discussion of the 
geography, physiography, geology, climate, human history, 
soils, vegetational zones, history of botanical work on the 
islands and several pages devoted to fauna. There follows 
the main body of the work arranged in the following order: 
Ferns and Relatives, Apetalae, Gamopetalae, Polypetalae, 


1973] Book Review — Hodgdon 145 


and finally Monocotyledons. The families, genera and spe- 
cies are arranged alphabetically within these 5 larger 
groupings. Many families and genera of the main body 
of the book were done by specialists and, as one might ex- 
pect, the treatments are somewhat uneven though presum- 
ably all are scholarly. Some groups are amply illustrated 
while others such as Mollugo would have benefited by more 
pictures. A few errors were noted. Some of the obvious 
ones were -pointed out by Weber in his review in Science. 

While the foregoing remarks may seem often to be 
peripheral to the intended theme — a review of the Flora 
of the Galápagos Islands — the comments are all occa- 
sioned directly or indirectly by my use of the book. This 
reviewer is probably prejudiced but he can think of no 
book in recent years that has given him more pleasure in 
the using. 


LITERATURE CITED 


JOHNSON, MICHAEL P. and PETER H. RAVEN. 1973. Science 179: 
893-895. 
WEBER, WILLIAM E. 1972. Science: 175: 290-291. 


A. R. HODGDON 
DEPARTMENT OF BOTANY 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, N.H. 03824 


UMALDY T. WATERFALL (1910-1971) 


U. T. Waterfall was born in Frederick, Oklahoma on Au- 
gust 13, 1910 and died in Stillwater, Oklahoma on October 
27,1971. He married La Clida Cotter, July 24, 1935 at Fair- 
field, Oklahoma. He is survived by his wife and three chil- 
dren. Dr. Waterfall received his public school education at 
Albion, Oklahoma where he graduated from high school in 
1931. He attended Arkansas Tech at Russelville, Arkansas 
for two years and received a B.S. from Oklahoma A & M 
(now Oklahoma State University) in Stillwater in 1935. He 
studied botany at the University of Oklahoma, Norman, and 
received his M.S. in 1942 and Ph.D. in 1956. His doctoral 
dissertation was a monographie treatment of the genus 
Physalis (Solanaceae) from which numerous papers have 
appeared in print. 

U. T. Waterfall held memberships in several professional 
organizations including: Southwestern Association of Nat- 
uralists (served as president in 1965), International So- 
ciety of Plant Taxonomists, American Society of Plant Tax- 
onomists, New England Botanical Club, Oklahoma Academy 
of Sciences (served as section editor until his death), Phi 
Kappa Phi, and Sigma Xi. In addition, he is listed in 
*Who's Who in the South and Southwest", “Leaders in Am- 
erican Science", and “World Who's Who in Science." 

Dr. Waterfall began his teaching career in 1934 at Hel- 
ena High School, moving to Central High School in Okla- 
homa City in 1936. His first five years at Central High 
School were spent teaching music. However, after receiv- 
ing his M.S. in 1942, he changed to teaching science. In 
1944, he left the teaching field for two years and worked as 
a range ecologist with the Soil Conservation Service. In the 
fall of 1946 he accepted an instructorship to teach botany 
at the University of Oklahoma. He joined the faculty at 
Oklahoma State University in 1949 where he remained un- 
til his death. During his earlier years at OSU, he com- 
muted to the University at Norman while working toward 
his Ph.D. Besides his regular duties as classroom teacher 


146 


1973] Waterfall (1910-1971) — Stanford 147 


and curator, Dr. Waterfall served on graduate committees 
of numerous students from botany and biological sciences 
to zoology, wildlife, agronomy and education. Additionally, 
he supervised the research of several masters degree candi- 
dates as well as six doctoral candidates. I was the last stu- 
dent to receive the Ph.D. under the direction of U. T. Wa- 
terfall, and consider myself fortunate to have been influ- 
enced by his knowledge and experience. 

U. T. Waterfall’s first publication was the result of num- 
erous collecting trips to various parts of Oklahoma prior to 
1939. Study of the state flora continued until his death, re- 
sulting in over 30 papers. Most of these appeared in *Rho- 
dora" and “The Proceedings of the Oklahoma Academy of 
Science", others in “Sida”, “Field and Lab" and “South- 
western Naturalist". Some of the more recent papers have 
been published under the title “Studies in the Composition 
and Distribution of the Oklahoma Flora". In 1952, “A Cat- 
alogue of the Flora of Oklahoma" was prepared by Dr. 
Waterfall and published by the Oklahoma Research Foun- 
dation. It listed 2542 taxa of specific or infraspecific status. 
Immediately following this work, he began his long study 
of all the vascular plants of the state. Parts of this study 
were finished and made available to his plant taxonomy 
classes in 1953, 1955, 1957, 1958, and 1959. These early 
works comprised his 1st edition, “Keys to the Flora of Ok- 
lahoma.” The Keys...” has been revised periodically and 
is now in the 4th edition. 

In accordance with an original plan, Dr. Waterfall was, 
for over twenty years, expanding the “Keys ...” into a 
flora of the state. He died before this work was finished. 
Because of the nature of the work and the difficulty of typ- 
ing the manuscript he typed over 700 pages of this later 
work himself. It is the desire of all who knew him that this 
work will not go unfinished. 

In 1965 Dr. Waterfall received a grant awarded by the 
National Science Foundation to study collections of Physa- 
lis from Mexico, Central America and the West Indies 
which are housed in the great European herbaria. This 


148 Rhodora [Vol. 75 


study was published in “Rhodora” in 1967. Because of this 
and earlier treatments of Physalis, Waterfall is widely ac- 
claimed as the foremost authority on this difficult genus. 


In addition to his European travels, Dr. Waterfall made 
several extended collecting trips in the United States and 
Mexico. His first collecting trip outside Oklahoma was to 
the various national forests of Colorado, Wyoming and 
Utah, but he did little with these collections because his in- 
terests turned to the Mexican flora and the flora of the 
desert Southwest. During the early years of World War II, 
he spent three summers studying poisonous plants along 
the Texas-Mexico border for Harvard University. Other 
valuable collections from “Southwestern Texas" are in the 
OSU Herbarium as well as other large herbaria in the 
United States. 


Dr. Waterfall made four trips into Mexico. In a letter 
from his wife (November 26, 1972), she stated: “I think 
we must have traveled every road and trail north of Mexico 
City — as well as many places where we made our own 
trails!” A fifth trip was started in 1967 but had to be 
terminated at Laredo, Texas due to his health. From these 
trips, Dr. Waterfall collected some 5,000 sheets which have 
been distributed to various United States herbaria. The 
originals are in a separate *Mexican Herbarium" at OSU. 
Several new species and one new genus have been described 
by Dr. Waterfall as a result of these four collecting trips 
into Mexico. 

During the last ten years of his life, his major problem 
was his health. During that period he had 5 major opera- 
tions, 2 heart attacks, and several bouts with serious 
kidney infections. After an extended illness, he died on 
October 27, 1971 in the Stillwater Municipal Hospital. 


In addition to his study of Oklahoma plants, some of his 
more noteworthy works include: 
Waterfall, U. T., 1951. The Genus Callirhóe (Malvaceae) 
in Texas. Field and Lab. 19 (3): 107-119. 
. 1958. A Taxonomic Study of the Genus Phy- 


1973] Waterfall (1910-1971) — Stanford 149 


salis in North America North of Mexico. Rhodora 60: 
107-114; 128-142; 152-173. 

. 1959. Separotheca, A new Genus (Commelina- 
ceae) from Mexico. Rhodora 61 (725): 136-139. 
Waterfall, U. T., and Mahler, W. F., 1964. Baccharis 
(Compositae) in Oklahoma, Texas and New Mexico. 
Southwestern Naturalist 9 (3): 189-202. 
Waterfall, U. T., 1967. Physalis in Mexico, Central Am- 
erica and the West Indies. Rhodora 69: 82-120; 203- 
239; 319-329. 


JACK W. STANFORD 
HOWARD PAYNE COLLEGE 
BROWNWOOD, TEXAS 76801 


THE ENVIRONMENT OF SCHISTOSTEGA PENNATI 
.(HEDW.) HOOK. & TAYL.: 
NEW VERMONT STATIONS 


DONOVAN R. BOWLEY! 


Schistostega pennati is the sole member of the moss 
family Schistostegaceae. It has attracted attention over 
the years primarily because of its characteristic of reflect- 
ing light back out of the crevices it grows in, with a 
chatoyant green-gold glow. Every six or eight years some- 
one reports a new station, ranging in size from pea-sized 
patches to the magnificent expanse of several yards under 
the barn at Oldfields, in Groton, New Hampshire (Thom- 
son, 1956). In most of the literature it will be found under 
S. osmundacea (Dicks.) Mohr. Most often it is discovered 
as a glowing mass of protonema on damp soil or rock in 
some dark crevice. Occasionally fertile and sterile fronds 


*Biology Department, Boston University, Boston 02116. This note 
is on part of the work done under NDEA Title IV Fellowship 
number 02022. 


150 Rhodora [Vol. 75 


are found as well, looking like diminutive ferns of the genus 
Osmunda. Grout (1935) gives a concise description of the 
plant. Marshal (1920) has a good description and several 
drawings. 

This plant is apparently circumboreal and tolerates a 
wide range of conditions. J. S. Erskine (1950) found it 
on slate a few yards from an arm of Halifax Harbour, N.S. 
D. S. Johnson (1925) found a maritime station at Mt. 
Desert Island, Maine. Here, the moss grew three or four 
feet above the high tide within the range of the salt spray, 
and in fact, on some Fucus that had washed into its cranny. 
In this particular station, the only light it received was that 
reflected into its cavern from the surface of the ocean. 
Schofield (1969) reported it “in shallow caverns, and on 
the raw earth of overturned tree roots in swampy parts of 
cedar-hemlock forests.” 

Early information suggested it was a calciphobe, but 
Grout (1933) found it in a crevice of a limestone cliff in a 
quarry. At this station, it was growing “in the open” in 
a crevice of a north-facing cliff. 

The more typical site is that described by Grout (1899, 
1902a, 1902b, 1906), Kaiser (1921), and Nichols (1933). 
Each of these sites is a moist crevice, usually in the back 
of a larger cavern or hole; or a moist cellar under a shed 
or a barn. Champlin (1969) found it inside the entrance 
to a graphite mine in Rhode Island. 

It appears that in nearly every case the moss is located 
on a condensing surface, rather than where water is 
actively flowing. Johnson (1925) reported that the moss 
in the seaside station in Maine obtained its water from 
condensation on the cooler rock, of fog, or of water evapo- 
rated from the surface of the ocean. 

A review article by J. M. Coulter (1918) of the work 
in Japan of Viscount Toda reveals some interesting notes 
on the physiology of the moss. The protonema can live 
seven months in culture without producing a leafy shoot. 
“Chromatophores” scatter in a day and change direction 
of orientation in seven to ten days. Spores germinate in 


1973] Schistostega — Bowley 151 


one month at 60-77°F. The leafy shoot dies at about 0°F; 
the protonema survives to —5?F. The optimum temperature 
for the leafy shoot is 60-77°F. Gistl (1926) reports lens- 
like cells in the protonema, which focus light within the 
cell. At the focus of the light are located the chloroplasts. 
This cell develops best when subjected to unidirectional 
light. He cultured Schistostega for three years in the labo- 
ratory and found that under optimum conditions the posi- 
tion of the chloroplasts could change in from one to three 
hours. 

So we have in this small plant an adaptation to low levels 
of unidirectional light, giving it a competitive edge in those 
situations. The readjustment to changes in light is rapid, 
and would enable it to survive in areas where its caverns 
under rocks and tree roots were shifted by storm or wind. 
As evidenced by its growing on raw earth in the open in 
British Columbia (Schofield, 1969), on the cliff where 
Grout found it (1933), and in full sunlight in Gistl’s 
laboratory (1926), it can survive in the open. It is most 
likely an early colonizer in humid, cool locations, but is 
rapidly overgrown by heartier vegetation, and survives in 
the hollows and crevices where the other plants cannot 
follow. 

During the summer of 1969 several new stations for the 
luminous moss were discovered by Duncan Galbraith of 
Essex Junction, Vermont, and by the author. Two more 
stations were discovered in the summer of 1971 by Ranger- 
guide Kenn Boyd of Burlington, Vermont. All of the sta- 
tions are mid-talus slope on the east side of Mt. Mansfield, 
Vermont. All are within half a mile of Cliff House (top 
station of the Gondola lift). Grout (1906) reported similar 
stations on the “Nose” cliffs of Mt. Mansfield. 

One station near Cliff House was observed through the 
winter of 1969-1970. Ground heat and warmer air rising 
through the talus from below passed through the crevice, 
melting all the snow out of the crevice and from in front 
of the opening of the rocks. A small hole opened through 
to the “outside world". As the temperature of the outside 


152 Rhodora [Vol. 75 


air rose, the opening became larger. As the outside air 
temperature fell, rime ice formed around the rim of the 
opening, closing it down — in effect, a natural thermostat. 
Measurements on a Taylor maximum-minimum thermome- 
ter suspended through the hole on a string showed a nearly 
constant internal temperature of 35°F, +1°F, while the 
outside temperature varied through a 31?F range. In addi- 
tion, the thin shell of snow that remained over the hollow 
acted as a sort of "greenhouse roof", allowing light to pass 


through. 
In summer, the temperatures of his *moss-hole" were 


lower than that of the surrounding woods because of cool, 
moist air draining down out of the talus above. Both 
summer and winter there was a slight excurrent of moist 
air. 


LITERATURE CITED 


British Bryological Society. 1946. “Annual meeting of the British 
Bryological Society, 1946." Bryologist 49: 99. 

CAMPBELL, DouGLAs H. 1918. The structure and development of 
mosses and ferns, 3rd edition. Macmillan Company, New York. 

CHAMPLIN, R. L. 1969. *A Rhode Island station for luminous 
moss." Rhodora 71: 305. 

CoNARD, HENRY S. 1938. “The foray in upper Michigan, 1937." 
Bryologist 41: 20. 

COULTER, J. M. 1918. Review in Bot. Gaz. 67: 278-279. 

ERSKINE, J. S. 1950. “More minute mosses from Nova Scotia." 
Bryologist 53: 54-56. 

GisTL, R. 1926. "Beziehung zwischen licht und Schistostega-vork- 
eim." Ber. d. Deut. Bot. Ges. 44: 483-492. 

GROUT, A. J. 1899. “An annotated list of rare or otherwise inter- 
esting mosses occurring in or near Plymouth, New Hampshire.” 
Rhodora 1: 54. 

1902a. “A new habitat for Schistostega.” Bryologist 
5: 103. I 

————————— 1902b. “Notes on Vermont Mosses.” Rhodora 4: 182. 

1906. “Notes on Vermont Bryophytes — 1906." Bry- 
ologist 10: 6, 7. 

1933. ‘Miscellaneous notes on mosses.” Bryologist 
36: 25. 

1935. Moss flora of North America. vol. 2, part 2. 
By the author. Newfane, Vermont, 


1973] Schistostega — Bowley 153 


1940. “Moss notes, 1940." Bryologist 43: 75. 

HUNTINGTON, J. W. 1902. *How I found Schistostega osmundacea." 
Bryologist 5: 52. 

JOHNSON, D. S. 1925. “A maritime station for Schistostega os- 
mundacea.” Bryologist 29: 18-19. 

KAISER, G. B. 1921. "Little journeys into moss-land. IV — the lu- 
minous moss." Bryologist 24: 48. 

MARSHALL, N. L. 1920. Mosses and lichens. Doubleday, Page, and 
Co. pp. 199-202. 

NICHOLS, G. E. 1933. “Notes on Michigan bryophytes — II" Bry- 
ologist 36: 75. 

SCHOFIELD, W. B. 1969. “Some common mosses of British Colum- 
bia." British Columbia Provincial Museum, Handbook No. 28. 

THOMSON, J. W. 1956. The 1956 foray of the American Bryological 
Society." Bryologist 60: 41. 


A NEW STATION FOR SAXIFRAGA RIVULARIS L. 
IN THE WHITE MOUNTAINS, NEW HAMPSHIRE 


On June 18, 1969 Dr. Rosemary Mackay of Montreal 
drew my attention to a small group of plants growing at 
the southwest corner of the Appalachian Mountain Club 
Lake-of-the-Clouds Hut located at 5,000 ft. on the slope of 
Mt. Washington (Fig. 1). The plants were Saxifraga 
rivularis L. primarily known in recent years from a small 
station on the summit of Mt. Washington at 6,288 ft. 
(Pease, 1964). The identification was confirmed by the 
late Dr. Stuart K. Harris of Boston University. A speci- 
men has been deposited in the herbarium of the University 
of New Hampshire. 

The station includes about 15 plants growing in a com- 
pact group. Most plants were in flower on June 18, and 
by July 3, 1969 most had set fruit. Further checks in June, 
1970, 1971 and 1972 showed the plants to be persisting 
and in 1971 and 1972 to be spreading slightly. 

Porsild (1957) notes that Saxifraga rivularis is found 
on wet and mossy areas and on wet cliffs and by brooks 
in the arctic. However, Britton (1957 ) suggests that it 
may be found on fairly well-drained substrates. Both the 
S. rivularis stations described here are comparatively dry, 


154 Rhodora [Vol. 75 


Fig. 1. Saxifraga rivularis L. growing in conjunction with Carex 
bigelowii Torr. at 5,000 ft. near the Lake-of-the-Clouds Hut on Mt. 
Washington, New Hampshire. 


certainly not as damp as would be suggested by the name 
*Alpine Brook Saxifrage". In past years the Mt. Wash- 
ington summit locality was in a position to be influenced by 
leaks from a nearby water tank, but recently the tank has 
been removed without apparent detriment to the plants. 
Both stations are characterized by thin winter snow cover 
and by early snow melt in the spring. Porsild (1957) states 
that S. rivularis is a pronounced nitrophile growing, 
“ . below bird cliffs and near human habitations". Both 
stations are in a position to be influenced by human wastes. 

It is well that this plant has spread to at least one new 
station as development plans for the summit of Mt. Wash- 
ington endanger the prime station for this relatively rare 
plant on the Presidential Range. 

I wish to thank the Mt. Washington Observatory for 
providing living facilities, and Bruce Tiffney and Lindsay 
Fowler for their assistance. 


1973] Saxifraga rivularis 155 


LITERATURE CITED 


BRITTON, M. E. 1957. Vegetation of the arctic tundra, p. 67-130 
in H. P. Hansen (ed.), Arctic biology. Oregon State Univ. Press 


(1967). 318 p. 

PEASE, A. S. 1964. A flora of northern New Hampshire. New 
England Bot. Club, Cambridge, Mass. 278 p. 

PORSILD, A. E. 1964. Illustrated flora of the Canadian Arctic 
Archipelago (Second ed.). National Mus. Canada Bull. 146, 
Ottawa. 218 p. 


WESLEY N. TIFFNEY, JR. 
BIOLOGY DEPT., UNIV. OF MASS./BOSTON 
100 ARLINGTON ST. 02116 


TWO INTERESTING PLANTS ON MT. CARDIGAN, 
ORANGE, NEW HAMPSHIRE. Many interesting occur- 
rences of plants are reported by Frank Seymour in his ex- 
cellent Flora of New England. One of the most unusual 
was his discovery of Carex capitata. L. on Mt. Cardigan in 
1960 at an elevation of 3200 feet. In New Hampshire, Carex 
capitata is known otherwise only on Mt. Washington where 
a few small colonies of it occur in moist peaty moss at an 
elevation of 5000 feet. It does not grow on Mt. Katahdin 
but is found in Newfoundland and the arctic. 

The report of it on Mt. Cardigan seemed so unlikely that 
the authors thought possibly some mistake had been made 
and decided to investigate. Accordingly in September, 1972 
they climbed the mountain. Above 2500 feet ledges and bare 
rock were encountered but no unusual plants. The west side 
of the summit consists of large expanses of bare sloping 
rock with occasional patches of soil or clumps of dwarf 
trees. In this area a colony of Carex capitata was soon lo- 
cated. It was growing in dry open soil along with Poten- 
tilla tridentata Ait. Eventually 3 colonies were found in 
similar situations, the largest of which was 20 feet in diam- 
eter. The plant was in abundance in each colony and seemed 
to be more vigorous than on Mt. Washington. 


156 Rhodora [Vol. 75 


All of the summit area was searched and eventually a 
small colony of Carex Bigelowti Torr. was located in a dry 
peaty area. lt later developed that this plant had been 
found there by Steele in 1955 but not reported. Various 
plants that might be expected in such a mountainous situ- 
ation were found but nothing else unusual. Carex Bigelowii 
is confined to alpine areas, except for this situation but has 
a wide distribution. It is common on the Presidential] and 
Franconia Ranges and also occurs on The Twin Range, and 
on Mt. Moosilauke. 

It is very difficult to account for the presence of these 
plants, especially Carex capitata which is rare even on Mt. 
Washington, and the authors cannot even agree between 
themselves as to the most likely explanation. 

About 100 years ago there was a fire on Mt. Cardigan 
which was so spectacular that it was seen for miles and an 
eastern promotory received the name Firescrew. Presum- 
ably the fire drastically altered the environment, creating 
new open habitats thus providing a suitable site for Carex 
capitata which then seeded in. The difficulty with this hy- 
pothesis is that Mt. Washington is 50 miles to the northeast 
and it is hard to see how the fruits could have been trans- 
ported. They are comparatively heavy and not windborne 
and, if eaten by birds, would presumably be digested, de- 
stroyed or lost. Furthermore the plant is of such limited 
occurrence on Mt. Washington that it seems too much of a 
coincidence to suppose that a bird had transported it to 
Cardigan, especially when there are many other suitable 
sites closer at hand. Even though Carex Bigelowii is a 
much commoner plant it is not easy to see how its fruits 
could have been transported. 

Another explanation is that Carex capitata existed on Mt. 
Cardigan as a relic since the retreat of the glacier. There 
undoubtedly were ledges and probably small peaty openings 
where it might have survived. If this were the case, how- 
ever, it is very difficult to explain why it or other more com- 
mon arctic alpines are not found on some of the numerous 
apparently suitable sites on other mountains. The reason 


1973] Mt. Cardigan — Steele & Hodgdon 157 


it is not found in such situations is undoubtedly that it can- 
not withstand competition with other plants. It is very 
difficult to see how it could have survived on Cardigan for 
thousands of years as would have to be the case if it were 
a true glacial relic. 

There is a third possibility, not very attractive, but not 
easy to disprove, and that is that some botanist for reasons 
best known to himself, transplanted the plants there from 
Mt. Washington. Two instances are known of transplants 
on Mt. Washington and there are many instances of plants 
being moved to lowland sites where they do not long per- 
sist. It is perhaps significant that although the plant is 
fairly conspicuous it was not discovered on Cardigan until 
Seymour found it in 1960. 

It will be very difficult to decide positively among these 
possibilities. This article is being published partly so that 
some botanist 50 years from now can compare the distri- 
bution of the plants with their present state. 


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


A. R. HODGDON 

DEPT. OF BOTANY 

UNIVERSITY OF NEW HAMPSHIRE 
DURHAM 08824 


NEW OR NOTEWORTHY NEW HAMPSHIRE PLANTS 
DAVID E. BOUFFORD 


In the course of field studies during 1971 and 1972 several 
plants previously unreported in New Hampshire were 
found. They are listed here along with some new stations 
for a few other noteworthy plants of the New Hampshire 
Flora. Collections were made of all and a specimen of each 
has been deposited in the Keene State College Herbarium. 
Duplicates have been sent to the University of Massachu- 
setts Herbarium in Amherst, Massachusetts and except 
for Spiranthes lucida, Dentaria laciniata and Dicentra 
canadensis to the University of New Hampshire Her- 
barium. All collections were made by me and the collection 
numbers are my own. 


Salix amygdaloides Anderss. Outlet of Meetinghouse 
Pond, Marlborough, Cheshire County, 5 May 1971 (flower- 
ing), and 1 July 1971 (mature foliage), Boufford 1938; 
tree about 8 m. tall. Seymour cites a specimen collected in 
Lenox, Massachusetts and deposited in the Pringle Herbari- 
um of the University of Vermont as the only verified col- 
lection from New England. He also says it is reported from 
Vermont. Gray’s Manual, 8th ed., gives the range as being 
from Vermont westward. 


Clematis dioscoreifolia Levl. & Vaniot. Steep roadside 
bank, Rte. 12 in Walpole, Cheshire County, 24 July 1972, 
Boufford 7268. Several well-established plants in rather dry 
soil. Seymour calls this plant rare. Including var. robusta 
he mentions just six New England towns where it has been 
collected, only one being in Vermont and none in New 
Hampshire or Maine. Fernald (1950) says, “occasionally 
spread from cult., Mass. to Va." 

Cassia nicitans L. Railroad ballast, approximately 0.5 
mile south of Swanzey line in Winchester, Cheshire County, 
25 August 1971, Boufford 4441. Seymour lists no collections 
from New Hampshire or Maine, but there are some from 


158 


1973] New Hampshire Plants — Boufford 159 


Vernon, Vermont. This appears to be the northeastern edge 
of the plant's range. 

Baptisia australis (L.) R. Br. Old field just south of the 
intersection of Rtes. 119 and 10 in Winchester, Cheshire 
County, 20 June 1972, Boufford 6041. Several clumps of the 
plant were well-scattered throughout the field. A house sev- 
eral hundred yards away has many plants around it and is 
the obvious source of this escape. 

Thermopsis villosa (Walt.) Fernald & Schubert. Road- 
side and railroad bed, Rte. 12 in Walpole, Cheshire County, 
19 June 1971, Boufford 2786. The majority of the plants are 
on a steep, dry roadside bank, but some are scattered up to 
a quarter mile away, indicating that the plant is spreading. 
This plant is growing with the previously mentioned Clema- 
tis dioscoreifolia. 

Digitalis lutea L. Roadside Rte. 10 just north of Beaver 
Brook Falls and junction with Rte. 9, Keene, Cheshire 
County, 14 July 1971, Boufford 3529. Over a hundred plants 
make up the colony. There is a specimen in the herbarium 
of the New England Botanical Club of D. lutea made from 
a garden planting and with the notation on the label stating, 
“Brought from Keene where growing wild." Wakefield, Ju- 
ly 2, 1944, Coll. R. C. Bean. Whether or not this is the same 
colony is not known. A proposed dam in the immediate 
area will no doubt destroy the colony if construction begins. 

Hieracium vulgatum Fries. Roadside along Spaulding 
Brook, Sullivan, Cheshire County, 31 July 1972, Boufford 
7433. Several plants. This plant has been collected in all 
the surrounding states, but not in New Hampshire. 

Hieracium umbellatum L. Beech Hill, Keene, Cheshire 
County, 23 September 1971, Boufford 4988. Large colony of 
several hundred plants mostly in thin, sandy woods, but ex- 
tending into deeper woods. There is a collection in the her- 
barium of the New England Botanical Club and another in 
the Gray Herbarium; both are from Massachusetts and ap- 
peared as transient waifs. The station in Keene is apparent- 
ly the first in New England where the plants are thoroughly 


160 Rhodora [Vol. 75 


established. Gray's Manual states that the plant is western 
and Eurasian and comes east only to Michigan. 

The following four species, collected on the shore of the 
Connecticut River below the Bellows Falls dam in Walpole, 
Cheshire County, have been collected elsewhere in New 
Hampshire but are unusual enough in that state to deserve 
mention. 

Tofieldia glutinosa (Michx.) Pers. 13 August 1971, Bouf- 
ford 4164. Previously reported only from Plainfield in New 
Hampshire. 

Spiranthes lucida (H. H. Eaton) Ames. 7 July 1971, 
Boufford 3313. Also known in New Hampshire only from 
Plainfield. Quite abundant. 

Cardamine pratensis L. 29 May 1971, Boufford 2122. Re- 
ported previously in New Hampshire only from Croydon. 

Lobelia Kalmii L. 18 August 1972, Boufford 4153. Sev- 
eral scattered plants. 

This area below the Bellows Falls Dam is rather unusual. 
The shore consists mostly of ledge and loose boulders. It is 
completely inundated by torrents of water in the spring ; 
during the summer several seepage streams flow from the 
upper banks to the main channel of the river. 

Overlooking the Connecticut River station on the dry 
south-facing slope of Fall Mountain in Walpole, Cheshire 
County, were found two more interesting plants: Asclepias 
quadrifolia Jacq., 5 June 1972, Boufford 5686, and Solidago 
odora Ait., 10 August 1972, Boufford 1603. The only New 
Hampshire station known previously for the Asclepias, ac- 
cording to Seymour, was in Charlestown. However, there 
is a specimen of this plant in the University of New Hamp- 
shire Herbarium, collected in 1890 by William S. Harris in 
Windham, Rockingham County. According to Gray's Man- 


ual, this station would be at about the northeastern limit of 
the range. 


Further south still, in the Connecticut River Valley and 


just across the Walpole line in Westmoreland, Cheshire 
County, was found a large colony of Dentaria laciniata 


1973] New Hampshire Plants — Boufford 161 


Muhl., 10 June 1971, Boufford 2527, previously reported 
from Rumney and Plainfield in New Hampshire. 

A station for Dicentra canadensis (Goldie) Walp. was 
found on a north-facing slope in Alstead, Cheshire County, 
near the Surry line, 11 May 1972, Boufford 5105. It had 
been reported by Seymour to occur from Hanover north- 
ward in New Hampshire. Dr. Albion Hodgdon of the Uni- 
versity of New Hampshire says that it is also growing in 
Plainfield, Sullivan County. 

At Sumner’s Falls in Plainfield, Sullivan County, was 
found Rhynchospora capillacea Torr., 22 August 1971, 
Boufford 4364. The only previous report for New Hamp- 
shire was in Freedom. 

Mirabilis nyctaginea (Michx.) MacM. was found in a 
weedy area behind a gas station on Main Street in Concord, 
Merrimack County, 25 August 1972, Boufford 8067. Al- 
though Seymour does not show a record of this in New 
Hampshire, there is one specimen in the herbarium of the 
University of New Hampshire from Rochester. 


I wish to thank Mr. Harry E. Ahles of the University of 
Massachusetts for the identification of Salix amygdaloides 
and Hieracium umbellatum. However, responsibility for 
the aecuracy of all identifications is solely my own. 


LITERATURE CITED 


FERNALD, M. L. 1950. Gray’s Manual of Botany, eighth edition. 
American Book Company, New York, 1632 p. 

SEYMOUR, FRANK C. 1969. The Flora of New England. The Charles 
Tuttle Company, Rutland, Vermont. 596 p. 


RFD 2 BOX 357 
KEENE, NEW HAMPSHIRE 03431 


CHIMAPHILA MACULATA (L.) PURSH 
IN MAINE AND NEW HAMPSHIRE 


A. R. HopGpDON AND LESLEY M. EASTMAN 


The Spotted Wintergreen, Chimaphila maculata was 
reported by Seymour (1969) from Lee and Goffstown in 
New Hampshire and from Rutland and Windham counties 
in Vermont, those being the only localities given for north- 
ern New England. In this article we are discussing several 
recently discovered stations and citing herbarium records 
of several others in New Hampshire that have been over- 
looked. 

In northern New England Chimaphila maculata tends 
to occur as disjunct colonies of a few individuals or some- 
times single plants, in this respect resembling some species 
of orchids. Even with careful searching it is usually im- 
possible 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. Thus one 
can assert with some confidence that the population of 
Spotted Wintergreen in northern New England is sparse. 
Not only do there seem to be few colonies but there appear 
to be few individuals as well. Although the species in our 
area is scarce it is quite possible that it is considerably 
more frequent that the few records would indicate. 

The recent discovery of it at the base of Mt Agamenticus 
in York, York County, Maine on October 3, 1971, by the 
junior author and C. Paul Wight is the first report from 
the state. The plant was growing in a large rocky depres- 
sion which was filled with matted beech and maple leaves. 
It had shown no signs of flowering and on a subsequent 
visit to the same area in the fall of 1972, the plant had not 
flowered. Only the one specimen was seen on Mt Agamen- 
ticus and in lieu of collecting the solitary specimen, photo- 
graphs were taken which are on deposit at the herbaria 
of the University of Maine and the New England Botanical 


162 


1973] Chimaphila maculata — Hodgdon & Eastman ` 163 


Club. Several years ago the senior author attended a field 
excursion of the New England Wild Flower Society at their 
well known stand of Rhododendron maximum in Sanford, 
York County, Maine. At the end of the tour, one partici- 
pant reported the presence of Spotted Wintergreen as a 
limited colony somewhere along the trail into the tract of 
Rhododendrons. Such a report should be supported by a 
photograph or an identifiable fragment of a plant but it 
does seem entirely probable that Chimaphila maculata 
occurs in Sanford. It should be sought farther north even 
into Oxford County where many species of southern affinity 
have been found. 

More evidence that Spotted Wintergreen is much more 
frequent than supposed comes from New Hampshire: on 
October 3, 1972 the senior author and some members of 
his class in aquatic vascular plants found scattered plants 
of it in dryish or at least well-drained deciduous woods on 
the western bank of the Oyster River in Durham, Strafford 
County, along that part of it known locally as the Mill 
Pond. Durham has received more botanical attention from 
the senior author than any other area of similar size any- 
where and the Chimaphila station is only about a mile 
from the campus of the University of New Hampshire. 
It might be argued that the species has invaded the area 
during the ten or twelve year period since the time when 
the area was carefully botanized. However, the fact that 
the plants, though few in number are scattered over a con- 
siderable area and indeed occur on both sides of the Oyster 
River where it empties into the “Mill Pond" suggests that 
the colony or colonies have been there for some time. Also 
having seen the Spotted Wintergreen a number of times 
during the past ten years the senior author had the advan- 
tage of experience on his more recent visit to the same 
area. 

A fragment sufficient for identification was collected at 
the Durham Station, October 3, 1972. A. R. Hodgdon, 
L. Loiselle and J. Wicks 19631. This is deposited in the 
herbarium of the University of New Hampshire. Earlier 


164 Rhodora [Vol. 75 


in the summer on August 9, 1972, the senior author in 
company with Frederic L. Steele and Edward Flaccus made 
a leisurely ascent of Sentinel Mountain in Tuftonboro, 
Carroll County, New Hampshire. At about the middle of 
the south facing rocky slope in deciduous forest a single 
specimen was sighted from which one leaf was removed 
as a voucher. A few yards away one or two other shoots 
were seen, one having been in flower. Although further 
search was made no more plants were found that day. This 
specimen, August 9, 1972. Hodgdon, Steele and Flaccus 
19632 is also deposited in the herbarium of the University 
of New Hampshire. 

These records of recent years extend the range of Chima- 

phila maculata into Maine and central New Hampshire but 
there are several other townships in New Hampshire from 
which specimens are present in the University of New 
Hampshire Herbarium. These are listed as follows: 
ROCKINGHAM CO. North Hampton, May 14, 1969. A. R. Hodgdon & 
E. J. Hehre 18314; Windham, July 30, 1898, W. S. Harris, s.n.; HILLS- 
BORO CO. Pelham, Gumpas Pond area, 1950, J. Straughan, s.n.; 
CHESHIRE CO. Marlboro, north side of Clapp Pond, Sept. 20, 1954, 
Carl Webber, s.n. 
The Harris Collection from Windham was reported in 
Rhodora (1949). In summary there are authentic records 
of Chiamaphila maculata from York, York Co., Maine, and 
the following New Hampshire stations: Tuftonboro, Car- 
roll Co., Lee and Durham in Strafford Co., North Hampton 
and Windham in Rockingham Co., Goffstown and Pelham 
in Hillsboro Co. and Marlboro in Cheshire Co. There is 
some evidence also that it grew a few years ago in Sanford 
in York County, Maine. Also an additional station in Lee 
was discovered several years ago and during the summer 
of 1972 it was found again in Windham by Mrs. Clotilde 
Straus. 

We would like to emphasize that care should be exercised 
to document stations for this rare species without destroy- 
ing entire shoots, photography being recommended for that 
purpose. 


1973] Chimaphila maculata — Hodgdon & Eastman 165 


Without some knowledge of the history of a colony it is 
impossible to determine whether the plant has arrived re- 
cently, or instead, has been in that particular area for a 
long time. Over a span of several years it should be possible 
to collect much useful information by repeated or at least 
annual visits to the same plant or colony, at the same time 
recording data on the spread or recession of the colony, the 
size and vigor of plants, the condition of flowering, etc. 
In this way we may be able to determine whether the 
species has recently migrated, is becoming extinct locally 
or is merely persisting. Those who argue for its recent 
migration to those places where it has recently been found 
must bear in mind that in northern New England there 
are few sources of the seeds and that the chance of their 
being lifted from near the surface of the protected forest 
floor and effectively transported many miles is highly 
improbable. 


LITERATURE CITED 


Hopepon, A. R. and HERBERT FRIEDLANDER. 1949. Rhodora 51: 


111. 
SEYMOUR, F. L. 1969. The Flora of New England, Charles E. 


Tuttle Co., Rutland, Vt.: 425. 


DEPARTMENT OF BOTANY 
UNIVERSITY OF NEW HAMPSHIRE, DURHAM 03824 


OLD ORCHARD BEACH, MAINE 04064 


SOME NEW LOCALS IN THE MAINE FLORA. Bo- 
tanical excursions in Maine during the seasons of 1971 and 
1972 have resulted in the discovery of some interesting and 
unusual plants. 


Aster dumosus L. North side of Lovewell Pond, Frye- 
burg, Oxford County, September 3, 1971. During a day 
visit in the Fryeburg area a friend led me to where exca- 
vation work was being done to uncover an ancient Abenaki 
campsite. It was at this digging that I found a large num- 
ber of these Asters growing in the damp yellow sand along 
the shore. This location is an extension of its known range, 
it never having been found north of York County. Another 
plant of interest growing in the same area was Panicum 
rigidulum. 


Isotria medeoloides (Pursh) Raf. North Sebago, Cum- 
berland County. Ninteen years ago Mrs. Iva Knight of 
Kennebunk found the only known station of this rare orchid 
in the state. During the preceding years she visited this 
location three or four times but never saw signs of this 
plant again. It was doubtful that after nineteen years 
Isotria could ever be found, but on June 7, 1972, following 
a well defined map which Mrs. Knight had drawn for me, 
I found the area, and within a very short time I came upon 
two of the plants. These orchids were growing under 
deciduous trees in leaf mould next to a small brook. It was 
interesting to note that a number of plants of Habeneria 
Hookeri were growing in the same area. 


Epipactis Helleborine (L.) Crantz. This European weed 
orchid was first reported in the state about ten years ago 
by Dr. A. E. Brower of Augusta. He found this orchid 
growing at Benton Falls, in Kennebec County. I have come 
across this plant numerous times in the past two years. 
On August 7, 1971, C. Paul Wight and I found two stations 
of Epipactis growing among white pines in Cornish, York 
County. A week later, I found another station under red 
maples in Saco, York County. The following week I found 


166 


1973] Maine Flora — Eastman 167 


it growing in a dense beechwood growth in Stow, Oxford 
County. 


On August 6th of 1972, while walking along a large 
glacial esker called “the Ridge” in Chesterville, Franklin 
County, I found Epipactis growing on the gravel slopes 
among grasses and ferns. I found it again on the same day, 
on Day Mountain in Strong, Franklin County, where it 
was growing in profusion from the base of the mountain 
to the summit. The eastern slope of the mountain is com- 
posed of calcareous slate in which many rare plants can 
be found, namely Panax quinquefolium, Carex plantaginea, 
Dryopteris Goldiana and Asplenium Trichomanes. An ac- 
count of the flora of Day Mountain by C. H. Knowlton 
can be found in one of the early numbers of this journal 
(Rhodora 6: 206, 1904). 


Conopholis americana (L.) Wallr. Day Mountain, Strong, 
Franklin County, August 30, 1972. While exploring the 
eastern part of the mountain I came across a number of 
these root parasites growing under red oaks. This is well 
north of previously reported stations at York, Cumberland, 
Oxford, and Androscoggin Counties. 


Specimens of the species mentioned have been deposited 
in the herbaria of the University of Maine and the New 
England Botanical club. Photographs were deposited in 
lieu of specimens in the case of /sotra medeoloides. 


LESLEY M. EASTMAN 
OLD ORCHARD BEACH, MAINE 04064 


Volume 75 No. 801, including pages 1-168, was issued March 30, 1973. 


168 Rhodora [Vol. 75 


INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA 


Manuscripts must be double-spaced or preferably triple- 
spaced (not on corrasable 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 indicate the style of type through the use of capitals 
or underscoring, particularly in the citations of specimens, 
except that the names of species and genera may be under- 
lined to indicate italics in discussions. Specimen citations 
should be selected critically especially for common species 
of broad distribution. Systematie revisions and similar 
papers should be prepared in the format of “The System- 
aties and Ecology of Poison-Ivy and the Poison-Oaks," 
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu- 
larly with reference to the indentation of keys and syno- 
nyms. Papers of a floristic nature should follow, as far as 
possible, the format of “Contribution to the Fungus Flora 
of Northeastern North America. V.," H. E. Bigelow & M. E. 
Barr, Rhodora 71: 177-203. 1969. For bibliographic cita- 
tions, a recommended list of standard journal abbreviations 
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. 
Club 85: 277-300. 1958. 


Readers of RHODORA will miss the names of three long- 
time editors on the front cover of this issue. As a matter of 
poliey we have decided not to continue the custom of recent 
years of including the inactive editors. This is being done 
now to make it clear who are responsible for the work of 
editing the Journal and to make it easy to simplify the list 
when more changes in editors will be necessary after the 
present year. 

We want to extend our appreciation for their many 
editorial contributions to Albert Frederick Hill who re- 
placed Ludlow Griscom on the Board in 1944, to Ralph 
Carleton Bean who became an editor in 1949 and to Richard 
Creighton Foster who has served as an associate editor 
since 1961. 

Albion R. Hodgdon 
Editor-in-Chief 


CONTENTS: — continued 
A Re-evaluation of Potamogeton fibrillosus Fern. 
(l'otamogetonaceae) 
Robert R. Haynes and James L. Reveal .......... ertt 75 
The Audouinella Complex (Rhodophyta) in the Western 
Sargasso Sea 
William J. Woelkerling vicccccccccccsecccsssscscsssesscccssesscessscecsesseesseeass 78 
Salt Concentrations in Ground Waters beneath Rhizophora 
mangle and Avicennia germinans 


Loring Morrow and Norton H. Nickerson .....................a.... 102 
Variation of Seed Size in Atriplex patula var. hastata (I.) 

Gray 

Frank R. Drysdale ......... eese eene nnne nan 106 


Floral Dimorphism in Rachicallis americana (Jacq.) Hitch. 
(Saltwater-Bush) 


Norton H. Nickerson and Joseph W. Tripp ses 111 
Book Review 

Rolla Trqom. .ecesccecerecssesosccsrsoscnncssesscccasscescevncesssveccsssenssnsecnsesgoneces 114 
A New Alaskan Species of Erigeron L. (Compositae) 

Stephen A. Spongberg |... neces 116 
Tribal Classification of Triosteum (Caprifoliaceae) 

Water H. Lewis and Paul R. Fantz .................................... 120 


Chromosome Pairing in Obligately Apogamous Ferns; Pel- 
laea atropurpurea and Pellaea glabella var. glabella 
Sylvia Jane Rigby vicccccccccccssccccsssscscccsscccsscssccssscssccssescceceseesssees 122 
The Past and Present Geographical Distribution of 
Petalostemon foliosus and Notes on its Ecology 


Jerry M. Baskin and Carol C. Baskin esses 132 
Book Review 

Albion R. Hodgdom cicccccccccccccccccccccccccsscecsevsseegesessecscesecausevecsueecs 140 
Umaldy T. Waterfall (1910-1971) 

Jack W. Stanford u u ui. U UU... .. U ULU L teens etenim ases gas 146 


The Environment of Schistostega pennata (Hedw.) Hook & 
Tayl: New Vermont Stations 
Donovan R. Bowley esset enne 149 
A New Station for Saxifraga rivularis L. in the White 
Mountains, New Hampshire 
Wesley N. Tiffney, Jt. essetis 153 
Two Interesting Plants on Mt. Cardigan, Orange, New 
Hampshire 


F. L. Steele and A. R. Hodadon .......... eee 155 
New or Noteworthy New Hampshire Plants 
David E. Boufford ....... sse 158 


Chimaphila maculata (L.) Pursh in Maine and New Hamp- 
shire 


A. R. Hodgdon and Lesley M. Eastman ..... eee 162 
Some New Locals in the Maine Flora 

Lesley M. Eastman ..... sse tete tete 166 
Instructions for Contributors ................ eee 168 


Editor's Note ¿ul L UU ineen nqa Aa QA uA 168 


Hodova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
ALBION REED HODGDON, Editor-in-Chief 


` 


ROLLA MILTON TRYON 
RADCLIFFE BARNES PIKE 
STEPHEN ALAN SPONGBERG Associate Editors 
GERALD JOSEPH GASTONY 

ALFRED LINN BOGLE J 


Vol. 75 June, 1973 No. 802 
CONTENTS: 


A Revision of North American and Caribbean Melanthera 
(Compositae) 
James C. Parks sss 169 


Senecio anonymus Wood, an Earlier Name for Senecio 
smallii Britton 
Robert R. Kowal and T. M. Barkley ................................ 211 


A Note on the Karyology of Haplopappus spinulosus 
(Pursch) DC. ssp. typicus 
L. S. Gill and C. C. Chinnappa. ..... eee 220 


Floral Structure, Hybridization and Evolutionary Relation- 
ship of Two Species of Mimulus 
Ta qu AT T n uA L E i La 225 


(Continued on Inside Back Cover) 


Che Nef England Botanical Club, Ine. 
Botanical Museum, Oxford St. Cambridge, Mass. 02138 


RHODORA. — A quarterly journal of botany, devoted primarily to the 
flora of North America and floristically related areas. Price $10.00 
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 
plants of North America, will be considered by the editorial com- 
mittee for publication. Articles concerned with systematic botany 
and cytotaxonomy in their broader implications are equally accept- 
able. All manuscripts 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. Extracted reprints, if ordered in advance, will be 
furnished at cost. 

Address manuscripts and proofs to Albion R. Hodgdon, 

Dept. of Botany, Nesmith Hall, 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. 
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QTRbooora 


JOURNAL OF THE 
NEW ENGLAND BOTANICAL CLUB 


Vol. 75 June, 1973 No. 802 


A REVISION OF NORTH AMERICAN AND 
CARIBBEAN MELANTHERA (COMPOSITAE) 


JAMES C. PARKS 


The genus Melanthera Rohr sensw stricto comprises a 
small group of helianthoid composites restricted to the 
Western Hemisphere. The center of distribution is Carib- 
bean. with representatives present in Mexico, Central 
America, South America, and the southeastern United 
States. The various species occupy beaches, pine flatwoods, 
and moist or dry forests from sea level to fairly high main- 
land elevations. Melanthera aspera exhibits weedy tenden- 
cies. This ecological and corresponding morphological vari- 
ability, coupled with the wide distribution of some species 
in habitats favorable for collecting, has led to a prolifera- 
tion of synonyms in the genus. Natural hybrids also have 
contributed to the confusion. 

Although Schulz (1911) published the only comprehen- 
sive revision of North American Melanthera, his ineffective 
reconciliation of biological and nomenclatural problems has 
caused most subsequent workers to rely on regional manu- 
als for identification. Standley (1938) cogently expressed 
the difficulties encountered in dealing with Melanthera but, 
I believe, offered a much too conservative solution in recog- 
nizing only one species. 

In the present treatment of Melanthera, I recognize five 
species and two varieties, and provide a morphological and 
nomenclatural basis for more intensive studies of all taxa 


169 


170 Rhodora [Vol. 75 


in the genus. During the course of this investigation, my 
personal collections and approximately 2200 herbarium 
specimens from 27 herbaria have been examined. (A com- 
plete list of specimens studied is available from the author 
upon request.) An important aspect of this revision has 
been the observation of living plants of Melanthera both in 
the field and in the greenhouse. Specimens of all taxa were 
either transplanted from the field or germinated from seed 
and grown in the Vanderbilt University greenhouses. Ob- 
servation for approximately two years was most helpful 
in discovering and evaluating taxonomic characters at all 
stages of development. 


HISTORICAL ACCOUNT 


Elements of the genus Melanthera were first described 
by Dillenius (1732) in his Hortus Elthamensis in which 
he mentioned that the description was based on live plants 
grown from seed shipped from Carolina. Linnaeus (1753) 
cites Dillenius’ polynomial under the name Bidens mivea. 
The second of Dillenius’ two excellent plates (f. 47), each 
representing leaf variations, best serves as the lectotype 
for Melanthera nivea (L.) Small. Patrick Browne (1756) 
described another species of Melanthera based on Jamaican 
material and placed it in his genus Amellus, giving it 
a polynomial epithet. Linnaeus subsequently acquired 
Browne's herbarium, and his treatment of Browne's genera 
Amellus and Santolina has resulted in confusion for stu- 
dents of Melanthera. Robert Brown (1817) correctly elu- 
cidated this problem, but a few points should be noted here. 

Linnaeus apparently concluded that the Browne speci- 
men, now housed in the Linnaean herbarium as Bidens 
nivea, was equal to B. nivea of Species Plantarum (1753), 
for he cited it in Amoen. Acad. 5: 381 (1759) as part of 
the Jamaican flora. However, each of these elements repre- 
sents a distinct species of Melanthera. Bidens nivea of the 
Species Plantarum (1753) is M. nivea, a species restricted 
to the mainland, and P. nivea of the Amoen. Acad. (1759) 
is M. aspera Jacq. As a partial result of the Linnaean con- 


1973] Melanthera — Parks dud 


fusion, the Browne specimen identified as Bidens mivea 
eventually came to represent the name, while the Dillenean 
plate was ignored. Michaux (1803) redescribed the true 
B. nivea as Melananthera hastata, the epithet commonly 
used today. 

The genus Amellus P. Bn. (1756) would be the valid 
name for Melanthera except that Linnaeus selected it for a 
distinct assemblage of species (Syst. Nat., 1759, p. 1225 
and 1377), and this latter usage has been conserved (Lan- 
jouw et. al., 1966). Melanthera was described by Rohr 
(1792) who used specimens from Martinique. Although 
Rohr did not include a specific epithet or cite specimens, 
his clear description, plus the fact that only one species of 
Melanthera grows on Martinique, soundly establishes M. 
aspera as the type species. Rohr's editor, Vahl, included a 
citation to Bidens nivea in Swartz (1791), which is Melan- 
thera aspera. 

Strother (1970) established that specimens in Copen- 
hagen likely represent material Rohr used in describing 
Melanthera. I examined photographs and drawings of 
these specimens and find them to represent material of M. 
aspera var. glabriuscula. I compared Rohr’s specimens to 
the Dillenean plates typifying M. nivea and to Jacquin’s 
description and plant of M. aspera and find them closer to 
the Jacquin work. I agree with Strother that M. nivea 
should not be the type species of Melanthera. Thus, I find 
it strange that he would select M. panduriformis H. Cass., 
which he correctly observes to be a taxonomic synonym of 
M. nivea based on a trivial leaf-form variant, to typify the 
genus. 

Adanson (1763) described the genus Ucacou based on 
material now distributed in Melanthera Rohr (1792) and 
Synedrella Gaertn. (1791). J. E. Dandy of the British 
Museum (1969, and personal correspondence, 1968) has 
found that Synedrella best typifies Ucacow and has pro- 
posed conservation of the former. 

Browne’s Amellus species was described by Jacquin 
(1788) as Calea aspera, and in 1803 Michaux described 


172 Rhodora [Vo]. 75 


the genus Melananthera with the epithets hastata and 
deltoidea, which are synonyms for nivea and aspera re- 
spectively. Cassini (1823) contributed a thorough morpho- 
logical discussion but added three superfluous epithets. In 
1850 A. Richard described M. angustifolia. Later, J. K. 
Small (1903, 1905) described the Florida endemics M. 
parvifolia and M. ligulata. 

In my opinion, Schulz (1911), in his revision of North 
American and Caribbean Melanthera, failed to understand 
the morphological variability within the genus and added 
many superfluous names to the literature. Other early 
twentieth century works which treated Melanthera were 
usually floristic studies of a limited Caribbean area (e.g. 
Britton & Wilson, 1925) or reports of botanical expeditions 
(e.g. Blake, 1922, 1924, 1932, and J. K. Small, 1905, 1909). 
While the type specimens of some of the species described 
in these works appear distinct, they are actually local 
variants or hybrids. In this treatment I have attempted to 
present a comprehensive view of the genus (sensu Rohr) 
whereby the 47 specific or varietal epithets are properly 
disposed and the taxa clearly delimited. 


INTERGENERIC RELATIONSHIPS 


Most synantherologists have placed Melanthera in the 
tribe Heliantheae and the subtribe Verbesininae. Obviously 
Melanthera is morphologically similar to Verbesina. A dif- 
ficult problem is the satisfactory delimitation of these gen- 
era, As originally defined by Rohr, Melanthera is said to 
have corollas “composita uniformis, tubulosa”; M. aspera, 
on which this description was based, is indeed discoid. In 
the strictest sense, then, Melanthera is easily distinguished 
from similar genera by its discoid capitulum of white 
florets. However, the description of Melanthera was 
broadened by later workers, e.g. Bentham & Hooker 
(1873), to include radiate representatives, and this con- 
cept was accepted by Wild (1965) in his treatment of the 
exclusively radiate African Melanthera. If this broadened 
circumscription is recognized, Melanthera intergrades with 


1973] Melanthera — Parks LES 


Echinocephalum and Perymenium and is often confused 
with Aspilia, Bainvillea, Eleutheranthera, Wedelia, Wulffia, 
and occasionally with Calea and Bidens of the subtribe 
Coreopsidinae. 

Possibly one or more of the genera noted should be com- 
bined with Melanthera. I have examined material of 
Echinocephalum latifolium Gardn. and find it nearly iden- 
tical with M. nivea except for the pale yellow rays of the 
former (cf. Blake, 1930). Since consideration of these 
generic limits is beyond the scope of the present work, I 
have eliminated South America, and hence Echinocepha- 
lum, from consideration. Therefore, I have chosen to treat 
only those entities which justifiably could be included in the 
original description. While there is some overlap in dis- 
tribution of these taxa, western hemisphere Melanthera is 
centered in the Caribbean, while Echinocephalum and 
Perymenium are distributed principally in South America. 
Until further study reveals the natural] relationships of 
these genera, Melanthera can be distinguished from the 
others, albeit somewhat artificially, by its discoid capitulum. 
Distinguishing morphological features of Melanthera are 
presented in Table I. 


CYTOLOGY 


Chromosome counts of n=15 for two species of Melan- 
thera have been reported by Turner et al. (1961), Powell 
& Turner (1963), and Turner & Flyr (1966). Upon exam- 
ining the vouchers of these plants, I find the specimen cited 
by Turner & Flyr (1966) is indeed M. aspera var. glabri- 
uscula, but those cited by Turner et al. (1961) and Powell 
& Turner (1963) are misidentified and represent M. nivea. 
I have studied meiotic material of all North American 
species of Melanthera using squash techniques and have 
determined a haploid number of n=15 for all of the species 
covered in this treatment (Table II). 


ECOLOGY AND HYBRIDIZATION 


Field and herbarium studies indicate that in the United 


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175 


Melanthera — Parks 


1973] 


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176 Rhodora [Vol. 75 
TABLE II 
CHROMOSOME VOUCHERS IN MELANTHERA 
Taxon Locality and Collection Repository 
M. nivea FLORIDA. WAKULLA CO.: VDB 
Newport, woods near road. 
Parks 135. 
LOUISIANA. TENSAS PAR.: VDB 
moist woods near Newellton. 
Parks 337. 
MEXICO. VERA CRUZ: TEX 
Cordoba, roadside, King 2361. 
cit. Turner et. al., 1961. 
Huatusco, roadside, Powell TEX 
642. cit. Powell and Turner, 
1963. 

M. aspera FLORIDA. COLLIER CO.: VDB 

var. aspera Naples, back of beach, sand, 
roadside. Parks 124. 
SARASOTA CO.: Longboat Key, VDB 
oolitic spoil bank. Parks 125. 

M. aspera FLORIDA. MONROE CO.: VDB 
var. Long Key, dry beach. Parks 
glabriuscula 98. Islamorada, Anderson TEX 

3226. cit. Turner and Flyr, 
1966. 

M. angustifolia FLORIDA. COLLIER CO.: VDB 
Monroe Sta., moist glade, on 
oolite. Parks 298. 

M. parvifolia FLORIDA. MONROE CO.: VDB 
Big Pine Key, dry pine flat- 
woods on oolite. Parks 289. 

M. ligulata FLORIDA. PALM BEACH CO.: VDB 


Loxahatchee, dry, sunny 
palmetto-pine flatwoods. 
Parks 89. 


1975] Melanthera — Parks 177 


States the various taxa of Melanthera are usually distinct 
morphologically and ecologically. Melanthera nivea inhabits 
fairly moist forests and borders in the lower Coastal Plain 
southward to central Florida. Melanthera ligulata fre- 
quents the low hammocks of south-central Florida but may 
exhibit weedy tendencies. Melanthera parvifolia is re- 
stricted to to oolitic “pine islands” of southern Florida and 
to Big Pine Key. Melanthera angustifolia is relatively rare, 
but is usually found in moist, open glades in southern 
Florida. Melanthera aspera var. aspera is often weedy, 
preferring dry, rocky places such as roadsides and cal- 
careous spoil banks fairly near the ocean in southern 
Florida, while M. aspera var. glabriuscula is a strand spe- 
cies growing high on beaches in extreme southern Florida 
and throughout the Caribbean. 

In Mexico and Central America, Melanthera nivea is 
usually found in the habitat already noted at altitudes 
above 150 m. When found at lower altitudes it may inter- 
grade into M. aspera var. aspera, possibly due to occasional 
hybridization and introgression. Such occurrences may 
explain Guatemalan collections by Skinner 1867 (NY) and 
Brenckle 47-242 (WIS), each of which contains plants 
best placed in either M. nivea or M. aspera. Hybridization 
between M. nivea and M. angustifolia can be seen in Blake 
7601 and 7601A (US), which were collected along a Gua- 
temalan railroad bank; M. linearis is typified by the latter 
number (Blake, 1922). 

A field study in southern Florida revealed a situation 
that seems representative of those found throughout the 
genus. In southern Florida, hammocks, or high spots of 
coquina rock run southwest from Miami into the ever- 
glades where they gradually disappear (Young, 1953). 
Sometimes called pine islands, these hammocks are domi- 
nated by Pinus elliottii and are distinct ecologically. Exam- 
ination of herbarium records, including appropriate type 
collections of J. K. Small, indicated that a morphological 
gradient exists between Melanthera parvifolia and M. an- 
gustifolia along these pine islands. Plants found on oolitic 


178 Rhodora [Vol. 75 


e M. ANGUSTIFOLIA 
° + INTERMEDIATE 


3 ° 
° M. PARVIFOLIA 
c? oo ^ + 
PHYLLARY Qe? + 
UN 2 1 o +, ++ 
RATIO F5 + ° 
1 + ° et 
+ ° ooo ° ° 


kaa 


1 2 3 4 5 6 7 8 9 101 12 13 14 
LEAF L/w RATIO 


Fig. 1. Three populations of Melanthera from southern Florida 
analyzed to show interpopulational relationships of phyllary and leaf 
length-width ratios. 


rock near Miami are much like M. parvifolia, while those 
on the lower everglades keys resemble strongly M. angusti- 
folia. Morphological intermediates were collected at geo- 
graphically intermediate localities. 

In order that the above-described variation be under- 
stood more fully, population samples were collected and 
analyzed. The data obtained were analyzed (Fig. 1) in the 
manner of Anderson (1949). The population sample of 
Melanthera parvifolia (Parks 289), taken from Big Pine 
Key, formed a closely knit group with an average leaf 
L/W ratio of 1.5 and an average phyllary L/W ratio of 2.5. 
The population sample of M. angustifolia (Parks 298), 
collected near Monroe Station, Collier Co., Fla., formed a 
group with an average leaf L/W ratio of 8.0 and an aver- 
age phyllary L/W ratio of 1.5. The other population plotted 
on the graph was collected on a pine island in Everglades 
National Park (Parks 280). It exhibits more variability in 
the traits considered, being intermediate morphologically 


1973] Melanthera — Parks 179 


between M. parvifolia and M. angustifolia. Representative 
specimens of these populations were grown in the Vander- 
bilt University greenhouses under uniform conditions for 
about a year where they maintained their morphological 
distinctiveness. 

This study seems to indicate that some gene exchange 
is occurring between Melanthera parvifolia and M. angusti- 
folia in the area studied. This knowledge is valuable in 
helping to delimit properly these taxa, to annotate unusual 
herbarium specimens, and to determine synonymies more 
accurately. For example, M. radiata Small is quite likely 
an introgressant between M. parvifolia and M. angustifolia. 
'The type collection is from oolitic rock near Miami and is 
similar to the Everglades Park collection noted above. As 
such material most closely resembles M. parvifolia, it is 
treated here as a synonym of that taxon. 

During the two year period when living Melantherae 
were grown in the greenhouse, numerous reciprocal crosses 
were conducted to see if the taxa recognized would hy- 
bridize and to note the morphology of any hybrids recov- 
ered. Quantitative data were not taken, but vigorous F, 
offspring were recovered from all interspecific crosses and 
many of these were grown to flower. As the potential for 
interbreeding exists among all species, ecological differ- 
ences appear to be most important in maintaining the 
identity of Melanthera taxa in nature. While my limited 
biosystematie studies have been most helpful in interpret- 
ing anomalous plants seen in both the field and herbarium, 
they indicate that more extensive work might be profitable 
in further elucidating evolutionary relationships in the 
genus. 


PHYLOGENETIC SPECULATIONS 


The origin, phylogeny, and evolution of the Compositae 
are lucidly discussed by J. Small (1917-1919) and Cron- 
quist (1955). Cronquist would choose a helianthoid genus, 
e.g. Wyethia, to represent the fundamental composite 
group. Upland South America is the present center of dis- 


180 Rhodora [Vol. 75 


tribution for the Heliantheae and the Senecioneae and con- 
tains many primitive composites. 

Echinocephalum certainly would be considered a primi- 
tive genus of composites. Its geographic range includes the 
uplands of South America. It has opposite leaves, an her- 
baceous perennial habit, and a ligulate, paleaceous capitu- 
lum. all of which are accepted by many authors as primi- 
tive characters in the Compositae. Melanthera differs from 
Echinocephalum in that it lacks ray florets and, as a dis- 
coid capitulum is considered an advanced trait, I regard 
Meianthera to have been derived from Echinocephalum. 

Present distributional data suggest that Melanthera 
nivea is the ancestral species of the genus. It resembles 
strongly Echinocephalum latifolium and is partially sym- 
patiic with it. Though the range of M. nivea is extensive, 
it is restricted to the mainland and, in the south, to higher 
elevations. Ecologically, this species prefers the stable 
habitat of forests and borders. 

Melanthera nivea probably migrated northward from 
South America, the postulated area of its origin, during 
Eocene time and came to be continuously distributed 
throughout the southern United States and Mexico. As 
increasing aridity sporadically occurred during Oligocene 
and Miocene time, M. nivea would have failed to survive 
in northern Mexico and southern Texas; this failure would 
account for its present disjunct distribution. Melanthera 
nivea may have migrated southward into the newly formed 
areas of Florida during the colder climate of the Pleisto- 
cene period. Differentiation to form M. parvifolia and M. 
ligulata may have occurred during this and more recent 
time. Other plant migrations which occurred during Ter- 
tiary time are believed to have taken place in a similar 
manner (Chaney, 1947). 

Perhaps Melanthera nivea migrated to the sea along the 
Orinoco or other local rivers, and in the process, differen- 
tiated into M. aspera which then was dispersed by water 
throughout the Caribbean. The pattern of Caribbean cur- 
rents (Cruxent & Rouse, 1969) and the observation of 


1973] Melanthera — Parks 181 


Millspaugh (1907) that M. aspera var. glabriuscula may 
readily inhabit newly formed strands or islets both sup- 
port this hypothesis. The varieties of M. aspera may rep- 
resent ecotypes developed in accordance with factors noted 
by Carlquist (1965). More recent differentiation, possibly 
from M. aspera, may have given rise to M. angustifolia. 
This could have occurred in one or several regions but 
most likely took place in southeastern Mexico or Yucatan. 


SYSTEMATIC TREATMENT 


Melanthera Rohr, Skriv. Nat. Selskabet (Copenhagen) 2: 
213. 1792. Type species: M. aspera (Jacq.) Small. 
Ucacou Adanson, Familles des Plantes 2: 131. 1763. (ex 

parte, non Synedrel/a propos. nom. cons.). 
Lipotriche R. Br., Trans. Linn. Soc. (London) 12: 118. 
1817. 
Melananthera Michx., Flora Bor. Amer. 2: 106. 1803. 


Suffrutescent perennials, 0.2-2.2 m tall, arising from a 
hemispheric or elongate, caudex-like rootstock. Stems one 
or several, quadrangular to sulcate, erect or sprawling, 
glabrescent to hispid-hirsute, green, sometimes with ma- 
roon mottling, rarely maroon. Leaves opposite, petiolate, 
triple-nerved, broadly ovate or deltoid to narrowly lan- 
ceolate-linear, glabrescent to hispid-hirsute; base attenu- 
ate, cuneate to nearly truncate, hastate or with basal lobes 
ascending in some species; apex mostly acute; margin 
crenate to serrate, sometimes irregularly so. Capitula 
terminal, solitary or arranged in very loose cymes, borne 
on rather elongate, sulcate, generally pubescent peduncles. 
Involucre 0.6-1.5+ em broad, of two subequal, often loosely 
imbricate series, grading somewhat into pales. Phyllaries 
ovate to lanceolate, equal to or much longer than florets at 
first anthesis, apices more or less herbaceous, pallid basally 
with or without green nerves. Pales oblanceolate, strongly 
keeled and membranaceous basally where they clasp flo- 
rets; apex thicker, aristate or caudate with cauda 0.4-1.5 + 
mm long, straight or recurved, sometimes pubescent api- 


182 Rhodora [Vol. 75 


caly and marginally. Florets 25-100 per capitulum, all 
discoid, regular, hermaphroditie; corolla white; tube 4-7 
mm long, constricted and pale yellow toward a nectarifer- 
ous base; limb 0.5-1 mm long, spreading or rarely ascend- 
ing, papillate adaxially, strigillose or glabrous abaxially. 
Anther sacs black, 1.5-2 mm long; apical appendages white, 
sagittate, 0.5 mm long; basal appendages absent, style 
branches white, 1.2-2.5 mm long, narrowing to a lanceolate, 
papillate or hairy tip, this partly or completely recurved 
upon senescence; pappus of 2 to 5+ delicately barbed, 
caducous awns. Achene indurate, obpyramidal, compressed- 
quadrangular (triangular in outer florets), 2-3 mm long, 
1-2.5 mm wide; apex usually truncate or slightly concave 
due to marginal cusps of crown, usually convex in M. as- 
pera var. glabriuscula, sparsely to moderately pubescent; 
surface usually brown with fine striations, warts and/or 
gold to silver mottling may be present. 


Key to the Species 


a. Aristae of pales greater than 1 mm, usually recurved 
when long; phyllaries narrowly lanceolate, the upper 
1% to 34 green, often recurved; leaf shape various but 
base often hastate. ........ MEM . b. 
b. Largest leaves 4 cm or less long; plants averaging 

0.3 m in height. Southern Florida oolitic “pine 


islands” and Big Pine Key. ... .. 5. M. parvifolia. 
b. Largest leaves greater than 4 cm long; plants aver- 
aging 0.8-1 m in height. .. . C. 


c. Peduncles and leaves hispid and pale aristae 
1-1.4 mm long; phyllaries and pales lanceolate, 
moderately longer than dise, never recurved ; 
leaves moderately coarse. Greater Antilles and 
tropical mainland, often weedy. .......--.+--- 

.....|.............. 2. M. aspera var. aspera. 

c. Peduncles and leaves strigose to glabrescent or 
if these hispid, the caudae of the pales 1.4 mm 
long; phyllaries and pales narrowly lanceolate, 
noticeably longer than disc, the longest often 


1973] Melanthera — Parks 183 


recurved; leaves moderately membranaceous 
(occasionally rugose in M. nivea) x me d. 

d. Plants erect, up to 2.2 m tall with several 
Stems arising from large, woody root; leaves 

ovate, very rarely lanceolate, rich, dark 

green, often rugose, base commonly hastate, 

vesture moderate, strigose; larger capitula up 

to 2 cm across the involucre. Central Florida, 

the coastal plain from S. Carolina to Louisi- 

ana; Mexico, and Central America. ........ 
EMI etel we ... 1. M. nivea. 

d. Plants decumbent to weakly erect, bushlike, 

with one main stem freely branched at the 

often jointed nodes; leaves ovate to lanceolate 

(quite variable), rarely rugose, trichomes 

sparse; larger capitula up to 1.5 cm across 

the involucre. South-central Florida at the 
approximate latitude of Lake Okeechobee. .. 
Deo ri EN EE 6. M. ligulata. 

a. Aristae of pales usually 1 mm or less (may be slightly 
longer in M. aspera var. aspera), commonly less than 
0.5 mm, never recurved; phyllaries ovate to broadly 
lanceolate, the upper l4 to l5 green never recurved; 
leaves various, rarely hastate..................... € 


TEMERE Ons ERE 4. M. angustifolia. 

€. Leaves ovate, deltoid or weakly hastate; pubescence 
various; sprawling or bushy plants av. 1 m in 
height; capitula various. ...................... Jy. 

f. Trichomes of peduncle appressed-ascending and 
moderately sparse; cauda of pale 0.5 mm long; 
phyllaries ovate; capitulum 1.3 em wide or less 
across the involucre; achenes usually 2.5 mm 

long or less, the apex often convex-hairy, or 


184 Rhodora [Vol. 75 


truncate. Beaches of the Caribbean. .......... 
VENE ... 3. M. aspera var. glabriuscula. 
f. Trichomes of peduncle spreading (hispid) and 
relatively abundant; cauda of pale 1 mm long; 
phyllaries broadly lanceolate; capitulum 1.8 cm 
or more across the involucre; achenes usually 
2.5 mm or more in length, the apex truncate to 
concave, sparsely pubescent. Greater Antilles 
and tropical mainland, often weedy. .........- 
s... l............. 2. M. aspera var. aspera. 


1. Melanthera nivea (L.) Small, Flora SE. United States 

1251, 1340. 1903. Fig. 2. 

Bidens nivea L., Sp. Pl. 2: 833. 1753. Lectotype: Dil- 
lenius, J. 1732 Hort. Elth. tab. 47(!) 

Athanasia hastata Walt., Flora Caroliniana 201. 1788. 
Type: ? 

Melananthera hastata Michx., Flora Bor. Amer. 107. 
1803. Type: Carolina-Virginia, Michaux (photo. GH!) 

Melananthera hastata Michx. vars. lobata and pandu- 
rata Pursh, Flora Amer. 2: 519. 1814. based on Dil- 
lenius, /.c. 

Melanthera panduriformis H. Cass. in Levrault, F. G. 
(ed.). Dict. Sci. Nat. 29: 483. 1823. based on Dil- 


lenius, /.c. 
Melanthera trilobata H. Cass., l.c. based on Dillenius, 
l.c. 


Melanthera oxylepis DC., Prodr. 5: 545. 1836. Type: 
ex. herb. Haenke (photo & fragment Us!) 

Elephantopus cuneifolius Fourn., Bull. Soc. Bot. France 
30: 186. 1883. Type: not ascertained. (photo. us!) 

Melanthera carpenteri Small, Flora SE. United States 
1251, 1340. 1903. Type: Louisiana: W. Feliciana 
Par., pine thickets, Carpenter s.n. (NY! isotype NO!) 

Melanthera lobata Small, l.c. Type: Florida; Lake Co., 
Eustis, lake edge, Nash 1141 (Ny! isotypes GH! MICH! 
MO! uc! US!) 

Melanthera oxycarpha Blake, Contr. U.S. Ntl. Herb. 22: 


1973] Melanthera — Parks 185 


Fig. 2. Melanthera nivea (L.) Small. A. habit sketch, B. capitu- 
lum, C. phyllary, D. pale, E. achenes. F. meiotic chromosomes, meta- 
phase one, X 1000. 


186 Rhodora [Vol. 75 


628. 1924. Type: Mexico; Vera Cruz: Zacuapan, 
fields, Purpus 2437 (US! isotypes F! MO! uc!) 

Melanthera hastata (Walt.) L. C. Rich. spp. lobata 
(Pursh) Borhidi. Botanikai Kozlemenyek 58: 177. 
1971. based on Pursh, l.c. 


Coarse, perennial, caespitose herb, 0.8-2.2 m tall arising 
from a woody, noticeably swollen, hemispheric, caudex-like 
rootstock. Stems erect, quadrangular to sulcate, green, 
often with maroon mottling in the hollows, branched apic- 
ally, puberulent to pubescent at the nodes. Leaves mostly 
ovate, 8-12 cm long, 3-8 em wide, triple-nerved, membrana- 
ceous, sometimes rugose, rich, dark green, strigose to 
hispidulous; base broadly acute to hastate, the lobes di- 
vergent or ascending, the petioles averaging 4.5 cm; apex 
acute to acuminate; margin serrate. Capitulum borne on a 
hispidulous, sulcate peduncle 4-12 cm long; involucre in 
two loosely imbricate series 1.5-2 cm broad; phyllaries 
lanceolate, green for 14 or more of the length apically, 
white with green nerves basally, strigillose, noticeably 
longer than florets at first anthesis; pales 5-7 mm long, 
2. mm wide, strongly keeled, translucent and nerved 
basally, thicker, often red-tipped, fimbriate to pubescent 
apically; apical aristae 1.4+ mm long, often recurved; 
corolla tube 5 mm at anthesis, 6 mm at style exsertion, 
limb 1-1.5 mm, spreading, color snow-white, pale yellow at 
constricted nectariferous base; anther sacs 2 mm long; 
style branches long, white, often completely recurved upon 
senescence; pappus of 2 to several caducous awns; achene 
obpyramidal, 2-3 mm long, 1.5-2 mm wide; apex truncate 
to slightly concave, corolla base sessile; surface finely 
striated, often verrucose; color golden brown to dark 
brown. 


Moist forests and borders from Lake Okeechobee, Flor- 
ida, north into the Coastal Plain from South Carolina to 
Louisiana; then at somewhat higher elevations (usually 
greater than 150 meters) in Mexico and Central America 
(Fig. 3). Schwegman (1968) reports Melanthera nivea 


187 


Melanthera — Parks 


1973] 


VIY1OJ3IAHVd 'W a 
VIV1n9l1'W e 


Iyoy v4owjupjopy jo uonnqrusiq 


A en 
YH3dSV MVA VH3OSV Wo o 


V3AIN + "e 


'g ‘Bld 


188 Rhodora [Vol. 75 


from Massac and Pulaski Counties, Illinois. Athey (per- 
sonal correspondence, 1972) says that he has collected it in 
McCracken, Ballard, and Livingston Counties, Kentucky. 
The sheets, nos. 403, 779, and 1160 respectively, are on 
deposit at Memphis State University. These would estab- 
lish Kentucky state records for Mr. Athey. I have not yet 
seen these collections. Dr. Kral assures me that Athey 
403, VDB is M. nivea. These are significant northern range 
extensions and establish this taxon in the Mississippi em- 
bayment. Flowers in summer and fall. 

The unambiguous Dillenean treatment was overlooked 
by many later workers (c.f. Historical Account) and the 
epithet nivea was usually applied to plants properly called 
Melanthera aspera. Perhaps as a result of this confusion, 
Dillenius’ species was renamed Athanasia hastata by Wal- 
ter (1788) and Melananthera hastata by Michaux (1803). I 
concur with Coulter's opinion (1891) that M. oxylepis DC. 
merely represents a leaf variant common in Mexico. 

In southeastern Mexico and adjacent Guatemala, Melan- 
thera nivea may intergrade with M. aspera var. aspera but 
in the United States it is distinct in morphology and in 
geographic range, being best recognized in the field by its 
habit and on herbarium sheets by its large, dark green. 
often rugose leaves, lanceolate phyllaries and long-caudate 
pales. 


REPRESENTATIVE SPECIMENS EXAMINED. EL SALVA- 
DOR: Morazan: stream bed in coffee grove, Montes de Cacaquatique. 
Tucker 723 ( F, MICH, NY, UC, US). GUATEMALA: Alta Verapaz: 
2 mi E of Coban, open sun, fields, King 3320 (DS, MICH, NY, TEX, UC, 
US); near Amatitlan, brushy bank, alt. 1170 m, Standley 61374 (F). 
izabal: 25 km SSW Puerto Barrios, weedy openings along river in 
rain forest, Raven & Gregors 615 (F, MICH). Soloa: Santa Barbara, 
Shannon 589 (us). HONDURAS: Cortes: E of San Pedro Sula, 
Bosque Iluvioso de Montana La Zona, alt. 190 m, Molina 3437 (F). 
El Paraiso: Yuscaran, moist thicket, alt. 980 m, Standley 25777 (F). 
MEXICO: Chiapas: 4 mi NE Bochil, slope with Pinus & Liquidam- 
bar, alt. 4500 ft, Breedlove 8838 (MICH); between Tumbala & El 
Salto, alt. 1500-4500 ft, Nelson 3389 (US). Guerro: 2 mi S Coahua- 
yultla, pine-oak forest, alt. 3600 ft, Rowell 3086 (sMU). Jalisco: 
hillsides S of Zapotlan, Goldsmith 107 (Ds, MO, NY, UC, US). Michoa- 


1973] Melanthera — Parks 189 


can: Aquila, coconut grove, Hinton 15848 (PH, TEX, US). Oaxaca: 
4 km NE Tehuantepec, grazed area in thorny legumes, alt. 50 m, 
King 425 (MICH). San Luis Potosi: 7-8 mi NE Xilitla, mountains, 
roadside weed, King 4944 (F, MICH, NY, TEX, UC, US). Tabasco: Teno- 
sique, second growth, Matuda 3450 (F, MICH, NY, UC). Tamaulipas: 
Tampico, alt. 15m, Palmer 174 (F, MO, NY, US). Vera Cruz: Jalapa, 
alt. 4000 ft, Pringle 8194 (F, MICH, MO, NY, PH, UC, US); Orizaba, 
open woods, Purpus 1161 (F, MO, UC). NICARAGUA: Matagalpa: 
Santa Maria de Ostuma, Cordillera Central, alt. 12-1500 m, Heller 
35, 39 (F). UNITED STATES: Alabama: BARBOUR CO., Spring 
Hill, common in swamp, Bush 169 (MO, NY). COLBERT CO., Muscle 
Shoals, Buckley Aug. 1826? (PH). MORGAN CO., Decatur, Shimek s.n. 
(F). MOBILE CO., Mobile, Mohr Aug. 1893 (Mo). Florida: BREVARD 
co., rich hammocks, Cape Canaveral, Burgess 684 (F, NY). DUVAL CO., 
Jacksonville, shore of St. John’s river, Curtiss 1414 (F, GA, GH, MICH, 
MISSA, MO, NY, PH, SMU, UC). HERNANDO CO., 1 mi N of Brooksville, 
moist roadside shade, Parks 314 (F, FSU, GA, NY, PAC, SMU, US, VDB, 
WIS). HIGHLANDS CO., Parker Is., 5 mi S of Lake Istokpoga, among 
saw palmetto, edge of hammocks, Brass 15690 (GH, US). MARION CO., 
Orange Lake, in open shade around sink hole, Parks 224 (EVG, F, 
FSU, GH, NY, PAC, SMU, US, VDB, WIS). PALM BEACH CO., loc. not spec., 
Hitchcock 1026 (F). POLK CO. 7 mi E of Haines City, pineland 
burned & cleared 2 yrs ago, Conard 20 Aug. 63 (FLAS). SUMTER CO., 
in swamp, 0.5 mi E of wahoo ch., 5 mi W of Bushnell, Beckner 1582 
(DUKE, FLAS). SUWANNEE CO., wooded stream bank, Ichetucknee 
Springs, Will 17 Sept. 60 (FLAS). WAKULLA CO., W side of Newport, 
rt. 98, roadside ditch, wooded border, Parks 185 (EVG, F, FSU, GH, PAC, 
MICH, MO, NY, SMU, US, VDB, WIS). WALTON CO., sandy, wooded ra- 
vine, rt. 81, Red Bay Springs, Hood 2982 (FLAS). Georgia: CHATHAM 
CO. sandy hammock, Ft. Pulaski Mon., Eyles 4634 (DUKE). LEE CO., 
rich woods along Mill Creek, Harper 1074 (GH, MO, NY, US). MCIN- 
TOSH CO., open live oak woods. Sapelo Is., Duncan 20511 (DUKE, FLAS, 
GH, LAF, MICH, NCU, SMU, TEX, US, USF, WIS). Louisiana: IBERIA PAR., 
disturbed, sandy soil along pipeline clearing, Weeks Is., Thieret 9721 
(FSU, LAF, NCU, SMU, VDB). RAPIDES PAR., Alexandria, Hale s.n. (GH). 
TENSAS PAR. 8.5 mi W of Newellton, along gravel rd. from Sikes 
grocery, mature sweet gum-hickory bottom, Parks 337 (Ds, DUKE, 
EVG, F, FLAS, FSU, GA, GH, LAF, MICH, MO, NCSC, NCU, NY, PAC, PH, 
SMU, TEX, UC, US, VDB, WIS). Mississippi: LOWNDES CO., bottom land 
woods, Bylers Lake, Ray 6683 (NCU, USF). NEWTON CO., wooded 
bluffs, chunky, Bailey 129 (MISSA). South Carolina: ALLENDALE CO., 
cypress swamp, Pipe Creek, Bell 18466 (NCU). BERKELEY CO., road- 
side ditch, 1.1 mi S. of Monck’s Corner, Bradley 3530 (FSU, NCU, NY. 
VDB). CHARLESTON CO., border of live oak woods, Edisto Is. Rd.. 
Godfrey 50475 (NCSC). 


190 Rhodora [Vol. 75 


2. Melanthera aspera (Jacq.) Small, Bull. Torrey Bot. Club 
36: 164. 1909. 

Suffrutescent perennial, 0.5-1.5 m tall, sprawling to sub- 
scandent arising from a woody, swollen, elongate or knobby 
root. Stem sulcate or rounded near base, 0.5-1.5 cm in 
diameter, green may be maroon tinged, glabrescent to 
hispid particularly at nodes. Leaves ovate to deltoid, 6-12 
cm long, 4-8 em wide, membranaceous, rarely rugose, may 
be triple-nerved, light green and drying dark green; in- 
dument below sparse, appressed-ascending to hispid with 
noticeable accumulation or webbing in vein axils, erect- 
ascending above; base broadly acute to truncate, becoming 
short attenuate; petiole 1.5-4.5 cm long; apex acute to 
acuminate; margin crenulate-crenate to serrulate. Capitu- 
lum borne on a slender to sturdy sulcate peduncle 2-14 cm 
long, which has sparse appressed-ascending or abundant 
erect hairs; involucre in two imbricated series, 1.2-1.5 cm 
in diameter; phyllaries broadly ovate and 3-5 mm long, 
3-4 mm wide or lanceolate and 5-7 mm long, 2-3 mm wide, 
tips green and equalling or surpassing the disc at anthesis 
by 1 mm; pales oblanceolate, 4-6 mm long, 1.5-2 mm wide, 
strongly keeled, translucent and nerved basally, thicker, 
pale green and puberulent apically, apical arista or cauda 
0.5-1.4 mm long, never recurved; corolla tube 5-7 mm long, 
basally constricted for 1-1.5 mm, strigillose; corolla limb 
spreading or ascending 0.8-1.5 mm long, papillate adaxi- 
ally; anther sacs 2 mm long; style branches about 2.5 mm 
long, partly or completely recurved when senescent; pappus 
of two to several caducous awns; achenes obpyramidal 
2-3 mm long, 1-2 mm wide, apex truncate to concave and 
glabrescent or convex and tan pubescent, corolla base stipi- 
tate; surface finely striated; color shiny to dul] tan. 


Much of the confusion in Melanthera centers on this 
widespread, often weedy and variable taxon; problems of 
typification and of synonomy are discussed in the historical 
treatment. Because criteria for delimitation of some Mel- 
anthera species are not always reliable, conservative work- 
ers have tended to recognize few species; in one case a sin- 


1973] Melanthera — Parks 191 


us aL 


Fig. 4. Melanthera aspera Jacq. var. aspera. A. flowering branch, 
B. capitulum, C. phyllary, D. pale, E. achenes, F. meiotic chromo- 
somes, late anaphase one, X 1000. 


192 Rhodora [Vol. 7 


gle taxon (Standley, 1938). I believe that the two varieties 
of M. aspera treated here are reasonably distinct morpho- 
logically and ecologically and that ignoring this variability 
would obscure the true character of this species. While 
intermediates do occur, close consideration of morphology 
and habitat will usually enable one to determine a specimen 
with confidence. 


2a. Melanthera aspera (Jacq.) Small var. aspera. Fig. 4. 

Calea aspera Jacq., Collectanea 2: 290. 1788. Type: 

Jacquin (BM, photo. US!) Plate in Icon. Plantarum 
Rariorum 3: 13, t. 583. 1789. 

Melananthera deltoidea Michx., Flora Bor. Amer. 107. 
1803. Type: based on O. Swartz, Obs. Bot. 296. 1791. 

Melananthera Linnaei H.B.K., Nov. Gen. & Sp. 4: 199. 
1820. Type: ? 

Melanthera corymbosa Spreng., Neue Entdeckungen 2: 
135. 1821. Type: ? 

Melanthera urticifolia H. Cass., in Levrault, F. G. 
(ed.), Dict. Sci. Nat. 29: 484. 1823. based on Dil- 
lenius, op. cit. 

Amellus asper O. Kuntze vars. normalis et canescens 
O. Kuntze, Rev. Gen. Pl. 305. 1891. Types: ? 
Melanthera  buchii Urban, Symb. Antillanae 3: 411. 

1903. Type: Haiti: ad Bilbaro, 700 m, Buch 364 n.v. 

Melanthera amethystina O. E. Schulz, in Urban, Symb. 
Antillanae 7: 127. 1911. Type: not designated. 
Lectotype: Santo Domingo, Turckheim 3199b (NY !. 
c.f. M. molliuscula) 

Melanthera canescens (O. Kuntze) O. E. Schulz, l.c. 
based on Amellus asper O. Kuntze var. canescens 
O. Kuntze, l.c. numerous cited specimens examined 

Melanthera hastata var. cubensis O. E. Schulz, l.c. 
Type: not designated, Lectotype: Cuba: prope Ha- 
bana, Wright 3608 (NY! dup. US!) 

Melanthera molliuscula O. E. Schulz, Le. Type: not 
designated, Lectotype: Santo Domingo, Turckheim 
3199 (NY!, cf. M. amethystina) 


1973] Melanthera — Parks 193 


Melanthera montana O. E. Schulz, l.c. Type: Porto 
Rico: San Juan, near Bayamon in montane woods, 
Sintenis 1145, ex parte, n.v. 

Melanthera hastata (Walt.) L. C. Rich. spp. cubensis 
(O. E. Schulz) Borhidi, Botanikai Kozlemenyek 58: 
177. 1971. based on Schulz, l.c. 


Melanthera aspera var. aspera is distinguished by its 
coarseness and by the presence of short, often stiff, erect 
trichomes on stems and leaves. Its leaves are usually sub- 
coriaceous and ovate with broadly acute bases and crenulate 
margins. In flower it is distinguished by broadly lanceolate 
phyllaries, pales with apical caudae 0.5-1.4 mm in length, 
corolla with a spreading limb, and by achenes with trun- 
cate to concave, sparingly pubescent apices. 

Variety aspera grows on dry old beaches, rocky banks, 
and weedy fields from near sea level to 100-150 m altitude 
in the Greater Antilles, Florida, Mexico, and Central Amer- 
ica (Fig. 3). Flowering in the summer and fall. 


REPRESENTATIVE SPECIMENS EXAMINED. BRITISH 
HONDURAS: Stann Creek railway, 50 ft alt., small shrub, Schipp 
282 (F, MICH, MO, NY, UC, US). COSTA RICA: prov. unknown, in 
brush, roadside, 17 km E of Turrialba, Cronquist 8833 (MICH, TEX, 
UC, USF). Guanacaste: Rio Liberia, NE of Liberia, alt. 100 m, Dodge 
6243 (F, MO). CUBA: Habana: Santiago de las Vegas, Baker 4983 
(F, UC, US). Oriente: Baracoa, Pollard 8 (F, MO, UC, us). DOMINI- 
CAN REPUBLIC: Barahona: Barahona, Fuertes 40B (F, MO, UC). 
La Vega: loc. unknown, Ekman 14122 (F, us). SANTO DOMINGO: 
vicinity of Ciudad Trujillo, alt. 0-25 m, Allard 13120, 13133, 14014, 
14463 (us). EL SALVADOR: La Union: openings, roadside, sea 
level, Beetle 26267 (SMU, US). Sonsonate: Armenia, Standley 23443a 
(us). GUATEMALA: Alta Verapaz: damp limestone thickets, 48 km 
NE of Coban, Standley 70118 (F). Huehuetenango: swampy places, 
below Miramar, alt. 300 m, Steyermark 51504 (F). Izabal: Puerto 
Barrios, sea level, Standley 24916 (US). Santa Rosa: wet forested 
quebrada, E of Taxisco, alt. 225 m, Standley 79007 (F). HATIM: 
dept. unknown: Cap Haitien, Leonard 5308 (us); Port au Prince, 
Morne l'Hospital, alt. 350 m, Holdridge 1037 (F, MICH, US). HON- 
DURAS: Atlantida: stream bank thicket, foothills of Ceiba, Yuncker 
8266 (MICH, MO, NY). Comayagua: moist thickets, Siguatepeque, 
alt. 1050 m, Standley 6717 (F). Olancho: mimosa thickets, 400 m alt., 
Juticalpa, Standley 17762 (F). JAMAICA: parish unknown, Lititz 


194 Rhodora [Vol. 75 


Savanna, alt. 300-900 ft, Harris 11739 (F, MO, US). MEXICO: 
Jalisco: cultivated areas W of airport, sea level, Feddema 2547 
(MICH). Oaxaca: near Totontepec, alt. 3700-5500 ft, Nelson 819 (US). 
Vera Cruz: near Cordoba, Greenman 152 (F). Yucatan: Izamal, 
Gaumer 404 (DS, F, MICH, MO, NY, PH, UC, US). NICARAGUA: dept. 
unknown: dry bushy slope, sierras de Managua, alt. 400 m, Grant 
950 (F, MICH). Matagalpa: Santa Maria de Ostuma, alt. 1200- 
1500 m, Heller 68 (F). PANAMA: Bocas del Toro: Almirante, 
roadside, Blum 1319 (FSU, MO). Canal Zone: Ft. Clayton, old Las 
Cruces Trail, Standley 29008 (vs). San Blas: Irandi airport, Duke 
6516 (Mo). San Jose Is.: J. Johnston 1100 (us). PUERTO RICO: 
Arecibo: roadside herb, Florida, Wagner 94 (DUKE, MO). Mayaguez: 
sandy soil near Mayaguez, alt. 350 m, Holm 170 (MO). TRINIDAD: 
St. Ann’s Cascade, Broadway 5147 (F, MO). UNITED STATES: 
Florida: BREVARD CO. Hammock, Merritt’s Island, Small 9437 (au, 
MICH). COLLIER CO., roadside by the gulf, Naples, Parks 124 (ps, 
DUKE, EVG, F, FLAS, FSU, GA, GH, MICH, MO, NCU, MCSC, NY, PAC, PH, 
SMU, UC, US, VDB, WIS). DADE CO., hammocks on sand dunes, Lemon 
City, Small 5821 (DUKE, FLAS, MO, NCU, NY, US). LEE CO. waste 
places, Sanibel Is., Cooley 2515 (FLAS, GH, NY, US, USF). MANATEE 
co., Palmetto, Nash 2433 (F, GH, MICH, NCU, MO, NY, US). MARTIN 
CO., one pt., open sand on new levee, Port Mayaca, Parks 83 (vpR). 
PINELLAS CO., sandy soil, Maximo Pt., Thorne 15430 (GH). 


2b. Melanthera aspera (Jacq.) Small var. glabriuscula (O. 
Kuntze) Parks, comb. nov. Fig. 5. 

Amellus asper O. Kuntze var. glabriusculus O. Kuntze, 
Rev. Gen. Pl. 305, 306. 1891. Lectotype: Panama: 
Colon, O. Kuntze (NY!) 

Melanthera brevifolia O. E. Schulz, in Urban, Symb. 
Antillanae 7: 123. 1911. Type: not designated, Lec- 
totype: Florida: Monroe Co., Elliot’s Key in areno- 
sis maritimis, Curtiss 1415 (Us! dup. F! MISSA! NY! 
PH !) 

Melanthera crenata O. E. Schulz, Lc. Type: not desig- 
nated, Lectotype: Bahamas, New Providence, J. & A. 
Northrop 58 (NY!) 

Melanthera calcicola Britton, Sci. Survey Puerto Rico 
and the Virgin Islands 6(1): 309. 1925. Type: 
Puerto Rico, Britton 6784 (NY!) 

Melanthera confusa Britton, lc. Type: Virgin Is.: 
Tortola, Fishlock 440 (NY!) 


1973] Melanthera — Parks 195 


Fig. 5. Melanthera aspera Jacq. var. glabriuscula (Ktze.) Parks. 
A. flowering branch, B. capitulum, C. phyllary, D. pale, E. achenes, 
F. meiotic chromosomes, metaphase one, 1000. 


196 Rhodora [Vol. 75 


This variety is glabrescent with appressed-ascending 
hairs on stems and leaves. The leaves are ovate to deltoid, 
membranaceous, light green, and with crenate margins. 
The capitulum is 1.2 cm or less in diameter; the phyllaries 
are broadly ovate, subequal and only slightly exceed the 
disc at first anthesis; the apical caudae of the pales is 
0.5 mm or less in length; the corolla limb is usually ascend- 
ing; the style branches usually recurve upon themselves 
when senescent; the achenes are smaller than in the typi- 
cal variety being 2.0-2.5 mm long, apex usually convex and 
tan-pubescent. 

Kuntze (1891) cited no specimens with his descriptions, 
but his itinerary fits well the location and time noted on 
the specimen I have designated as lectotype. Possibly, 
duplicates of this specimen were deposited at Berlin and 


Kew. 
This variety grows high on beaches throughout the 


Caribbean and on the mainland of southern Florida, Mex- 
ico, and Central America (Fig. 3). 


REPRESENTATIVE SPECIMENS EXAMINED. BAHAMAS: 
mangrove bay, Andros, Brace 4886 (F); Long Is, beach near dock, 
Clarence Town, Hackett 136 (DUKE, MICH, UC) ; South Bimini, cleared 
land near dock, Hackett 87 (DUKE, LAF, MICH, UC, Us, USF). CUBA: 
Camaguey, near Pueblo Romano, Cayo Romano, Shafer 2453 (F, Us). 
Habana: high on sandy beach, Playa Baracoa, Sauer 1770 (wis). 
HONDURAS: Atlantida: near beach, Salado, Ywncker 8329 (F, 
MICH, MO). JAMAICA: Portland: Long Bay, Adams 7586 (DUKE). 
Westmoreland: scrub woodland near lighthouse, West Pt., Webster 
6067 (MicH). LESSER ANTILLES: Antigua, seashore, Box 1157 
(F, MICH, MO, US); Barbados: Christchurch Parish, in dunes, 
Sauer 2185 (F); Grand Caymen, Georgetown, Armour 1395 (F); 
Grenada, wayside, levera, ign. 1268 (US); Guadeloupe, Griesbach 
Dec. 1857 (Mo); Martinique, alt. 40 m, Stelle 5125 (US); Virgin Is., 
St. Thomas, Delanses 1845 (F). MEXICO: Quintana Roo: Cozumel 
Is., Gaumer Aug. 1886 (US). Yucatan: Izamal, Gaumer 15543 (wis). 
PANAMA: Bocas del Toro: Bocas del Toro, Carleton 154 (US). 
Canal Zone: Ft. Sherman, bathing beach, Tyson 2263 (FSU, MO). 
PUERTO RICO: Aguadilla: roadside, Lares, Liogier 9956 (DUKE, 
FPDB, US). Culebra Is.: Southshores, Armour 627 (F). Humacao: 
coastal bank, Playa de Fajordo, Britton 1592 (F, US). TRINIDAD: 
upper beach, Manzanilla Bay, Sauer 2249 (F). UNITED STATES: 


1973] Melanthera — Parks 


197 
M 
1 à | ¥ 
W % Ao NY ,⁄ 
M | 1 cm 
W Y 


1 dm 


| 1 cm 

i / 

| f as 

E | A BA 

M 

NY RA 4s 

ASA 4 yey " F 
E [9 
E 


Fig. 6 Melanthera angustifolia A. Rich. A. habit sketch, B. ca- 
pitulum, C. phyllary, D. pale, E. achenes, F. meiotic chromosomes, 
diakinesis, X 1000. 


198 Rhodora [Vol. 75 


Florida: BROWARD CO., Sandy soil, Lauderdale beach, Moldenke 262 
(DUKE, MO, NY). Collier Co., sand island, Everglades City, Parrott 
189 (DUKE). DADE CO., coral rock, Totten Key, Cooley 9343 (USF). 
LEE CO., weedy on shell mound, Bird Is., Brass 18092 (US). MONROE 
co., beach, Bradley Key, off East Cape, Parks 274 (EvG, F, US, VDB) ; 
Long Key, common high on ocean beach, Parks 283 (GA, MICH, NCSC, 
NCU, PAC, VDB); Marquesas Key, Lansing 2116 (F, NY). 


3. Melanthera angustifolia A. Rich., in Sagra, R., Hist. 
Isla Cuba 11: 54. 1850. Fig. 6. Lectotype: Cuba: 
Isla de los Pinos: La Sagra (P) 

Melanthera lanceolata Benth. ex Oerst., Kjoeb., Vidensk. 
Meddel. 88. 1852. Type: Coll. & date not ascertained 
(BM, photo & fragment us!) 

Melanthera microphylla Steetz, Bot. Voy. Herald 156. 
1854. Type: Panama, Seemann 254 (photo US!) 

Melanthera linearis Blake, Contr. U.S. Ntl. Herb. 24: 
30. 1922. Type: Guatemala: Izabal: Cristina, R.R. 
ditch, Blake 7601 (us!) 

Melanthera purpurascens Blake, Contr. U.S. Ntl. Herb. 
22: 629. 1924. Type: Mexico: Chiapas: tableland 
about Ocuilapa, Nelson 2995 (us!) 

Melanthera angustifolia A. Rich. var. subhastata O. E. 
Schulz, Report Sp. Nov. 26: 109. 1929. Type: Cuba: 
Ekman 12359 (NY!) 


Weakly erect perennial herb, 3-9 dm tall, arising from 
a woody usually buried rootstock. Stems several, slender, 
3-9 dm tall, 2-3 mm in diameter, quadrangular, mostly 
red-maroon. Leaves lanceolate-oblanceolate-linear, rarely 
narrowly ovate-elliptical, 3-8 cm long, 0.5-1.5 cm wide, 
length-width ratio av. 8; base cuneate to attenuate; apex 
acute; margin irregularly serrate-serrulate; pubescence 
sparse, appressed-ascending. Capitulum solitary, terminal 
borne on a slender, sulcate, glabrescent, usually long (9-20 
em) peduncle; involucre of one to two subequal series, 
0.6-1.2 cm in diameter; phyllaries ovate, 4-5 mm long, 
2-3 mm wide, length-width ratio av. 1.5, the apica] 1/4-1/3 
green, pubescence appressed-ascending; pales oblanceolate, 
4-5 mm long, 1 mm wide, aristae 0.5 mm long. Neither 


1973] Melanthera — Parks 199 


floret nor fruit is distinctive. They resemble those of 
Melanthera ligulata but fewer of each are present in the 
M. angustifolia capitulum. 


Melanthera angustifolia seems to frequent low, moist to 
swampy, lightly shaded areas but is apparently found in 
drier situations in parts of its range. Distributed from 
southern Florida, through western Cuba and the Isle of 
Pines into Yucatan, southeastern Mexico, and Central 
America (Fig. 3), it is sparsely collected throughout its 
range and is nowhere a common plant. It flowers from 
mid to late summer. 

Achille Richard did not designate a type specimen for 
this taxon. However, Dr. Alicia Lourteig of the Museum 
of Natural History, Paris (P), has informed me (personal 
correspondence) that a specimen exists there labeled in the 
author's hand as noted above. In light of the supporting 
evidence, it seems reasonable to designate this sheet as the 
lectotype of the distinctive Melanthera angustifolia. (For 
additional discussion see Jennings, 1917.) 

Melanthera angustifolia is usually considered one of the 
most distinct species in the genus. Unfortunately, this 
assumption is based solely on the unstable character of leaf 
shape. Occasional specimens of M. nivea exhibit lanceolate 
leaves, as do M. parvifolia and M. ligulata. While the type 
specimens of M. lanceolata and M. purpurascens are some- 
what extreme in leaf characters, other traits indicate that 
they are best considered synonyms of M. angustifolia. 


REPRESENTATIVE SPECIMENS EXAMINED. CUBA: Cama- 
guey: Airoyo, Britton 13133 (us). Isle of Pines: savannahs, road to 
San Francisco de las Piedros, Killip 45440 (us). DOMINICAN 
REPUBLIC: Puerto Plata: coastal swamp, Cabarete, Ekman 14537 
(F, US). SANTO DOMINGO: in savannas, Ekman 13323 (F). 
GUATEMALA: Alta Verapaz: slopes near coves, SW of Lanquin, 
Steyermark 44027 (F, US). Izabal: dry pine slopes, Izabal, alt. 65- 
600 m, Steyermark 38559 (F). Peten: La Libertad, Aguilar 4 (MICH, 
MO, NY, US). HAITI: prov. unknown, Pearice, Ramielle, Ekman 
H6244 (us). MEXICO: Tabasco: Tenosique, in savanna, Matuda 
3502 (F, MICH, MO, NY, UC). Vera Cruz: meadow in oak forest, SE 
of Catemaco, Dressler 122 (F, MICH, MO, NY, UC, us); E of Tuxpan, 


200 Rhodora [Vol. 75 


toward La Playa, Strother 541 (SMU, TEX). PANAMA: Panama: 
Los Sabanas, open slope near sea level, Standley 40777 (vs). 
UNITED STATES: Florida: BROWARD CO., rocky, calcareous land, 
Ft. Lauderdale, Curtiss 5846 (DS, F, FLAS, GA, GH, MO, NCU, NY, 
PAC, SMU, UC, US). COLLIER CO., pine-palmetto flatwood on lime rock, 
Monroe Sta., Parks 298 (EVG, F, FSU, MICH, NY, PAC, SMU, US, VDB, 
wis); Tamiami Trail near Naples, Horner 22 Dec. 55 (WIS). DADE 
co., everglades, Camp Longview, Small 1790 (NY); burned pine 
flatwoods on limestone, 3 mi N of Rock Reef Pass, Godfrey 63420 
(FLAS, FSU); 3 mi W of Everglades Ntl. Park entrance, open glades, 
R. Smith 643 (NCU) ; wet, marly prairie, Grossman Hammock, Web- 
ster 10261 (DUKE); rocky pineland, Long Pine Key, Scull 18 Mar. 
40 (FLAS). MONROE CO., Pinecrest rd., Cypress ponds, pinelands, 
ditches, Lakela 29894 (USF). PALM BEACH CO., sandy peat in pine 
flatwoods, NW of Loxahatchee, Kral 5642 (FSU, SMU, VDB). 


4. Melanthera parvifolia Small, Flora SE. United States 
1251, 1370. 1903. Fig. 7. Type: Florida: (Monroe 
Co.), Big Pine Key, Blodgett, s.n. (NY! isotype GH!) 

Melanthera radiata Small, Bull. Torrey Bot. Club 36: 
163. 1909. Type: Florida: Dade Co., Camp Long- 
view, Small & Wilson 1575 (NY!) 

Melanthera hastata var. parvifolia (Small) O. E. 
Schulz, in Urban, Symb. Antillanae 7: 126. 1911. 
based on Small /.c. 


Suffrutescent perennial, 3-6 dm tall, sprawling to weakly 
erect with numerous, sparingly branched stems arising 
from large hemispheric, caudex-like, woody rootstock. 
Stems slender, obscurely 4 angled, green with a dark red 
tinge, rarely red; indument strigose to hirsute. Leaves 
ovate, 1.5-4 cm long, 1-1.5 cm wide, moderately coreaceous, 
rich, dark green; base broadly acute to obtuse, hastate or 
ascending basal lobes common, petiole very short; apex 
acute; margin coarsely serrate, sometimes undulate, basal 
teeth may be large, salient or ascending ; indument strigose 
to hispid. Capitulum solitary, terminal, borne on a slender, 
suleate, hispid to strigose peduncle which is 2.5-10 cm 
long; involucre in two loosely imbricate series, 1-1.4 cm in 
diameter; phyllaries lanceolate, 5-7 mm long, 1.5-3 mm 
wide, length-width ratio av. 2.5, tip often recurved, notice- 


1973] Melanthera — Parks 201 


1 dm 


Fig. 7. Melanthera parvifolia Small. A. habit sketch, B. capitu- 
lum, C. phyllary, D. pale, E. achenes, F. meiotic chromosomes, late 


anaphase one, X 1000. 


202 Rhodora [Vol. 75 


ably longer than florets at first anthesis, green for 3⁄4, 
length apically, pallid basally, pubescence of short, erect- 
ascending trichomes, abundant marginally and abaxially; 
pales oblanceolate 6-7 mm long, 2 mm wide, strongly 
keeled, apical aristae 1-1.5 mm long, often recurved, nu- 
merous hairs present on the upper, abaxial surface. Corolla 
tube 4 mm long, some hairs present; corolla limb 0.5 mm 
long, spreading, papillate adaxially, strigillose abaxially; 
anther sacs 1.5 mm long; style branches 1.2 mm long, apex 
slightly sagittate, drawn out into an unusually long, slen- 
der, pubescent tip. Achene obpyramidal, 2.5-3 mm long, 
1.5-2 mm wide; apex truncate, margin may be cuspidate or 
crowned, apical hairs sparse; surface finely striated, lus- 
trous, dark brown. 


Old coral reefs or porous oolitie rock, Big Pine Key and 
the everglades keys, southern Florida. Flowers throughout 
the summer. Fig. 3. 

This rather diminutive, endemic species is quite distinct. 
It is perhaps most easily recognized by its sprawling or 
bushlike habit and its large, knobby root. The achenes and 
capitula of this species are, proportionally, fairly large 
while the number of florets per capitulum is small. Anoma- 
lous plants, likely a result of introgression from Melan- 
thera angustifolia, may exhibit slightly smaller capitula 
with broadly lanceolate phyllaries, Plants growing on Big 
Pine Key are a bit different morphologically, having 
spreading rather than flat margins. 


REPRESENTATIVE SPECIMENS EXAMINED. UNITED 
STATES: Florida: BROWARD C0., near new River Canal, S of Lake 
Okeechobee, Small 4441 (DUKE, FLAS, GH, NCU, NY). COLLIER CO., 
dry sandy pineland, E Naples, Moldenke 24160 (WIS). DADE CO., 
pinelands, Addison’s Hammock, Small 6639 (NY); cut pineland over 
limestone, Bauer Hammock, Webster 10115 (DUKE); hammocks, 
Black Pt. Creek, Small 5970 (mo, NCSC, NY); dry, sandy soil, Buena 
Vista, Moldenke 309 (DUKE, MO, Nv); 8 mi W of Everglades Ntl. 
Park inf. center, pinewoods, Parks 112 (EVG, F, FSU, GH, MO, NY, PAC, 
SMU, US, VDB, WIS) ; thin soil over old coral reef, Florida City, Kral 
12044 (VDB); pine-palmetto woods, Grossman Rd, Parks 291 (EVG, 
PAC, US, VDB); pinelands over limestone, Homestead, Brass 21149 


1973] Melanthera — Parks 203 


(GH), Godfrey 65609 (LAF, NCU, USF); vacant lot behind Tradewinds 
Motel, Parks 109 (DUKE, EVG, F, FSU, GH, MO, NY, PAC, SMU, US, VDB, 
WIS); pinelands, Inlikita, 8 mi N of Homestead, Parks 118 (EvG, F, 
FSU, NY, PAC, SMU, US, VDB); rocky land, Miami, Curtiss 5851 (Ds, 
F, FLAS, GA, GH, MO, NCU, NY, PAC, SMU, UC, US); NW 54th St. & 
27th Ave. Miami, C. & J. Javish 535 (ps, MO); pinelands, Murden 
Hammock, Small 6439 (Ny), 6450 (Mo, NY); Nixon-Lewis Hammock, 
Small 7351 (DUKE, NCU, NY). MONROE CO., open hammock, Big Pine 
Key, Brown 128 (NY, PH); pine-palmetto woods, Big Pine Key, 
Killip 40217 (ps, GA, US); pine-palmetto woods, end of Rt. 940, Big 
Pine Key, Parks 289 (EVG, SMU, US, VDB); hammocks, Boca Chica 
Key, Small 3918 (NY); Key West, Chapman s.n. (NY); Long Key 
Frost 8 Feb. 1931 (pac); Old Rhode’s Key, Small 6 Nov. 1901 (Ny). 


5. Melanthera ligulata Small, Bull. New York Bot. Gard. 3: 
439. 1905. Fig. 8. Type: Florida: Broward Co., 
pinelands below Ft. Lauderdale, 1904, Small & Wilson 
1775. (NY!) 


Suffrutescent, often luxurious perennial, 3-9 dm tall, 
arising from a slightly swollen root. Main axis decumbent, 
freely branched at the jointed nodes, sulcate to obscurely 
quandrangular, usually reddened or red-mottled on upper 
surfaces, glabrescent. Leaf shape highly variable, ovate- 
elliptical-lanceolate forms often on the same plant, 4-12 cm 
long, 1-4 em wide, noticeably triple-nerved, commonly light 
green, glabrescent; base acute-cuneate, obtuse or hastate, 
commonly basal lobes strongly ascending; apex acute, mar- 
gin serrate, lanceolate leaves saliently and irregularly 
toothed. Capitulum mostly solitary, terminal, borne on a 
suleate, glabrescent peduncle 6-14 em long; involucre in 
two loosely imbricate series 1.2-1.4 cm in diameter; phyl- 
laries lanceolate, 5-7 mm long, 2 mm wide, dark green for 
2/3-3/4 length apically, pallid basally with obscure nerves, 
pubescence short, appressed-ascending, phyllaries often 
noticeably longer than florets at first anthesis and re- 
curved; pales oblanceolate, 6-7 mm long, 2 mm wide, apical 
aristae about 1 mm long, may be recurved. Corolla tube 
4.5 mm long; limb spreading, 1 mm long, papillate adax- 
ially. Anther sacs 2 mm long; style branches 1.5 mm long, 
may completely recurve when senescent. Achene not dis- 


[Vol. 75 


Rhodora 


204 


+ x 
“+ * w 
hears xx 


Fig. 8. Melanthera ligulata Small. A. habit sketch, B. capitulum, 
C. phyllary, D. pale, E. achenes, F. meiotic chromosomes, late ana- 


phase one X 1000. 


1973] Melanthera — Parks 205 


tinctive, obpyramidal, 2.5 mm long, 1.5 mm wide; apex 
truncate to slightly concave, with short, tan to white hairs 
at maturity ; color dull tan. 


Pine-palmetto flatwoods, usually in sun, from central to 
southern Florida. Flowers in early summer. Fig. 3. 

This species of Melanthera is sparingly represented in 
herbaria and most of the specimens available have not 
been properly identified. It has not been well understood 
partly because it is restricted, largely to inaccessible areas 
such as the region south of Lake Okeechobee, and partly 
because the original circumscription is too narrow. As a 
result of my field observations in southern Florida, it 
became evident that certain anomalous plants seen were 
probably M. ligulata. I have included the considerable 
variability expressed by the leaves in the description. 

Melanthera ligulata is easily recognized in the field by 
its decumbent stem which is profusely branched from the 
slightly swollen nodes. On herbarium specimens, one must 
note the stem characters, a generally glabrous vesture, 
and the capitulum which is similar to, but a bit smaller 
than that of M. nivea. 


SPECIMENS EXAMINED. UNITED STATES: Florida: county 
unknown, South Florida, Chapman 63 (US). BROWARD CO., roadside, 
Ft. Lauderdale, Crevasse 23 July 1940 (FLAS); low pineland, Ham- 
mondville, Brown 16 March 1928 (FLAS). CHARLOTTE CO., Cabbage 
Hammock, Frye 181 (FLAS); pasture, Parrott 90 (DUKE). CITRUS CO., 
moist soil on limestone, 8 mi SW of Inverness, Kral 7891 (FLAS, GH, 
VDB). COLLIER CO., roadside spoil heap, 5.5 mi N of Rt. 41 on Rt. 79, 
Parks 308 A&B (EVG, F, FSU, GA, MICH, NCSC, NCU, NY, PH, SMU, US, 
VDB); roadsides in glades, Brass 15408 (vs); East Henson Marsh, 
Big Cypress, Brass 15960 (Uus); bank sloping to swamp, intersection 
Rts. 29 & 82, Immokalee, Lakela 27442 (SMU, USF), Parks 310 (Ds, 
DUKE, EVG, F, FSU, GH, MO, NY, PAC, SMU, UC, US, VDB, WIS) ; between 
Miles City & Immokalee Seymour 5177 (FLAS); ranchlands with 
cypress heads, Collier-Hendry Co. line, Rt. 846, Lakela 28959 (USF); 
railroad gravel, 10 mi N of Jerome, Godfrey 61003 (FSU); wet glade, 
W of Miles City, Lakela 29344 (usF); roadside, Tamiami Trail, 
Atwater M237 (EVG, FLAS). DADE CO., pinelands near Camp Longview, 
Small 18 May 1904 (GH, Us) (Sheet at US later labeled as no. 1575, 
type of Melanthera radiata is M. ligulata), Small 1807 (NY); Miami, 


206 Rhodora [Vol. 75 


Garber 1 May 1877 (FLAS, NY, PAC); everglades, Miami, Small 1764 
(NY); Tamiami Trail, W. M. R. 15 Feb. 1985 (NY). GLADES CO., 
roadsides on prairie, Palmdale, Brass 15449 (GH, US). HENDRY CO., 
weedy in burned live oak hammock, Ft. Denaud, Brass 20623 (US); 
roadside, 4 mi W of Labelle, Henderson 63-1608 (FSU, TEX). HILLS- 
BOROUGH CO., border, wet woods, Tampa, Britton 39 (NY). LEE CO., 
near swamp, Alva, Francis 152 (Us); flatwood ponds, Ft. Myers, 
Hilcheock 157 (F, GH, MO, NY, US); pineland, 8 mi SE of Ft. Myers, 
J. Standley 452 (F, GH, MO, US); dcoryard, near Ft. Myers, Standley 
19024 (US); prairies near Okaloacoochee slough, Small 8805 (NY). 
OSCEOLA CO. swamp, Fredholm 6100 (NY); pineland, Kissimmee, 
Singeltary 167 (DUKE, NCSC). PALM BEACH CO., sandy soil, 2 mi N of 
Canal Pt, Moldenke 252 (NY); Lake Worth, Beardslee Dec. 1928 
(uc), O'Neill 831 (us); dry, sandy, pine-palmetto flatwoods, Loxa- 
hatchee, Parks 89 (DS, DUKE, EVG, F, FLAS, FSU, GA, GH, MICH, MO, 
NCSC, NCU, NY, PAC, PH, SMU, UC, US, vDB, WIS), Parks 90 (SMU, 
VDB), 91 (VDB), 92 (EVG, PAC, SMU), 98 (FSU, US), 237 (DS, DUKE, 
F, GA, MO, NCSC, NY, PAC, PH, SMU, US, VDB, WIS). PINELLAS CO., sandy 
soil, rare, St. Petersburg, Mrs. C. Deam 2888 (us), 2898 (US). POLK 
co., moist, open woods, Kissengens Spring, Correll 6295 (DUKE, 
Ncsc), McFarlin 3033 (MICH). SUMTER CO., Esperawell, Smith 18 
March 1879 (vs). 


DOUBTFUL OR EXCLUDED SPECIES 


Melanthera fruticosa Brandgee, Univ. California Publ. 
Bot. 10: 421. 1924. = Philactis Liebmanii (Klatt) Blake, 
Contr. Gray Herb. n.s. 52: 35. 1917. Type: Mexico: Chi- 
apas: near Tuxtla along rocky road, Purpus 9117 (US!, 
isotypes Mo! NY! US!) Having examined the type collec- 
tion, I concur with the treatment of Torres (1969). 


Melanthera hastifolia Blake, Contr. U.S. Ntl. Herb. 24: 
29. 1922. Type: Guatemala: Izabal: near Cristina in ditch 
along railway, Blake 7601A (Us!, isotype US!). Only one 
collection other than the type collection was available for 
examination, Lundell 4928 at MICH, US, NY. These were 
determined by Blake, but are not like the type material 
and seem to represent M. aspera. I believe the type collec- 
tion of M. hastifolia probably represents elements of a 
hybrid swarm, with the putative parents being M. angusti- 
folia and M. nivea. 


Melanthera parviceps Blake, Jour. Washington Acad. 


1973] Melanthera — Parks 207 


Sci. 22: 384. 1932. Type: British Honduras: El Cayo 
District: in ravine, Little Mountain Pine Ridge, Bartlett 
11882 (us!, isotypes F! UC!) I have seen only four her- 
barium specimens of this taxon, all from the same locality 
and three from the type collection. Melanthera parviceps, 
on the basis of these few specimens, appears rather distinc- 
tive. Although similar to M. aspera var. glabriuscula in 
capitulum characters, the heads of M. parviceps are more 
numerous, smaller, and bear fewer florets. Also the pe- 
duncles and phyllaries are strongly hispid to hirsute. Veg- 
etatively the plant seems to resemble M. nivea, with the 
leaves being mostly narrowly ovate with divergent hastate 
lobes. Until more collections become available, I prefer to 
place this as a dubious species, rather than to recognize it, 
or reduce it to M. aspera var. glabriuscula. 


ACKNOWLEDGEMENTS 


I wish to thank Dr. Robert Kral of Vanderbilt Univer- 
sity for suggesting this problem and, as major professor, 
for offering suggestions and guidance instrumental in its 
solution. I also thank the late Dr. Lloyd H. Shinners of 
Southern Methodist University, Drs. William T. Stearn 
and J. E. Dandy of the British Museum, Dr. Alicia Lour- 
teig of Paris, and Dr. Richard Howard of the Arnold Ar- 
boretum for their aid with various aspects of the nomen- 
clature. Dr. Frank C. Craighead Sr. helped with field work 
in Everglades National Park. The graciousness of the 
various curators who provided valuable herbarium materi- 
als for study is also acknowledged. I am indebted to my 
wife Vicki for her continuous and real help in all aspects 
of this investigation. This work was based, in part, on a 
dissertation submitted to the Department of Genera] Bi- 
ology, Vanderbilt University, in partial fulfillment of the 
requirements for the Ph.D. degree. It was supported by a 
National Defense Education Act Graduate Fellowship to 
the author. 


LITERATURE CITED 


ADANSON, M. 1763. Familles des Plantes. Paris: Vincent, 2: 131. 


208 Rhodora [Vol. 75 


ANDERSON, E. 1949. Introgressive Hybridization. New York: John 


Wiley & Sons. 109 pp. 
BENTHAM, G. & J. D. HOOKER. 1873. Genera Plantarum, London: 
Reeve & Co. 2: 377. 
BLAKE, S. F. 1922. New plants from Guatemala and Honduras. 
Contr. U.S. Ntl. Herb. 24: 29-30. 
. 1924. New American Asteraceae. Contr. U.S. Ntl. 
Herb. 22: 628-629. 
1930. Notes on certain type specimens of American 
Asteraceae in European herbaria. Contr. U.S. Ntl. Herb. 26: 
254. 


1932. New Asteraceae, Jour. Washington Acad. Sci. 
22: 384-385. 

Britton, N. L. & P. WILSON. 1925. in Scientific Survey of Porto 
Rico and the Virgin Islands. New York Acad. Sci. 6: 308-310. 

Brown, R. 1817. Observations on the natural family of plants 
called Compositae. Trans. Linn. Soc. London, 12: 107-112. 

BnowNE, P. 1756. The Civil and Natural History of Jamaica 
(ed. 1). London: Osborne. 503 pp. 
CaRLQUIST, S. 1965. Island Life. A Natural History of the Islands 
of the World. Garden City: Natural History Press. 451 pp. 
CASSINI, H. 1823. in Levrault, F. G. (editor), Dictionnaire des 
Sci. Nat. Strasbourg & Paris. 29: 483-497. 

CHANEY, R. 1947. Tertiary centers and migration routes, Ecol. 
Monogr. 2: 139-148. 

COULTER, J. 1891. New or noteworthy Compositae from Guatemala. 
Bot. Gaz. 16: 100. 

CRONQUIST, A. 1955. Phylogeny and taxonomy of the Compositae. 
Amer, Midland Nat. 53: 478-511. 

CRUXENT, J. & I. Rouse. 1969. Early man in the West Indies. Sci. 
Amer. 221: 42-52. 

Danpy, J. 1969. Nomina conservanda proposita. (proposal 286). 
Taxon. 18: 470. 

DiLLENIUS, J. 1732. Hortus Elthamensis. London. 437 pp. 

GAERTNER, J. 1791. Fruct. 2: 456. 

Jacquin, N. 1788. Collectanea —. Vindobonae: Wappler. 2: 290. 

JENNINGS, O. 1917. Contributions to Botany of the Isle of Pines, 
Cuba. Annals Carnegie Museum, 11: 286. 

KUNTZE, O. 1891. Reviso Generum Plantarum. Leipzig. 1011 pp. 

LaANJOUW, J. (Chief ed.) & F. STAFLEU (Sec.) 1966. International 
Code of Botanical Nomenclature. Utrecht: Kemink & Zoon. 
402 pp. 


1973] Melanthera — Parks 209 


LINNAEUS, C. 1753. Species Plantarum. (reprint 1957, Ray So- 
ciety, London). 2: 833. 

1759. Systema Natura (ed. 10). Stockholm. 

1759. Amoenites Academie. Uppsala. 5: 371-388. 

MicHAUX, A. 1803. Flora Boreali-Americana. Paris & Argento- 
rati: Levrault. 2: 107. 

MiLLsPAUGH, C. 1907. Flora of the sand keys of Florida. Publ. 
Field Museum Nat. Hist. 2: 191-245. 

POWELL, A. & B. TURNER. 1963. Chromosome numbers in the Com- 
positae VII. Additional species from the southwestern United 
States and Mexico. Madrono. 17: 128-140. 

RICHARD, A. 1850. in La Sagra, de Ramon. Historia Fisica, Poli- 
tica y Natura de la Isla de Cuba. Paris: Maulde & Renou. 11: 
54. 

Ronm, J. von. 1792. Plantae-Slaegter poa St. Croix, med tilfoiede 
Anmaerkninger af Vahl. Skriv. Nat. Selsk, (Kiobenhaven). 2: 
205-227. 

ScHULZ, O. E. 1911. Compositarum Genera Nonnulla. i» Urban, 
I., Symbolae Antillanae. 7: 115-127. 

ScHWEGMAN, J. 1968. New plant records from southern Illinois. 
Transactions — Ilinois State Academy of Science. 61: 314. 
SMALL, J. 1917-19. The origin and development of the Compositae. 
New Phytologist. 16: 157-177; 198-221; 253-276. 1917. 17: 
13-40; 69-94; 114-142; 200-230. 1918. 18: 1-35; 65-89; 129-176; 

201-234. 1919. 

SMALL, J. K. 1903. Flora of the southeastern United States. New 
York: published by the author. 1370 pp. 

1905. Additions to the flora of subtropical Florida. 

Bull. New York Bot. Gard, 3: 439. 

1909. Additions to the flora of peninsular Florida. 
Bull. Torrey Bot. Club. 36: 163. 

STANDLEY, P. E. 1938. Flora of Costa Rica. Contr. Field Museum 
Nat. Hist. 18: 1493-1494. 

STROTHER, J. 1970. Typification of Melanthera Rohr (Compositae: 
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424 pp. 

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210 Rhodora [Vol. 75 


. & D. FLYR. 1966. Chromosome numbers in the Com- 
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Hist. 1: 103-112. 


DEPARTMENT OF BIOLOGY 
MILLERSVILLE STATE COLLEGE 
MILLERSVILLE, PA. 17551 


SENECIO ANONYMUS WOOD, AN EARLIER NAME 
FOR SENECIO SMALLII BRITTON' 


ROBERT R. KOWAL AND T. M. BARKLEY 


Taxonomists regret the occasional necessity to change 
long established specific epithets. While we firmly believe 
that adherence to the code of nomenclature is a practical 
necessity, we offer the present paper with some trepida- 
tion. 

Senecio anonymus Wood is a widespread and often weedy 
plant of the southeastern United States. It has been known 
in the past almost exclusively as Senecio smallii Britt. 

When Alphonso Wood introduced the epithet anonymus 
into the literature in 1861 (Class-book, p. 464), he pre- 
sented it without citing an author’s name. As Wood cited 
authors for other species, one might conclude that he 
merely noted an unnamed species of Senecio. However, 
in his American Botanist and Florist (1870, p. 187), Wood 
cited the name as Senecio anonymus Wood, thereby indi- 
cating his intent to use anonymus as a specific epithet. 
Article 23, Note 1, of the current International Code of 
Botanical Nomenclature (Stafleu, 1972) provides for the 
rejection of words not intended as names, but here anony- 
mus must be regarded as an intentional name. The later 
publication date for S. anonymus (with the author’s cita- 
tion) antedates S. smallii Britt. (1894). 

The name S. anonymus Wood has been previously found 
and indexed, but on the basis of the description in Wood’s 
Class-book, Greenman treated it as a synonym for S. to- 
mentosus Michx. (1803) (Merrill, 1948). Wood’s (1861) 
descriptions of S. tomentosus and S. anonymus are as 
follows: 


4 S. tomentosus Mx. Clothed with soft, cotton-like, 


‘Contribution No. 1167, Division of Biology, Kansas Agr, Expt. 
Station, Manhattan. Support by National Science Foundation Grant 
GB 5449 is gratefully acknowledged. 


211 


212 Rhodora [Vol. 75 


nearly persistent tomentum; root lvs. oblong or ob- 
lanceolate or ovate, obtuse, tapering to a long, slender 
petiole, crenate, the upper sessile; hds. fastigiate, rays 
12 to 15; ach. pubescent. — 4 Va. to Fla. and La. 
St. 1 to 2f high, often nearly leafless above. Corymb 
simple, subumbellate. Root lvs. with their petioles 
6 to 9’ long, 1 to 3’ wide. Rays spreading 16". Apr. 
— Jn. — The leaves are exceedingly variable. A 
variety (on Stone Mt., Ga.) is low, densely tomentous 
[sic], with the lvs. all radical. 


5 S. anonymus. Plant clothed with a white, partly 
deciduous tomentum; root lvs. small, oblong, obtuse, 
crenate-serrate, some of them slightly lobed, taper- 
ing to a petiole, cauline lvs. long and narrow, re- 
motely sinuate-pinnatifid, the segm. cut-dentate; hds. 
subumbellate, small, ach. pubescent. —4 ? Mont- 
gomery, Ala. St. 16 to 24’ high. Root lvs. 14’ wide 
and with their petioles 2 to 3' long. St. lvs. 6' long, 
the upper 1’, almost bipinnatifid. Rays 8 to 10, 
spreading about 7". May., Jn. 


Except for the tomentum mentioned, S. anonymus is clear- 
ly described in the original description as having the nar- 
row basal leaves, pinnatifid cauline leaves and small heads 
regularly associated with the entity long-called S. smallii. 

Alfonso Wood's herbarium eventually came to the College 
of Pharmaceutical Sciences, Columbia University (Mer- 
rill, 1948) and from there most of it went to the New York 
Botanical Garden. A search of the latter institution's her- 
barium produced no specimen resembling or bearing the 
name S. anonymus Wood. However, Dr. Frank Pokorny 
of the College of Pharmaceutical Sciences kindly located 
for one of us (R.R.W.) what we take to be the holotype 
among specimens left in the teaching collection. The holo- 
type will now be permanently deposited in the New York 
Botanical Garden. The holotype bears the following label 
data: “Ex herbario Alfonso Wood / Senecio anonymus / 
Montgomery, Ala. / Legit ipse." No other data accompany 


1973] Senecio anonymus — Kowal and Barkley 213 


ve 


weed (IDEI), a name arta- 
datin 2- ima Gettin 0. 
(eq), the latter nse 
being the na^ few vita 
for the spec thet flee 
Spem repre sets 
Robert Ñ. Kowal, 
Sex. 1,1967. 


Figure 1. Holotype of Senecio anonymus Wood 


[Vol. 75 


Rhodora 


214 


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215 


Senecio anonymus — Kowal and Barkley 


1973] 


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216 Rhodora [Vol. 75 


the specimen, which consists of but two flowering stalks 
and no basal rosettes (fig. 1). 

The holotype appears to be a rather unexceptional speci- 
men of the species. The tomentum mentioned in the ori- 
ginal description is presumably what led Greenman to 
conclude that S. anonymus was a synonym for S. tomen- 
tosus, but the holotype is glabrescent except for a per- 
sistent tomentum in leaf and branch axils. This is a 
common feature in the species now called S. anonymus. 
A particularly noteworthy point is that S. tomentosus is 
not known to oecur in the type locality (ie., the vicinity 
of Montgomery, Alabama) where individuals identified 
as S. smallii recently have been seen (Joab L. Thomas, 
John D. Freeman, and Wilbur H. Duncan, Zn lit.). 

Because many aureoid senecios appear to introgress 
freely with each other (Barkley, 1962; Kowal, 1968), there 
is always room for some doubt as to the biological nature 
of any specimen. Indeed, Chapman and Jones (1971) re- 
port limited natural hybridization between S. tomentosus 
and S. anonymus (as S. smallii) and, on the basis of arti- 
ficial crosses, discuss the potential for introgression be- 
tween them. In order to strengthen our argument, the 
holotype of S. anonymus was compared with materials of 
S. tomentosus and S. anonymus that were collected on 
May 11 and 18, 1969, along a roadside 4.1 miles east of 
Castalia, Nash Co., N. C. (Table 1; vouchers: Kowal, s. n., 
at wis). Here the two species grew together with limited 
but evident putative hybridization. 

Diseriminant analysis was used to confirm our inter- 
pretation of the holotype. This multivariate statistical 
technique can be used to objectively identify unknown 
individuals as belonging to one of two taxa or as being 
intermediate (Fisher, 1936; Morrison, 1967). Given 
means, variances, and convariances for a set of characters 
from each of the two taxa, discriminant analysis determines 
the weights for the weighted average (or coefficients for 
the linear combination) of the original characters that 
most efficiently separates the two taxa. It results in a 


_ 


1973] Senecio anonymus — Kowal and Barkley 217 


discriminant function, Y = a,X, + a, X, + ... + ap Xp 
— e, where the Xi's (i = 1,..., p) are measurements on 
p characters on an individual, the ai's (i = 1,... p) are 
the weights (or coefficients), and c is an arbitrary constant. 
Entering actual measurements (X; X, ..., Xp) from an 
individual into the function gives that individual's dis- 
criminant score, Y. Comparing the discriminant score of 
an unknown individual with corresponding scores of indi- 
viduals known to belong to the two taxa enables one to 
classify the unknown as one of the taxa or as an inter- 
mediate. In the present analysis logarithms of nine char- 
acters that could be measured on the type or derived from 
Wood's description were used to construct the discriminant 
function separating the North Carolinian local populations 
of S. tomentosus and S. anonymus. The coefficients and 
arbitrary constant of the discriminant function were cal- 
culated such that the pooled intraspecific variance of scores 
equalled one and such that the mean score for the two 
species were equal in absolute size but opposite in sign. 
Table 2 lists characters, coefficients, and coefficients stand- 
ardized to facilitate evaluating the importance of characters 
to the discriminant function (as the original coefficients are 
dependent on arbitrary units of measurement). The holo- 
type’s score (2.4) is more than five standard deviations 
from the mean score of S. tomentosus (—93.3) but within 
one standard deviation of the mean score for S. anonymus 
(3.3). Thus, discriminant analysis substantiates treating 
the holotype of S. anonymus as a perfectly good member 
of the species heretofore called S. smallii Britt. 
The synonomy for Senecio anonymus now becomes: 


Senecio anonymus Wood, Class-book of Botany (ed. 1861), 
p. 464. 1861. Type coll.: Wood, s.n., Montgomery, Ala. 
(NY). 

Senecio smallii Britton, Mem. Torr. Bot. Club 4: 132. 1894. 
Senecio aureus var. angustifolius Britt. Mem. Torr. Bot. 
Club 2: 39. 1890. Type coll.: Brown et al., “The Pass." 
Peaks of Otter, Va. (NY, PH). 


218 Rhodora [Vol. 75 


Table 2 


Coefficients of discriminant function separating Senecio 
tomentosus from S. anonymus using characters measur- 
able on the holotype of S. anonymus Wood. 


Standardized 

Character Coefficients — coefficients* 
1. Stalk height 6.0 .67 
2. Number of cauline leaves 0.2 04 
3. Inflorescence — no. of heads -1.5 —.55 
4. Inflorescence — no. of branches 2.4 54 
5. Involucre — length 17.7 1.00 
6. Involucre — width at base -4.5 —.41 
7. Ray floret — number 3.0 7 
8. Ray floret — ligule length -1.0 —.11 
9. Ray floret — ligule width 0.2 .03 
Constant 43.2 29.00 


*Standardized coefficients are those that would be used on 
data standardized such that the pooled intraspecific vari- 
ance of each character equals one. The coefficient that is 
largest in absolute value is arbitrarily set equal to one. 


1973] Senecio anonymus — Kowal and Barkley 219 


Senecio earlei Small, Bull. Torr. Bot. Club 25: 147. 1898. 
Type coll.: Earle & Underwood, Auburn, Lee Co., Ala. 
(NY). 

Senecio smallii f. tristis Fernald, Rhodora 47: 302. 1945. 
Type coll.: Fernald 14,859, Ram-Hole Swamp, Seward 
Forest, Near Triplett, Brunswick Co., Va. (GH). 


LITERATURE CITED 


BARKLEY, T. M. 1962. A revision of Senecio aureus Linn. and allied 
species. Trans. Kan. Acad. Sci. 65: 318-408. 

FisHer, R. A. 1936. The use of multiple measurements in taxo- 
nomic problems. Ann. of Eugenics 7: 179-188. 

KowaL, R. R. 1968. Senecio aureus and allied species on the Gaspé 
Peninsula of Quebec. Ph.D. thesis, Cornell University, Ithaca, 
N. Y. 331 p. 

MERRILL, E. D. 1948. Unlisted new names in Alphonso Wood's 
botanical publications. Rhodora 50: 101-130. 

Morrison, D. F. 1967. Multivariate statistical methods. McGraw- 
Hill, N. Y. 338 p. 

STAFLEU, F. (ed.) 1972. International code of botanical nomen- 
clature. Regnum Vegetabile 88: 1-426. 

Woop, A. 1861. Class-book of botany. Barnes and Burr, N. Y. 
832 p. 

1870. The American botanist and florist. A. S. Barnes 

and Co., N. Y. and Chicago. 172 + 392 p. 


DEPARTMENT OF BOTANY 
UNIVERSITY OF WISCONSIN 
MADISON, WISCONSIN 53706 


HERBARIUM, DIVISION OF BIOLOGY 
KANSAS STATE UNIVERSITY 
MANHATTAN, KANSAS 66502 


A NOTE ON THE KARYOLOGY OF HAPLOPAPPUS 
SPINULOSUS (PURSCH) DC. SSP. TYPICUS HALL 


L. S. GILL AND C. C. CHINNAPPA 


Karyotypic studies and chromosome behaviour at meiosis 
are important to understand the genetic systems. Though 
some species of the genus Haploppus have been studied in 
this manner, no such information is available for Haplo- 
pappus spinulosus spp. typicus Hall. This is a small peren- 
nial Composite which ranges from Alberta to Minnesota 
and southwards to Texas and as far as Mexico. Several 
chromosome counts for this species have been recorded 
(Jackson 1957, 1962; Li and Jackson 1961; Dejong and 
Longpre 1963; Solbrig et al 1969). All of them, however. 
are from the southern range of its distribution. 

The purpose of this note is to report the karyology of 
this species from the northernmost part of its distribution. 

Seeds were collected from Brooks, Alberta in May 1971. 
Plants were raised in the greenhouse at the University 
of Waterloo. Meiotic and mitotic preparations were made 
following Gill (1971 a, b). Voucher (L. S. Gill 353) 
specimens are preserved in the University of Waterloo 
herbarium. 

One hundred well spread metaphase cells from each of 
two plants were analyzed. Root tip cells of both plants 
have 8 chromosomes. One plant has 0-2 B— chromosomes 
(Figures 5, 6, 7). A haploid chromosome set is shown 
diagramatically in Figure 8. The smallest pair of chromo- 
somes has almost median centromeres. Two pairs have 
submedian centromeres and the longest pair have sub- 
terminal centromeres. The smallest pair with the near 
median centromere is designated as ‘A’, and the rest as 
B, C, D respectively with the size increase. The length 
of Chromosome A is 8.2 microns and D is 13.1 microns. 
The arm length ratio are A- 0.87, B- 0.54, C- 0.80, and 
D- 0.15. Chromosome C has a distinct secondary constric- 
tion on its short arm. Table 1 presents a few more details. 


220 


1973] Haplopappus — Gill and Chinnappa 221 


The relative length and the centromere index are deter- 
mined following Anderson (1970). The relative length 
reflects the amount of size homogeniety in each chromo- 
some complement and centromere index shows the degree 
of terminalization of the centromere. 


Table 1 


Chromosome morphology of H. spinulosus spp. typica. 
Absolute chromosome 


length in microns. Relative Centromere 
Plant longest average length index 
1 13.2 10.3 0.61 1.63 
2 13.1 10.1 0.68 1.65 


One hundred meiotic cells in each plant were examined 
and all had a consistent haploid chromosome number of 4. 
with no B— chromosomes. The pair of A form a regular 
ring bivalent, pair of D as a rod bivalent. The other two 
submedian pairs are seen to form as two rings, but less 
frequently (Figures 1-4). 

Li and Jackson (1961) mention that in H. spinulosus 
spp. cotula the smallest two regular chromosomes have an 
almost median centromere and are about 8.4 microns in 
length. There are no details mentioned about the other 
sets. They report the occurrence of differences in B- 
chromosome number in the cells of roots and flowers. B- 
chromosome number was not only different between root 
and shoot, but also existed from cell to cell within the 
same flower. The elimination of supernumerary chromo- 
somes was not complete in the root as one plant was found 
to have 2n = 8 + 1. 

In view of the above, it is interesting to note the elimi. 
nation of supernumerary chromosome in some of the plants 
of the subspecies typicus in its northern part of distribu- 
tion. Earlier records of chromosome counts for ssp. typicus 
are n — 4 (Jackson, 1957), but different counts such as 
2n — 8 + 2,8 + 3,8 + 4, were reported in ssp. cotula 


[Vol. 75 


Rhodora 


N 


N 


1973] Haplopappus — Gill and Chinnappa 223 


Fig.a8 


Figure 8. Idiogram of haploid chromosome set for Haplopappus 
spinulosus ssp. typicus. X 4000. 


Figure 1. Two ring and two rod bivalents at diakinesis. 2500. 

Figure 2. Three ring and one rod bivalents in diakinesis. X 2500. 

Figure 3. Three ring and one rod bivalents in diplotene. X 2500. 

Figure 4. Late disjunction of one ring bivalent at anaphase one. 
x 2500. 


Figure 5. Mitotic metaphase 2n = 8 + 2B chromosomes. > 2500. 

Figure 6. Mitotic metaphase with 2n — 8 + 1B chromosome. 
>< 2500. 

Figure 7. Four mitotic cells with 8 chromosomes in each cell. 
> 1000. 


224 Rhodora [Vol. 77 


(Li and Jackson, 1963). More recently Solbrig et al. 
(1969) reported n — 8 for ssp. spinulosus, which repre- 
sents a new ploidy level for this species. 

Chromosomal diversity, number and karyology, is evi- 
dent within the species. A comprehensive study of the 
species and related ones from their entire range of dis- 
tribution in North America is being undertaken to under- 
stand the evolutionary divergence in this interesting group 
of species. 


LITERATURE CITED 


ANDERSON, L. C. 1970. The Karyotype of Chrysothamnus parry? 
ssp. parryi and its implications. Trans. Kansas Acad. Sciences 
72(3): 399-401. 

DaRLINGTON, C. D. 1963. Chromosome botany and the origin of 
cultivated plants. George Allen and Unwin Ltd. London. p. 231. 

DEJoNG, D. C., and E. K. LONGPRE. 1965. Chromosome studies in 
Mexican Compositae. Rhodora 65(763): 225-240. 

GILL, L. S. 1971a. Cytology of West-Himalayan Labiatae: Tribe 
Satureineae. Caryologia 24(2): 203-207. 

1971b. Chromosome number of Lysimachia ciliata L. 
Rhodora 73(796): 556-557. 

JACKSON, R. C. 1957. Documented Chromosome numbers of plants. 
Madrono 14: 111. 

1962. Interspecifie hybridization in Haplopappus 
and its bearing on chromosome evolution in the Blepharodon 
Section. Am. J. Bot. 49(2): 119-182. 

li, N., and R. C. Jackson, 1961. Cytology of supernumerary 
chromosomes in Haplopappus spinulosus ssp. cotula Am. J. Bot. 
48(5): 419-426. 

SoLBRIG, O. T. L. C. ANDERSON, D. W. Kynos and P. H. RAVEN 
1969. Chromosome numbers in Compositae VII: Asterae IH 
Am. J. Bot. 56(3) : 348-353. 


DEPARTMENT OF BIOLOGY, 
UNIVERSITY OF WATERLOO, 
WATERLOO, ONTARIO 


FLORAL STRUCTURE, HYBRIDIZATION AND 
EVOLUTIONARY RELATIONSHIP OF 
TWO SPECIES OF MIMULUS' 


Y. T. KIANG 


Floral characters of angiosperms have long been recog- 
nized as a useful guide in plant systematics. The diverse 
floral structures represent results of long periods of natu- 
ral selection and often reflect a close relationship with 
breeding systems of plant species (Stebbins, 1970). Fur- 
thermore, they also play an important role in reducing 
interspecifie crossing (Grant & Grant, 1965). Therefore, 
studies of floral characters in relation to breeding systems 
and ecological adaptation of plant species would contribute 
toward the understanding of processes of plant speciation. 


Mimulus guttatus D. C. (Yellow monkey flower) and 
M. nasutus Green (Fig. 1) are two closely related species 
abundantly occurring in moist areas in California (Grant, 
1924). The flowers of both species have yellow corollas 
with variable numbers and sizes of red blotches (Abrams, 
1951). Mimulus guttatus is a perennial, self-compatible, 
but normally outcrossing species (Kiang, 1972), while 
M. nasutus is an annual and predominantly autogamous 
species. Since both species have similar geographical dis- 
tribution, yet adopt distinctly different growth and breed- 
ing habits, they are desirable for the study of phylogenetic 
relationship by examining floral morphologies, hybridiza- 
tion and phenology in relation to their ecologica] adapta- 
tion. This paper reports the floral structure, breeding 
systems, hybridization and the possible evolutionary rela- 
tionship of the two species. The studies indicate that the 
two species are phylogenetically closely related and M. 
nasutus is probably a daughter species of M. guttatus. 


‘Published with the approval of the Director of the New Hampshire 
Agricultural Experiment Station as Scientific Contribution No. 622. 


225 


Rhodora 


[Vol. 75 


1973] Mimulus — Kiang 227 


MATERIALS AND EXPERIMENTAL PROCEDURE 


Populations of Mimulus guttatus and M. nasutus were 
established in the greenhouse from seeds collected from 10 
randomly chosen plants from a single population of each 
species. The M. guttatus population was growing in gran- 
itic sandy soil along shallow channels of a creek about 
800 m west of the entrance of Eureka State Park, Cali- 
fornia (120.70°W., 39.37°N.; elevation about 1820 m), 
while the M. nasutus population was in a roadside ditch 
watered by a small seep in the road cut of State Highway 
70, Feather River Canyon near Rock Creek, California 
(121.28°W., 39.98°N.; elevation about 610 m). 

Crosses between the two species were made on twenty 
plants of each species. In order to avoid contamination 
caused by the precocious dehiscence of anthers of Mimulus 
nasutus, emasculation had to be performed on the flower 
buds two or three days before they opened. 

Since flowers of both species exhibit some degree of 
intraplant variation, only the third or the fourth node 
flowers on main stems were used for measuring the size 
of floral parts as well as estimating pollen viability by 
staining pollen with lactophenol aniline blue. The number 
of seeds from the mature capsules was counted, and the 
seed viability tested by germination. Total fertility and 
productivity of the two species and the hybrids were 
assessed. Chromosome counts were made from pollen 
mother cells following the techniques used by Mia et al. 
(1964). Both species were found to be diploid with n — 14. 


RESULTS AND DISCUSSION 


1. Floral characteristics and breeding system. 


Viable seeds were harvested from all capsules on which 
interspecific pollination was attempted. Mimulus guttatus 


Fig. 1. Mimulus guttatus (left) and M. nasutus (right) grown 
in the same pot (4") for about 8 weeks in the greenhouse. Many 
capsules can be seen on M. nasutus, while M. guttatus is just coming 
into bloom. 


[Vol. 75 


Rhodora 


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1973] Mimulus — Kiang 229 


has smaller calyces and larger corollas than M. masutus 
(Table 1). Floral parts of M. guttatus are relatively stable 
with respect to size. The stability of the flora] character- 
istics is essential for the insect-pollinated plant species in 
achieving successful seed production. Any drastic change 
in floral parts may result in failure of pollination by pol- 
linators. Contrary to the rather stable flower size of M. 
guttatus, M. nasutus exhibits morphological plasticity of 
the flower, particularly corolla size. Flowers are progres- 
sively reduced in size on a plant, and after ten nodes or so 
on the main stem, flowers often become so small as to be 
inconspicuous. Since M. nasutus is predominantly self- 
pollinated, there is no necessity to attract insect visitors 
by having large, showy flowers. 

Both species have two pairs of stamens, one short and 
one long, in each flower. The two pairs of stamens in 
Mimulus nasutus are more or less similar in length and 
the anthers are clustered around the stigma. The bilabiate 
stigma, small and insensitive, always remains open, even 
in the flower bud. Thus, the structural arrangement of 
floral parts of M. masutus greatly facilitates successful 
self-pollination. 

In contrast, the pistil in Mimulus guttatus is much 
longer than the stamens, which themselves differ in length. 
The different stamen lengths may aid the visiting insects 
to obtain pollen. The large and sensitive bilabiate stigma 
opens up only when it becomes receptive. 

The floral morphologies of the F, hybrids are somewhat 
intermediate between those of the parent species. Although 
there is a slight difference in the length of stamens and 
pistils, in the pollinator-free greenhouse every capsule 
produced by the hybrid is fully loaded with seeds. Self- 
pollination is facilitated by the lower lip of the stigma 
curling all the way back toward the anthers. Except for 
its autogamy, the morphology of the F, hybrid flower is 
more like that of Mimulus guttatus. The frequency dis- 
tributions of the sum of three ratios (calyx to corolla, 
ovary to style and stamen to pistil) for the parent species, 


230 Rhodora [Vol. 77 


M.guttatus M. nasutus 


4 
Fi x M.nasutus 


04 
> 2 F, x M. guttatus 
= 
x 
= 
= 14 
« 
a— 
ka 
o T T T T Y T T T T T 
1.4 1.6 18 20 2.2 24 2.6 2.8 3.0 


Sum of the ratios 


Fig. 2. Frequency distribution of the sum of ratios (calyx/corolla, 
ovary/style, stamen/pistil) of M. guttatus, M. masutus and their 


hybrids. 


1973] Mimulus — Kiang 231 


the F,, F, and backcross generations are plotted as histo- 
grams (Fig. 2). The distributional patterns suggest that 
the sum of the three ratios may be useful in studying hy- 
bridization and introgression in natural populations of 
Mimulus. 


The time of anther dehiscence and occurrence of self- 
pollination in Mimulus nasutus may be a significant factor 
in seed setting and in reducing gene exchange between the 
two species. Twenty flowers each from one plant were 
emasculated by pulling off the corollas within 10 hours 
after the flowers were open. No seed was produced by the 
emasculated flowers of M. guttatus. In the case of M. 
nasutus, all emasculated flowers yielded on the average as 
many seeds as from non-emasculated ones. In the F, 
hybrid, 8 out of 20 emasculated flowers (or 40%) produced 
seed, and the average number of seed per capsule was only 
49, much smaller than that produced by a non-emasculated 
flower. 


When flower buds of the F, and Mimulus nasutus were 
carefully dissected to examine anthers, no pollen shedding 
was noticed in the F, hybrid, but in M. nasutus, the an- 
thers had dehisced, and the stigmas were full of pollen 
grains. One day or two prior to the opening of the flower, 
the style of M. nasutus bends inward slightly, bringing the 
open stigma in contact with the dehiscent anthers to ensure 
self-pollination. Therefore, in M. nasutus pollination nor- 
mally occurs before flowers open. Pseudocleistogamy has 
been used to describe this phenomenon (Vickery, 1964). 


In annual plants, seed set in a season is one of the most 
crucial factors for the continuing survival of populations. 
Plants of Mimulus nasutus usually start to produce cap- 
sules when plants are still at the 2 true leaf stage. The 
first two or four precociously produced capsules are smaller 
than normal ones, and usually set seeds by cleistogamy. 
Several larger chasmogamous flowers very similar to those 
of M. guttatus will follow the first four flowers; then, as 
the blooming progresses, the flowers rapidly reduce in size. 


232 Rhodora [Vol. 75 


2. Fertility and biomass 


Pollen fertility. 


On the average, 98% of Mimulus nasutus pollen and 
93% of M. guttatus pollen were stained. The F, pollen 
(88% stained) was about 10% less fertile relative to that 
of M. nasutus and the difference is statistically significant. 

Mimulus guttatus on the average produced more pollen 
grains per flower than M. nasutus (30368 + 218 vs. 20958 
+ 44). The amount of pollen produced by the F, hybrid 
(24006 + 135) was approximately intermediate between 
the parental species. In M. nasutus, less than one-third of 
pollen grains remained in the anthers of newly open flow- 
ers (6775 + 65), because pollen shedding occurs before 
flowers open. The precocious dehiscence of anthers in M. 
nasutus may reduce pollen flow within and between spe- 
cies. There is no evidence of nectar glands in Mimulus 
(Grant, 1924); thus, insects visiting the flowers are for- 
aging for pollen. Generous pollen production would be 
advantageous for M. guttatus in achieving outcrossing. 


Seed production and germination. 


Mimulus guttatus produces on the average 401.5 + 94 
seeds per capsule, M. nasutus 256.4 + 39 and the F, 
492. + 96. The data indicate that the F, can potentially 
produce more seeds per capsule than the parental species. 

In order to test viability, 200 seeds from each species, 
F, and the F, backcross were sown on moist filter paper. 
The germination percentages were Mimulus guttatus, 
90.5% ; M. nasutus, 84.096; F, 84.4% and F, backcross, 
74.5%. There was no significant difference in seed via- 
bility between the parental species and the F, hybrid. 
However, the proportion of seed germination declined 
appreciably in the seed of the F, backcross. 


Production of flowers, capsules and biomass. 

To assess productivity, plants were harvested after 
growing in the greenhouse for three months. The average 
number of flowers, capsules, stolons, and the dry biomass 


233 


Mimulus — Kiang 


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234 Rhodora [Vol. 75 


are presented in Table 2. When the productivity of the 
two species was compared, the differences between them 
were found to be statistically significant. Mimulus nasutus 
started to bloom early and terminated early. It usually 
branches from the base of the stem, and soon all branches 
become ascending with flowers. 

When the reproductive effort of the plants was exam- 
ined, Mimulus nasutus invested more energy in seed pro- 
duction than M. guttatus and the F,. The flowers and 
capsules consisted of 36.696 of the total biomass in M. 
nasutus, but only 4.5% in M. guttatus. Mimulus nasutus 
is an annual, and seed production is crucial for continua- 
tion of the annual species. On the other hand, vegetative 
propagation is more important to M. guttatus, a perennial, 
for maintaining its population. In fact, M. guttatus showed 
more vegetative growth than M. nasutus, and produced a 
considerable amount of stolons, or 14.5% of the total bio- 
mass (Table 2). 

In order to determine whether hybridization between 
the two species is generally possible, three additional popu- 
lations of the two species and plants from Feather River 
Canyon and Eureka State Park were intercrossed. They 
were all interfertile. 

All the experimental data indicate that barriers do not 
exist to prevent gene exchange either between Mimulus 
guttatus and M. nasutus, or between the parental species 
and the F, hybrids. The F, plants under greenhouse con- 
ditions were quite vigorous and showed no sign of sterility 
or breakdown. 


3. Relationship of Mimulus guttatus and M. nasutus. 


It has been well established that self-fertilized plant 
species are always derived from outbreeding ancestors, and 
annuals are most likely derived from perennials (Stebbins, 
1957; Baker, 1959). The evidence from floral structure, 
breeding systems and fertility of hybrids strongly sup- 
ports the view that Mimulus guttatus and M. nasutus are 
genetically closely related species and the latter is newly 


1973] Mimulus — Kiang 235 


derived from the main species, M. guttatus (Vickery, 
1964). It has been demonstrated with many closely re- 
lated pairs of species in Clarkia that one member of the 
pair is derived from the other (Vasek, 1958; Lewis and 
Raven, 1958). Lewis (1962) has suggested that the 
daughter species in Clarkia are derived from marginal 
populations which are often subject to catastrophic selec- 
tion leading to a rapid genetic reorganization and forma- 
tion of the daughter species. The evolutionary relationship 
of Mimulus guttatus and M. nasutus may represent such 
a rapid evolution as a consequence of the joint action of 
mutation, selection and genetic drift. 

Changes of breeding systems from outcrossing to self- 
pollination under the unusual ecological environments have 
been reported. Gilia achilleaefolia subsp. multicaulis, pre- 
dominantly autogamous, is derived from normally outcross- 
ing, but self-compatible G. a. subsp. achilleaefolia (Grant 
and Grant, 1965). A similar change has been observed in 
Mirabilis longiflora, a self-compatible species in a genus 
where self-incompatibility is known, which in the absence 
of successful insect visitation sets seeds by self-pollination 
(Baker, 1961a). A change of breeding system from out- 
breeding to inbreeding in plant species could trigger the 
rapid formation of a new species (Baker, 1961b). Al- 
though a direct observation of the formation of Mimulus 
nasutus from M. guttatus is not possible, one is tempted to 
reconstruct a hypothetical evolutionary process of M. nasu- 
tus based on the evidence gathered in this study, and on the 
available examples cited above. 

Mimulus guttatus is a riparian species which maintains 
its populations largely by long life span and by growing 
profuse stolons as propagules in nature (Kiang and Libby, 
1972). However, only in a moisture-stable environment 
can it grow as a perennial and reproduce vegetatively. 
Therefore, if environmental conditions of a peripherally 
isolated population happen to become dry and unstable, 
the population may no longer be able to maintain itself by 
vegetative propagation. Thus, seed production becomes a 


236 Rhodora [Vol. 75 


decisive factor for continuation of the population under 
arid conditions, which would stimulate a rapid evolution 
(Axelrod, 1967). Moreover, it is probable that insects may 
fail to visit the small number of flowers produced by one 
or a few plants surviving from the catastrophic selection 
(Lewis, 1962). Therefore, under conditions of unstable 
moisture availability, only individuals with successful self- 
pollination can produce seeds, and autogamy has obvious 
immediate high selective value (Stebbins, 1950; Grant & 
Grant, 1965). 


When the environment becomes favorable, a new popu- 
lation may be established from the seeds produced by self- 
pollination. The newly founded colony with self-pollination 
will lead to genetic change, and those new, well adapted 
individuals would become the progenitors of the daughter 
species. Thus, the daughter species, Mimulus nasutus may 
have been derived from one or several plants which ob- 
tained a new adaptive mode under the favorable conditions. 


The greenhouse and field observations strongly support 
the hypothesis that Mimulus nasutus is derived from M. 
guttatus under moisture unstable conditions. In the green- 
house 30 seeds of each species were sown in each of the 5 
moisture saturated 8” pots. One week after the seeds were 
sown, irrigation was completely discontinued. About 86% 
of Mimulus guttatus and 83% M. nasutus germinated. No 
M. guttatus survived to produce flowers, but 8 M. nasutus 
per pot survived to produce on the average 3.6 mature 
capsules. In nature, Mimulus nasutus occurs more com- 
monly in the southern parts of the Sierra, where moisture 
supply is less dependable. Under normal conditions, it 
reaches the peak of bloom in relatively short days, April 
and May when moisture is most abundant (J. L. Hamrick, 
personal communication). The phenology of Mimulus nasu- 
tus, larger cotyledons with a rapid seedling growth and 
precociously produced capsules by self-pollination, are 
clearly indicative of its adaptation to a moisture unstable 
environment. 


1973] Mimulus — Kiang 201 


CONCLUSION AND SUMMARY 


The breeding systems, floral structure, and phenology of 
Mimulus guttatus and M. nasutus were compared. The 
former is an outcrossing perennial while the latter is a 
self-pollinated annual. Mimulus guttatus has large, showy 
corollas and long pistils with sensitive labiate stigmas, 
whereas M. nasutus has small corollas which are variable 
in size, and short pistils with anthers clustered around 
stigmas. The anthers of Mimulus nasutus dehisce pre- 
cociously and, therefore, self-pollination occurs before flow- 
ers open. Greater variation with regard to severa] quanti- 
tative traits was observed in M. guttatus than in M. nasu- 
tus. Mimulus nasutus has a higher reproductive effort than 
M. guttatus, which exhibits more vegetative growth. 


Fully fertile hybrids were obtained experimentally. The 
growth pattern and floral characters of the F, hybrids are 
intermediate between the parental species. The hybrids 
can set seed by self-fertilization and grow as perennials 
with stolons. 


A comparison of fertility and productivity indicates that 
there is no complete postmating isolating mechanism be- 
tween M. guttatus and M. nasutus. The evidence suggests 
that the two species are genetically closely related and M. 
nasutus may be a daughter species newly derived from 
M. guttatus. Mimulus nasutus may represent the product 
of quantum evolution (Grant, 1971) as a consequence of 
the joint action of mutation, selection and genetic drift. 


ACKNOWLEDGMENTS 


I gratefully acknowledge the generosity of Professors 
James L. Hamrick and Robert K. Vickery, Jr., in providing 
the seeds for this investigation. I wish to thank H. G. 
Baker, and J. L. Hamrick for reading the early draft of the 
manuscript, and their valuable suggestions. This investiga- 
tion was partly supported by a UNH 1972 Faculty Summer 
Fellowship. 


238 Rhodora [Vol. 75 


LITERATURE CITED 


ABRAMS, LEROY. 1951. Illustrated flora of the Pacific states. Vol. 
III. Stanford Univ. Press. 866 p. 

AxELROD, D. I. 1967. Drought, diastrophism, and quantum evolu- 
tion. Evolution 21: 201-209. 

BAKER, H. G. 1959. Reproductive methods as factors in speciation 
in flowering plants. Cold Spring Harbor Symp. Quant. Biol. 
24: 177-191. 

1961a. The adaptation of flowering plants to noc- 
turnal and crepuscular pollinators. Quart. Rev. Biol. 36: 64-73. 

1961b. Rapid speciation in relation to changes in 
breeding systems of plants. (Pp. 881-885) Im Recent advances 
in Botany. Toronto Univ. Press. Toronto. 

GRANT, ADELE L. 1924. A monograph of the Genus Mimulus. Ann. 
Misso. Bot. Garden 11: 99-389. 

GRANT, V. 1971. Plant speciation. Columbia Univ. Press. New 
York. 485 p. 

, and K, A. GRANT. 1965. Flower pollination in the Phlox 
family. Columbia Univ. Press. New York. 

KiawG, Y. T. 1972. Pollination study in a natural population of 
Mimulus guttatus. Evolution, 26: 308-310. 

. and W. J. Lippy. 1972. Maintenance of a lethal in 
a natural population of Mimulus guttatus. Amer. Nat. 106: 
351-367. 

Lewis, H. 1962. Catastrophic selection as a factor in speciation. 
Evolution 16: 257-271. 

, and P. H. Raven. 1958. Rapid evolution in Clarkia. 
Evolution 12: 319-336. 

Mra, M. M., B. B. MUKHERJEE, and R. K. VICKERY, JR. 1964. 
Chromosome counts in the section simiolus of the Genus Mimulus 
(Scrophulariaceae). IV. New numbers in M. guttatus, M. tigri- 
nus, and M. glabratus. Madrono 17: 156-160. 

STEBBINS, G. L., JR. 1950. Variation and evolution in plants. Co- 
lumbia Univ. Press. New York. 643 p. 

1957. Self-fertilization and population varia- 

bility in the higher plants. Amer. Nat. 91: 337-354. 

1970. Adaptive radiation of reproductive 
characteristics in Angiosperms, I: Pollination mechanisms. In 
Annual Rev. of Ecol. and Systematies. Vol. 1: 307-326. 

VASEK, F. C. 1958. The relationship of Clarkia exilis to Clarkia 
unguiculata, Amer. Jour. Bot. 45: 150-162. 

Vickery, R. K., JR. 1964. Barriers to gene exchange between 
members of the Mimulus guttatus complex (Scrophulariaceae). 
Evolution 18: 52-69. 


DEPARTMENT OF PLANT SCIENCE 
UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, NEW HAMPSHIRE 03824 


PALYNOLOGICAL EVIDENCE FOR THE 
LATE GLACIAL OCCURRENCE OF PRINGLEA 
AND LYALLIA ON KERGUELEN ISLANDS: 


STEVEN B. YOUNG? 


EILEEN K. SCHOFIELD’ 


Kerguelen Islands are an isolated subantarctic archi- 
pelago situated in the southern Indian Ocean (Fig. 1). The 
nearest continental land masses other than Antarctica are 
Africa and Australia, each some 4,000 km distant. The 
native vascular flora of Kerguelen includes only about 30 
species (Lourteig and Cour, 1963; Aubert de la Rüe, 1964). 
The majority of the non-endemic species have their affinities 
with the Magellanie flora of southern South America. A 
lesser number of species also occur in New Zealand and 
southern Australia. A few of the species found on Ker- 
guelen, such as Cystopteris fragilis (L.) Bernh. are 
nearly worldwide in their distribution. 

Six species are endemic to Kerguelen or to Kerguelen 
and such nearby islands as Heard and Crozet. Both the 
quantity and the nature of the endemic species are unique 
in the subantarctic, and this region has been considered 
to be a distinct phytogeographic province (Good, 1964), 
although most other provinces support floras of over 100 
times as many species. 

The validity of the concept of the Kerguelen phytogeo- 
graphic province depends upon the assumption that the 
flora has been isolated for a long period of time. However, 
much of the surface of Kerguelen is presently covered with 
ice, and there is some reason to believe that virtually all 
of the island may have been glaciated, eliminating all sig- 


"Contribution No. XXX of the Institute of Polar Studies, The Ohio 
State University. 
"Present address: Center for Northern Studies, Wolcott, Vermont 


05680. 
"Present address: New York Botanical Garden, Bronx, New York 


10458. 


239 


240 Rhodora [Vol. 75 


T 
89 E 
o ° 
a 
D i 
*KERGUÉLEN 
Q 
N ç 
9 N 
fy T? 10KM 
F 49's 49?s -] 
c eo 
90 
c 
Bo ° 
c» 


(44 
o 
o, oo 
s o7 
GEN CORE2 
A š 9 CORE! 


69°E 
1 sE | 


Fig. 1. Map of Kerguelen Archipelago with stars marking sites 
of cores. Inset shows position of Kerguelen in relation to Africa 
and Australia. 


nificant vegetation during the Pleistocene. We now offer 
evidence that the most characteristic members of the Ker- 
guelen endemic flora were present on the islands during 
the latter part of the last major glacial episode. 

Four of the six species endemic to the Kerguelen region 
(Poa Cookii Hook, f., P. kerguelensis (Hook. f.) Steuder, 
Colobanthus kerguelensis Hook. f., and Ranunculus Mosleyi 
Hook. f.) belong to large genera which are well represented 
throughout most of the far south. Although all four species 
are reasonably distinct, there is no intrinsic reason to 
postulate a long period of isolation on Kerguelen for them. 


1973] Kerguelen Islands — Young and Schofield 241 


The remaining two endemic species, Lyallia kerguelensis 
Hook, f. and Pringlea antiscorbutica R. Br. ex Hook. f., 
belong to monotypic genera which have no close relatives. 
In the subantarctic flora these two genera are unique with 
regard to their taxonomically isolated position, and they 
provide the main support for the contention that the Ker- 
guelen region can properly be considered to be distinct 
phytogeographic province. It is known that Kerguelen 
supported a complex flora, including coniferous trees, 
during the mid-Miocene (Cookson, 1947; Nougier, 1970). 
If Kerguelen has never been completely denuded of its 
flora by ice advances, it is probable that Lyallia and 
Pringlea are the last living remnants of a Tertiary Ker- 
guelenian flora. 

Pringlea antiscorbutica, the well-known “Kerguelen 
cabbage", was used by seamen during the last century 
for the control of scurvy. It occurs on Kerguelen, Heard, 
Crozet, Marion, and Prince Edward Islands (Lourteig and 
Cour, 1963). Pringlea (Fig. 2) bears a superficial resem- 
blance to the cultivated cabbage. The vegetative portion 
of the plant consists of a rosette of blue-green leaves which 
may lie nearly flat on the ground or may be borne atop a 
fleshy rootstock to 1 m tall and 10 em in diameter. The 
inflorescence consists of one to several spikes which arise 
from the base of the rosette at its junction with the root- 
stock. The petals of the numerous flowers are reduced or 
absent. 

Rabbits, which were introduced on Kerguelen in 1874 
(Aubert de la Riie, 1964) have apparently eliminated 
Pringlea from many of its former habitats, and it is now 
difficult to reconstruct the former natural distribution of 
the species. In areas not presently inhabited by rabbits, 
Pringlea is common to abundant in mesic situations near 
sea level, particulary where the soil has been enriched by 
sea-bird colonies. Pringlea will tolerate salt spray, and it 
is often found growing with Cotula plumosa Hook. f. in 
coastal situations. Plants of Pringlea are also found in 
high-elevation vegetation dominated by Azorella selago 


242 Rhodora [Vol. 75 


Fig. 2. Pringlea antiscorbutica R. Br. ex Hook. f. showing basal 
rosette of leaves and fruiting stalks. Although this specimen was 
found at an elevation of about 600 m, the leaves show evidence of 
rabbit grazing. 


1973] Kerguelen Islands — Young and Schofield 243 


Fig. 3. Lyallia kerguelensis Hook. f. 


Hook. f. where there is abundant moisture from seepage 
or meltwater. It is doubtful that Pringlea was ever a 
significant component of the vegetation of the upland wind 
deserts of Kerguelen. The species is never seen in boggy 
areas. It appears that Pringlea is tolerant of low tempera- 
tures and high winds, but that it requires a continuous 
supply of moisture and good drainage. 

Lyallia kerguelensis occurs only on the Kerguelen archi- 
pelago. It is a dwarf cushion plant (Fig. 3) which is 
rather rare over most of the main island, although it is 
common on Long Island in the Gulf of Morbihan (H. A. 
Imshaug, personal communication). Lyallia grows mainly 
on stony barrens at low to moderate elevations. 

Lyallia, with its small and highly reduced flowers, has 
been something of a puzzle to taxonomists since its dis- 
covery. First placed in the Caryophyllaceae by Hooker 
(1847), it was later transferred to the Portulacaeae ( Kid- 
der, 1876). More recently, Lyallia has been linked with 
the genus Hectorella, which is native to alpine areas in 
New Zealand, and both genera are considered to be in a 


244 Rhodora [Vol. 75 


separate family, the Hectorellaceae (Philipson and Skip- 
worth, 1961). The pollen of Lyallia, which is clearly 
neither portulacaceous or caryophyllaceous, supports the 
placement of the genus in another family (Cranwell, 1963) . 


PALYNOLOGICAL STUDIES 


In February 1971, columns of peat and buried soils were 
collected from the sides of stream cuts in two locations on 
the south shore of the Gulf of Morbihan (Fig. 1). These 
cores have since been analyzed for pollen content and dated 
by radiocarbon techniques. 

Core 1 is 308 cm long. A C-14 date at a depth of about 
240 cm was 9230 + 150 years B.P. (I-6661), while a date 
from a depth of about 250 cm was 8595 + 125 years B.P. 
(I-6154). This small anomaly in the dates is most probably 
a result of contamination of the lower sample with recent 
material, but it could also be caused by the presence of 
reworked older sediments in the upper sample. If the 
240-cm date is taken to be the most nearly correct, the 
lowermost strata of core 1 would probably have been 
deposited 12,000 to 12,500 years B.P. If the 250 cm date 
is accepted, the oldest strata would be 1,000 to 1,500 years 
younger. The strata below the bottom of core 1 contained 
large, rounded boulders embedded in blue clay-silt. These 
strata may be of glacial origin. 

Core 2, which was 550 cm long, did not penetrate to any 
till.like strata. Buried organic soils were found below the 
bottom of the core, but seepage problems made sampling 
impossible. A sample from a depth of approximately 525 
em was dated at 11,010 + 160 years B.P. (1-6284). If we 
consider the late glacial-postglacial boundary in the far 
southern hemisphere to be about 10,000 years B.P. (Heus- 
ser, 1966; Flint, 1971), both cores include late glacial 
sediments. 

There can hardly be doubt that the glaciers of Kerguelen 
were more extensive during parts of the Pleistocene than 
they are now, but evidence for total glaciation of the islands. 


1973] Kerguelen Islands — Young and Schofield 245 


b 
PRINGLEA J = 


| asa a o o o TI e e pu | 


CORE 2 


LYALLIA 


CORE 1 


oo8, S 
PRINGLEA nenne 
LL + > 


——À o oo © i onthe 


DEPTH,CM >o 


5004 


o eo 
$ 
= E: š 


Fig. 4. Partial pollen diagram showing distribution of Pringlea 
and Lyallia pollen in cores 1 and 2. The percentages are based on a 
total count of 150 grains per sample. Each division at the top repre- 
sents 10% and a dot represents 2% or less. 


is lacking (Bellair, 1965). As the site of core 2 was ice-free 
well into glacial times, we suggest that it may not have 
been glaciated during the Pleistocene. 

The pollen and spore content of the Kerguelen cores is 
presently being analyzed as a means of reconstructing the 
late Quaternary climatic history of the islands. Neither 
Pringlea nor Lyallia was found to be a major constituent 
of the pollen rain, but some pollen of both species was 
found in the cores (Fig. 4). Pringlea pollen was present 
in the lowermost samples of both cores. At the bottom 
of core 2, Pringlea pollen amounted to 6% of the total 
pollen counted, a percentage as high as was found in any 
other sample in either core. Pringlea was a constituent 
of the vegetation near the site of core 2 more than 11,000 
years ago. If the relative lack of Pringlea pollen in some 
of the upper levels of the cores has any significance, it 
may indicate that Pringlea was replaced at low elevation 
by less cold tolerant species. The Pringlea pollen content 
of core 1 was similar to that of core 2 except that there 
was no peak at the lowest levels. 

Lyallia pollen was much rarer than that of Pringlea; it 
was never found in core 2 and was found in only three 
samples from core 1. However, the presence of clearly 
identifiable Lyallia pollen at the bottom of core 1 shows 
that the species did grow on the island more than 10,000 
years ago. Lyallia is neither an abundant plant nor does 
it flower heavily on Kerguelen at present; thus, its com- 
parative rarity in the pollen record is not surprising. 


246 Rhodora [Vol. 75 


In view of the evidence presented here, we may state 
with certainty that both Pringlea and Lyallia were present 
on Kerguelen during late-glacial times. The presence of 
pollen of these species in our oldest samples, coupled with 
the lack of evidence for complete glaciation of Kerguelen, 
suggests that these species did indeed “overwinter” the 
Pleistocene epoch on Kerguelen. It is not necessary to 
postulate an extra-insular refugium for these species, with 
subsequent migration to Kerguelen and extinction in the 
refugial area. Pringlea and Lyallia are probably true 
relicts of an otherwise extinct Tertiary Kerguelenian flora. 


ACKNOWLEDGMENTS 


This research was supported by National Science Foun- 
dation grant GV-26139. Logistic support for field work 
on Kerguelen was supplied by TAAF (Terres Australes 
et Antarctiques Francaises). We thank E. D. Rudolph 
and J. F. Splettstoesser for commenting on the manuscript 
and P. A. Colinvaux for use of his laboratory facilities. 

LITERATURE CITED 

AUBERT, DE LA RUE, E. 1964. Observations sur les caractéres et la 
répartition de la végétation des Iles Kerguelen. Com. Nat. 
Franc. Rech. Antarct. No. 10, Biologie 1: 1-60. 

BELLAIR, P. 1965. Un exemple de glaciation aberrante: les Iles 
Kerguelen. Com. Nat. Franc. Rech. Antarct., No. 11: 1-27. 
Cookson, I. C. 1947. Plant microfossils from the lignites of 
Kerguelen Archipelago. Brit. Aust. N. Z. Antarct. Res. Exped. 

1924-31. II (Geology): 127-142. 

CRANWELL, L. M. 1963. The Hectorellaceae: pollen type and taxo- 
nomic speculation. Grana Palynol. 4: 195-202. 

FLINT, R. F. 1971. Glacial and Quaternary Geology. John Wiley, 
N. Y. 892 p. 

Goop, R. 1964. The Geography of Flowering Plants. 3rd ed. J. 
Wiley, N. Y. 518 p. 

HEUssER, C. J. 1966. Polar hemisphere correlations of palynological 
evidence from Chile and the Pacifie northwest of America. In: 
Sawyer, J. S. (ed.) World Climate from 8000 to 0 B.C. Proc. 
Roy. Meteorol. Soc. Int. Symp. 124-141. 

Hooker, J. D. 1847. Botany of the Antarctic Voyage of H. M. 
Discovery Ships Erebus and Terror .... II: 548. Reeve Bros., 
London. 


1973] Kerguelen Islands — Young and Schofield 247 


KIDDER, J. H. 1876. Contribution to the natural history of Ker- 
guelen Island, made in connection with the United States Tran- 
sit-of-Venus Expedition, 1874-75. Bull. U.S. Nat. Mus. 3: 1-122. 

LourTEIG, A. and P. Cour. 1963. Essai sur la distribution géogra- 
phique des plantes vasculaires de l'archipel de Kerguélen. Com. 
Nat. Franc. Rech. Antarct. No. 3: 65-78. 

NOUGIER, J. 1970. Contribution a l'étude géologique et géomorpho- 
logique des Iles de Kerguelen. Com. Nat. Franc. Rech. Antarct. 
27: 1-440. 

PHILIPSON, W. R. and J. B. SKIPWoRTH. 1961. Hectorellaceae: 
a new family of dicotyledons. Trans. Roy. Soc. N.Z., Botany 
1(4): 32. 


INSTITUTE OF POLAR STUDIES 
THE OHIO STATE UNIVERSITY 
COLUMBUS, OHIO 43210 


PHYTOGEOGRAPHY OF THE CARICES 
OF VIRGINIA 


A. M. HARVILL, JR. 


The carices growing in Virginia include Carex, a large 
and cosmopolitan genus with a preponderance of species 
in temperate regions of the Northern Hemisphere, and the 
monotypic genus, Cymophyllus, endemic to the Southern 
Appalachians. Carex is by far the largest genus of vascu- 
lar plants in Virginia, with 121 species recognized in this 
study. Each species of the carices is, as a rule, well-defined, 
has its own distributional pattern, and has particular eco- 
logical requirements. There are woodland, meadow, bog, 
marsh, swamp, rock, calcicolous, and arenaceous species. 
The geographic affinities of the native Virginia carices are 
circumboreal, Asiatic-American, and strictly American. 
Within the state of Virginia these species show most of the 
patterns occurring in other vascular plants. 

During the course of this study, counties’ records from 
voucher specimens were plotted on outline maps of Vir- 
ginia, and these data will be published at a later date. 
Collections were studied at the College of Wiliam and 
Mary, Gray Herbarium, Lynchburg College, National Ar- 
boretum, New York Botanical Garden, University of North 
Carolina, Old Dominion University, Philadelphia Academy 
of Science, Smithsonian Institution, West Virginia Uni- 
versity, Virginia Commonwealth University, and Virginia 
Polytechnie Institute. The writer is indebted to the respec- 
tive curators for their generosity. 

In general, I have taken a rather wide view of some 
species, for, until some of the closely related taxa are better 
understood, Joseph Hooker’s rationale in his work on 
arctic plant distribution (1862) still seems most useful 
for phytogeographic purposes: *My main object is to show 
affinities of the polar plants, and | can best do this by keep- 
ing the specific idea comprehensive. It is always easier to 
indicate differences than to detect resemblances . . D 


248 


1973] Carices of Virginia — Harvill 249 


This paper would have been much less complete but for 
the outstanding collections of Dr. Henry K. Svenson, mostly 
from our coastal plain; and those of Mr. Charles E. Stevens 
from many areas of the state, especially the higher levels 
on mountains of western Virginia. Four of Stevens' col- 
lections were new records for the state: Carex brevior, C. 
pallescens, C. polymorpha, and C. rostrata. Also, Dr. John 
Wurdack gave many helpful suggestions, Dr. Sydney 
McDaniel pointed out the Fernald-Long collection of C. 
chapmanii, and Dr. Svenson identified our specimen of 
C. pallescens. 

For the most-frequently collected species, C. lurida, l 
have examined specimens from 83 of the 100 old counties 
of Virginia, counties before the incorporation of Elizabeth 
City, Norfolk, Princess Anne, and Warwick counties as 
the cities of Hampton, Norfolk and Chesapeake, Virginia 
Beach, and Newport News. On the other hand, eight spe- 
cies have apparently been collected in but a single county : 
C. chapmanii, Greensville; C. extensa, Norfolk; C. palles- 
cens, Grayson; C. polymorpha, Rockingham; C. rostrata, 
Bath; C. tetanica, Sussex; C. trichocarpa, Washington; and 
C. woodii, Shenandoah County. 

I have seen specimens for all included records with the 
exceptions of C. biltmoreana and C. cherokeensis, and 
these are based on the authority of the Manual of the 
Vascular Flora of the Carolinas (Radford, A. E., et al. 
1968). 

The 121 Virginia species of the carices grow in the fol- 
lowing patterns: 


1. Circumboreal. 


C. brunnescens (Pers.) C. lasiocarpa, Ehrh. 
Poir. C. muricata L. 

C. buxbaumii Wahl. C. pallescens L. 

C. canescens L. (also C. rostrata Stokes 
Australia) 


2. North America and eastern Asia. 
C. pedunculata Muhl. C. pensylvanica Lam. 


250 Rhodora [Vol. 75 


3. Ranging west to the Pacific Coast. 
C. brevior (Dewey) C. leptalea Wahl. 
Mackenz. C. nigromarginata Schwein. 
C. comosa Boott C. scoparia Schk. 
C. eburnea. Boott C. stipata Muhl. 
C. hystricina Muhl. C. umbellata Schk. 
C. interior Bailey C. vulpinoidea Michaux 


4. Ranging southward beyond the United States. 
C. albolutescens Schwein. C. lurida Wahl. 
C. bromoides Schk. C. physorhyncha Lieb. 


5. Extending inland to the Middle West. 
a. Generally distributed in the north and south. 


C. amphibola Steudel C. jamesii Schwein. 

C. artitecta Mackenz. C. laevivaginata (Küken.) 

C. atlantica Bailey Mackenz. 

C. blanda. Dewey . laxiflora Lam. 

C. caroliniana Schwein. . leavenworthii Dewey 

C. cephalophora Muhl. . lupuliformis Sartwell 

C. complanata Torrey & . lupulina Schk. 
Hooker muhlenbergii Schk. 

C. debilis Michaux . oligocarpa Schk. 

C. digitalis Willd. . retroflexa Muhl. 

C. emmonsii Dewey rosea Schk. 


C. festucacea Schk. 

C. folliculata L. 

C. frankii Kunth 

C. granularis Muhl. 
C. grisea Wahl. 

C. howei Mackenz. 

C. intumescens Rudge 


sparganoides Muhl. 
. Striatula Michaux 

. stricta Lam. 

. styloflexa Buckley 
. tribuloides Wahl. 

. typhina Michaux 

. willdenowii Schk. 


aaaaaaaaaaaaaaa 


b. Generally northern in distribution. 


C. albursina Sheldon C. communis Bailey 
C. annectens Bicknell C. conjuncta, Boott 
C. argyrantha, Tuckerm. C. crinita Lam. 

C. bushii Mackenz. C. cristatella Britton 


C. careyana Torrey C. gracilescens Steudel 


1973] Carices of Virginia — Harvill 251 


C. gracillima Schwein. C. shortiana Dewey 
C. grayii Carey C. squarrosa L. 
C. hitchcockiana Dewey C. suberecta (Olney) Britton 
C. laxiculmis Schwein. C. swanii (Fernald) Mackenz. 
C. leptonervia Fernald C. tenera, Dewey 
C. normalis Mackenz. C. tetanica Schk. 
C. plantaginea Lam. C. torta Boott 
C. platyphylla Carey C. trichocarpa Muhl. 
C. prasina Wahl. C. trisperma Dewey 
C. scabrata Schwein. C. virescens Muhl. 
c. Generally southern in distribution. 
C. cherokeensis Schwein. C. flaccosperma Dewey 
C. crebriflora Weigand C. oxylepis Torrey & Hooker 


d. Coastal plain and Mississippi Valey, extending west- 
ward in the northern states. 
C. alata Torrey C. hyalinolepis Steudel 
C. decomposita Muhl. C. seorsa Howe 


e. Coastal plain and Mississippi Valley, extending west- 
ward in the south. 


C. abscondita Mackenz. C. joori Bailey 
C. crus-corvi Shuttl. C. louisianica Bailey 
C. gigantea Rudge C. reniformis (Bailey) Small 


C. glaucescens Elliott 


6. Limited to eastern North America. 
a. Generally distributed north and south. 
(1) Mostly on the coastal plain. C. walteriana 
Bailey. 
(2) Mostly on the coastal plain but extending in- 
land in the south. 


C. barrattii Schwein. & C. collinsii Nuttall 
Torrey 
C. bullata Schk. C. venusta, Dewey 


b. Plants mostly of northern distribution. 
C. aestivalis M. A. Curtis C. vestita, Willd. 
C. baileyi Britton C. woodii Dewey 
C. polymorpha Muhl. 


252 Rhodora [Vol. 75 


c. Southern species. C. chapmanii Steudel 


7. Southern Appalachian endemics. 
C. biltmoreana Mackenz. Cymophyllus fraseri 
(Andrz.) Mackenz. 


8. Introduced species. 

C. arenaria L. Europe C. kobomugi Ohwi e. Asia 
C. divisa Hudson Europe C. spicata Hudson Eurasia 
C. extensa, Good. Europe 


It is clear from this analysis that the vast majority of 
Virginia species are limited in range to the eastern part 
of North America, and it is probable that most of them 
had their origin here. This picture is not a true repre- 
sentation of the Virginia flora as a whole, however, be- 
cause, as stated earlier, the carices are predominantly 
species of temperate regions of the Northern Hemisphere. 
If the other genera of sedges were included in this study, 
the picture would more nearly approximate that of the 
entire flora because most of those genera have strong 
tropical affinities. Among their species are many aggres- 
sive, opportunistic types which are, nevertheless, virtually 
confined to our coastal plain. The piedmont here becomes 
a formidable barrier to the spread of these species, ap- 
parently because of topography (and lack of many types 
of habitats), soils, and even the vegetation itself. The 
closed oak-hickory-pine communities, growing on “tight,” 
clayey soils, are not communities through which many 
species can spread (Braun, 1950, p. 510). Some of these 
relationships are strikingly brought out by the distribu- 
tional patterns within Virginia. 

Within the state of Virginia, Carex intumescens has the 
widest range of all the 121 species, growing not only at sea 
level near the coast, but across the state and to the top of 
Mt. Rogers at 5719 feet. At the other extreme are the 
seven native species, each known from a single county; 
and although some of them may very well turn up else- 
where, we can be sure that they have very narrow environ- 
mental tolerances. 


1973] Carices of Virginia — Harvill 253 


The 116 native species show the following patterns 
within the state: 


1. Generally distributed across the state. (44 species) 

2. Coastal plain. (20 species) 

C. barrattii C. hyalinolepis 

C. bullata C. joori 

C. chapmanit C. louisianica 

C. cherokeensis C. lupuliformis 

C. collinsit C. oxylepis 

C. erus-corvi C. reniformis 

C. decomposita C. tetanica 

C. gigantea C. venusta 

C. glaucescens C. vestita 

C. howei C. walteriana 

3. Piedmont. C. bushii 

4. Coastal plain and piedmont. (7 species) 

C. alata C. grayi 

C. comosa C. typhina 

C. conjuncta C. umbellata 

C. emmonsii 

5. Mountains. (20 species) 

C. aestivalis C. muricata 

C. albursina C. pallescens 

C. argyrantha C. pedunculata 

C. biltmoreana C. polymorpha 

C. brevior C. rostrata 

C. brunnescens C. suberecta 

C. careyana C. trichocarpa 

C. eburnea C. trisperma 

C. hystricina C. woodii 

C. leptonervia Cymophyllus fraseri 
. Mountains and piedmont. (9 species) 


6 
C. communis C. interior 

C. cristatella C. plantaginea 
C. hitchcockiana C. shortiana 


254 Rhodora [Vol. 75 


C. sparganoides C. willdenowu 
C. torta 


7. Disjunct. (15 species) 
a. Coastal plain — mountains. 


C. albolutescens C. lasiocarpa 
C. baileyi C. leptalea 
C. bromoides C. oligocarpa 
C. buxbaumii C. seorsa 

C. canescens C. virescens 


C. folliculata 


b. Mountains — piedmont outlier (calcareous soils) 
C. scabrata 


c. Mountains — piedmont outlier (calcareous soils) — 
coastal plain (on marl) C. jamesii and C. prasina 


d. Mountains — eoastal plain (connected via the 
northern piedmont) C. swamnii 


These data show that almost 13% of the 116 native spe- 
cies have ranges with major discontinuities. Those with 
the most pronounced breaks are the coastal plain — moun- 
tain disjuncts. Of these eleven species, three have their 
larger populations on the coastal plain: C. albolutescens, 
C. folliculata, and C. seorsa. Four are more widespread in 
the mountains: C. baileyi, C. leptalea, C. oligocarpa, and 
C. virescens. The distribution of the four other species is 
about evenly balanced between the two areas: C. bromoides, 
C. buxbaumii, C. canescens, and C. lasiocarpa. For two of 
the last group, C. buxbaumii and C. lasiocarpa, we have 
collections from only two counties for each in the state, 
and oddly enough, in both cases they are at Big Meadows 
on the Blue Ridge in Madison County and the savanna-like 
swales of Sussex County on the coastal plain. It is also 
noteworthy that Big Meadows is one of the few locales in 
the state for Campanula aparinoides Pursh. The rare col- 
lections of the marsh-bellflower all come from our moun- 
tains except one from an amazingly isolated pocket, dis- 


255 


Harvill 


irginia — 


Carices of V 


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covered by Fernald, around a sandy, sphagnous springhead 
almost at sea level and, again, in Sussex County. (Fig. 1). 

Big Meadows is now the only station in Virginia for 
Sanguisorba canadensis L., yet, 10,000 to 12,000 years ago 
the Canadian burnet was a significant member of the veg- 
etation in the Dismal Swamp area when spruce, pine, birch, 
alder, Lycopodium lucidulum Michaux, and L. clavatum 
(Whitehead, 1965) were other prominent members of this 
vegetation. 

In his phytogeographie considerations of the coastal 
plain flora of Virginia, Fernald (1937) states, “Some stu- 
dents maintain that Coastal Plain species are moving into 
the ancient uplands. . . . however, it seems to me more 
probable that plants and animals of long-established and 
conservative groups should have moved out from the an- 
cient lands . . ." Later he quotes Pennell on these disjunc- 
tions: “Certain it is that our study of the relations of the 
Southern Appalachians and Coastal Plain have shown that 
migrations may occur in either direction." Since Fernald, 
Pennell, and many more, in fact, considered this problem, 
the paleobotanical work of Craig (1969), Knox (1969), 
Whitehead, and others has thrown considerable light on 
plant migrations in this region (Harvill, 1972). "These 
records of fossils clearly indicate that climatic oscillations 
of Pleistocene and Recent times brought plants migrating 
to and fro across the state. And it is reasonable to believe 
that with such climatic reversal, some species with narrow 
environmental tolerances would lose major portions of 
their ranges, whereas other species could persist in a few 
isolated areas where there are some unusual combinations 
of environmental factors. Therefore, it appears that the 
odd and disjunct ranges of these plants have most probably 
come about through the interplay between widespread 
climatie oscillations and local environmental conditions 
related to soils and sites, with both of these complexes of 
factors acting upon the tolerances of populations of these 
species which now have such discontinuous distributional 
patterns. 


1973] Carices of Virginia — Harvill 251 


LITERATURE CITED 


BRAUN, E. Lucy. 1950. Deciduous forests of eastern North Amer- 
ica. Philadelphia, Blakiston Co. 596 pp. 

CRAIG, A. J. 1969. Vegetational history of the Shenandoah Valley, 
Virginia. Geol. Soc. Am. Spec. Paper 123: 283-296. 

FERNALD M. L. 1937. Local plants of the inner coastal plain of 
southeastern Virginia. Rhodora 39: 465-491. 

HARVILL, A. M. 1972. The historical significance of some disjunct 
distributional patterns in Virginia. Castanea 37: 137-140. 
Hooker, J. D. 1862. Outlines of the distribution of arctic plants. 

Trans. Linn. Soc. London 23: p. 283. 

Knox, A. S. 1969. Glacial age marsh, Lafayette Park, Washing- 
ton, D. C. Science 165: 795-797. 

WHITEHEAD, D. R. 1965. Palynology and Pleistocene phytogeogra- 
phy of unglaciated eastern North America. In Wright, H. E. 
and D. G. Frey (eds.), The Quaternary of the United States. 
Princeton Univ. Press Princeton pp. 417-432. 


LONGWOOD COLLEGE 
FARMVILLE, VIRGINIA 23901 


THE MARINE ALGAE OF 
VIRGINIA AND MARYLAND 
INCLUDING THE CHESAPEAKE BAY AREA: °: 


FRANKLYN D. OTT 


INTRODUCTION 


Until 1965 there existed no publication treating any 
aspect of the benthic or attached marine algae of the coast 
of Virginia and Maryland including the Chesapeake Bay 
area. Prior to this date only minor references to particular 
species for this area can be found in the literature. This 
paucity of research can be attributed, in the main, to the 
lack of solid substrata permitting prolonged and extensive 
development of algal populations. Time spent on more 
favorable habitats such as the rocky shores of the New 
England coast offer, to the phycologist, far more rewarding 
results for his endeavors. 

The areas of the Virginia and Maryland coast facing 
directly upon the Atlantic are unstable, constantly moving, 
wave washed, sandy beaches. Behind the so-called Barrier 
Islands lie thousands of acres of Spartina salt marshes, In 
the shallow bays, between the Barrier Islands and the main- 
land, are abundant oyster reefs which offer a more stable 
substratum for algal growth. Since, however, the waters 
of these bays may reach temperatures in excess of 30°C 
during the summer only a limited number of algal taxa are 
to be found. The muddy bottoms of these bays, among the 
oyster reefs, offer a continuous supply of sediments which 
are constantly kept in suspension by the tidal currents. 
Consequently, light penetration is not great. Even in shal- 
low waters it is virtually impossible to visually locate the 
algal populations. 


"This work is a result of research sponsored by NOAA Office of 
Sea Grant, Department of Commerce, under Grant #1-36032 to the 
Virginia Institute of Marine Science. 

'Virginia Institute of Marine Science Contribution No. 429. 


258 


1973] Marine Algae — Ott 259 


In contrast to the higher salinities of the Atlantic coast, 
the Chesapeake Bay is a considerably more brackish area 
with salinities dropping significantly as one proceeds north- 
ward. Some stretches of sandy beaches do exist but they 
are subjected to less wave action than those of the Atlantic. 
Again Spartina salt marshes border extensive areas of the 
bay. The lower salinities, especially in the upper bay, ex- 
clude a great number of marine algal taxa. Some areas of 
the bay locally support a typical freshwater algal flora 
because of the steady and constant influx of fresh water. 
The upper reaches of all the estuaries are continually pour- 
ing into the water system a freshwater algal population. 
While such freshwater algal populations are not actively 
growing and will not long survive at the higher salinities, 
they often will be encountered on the microscopical exam- 
ination of water samples. Their contribution to the overall 
ecology of the area can only be speculative at the present 
state of our knowledge. 


Into this inhospitable habitat for marine algae man has 
constructed wood and rock jetties, breakwaters, piers, re- 
taining walls, groins, and other structures. These offer a 
satisfactory, however unnatural, substrate for algal growth 
which are almost immediately colonized by algae. These 
types of man-made structures offer to the investigator an 
opportunity, with a reasonable expenditure of energy, to 
secure some insight into the composition of the algal popu- 
lation. 


In 1965 the first significant publication on algae in this 
area appeared. Zaneveld and Barnes (1965) reported on 
the annual periodicity of the marine algae in the lower 
Chesapeake Bay. In the six following years, several papers 
have appeared reporting the algal flora of a particular rock 
jetty, geographical area, or habitat. In no case, however, 
does any of these published papers give an accurate picture 
of the wide variety of marine algae to be encountered in 
the states of Virginia and Maryland. In fact, many papers 
because of their limited scope leave the reader under the 


260 Rhodora [Vol. 75 


false impression that the algal flora is far less diverse than 
it actually is. 

This paper presents the results of a survey I initiated in 
September 1970, to establish the composition, abundance, 
location and periodicity of marine algae in Virginia and 
Maryland. These types of information were particularly 
desirable for us to accrue in view of: 1) the economic im- 
portance of the area's fishing industries which ultimately 
depend upon, and are affected by, the photosynthetic enti- 
ties of the marine environment; 2) the ever-increasing pol- 
lution of the Chesapeake Bay and its estuaries; pollution 
which may affect the absolute and relative composition of 
the photosynthetic entities; and 3) the need to have at hand 
a ready store of pertinent information to investigate and 
to consider more intelligently the various ecological factors 
affecting the local marine environment. This paper repre- 
sents a small step in the realization of these goals. 

The following treatment presents a synoptic survey of 
the marine algae species reported for the states of Virginia 
and Maryland. No attempt has been made to describe mor- 
phologically or taxonomically the various species in the area 
under study. However, an attempt has been made to bring 
together, in a single work for the investigator, a summa- 
tion of our eurrent knowledge concerning the composition 
of the marine algal populations and to give their geographi- 
eal distribution. 

In the following the names for the plant divisions have 
been taken from Bold (1967). The names of families have 
been drawn, with a single minor exception, from Papen- 
fuss (1955). 

In the Division Cyanophycophyta no attempt has been 
made to relegate to synonymy the algal names as deter- 
mined by the classical, standard works with those new 
names recently proposed by Drouet and Daily (1965) and 
Drouet (1968), nor does it seem desirable to do so in view 
of the fact that the authors have not given detailed opin- 
ions for their reduction of literally thousands of taxa to 
less than 20 genera. This has resulted in several cyanophy- 


1973] Marine Algae — Ott 261 


cean algae being entered under two different names. This 
situation is, however, compatible with the basic aim of this 
paper, namely a summation of the algal taxa reported for 
the area. 

All specimens cited, without reference to a publication, 
are deposited in the herbarium of the Virginia Institute of 
Marine Science. Specimen citations taken from a publica- 
tion follow the reference. 


DIVISION CHLOROPHYCOPHYTA 


Family ZYGNEMATACEAE (Meneghini) Kützing orth. mut. 
Engler (essentially freshwater species) 

Spirogyra sp.: GENERAL: Griffith (1961), Chesapeake 
Bay, R. C. Whaley. 

Family DESMIDIACEAE Kützing ex Ralfs orth. mut. 
Stizenberger 

Closterium gracile Brébisson: GENERAL: Griffith (1961), 
Chesapeake Bay, R. C. Whaley. 

Staurastrum sp.: GENERAL: Griffith (1961), Chesapeake 
Bay, R. C. Whaley. 

Family ULOTRICHAECEAE Kützing orth. mut. Rabenhorst 

Ulothrix endosporangia Humm nom. prov.: VIRGINIA: 
H. J. Humm, York River at Gloucester Point and through- 
out the immediate general area. 

Ulothrix flacca (Dillwyn) Thuret: VIRGINIA: F. D. Ott, 
York River at Gloucester Point. Wulff (1967), York River 
at Gloucester Point. Wulff and Webb (1969), York River at 
Gloucester Point. MARYLAND: F. D. Ott, on piling at Ocean 
City. Mathieson and Fuller (1969), pilings at Chesapeake 
Biological Laboratory boathouse, mouth of Patuxent River 
at Drum Point, abandoned Cedar Point Light House. 

Ulothrix subflaccida Wille: VIRGINIA: S. Ramsey, York 
River at Gloucester Point. 

Ulothrix sp.: GENERAL: Griffith (1961), Chesapeake Bay, 
R. C. Whaley. 

Family CHAETOPHORACEAE Harvey orth. mut. 
Stizenberger 
Entocladia viridis Reinke: VIRGINIA: H. J. Humm, on 


262 Rhodora [Vol. 75 


Grinnellia americana from the northern end of Willoughby 
Spit near Hampton Roads; F. D. Ott, on Grinnellia ameri- 
cana from the York River at Gloucester Point. 

Entocladia wittrockii Wille: VIRGINIA: J. Vogel et H. J. 
Humm, on Fucus vesiculosus from salt marshes along Hum- 
mock Channel, Wachapreague. 

Protoderma marinum Reinke: VIRGINIA: Leg. H. J. 
Humm, York River in Guinea Marshes. 

Pseudodendroclonium marinum (Reinke) Aleem et 
Schulz: MARYLAND: Mathieson and Fuller (1969), aban- 
doned Cedar Point Light House, beach between Cedar 
Point and Point No Point south of Patuxent River mouth. 

Family MONOSTROMACEAE Kunieda ex Suneson 

Monostroma leptodermum Kjellman: VIRGINIA: Wulff 
(1967), York River at Gloucester Point. Wulff and Webb 
(1969), York River at Gloucester Point. 

Monostroma oxyspermum (Kiitzing) Doty: VIRGINIA: 
H. J. Humm, York River at Gloucester Point; F. D. Ott, 
Chincoteague Inlet at Chincoteague. MARYLAND: Mathieson 
and Fuller (1969), shore of Patuxent River on rocks of 
retaining wall of Naval Base Property. 

Monostroma sp.: VIRGINIA: Rhodes (1970), Burton’s Bay 
near Wachapreague. 

Family ULVACEAE Lamouroux orth. mut. Dumortier 

Enteromorpha clathrata (Roth) J. Agardh: VIRGINIA: 
H. J. Humm, York River near Gloucester Point; F. D. Ott, 
Burton's Bay near Wachapreague. Mangum, Santos and 
Rhodes (1968), York River at Sandy Point. MARYLAND: 
F. D. Ott, on Spartina stems in marsh area south of jetty. 
Mathieson and Fuller (1969), beach on south shore of 
Patuxent River west of entrance to Naval Seaplane Harbor. 

Enteromorpha compressa (Linnaeus) Greville: VIRGINIA: 
M. Lynch, York River at Gloucester Point; Leg. B. L. 
Wulff, York River at Yorktown; Leg. F. D. Ott, Chesapeake 
Bay at Cape Charles. Wulff (1967), York River at Glou- 
cester Point; Mangum, Santos and Rhodes (1968), York 
River at Sandy Point; Wulff and Webb (1969), York River 
at Gloucester Point. MARYLAND: J. K. Lowry, B. H. Robi- 


1973] Marine Algae — Ott 263 


son, and B. L. Wulff, north jetty at Ocean City; Leg. F. D. 
Ott, on shells in marsh area just south of jetty. Wulff et al 
(1968), jetty at Ocean City. 

Enteromorpha erecta (Lyngbye) J. Agardh: VIRGINIA: 
Wulff (1967), York River at Gloucester Point; Mangum, 
Santos and Rhodes (1968), York River at Sandy Point; 
Wulff and Webb (1969), York River at Gloucester Point. 

Enteromorpha intestinalis (Linnaeus) Link: VIRGINIA: 
Zaneveld and Barnes (1965), lower Chesapeake Bay; Zane- 
veld (1966-67); Wulff (1967), York River at Gloucester 
Point; Wulff and Webb (1969), York River at Gloucester 
Point; Marsh (1970), York River; Rhodes (1970), Bur- 
ton’s Bay near Wachapreague. MARYLAND: B. H. Robison, 
J. K. Lowry and B. L. Wulff, north jetty at Ocean City. 
Zaneveld (1966-67), sine locus; Wulff et al (1968), jetty 
at Ocean City; Mathieson and Fuller (1969), beach on 
south shore of Patuxent River just west of entrance to 
Naval Station seaplane harbor, Drum Point at mouth of 
Patuxent River, #5 beacon at mouth of Patuxent River, 
rock retaining wall on Naval Base property on south shore 
of Patuxent River, abandoned Cedar Point Light House, 
Cover Point just north of Patuxent River mouth, beach 
north of Cove Point, 1.5 miles east of Bloodsworth Island, 
channel] between James Island and Red Nun #2, Kent Nar- 
rows and Piney Narrows Marina, Chester River off Cedar 
Point, Patuxent River off Half Pone Point, Patuxent River 
west of Broomes Island. 

Enteromorpha lingulata J. G. Agardh: VIRGINIA: M. 
Wass, York River at Gloucester Point. 

Enteromorpha linza (Linnaeus) J. Agardh: VIRGINIA: M. 
Wass, York River at Gloucester Point; Leg. M. Wass, York 
River at Gloucester Point; Leg. H. J. Humm, York River 
at Gloucester Point. Zaneveld and Barnes (1965), lower 
Chesapeake Bay; Zaneveld (1966-67) ; Wulff (1967), York 
River at Gloucester Point; Wulff and Webb (1969), York 
River at Gloucester Point; Marsh (1970), York River; 
Rhodes (1970), Burton’s Bay near Wachapreague. MARY- 
LAND: J. K. Lowry, B. H. Robison and B. L. Wulff, north 


264 Rhodora [Vol. 75 


jetty at Ocean City; Leg. J. K. Lowry, B. L. Wulff and 
B. H. Robison, north jetty at Ocean City; Zaneveld (1966- 
67) ; Wulff et al (1968), jetty at Ocean City. 

Enteromorpha marginata J. Agardh: VIRGINIA: Zaneveld 
(1966-67). 

Enteromorpha micrococca Kützing: MARYLAND: J. K. 
Lowry, B. L. Wulff, and B. H. Robison, north jetty at Ocean 
City. Wulff et al (1968), jetty at Ocean City. 

Enteromorpha minima Nügeli: VIRGINIA: M. Wass, York 
River at Gloucester Point; Leg. H. J. Humm, York River 
at Gloucester Point. Zaneveld and Barnes (1965), lower 
Chesapeake Bay; Zaneveld (1966-67) ; Wulff (1967), York 
River at Gloucester Point; Wulff and Webb (1969), York 
River at Gloucester Point. MARYLAND: Wulff et al (1968), 
jetty at Ocean City, Mathieson and Fuller (1969), mouth 
of Patuxent River at Drum Point, shore of Patuxent River 
at Naval Base property. 

Enteromorpha plumosa | Kützing: VIRGINIA: Marsh 
(1970), York River; Rhodes (1970), Burton's Bay near 
Wachapreague. 

Enteromorpha prolifera (Müller) J. Agardh: VIRGINIA: 
M. Wass, York River at Gloucester Point; W. Dillon, York 
River at Gloucester Point; H. J. Humm, Hummock Channel 
near Wachapreague; F. D. Ott, Chincoteague Inlet at Chin- 
coteague. Zaneveld (1966-67) ; Wulff (1967), York River 
at Gloucester Point; Wulff and Webb (1969), York River 
at Gloucester Point. MARYLAND: J. K. Lowry, B. H. Robi- 
son, and B. L. Wulff, north jetty at Ocean City; F. D. Ott, 
on oyster shells in marsh area just south of jetty. Zaneveld 
(1966-67) ; Wulff et al (1968), jetty at Ocean City; Mathie- 
son and Fuller (1969), Kent Narrows and Piney Narrows 
Marina. 

Percursaria percursa (C. Agardh) J. Agardh: MARY- 
LAND: Zaneveld (1966-67). 

Ulva lactuca Linnaeus; VIRGINIA: F. D. Ott, Chincoteague 
Inlet at Chincoteague. Zaneveld and Barnes (1965), lower 
Chesapeake Bay; Wulff (1967), York River at Gloucester 
Point; Mangum, Santos and Rhodes (1968), York River 


1973] Marine Algae — Ott 265 


at Sandy Point; Wulff and Webb (1969), York River at 
Gloucester Point; Marsh (1970), York River; Rhodes 
(1970), Burton's Bay near Wachapreague. MARYLAND: 
B. L. Wulff, J. K. Lowry and B. H. Robison, north jetty at 
Ocean City; F. D. Ott, on shells in shallow water at Ocean 
City. Wulff et al (1968), jetty at Ocean City; Mathieson 
and Fuller (1969), Patuxent River west of entrance to 
seaplane harbor, mouth of Patuxent River at Drum Point, 
Patuxent River on retaining wall of Naval Base property, 
beach midway between Cedar Point and Point No Point, 
south of Patuxent River mouth, beach north of Cove Point, 
Holland Strait off South Marsh Island, mid-channel in 
Broad Creek connecting Little Annemessex River and Po- 
comoke Sound, 1.5 miles east of Bloodsworth Island, 1.25 
miles southwest of Red Nun £2, between Poplar Island 
Narrows Light and Can £1, 900 yards east of Coaches Is- 
land, mouth of Crab Alley Bay off Parson and Bodkin 
Island, channel between James Island and Red Nun #2, 
east and west of Knapps Narrows, Kent Narrows and 
Piney Narrows Marina. 

Ulva lactuca Linnaeus var. latissima (Linnaeus) de Can- 
dolle: VIRGINIA: K. M. S. Aziz, York River at Gloucester 
Point; F. D. Ott, Chincoteague Inlet at Chincoteague. Zane- 
veld (1966-67). MARYLAND: F. D. Ott, adrift at Assateague 
Beach. Zaneveld (1966-67). 

Ulva lactuca Linnaeus var. rigida (C. Agardh) Le Jolis: 
VIRGINIA: F. D. Ott, Chesapeake Bay at Cape Charles. Zane- 
veld (1966-67). MARYLAND: F. D. Ott, on rock at jetty in 
Ocean City. 

Family CHLORELLACEAE (Wille) Brunnthaler 

Chlorella sp.: MARYLAND: Morse (1947), mouth of Pa- 
tuxent River. GENERAL: Griffith (1961), Chesapeake Bay, 
D. C. Morse. 

Family GOMONTIACEAE Bornet et Flahault ex De Toni 

Gomontia polyrhiza (Lagerheim) Bornet et Flahault: 
VIRGINIA: H. J. Humm, York River at Gloucester Point; 
K. M. S. Aziz, York River at Yorktown; F. D. Ott, Chesa- 
peake Bay at Cape Charles. MARYLAND: B. H. Robison, 


266 Rhodora [Vol. 75 


J. K. Lowry and B. L. Wulff, north jetty at Ocean City; 
F. D. Ott, Ocean City growing on Spartina stems. 
Family HYDRODICTYACEAE (S. F. Gray) Dumortier orth. 
mut Cohn (essentially freshwater species) 

Pediastrum boryanum (Turpin) Meneghini: GENERAL: 
Griffith (1961), Chesapeake Bay, R. C. Whaley. 

Pediastrum duplex Meyen: GENERAL: Griffith (1961), 
Chesapeake Bay, R. C. Whaley. 

Pediastrum simplex Meyen: GENERAL: Griffith. (1961), 
Chesapeake Bay, R. C. Whaley. 

Family SCENEDESMACEAE Oltmanns 
(essentially freshwater species) 

Actinastrum sp.: GENERAL: Griffith (1961), Chesapeake 
Bay, R. C. Whaley. 

Scenedesmus acuminatus (Lagerheim) Chodat: GEN- 
ERAL: Griffith. (1961), Chesapeake Bay, R. C. Whaley. 

Scenedesmus opaliensis P. Richter: GENERAL: Griffith 
(1961), Chesapeake Bay, R. C. Whaley. 

Scenedesmus quadricauda (Turpin) in de Brébisson et 
Godey: VIRGINIA: Marshall (1967), Elizabeth River. 

Scenedesmus sp.: VIRGINIA: Marshall (1967), Elizabeth 
River. 

Family CLADOPHORACEAE (Hassell) Cohn 

Chaetomorpha aerea (Dillwyn) Ktitzing: MARYLAND: 
Zaneveld (1966-67) ; Wulff et al (1968), jetty at Ocean 
City. 

Chaetomorpha linum (Müller) Kützing: VIRGINIA: H. J. 
Humm, York River in the Guinea Marshes; B. L. Wulff, 
York River off Wormley Creek. Marsh (1970), York River. 

Cladophora albida (Hudson) Kützing: VIRGINIA: F. D. 
Ott, Chesapeake Bay at Cape Charles. Zaneveld (1966-67). 

Cladophora crystallina (Roth) Kützing: VIRGINIA: Zane- 
veld (1966-67). 

Cladophora expansa (Mertens) Kützing: VIRGINIA: Zane- 
veld (1966-67). 

Cladophora fascicularis (Mertens) Kiitzing: VIRGINIA: 
H. J. Humm, Hampton Roads at Norfolk; M. Wass, York 
River at Gloucester Point. 


1973] Marine Algae — Ott 267 


Cladophora flexuosa (Dillwyn) Harvey: VIRGINIA: F. D. 
Ott, Chincoteague Inlet at Chincoteague. Zaneveld and 
Barnes (1965), lower Chesapeake Bay. MARYLAND: Mathie- 
son and Fuller (1969), Patuxent River at Broomes Island. 

Cladophora flexuosa (Dillwyn) Harvey forma densa 
Collins: VIRGINIA: Zaneveld (1966-67). MARYLAND: Math- 
ieson and Fuller (1969), Abandon Cedar Point Light 
House. 

Cladophora gracilis (Griffith ex Harvey) Kützing: VIR- 
GINIA: Zaneveld and Barnes (1965), lower Chesapeake 
Bay; Marsh (1970), York River. MARYLAND: F. D. Ott, 
in quiet water in marshes south of Ocean City. Mathieson 
and Fuller (1969), Cove Point just north of Patuxent 
River mouth, beach north of Cove Point, mid-channel in 
Broad Creek connecting Little Annemessex River and Po- 
comoke Sound. 

Cladophora magdalenae Harvey: VIRGINIA: H. J. Humm, 
York River in the Guinea Marshes. 

Cladophora rupestris (Linnaeus) Kützing: VIRGINIA: 
M. Wass, York River at Gloucester Point; F. D. Ott, Bur- 
ton’s Bay near Wachapreague. 

Rhizoclonium | kockianum Kiitzing: VIRGINIA: H. J. 
Humm, York River at Gloucester Point. 

Rhizoclonium riparium (Roth) Harvey: VIRGINIA: H. J. 
Humm, York River at Gloucester Point. Zaneveld (1966- 
67). MARYLAND: Mathieson and Fuller (1969), pilings at 
Chesapeake Biological Laboratory boathouse. 

Rhizoclonium tortuosum Kützing: VIRGINIA: H. J. Humm, 
Hummock Channel at Wachapreague. 

Family BRYOPSIDACEAE Bory de Saint-Vincent orth. mut. 
De Toni 

Bryopsis hypnoides Lamouroux: VIRGINIA: F. D. Ott, 
James River at north end of James River Bridge. Zaneveld 
and Barnes (1965), lower Chesapeake Bay; Zaneveld 
(1966-67); Mangum, Santos and Rhodes (1968), York 
River at Sandy Point. MARYLAND: F. D. Ott, on pilings at 
Ocean City. Zaneveld (1966-67). 

Bryopsis plumosa (Hudson) C. Agardh: VIRGINIA: W. 


268 Rhodora [Vol. 75 


Dillon, York River at Gloucester Point; P. E. Hargraves, 
York River at Gloucester Point; F. D. Ott, Hampton Roads 
at Fort Monroe. Zaneveld and Barnes (1965), lower Ches- 
apeake Bay; Wulff (1967), York River at Gloucester Point; 
Zaneveld (1966-67) ; Wulff and Webb (1969), York River 
at Gloucester Point; Marsh (1970), York River; Rhodes 
(1970), Burton's Bay near Wachapreague. MARYLAND: 
F. D. Ott, on pilings at Ocean City. Zaneveld (1966-67). 


DIVISION PHAEOPHYCOPHYTA 


Family ECTOCARPACEAE (C. Agardh) Kützing orth. mut. 
Harvey 


Ectocarpus confervoides (Roth) Le Jolis: VIRGINIA: M. 
Wass, Sarah's Creek below Gloucester Point; F. D. Ott, 
York River at Sandy Point. Zaneveld (1966-67); Wulff 
and Webb (1969), York River at Gloucester Point. MARY- 
LAND: F. D. Ott, Ocean City, in marsh area just south of 
jetty. Zaneveld (1966-67). 

Ectocarpus elachistaeformis Heydrich: vIRGINIA: H. J. 
Humm, York River at Gloucester Point. 

Ectocarpus penicillatus (C. Agardh) Kjellman: VIRGINIA : 
Mangum, Santos and Rhodes (1968), York River at Sandy 
Point. 

Ectocarpus siliculosus (Dillwyn) Lyngbye: VIRGINIA: 
M. Wass, York River at Sandy Point; M. Wass, York River 
at Gloucester Point; F. D. Ott, Hampton Roads at Lynn- 
haven Inlet. Rhodes (1970), Burton's Bay near Wacha- 
preague. MARYLAND: F. D. Ott, Ocean City, in marsh area 
just south of jetty. 

Giffordia duchassaigniana (Grunow) Taylor: MARYLAND: 
B. H. Robison, J. K. Lowry and B. L. Wulff, north jetty at 
Ocean City. Wulff et al (1968), jetty at Ocean City. 

Pylaiella littoralis (Linnaeus) Kjellman: VIRGINIA: Zane- 
veld (1966-67) ; Mangum, Santos and Rhodes (1968), York 
River at Sandy Point. MARYLAND: Zaneveld (1966-67). 

Family SPHACELARIACEAE J. Agardh orth. mut. Cohn 

Sphacelaria fusca (Hudson) C. Agardh: VIRGINIA: F. D. 


1973] Marine Algae — Ott 269 


Ott, York River at Gloucester Point. MARYLAND: Mathie- 
son and Fuller (1969), Patuxent River 200 yards west of 
Broomes Island. 
Family DICTYOTACEAE Lamouroux orth. mut. Dumortier 
Dictyota dichotoma (Hudson) Lamouroux: VIRGINIA: 
H. J. Humm, Hummock Channel near Wachapreague. 
Humm (1963), Hummock Channel near Wachapreague; 
Rhodes (1970), Burton’s Bay near Wachapreague. 
Family MYRIONEMATACEAE (Nägeli) Foslie orth. mut. 
Skottsberg 
Ascocyclus orbicularis Magnus: VIRGINIA: H. J. Humm, 
York River at Gloucester Point. Mangum, Santos and 
Rhodes (1968), York River at Sandy Point. 
Myrionema stragulans Greville: VIRGINIA: Mangum, San- 
tos and Rhodes (1968), York River at Sandy Point. 
Family ELACHISTACEAE Kjellman 
Elachistea facicola (Velley) Areschoug: VIRGINIA: J. L. 
Wood, adrift and epiphytic on Fucus vesiculosus, collected 
ca. 60 miles offshore from the mouth of the Chesapeake 
Bay. 
Elachistea sp.: VIRGINIA: Marsh (1970), York River. 
Family CORYNOPHLAEACEAE Oltmanns 
Leathesia difformis (Linnaeus) Areschoug: VIRGINIA: 
F. D. Ott, Chincoteague Inlet at Chincoteague. Zaneveld 
(1966-67); Rhodes (1970), Burton’s Bay near Wacha- 
preague. 
Family SPERMATOCHNACEAE Kjellman 
Stilophora rhizodes (Ehrhart) J. Agardh: VIRGINIA: 
B. H. Wulff, York River at Gloucester Point. 
Family STRIARIACEAE Kjellman 
Striaria attenuata (C. Agardh) Greville: VIRGINIA: 
F. D. Ott, Chincoteague Inlet at Chincoteague. Rhodes 
(1970), Burton's Bay near Wachapreague. MARYLAND: 
F. D. Ott, Ocean City, in marsh area just south of jetty. 
Family MYRIOTRICHIACEAE Kjellman 
Myriotrichia subcorymbosa (Farlow emend. Holden) 
Blomquist: VIRGINIA: H. J. Humm, York River at Glouces- 
ter Point; H. J. Humm, York River at Gloucester Point. 


270 Rhodora [Vol. 75 


Family PUNCTARIACEAE (Thuret) Kjellman 

Desmotrichum undulatum (J. Agardh) Reinke: VIR- 
GINIA: F. D. Ott, on Zostera at Cape Charles. Mangum, 
Santos and Rhodes (1968), York River at Sandy Point; 
Marsh (1970), York River; Rhodes (1970), Burton’s Bay 
near Wachapreague. MARYLAND: F. D. Ott, on bent stems 
of Spartina in marsh area just south of jetty. 

Petalonia fascia (O. F. Müller) Kuntze: VIRGINIA: F. D. 
Ott, on rocks at Chincoteague Inlet at Chincoteague. Zane- 
veld and Barnes (1965), lower Chesapeake Bay; Zaneveld 
(1966-67). MARYLAND: J. K. Lowry and B. L. Wulff, north 
jetty at Ocean City; F. D. Ott, Ocean City, on stones of 
jetty. Zaneveld (1966-67); Wulff et al (1968), jetty at 
Ocean City. 

Punctaria latifolia Greville: VIRGINIA: F. D. Ott, James 
River just north of James River Bridge. Rhodes (1970), 
Burton’s Bay near Wachapreague. MARYLAND: F. D. Ott, 
Ocean City, in marsh area just south of jetty. 

Punctaria plantaginea (Roth) Greville: VIRGINIA: Zane- 
veld and Barnes (1965), lower Chesapeake Bay. MARY- 
LAND: Zaneveld (1966-67). 

Family SCYTOSIPHONACEAE (Thuret) Hauck 

Asperococcus echinatus (Mertens) Greville: VIRGINIA: 
F. D. Ott, York River at Yorktown Beach. MARYLAND: 
F. D. Ott, Ocean City, in marsh area just south of jetty. 

Scytosiphon lomenataria (Lyngbye) C. Agardh: VIR- 
GINIA: F. D. Ott, Burton’s Bay near Wachapreague. Zane- 
veld and Barnes (1965), lower Chesapeake Bay; Zaneveld 
(1966-67); Marsh (1970), York River; Rhodes (1970), 
Burton’s Bay near Wachapreague. MARYLAND: B. H. Robi- 
son, J. K. Lowry and B. L. Wulff, north jetty at Ocean 
City; F. D. Ott, Ocean City on stones of jetty. Zaneveld 
(1966-67) ; Wulff et al (1968), jetty at Ocean City. 

Family DICTYOSIPHONACEAE Kützing orth. mut. Kjellman 

Dictyosiphon foeniculaceus (Hudson) Greville: VIRGINIA: 
Rhodes (1970), Burton’s Bay near Wachapreague. 

Family FUCACEAE Lamouroux orth. mut. Dumortier 

Fucus vesiculosus Linnaeus: VIRGINIA: J. Vogel, Haven 


1973] Marine Algae — Ott 271 


Beach, just south of Milford Haven facing Chesapeake 
Bay; H. J. Humm, adrift in Little Creek, Norfolk; K. de 
Witt, washed ashore at Virginia Beach; H. J. Humm, Hum- 
mock Channel near Wachapreague; F. D. Ott, on rocks at 
Chincoteague Inlet at Chincoteague. Zaneveld (1966-67). 
Rhodes (1970), Burton’s Bay near Wachapreague. MARY- 
LAND: F. D. Ott, Ocean City on rocks and at edge of salt 
marshes. Zaneveld (1966-67) ; Wulff et al (1968), jetty at 
Ocean City. 

Fucus vesiculosus Linnaeus var. sphaerocarpus J. 
Agardh: MARYLAND: Zaneveld (1966-67). 

Ascophyllum nodosum (Linnaeus) Le Jolis: VIRGINIA: 
F. D. Ott, adrift in Chincoteague Inlet at Chincoteague. 
Zaneveld (1966-67), sine locus. MARYLAND: F. D. Ott, 
Ocean City, abundant in the wash in October 1971. Zane- 
veld (1966-67). 

Ascophyllum nodosum (Linnaeus) Le Jolis forma scor- 
pioides (Hornemann) Reinke: MARYLAND: J. K. Lowry and 
B. H. Robison, found attached to the north jetty at Ocean 
City. 

Family SARGASSACEAE (Decaisne) Kiitzing orth. mut. 

De Toni 

Sargassum natans (Linnaeus) J. Meyen: VIRGINIA: F. D. 
Ott, Burton’s Bay near Wachapreague. Zaneveld (1966- 
67). MARYLAND: F. D. Ott, Ocean City, abundant in the 
wash during November-December 1971. Zaneveld (1966- 
on 

Sargassum hystrix J. Agardh var. buxifolium (Chauvin) 
J. Agardh: VIRGINIA: F. D. Ott, adrift at Hampton Roads 
at Lynnhaven Inlet. Zaneveld (1966-67). MARYLAND: 
F. D. Ott, Assateague Beach, abundant in the wash during 
November-December 1971. 


DIVISION RHODOPHYCOPHYTA 


Family GONIOTRICHACEAE Skuja 
Goniotrichum alsidii (Zanardini) Howe: VIRGINIA: 
Rhodes (1970), Burton’s Bay near Wachapreague. MARY- 
LAND: Wulff et al (1968), jetty at Ocean City. 


272 Rhodora [Vol. 75 


Family ERYTHROPELTIDACEAE Skuja 
Erythrocladia subintegra Rosenvinge: VIRGINIA: H. J. 
Humm, York River at Gloucester Point. 
Erythrotrichia rhizoides Cleland: VIRGINIA: Rhodes 
(1970), Burton’s Bay near Wachapreague. 
Family BANGIACEAE (S. F. Gray) Nägeli 


Bangia ciliaris Carmichael: MARYLAND: B. L. Wulff. 
J. K. Lowry and B. H. Robison, north jetty at Ocean City; 
mixed in with herbarium sample of Acrochaetium trifilum. 

Bangia fuscopurpurea (Dillwyn) Lyngbye: VIRGINIA: 
F. D. Ott, Virginia Beach on stones of jetty south of resort 
area. Zaneveld and Barnes (1965), lower Chesapeake Bay ; 
Zaneveld (1966-67) ; Wulff (1967), York River at Glouces- 
ter Point; Wulff and Webb (1969), York River at Glouces- 
ter Point. MARYLAND: B. L. Wulff, B. H. Robison and 
J. K. Lowry, north jetty at Ocean City; F. D. Ott, Ocean 
City on rocks in marsh area just south of jetty. Zaneveld 
(1966-67) ; Wulff et al (1968), jetty at Ocean City. 

Porphyra leucosticta 'Thuret: VIRGINIA: K. de Witt, Lynn- 
haven Inlet; P. E. Hargraves, York River at Gloucester 
Point. Zaneveld and Barnes (1965), lower Chesapeake 
Bay; Marsh (1970), York River; Rhodes (1970), Burton's 
Bay near Wachapreague. MARYLAND: B. L. Wulff, B. H. 
Robison and J. K. Lowry, north jetty at Ocean City; F. D. 
Ott, Ocean City on stones of jetty. Wulff et al (1968), 
jetty at Ocean City. 

Porphyra miniata (Lyngbye) C. Agardh: VIRGINIA: H. J. 
Humm, Hummock Channel near Wachapreague. Rhodes 
(1970), Burton’s Bay near Wachapreague. 

Porphyra umbilicalis (Linnaeus) J. Agardh: VIRGINIA: 
F. D. Ott, Chincoteague Inlet at Chincoteague. Zaneveld 
and Barnes (1965), lower Chesapeake Bay; Zaneveld 
(1966-67) ; Wulff (1967), York River at Gloucester Point; 
Wulff and Webb (1969), York River at Gloucester Point. 
MARYLAND: Zaneveld (1966-67) ; Wulff et al (1968), jetty 
at Ocean City. 


1973] Marine Algae — Ott 219 


Family ACROCHAETIACEAE Fritsch 

Acrochaetium aleyonidii Jao: VIRGINIA: K. M. S. Aziz, 
York River at Gloucester Point. 

Acrochaetium dasyae Collins: VIRGINIA: D. Dilliman, 
James River above Gum Rock. 

Acrochaetium flexuosum Vickers: MARYLAND: Mathieson 
and Fuller (1969), Patuxent River west of Broomes Island. 

Acrochaetium radiatum Jao: MARYLAND: B. L. Wulff, 
J. K. Lowry and B. H. Robison, jetty at Ocean City. 

Acrochaetium trifilum (Buffham) Batters emend. Aziz: 
MARYLAND: B. L. Wulff, J. K. Lowry and B. H. Robison, 
north jetty at Ocean City; B. L. Wulff, J. K. Lowry and 
B. H. Robison, north jetty at Ocean City; B. H. Robison, 
J. K. Lowry and B. L. Wulff, north jetty at Ocean City. 
Wulff et al (1968), jetty at Ocean City. 

Acrochaetium virgatulum (Harvey) J. Agardh: MARY- 
LAND: Wulff et al (1968), jetty at Ocean City. 

Acrochaetium virgatulum (Harvey) J. Agardh forma 
luxurians (J. Agardh) Collins: VIRGINIA: M. Wass, York 
River at Gloucester Point. 

Acrochaetium spp.: MARYLAND: Wulff et al (1968). jetty 
at Ocean City. 

Family HELMINTHOCLADIACEAE (J. Agardh) Harvey 

orth. mut. Hauck 

Nemalion multifidum (Weber et Mohr) J. Agardh: 
MARYLAND: J. K. Lowry and B. H. Robison, north jetty at 
Ocean City; E. M. and B. H. Wulff, north jetty at Ocean 
City. Wulff et al (1968), jetty at Ocean City. 

Family BONNEMAISONIACEAE Schmitz in Engler 

Asparagopsis hamifera (Hariot) Okamura: It has been 
established that the two independently described algae, 
Asparagopsis hamifera and Trailliella intricata, are re- 
spectively the gametophytic and tetrasporophytic plants 
of an alga having a heteromorphic alternation of genera- 
tions. While the gametophytic Asparagopsis hamifera has 
not been recorded for the geographical area under study, 
the tetrasporophytic Trailliella intricata has been found. 
This plant has been entered in this compilation under the 


274 Rhodora [Vol. 75 


Ceramiaceae where it was assigned prior to the establish- 
ment of its relationship with Asparagopsis hamifera. 
Family GELIDIACEAE Kützing orth. mut. Harvey 

Gelidium crinale (Turner) Lamouroux: VIRGINIA: M. 
Lynch, jetty at Little Creek in Norfolk; H. J. Humm, York 
River at Yorktown; F. D. Ott, Chesapeake Bay at Cape 
Charles on stones of jetty. Zaneveld and Barnes (1965), 
lower Chesapeake Bay; Rhodes (1970), Burton's Bay near 
Wachapreague. MARYLAND: F. D. Ott, Ocean City on rocks 
of jetty. 

Family CORALLINACEAE (Lamouroux) Harvey 

Fosliella farniosa (Lamouroux) Howe: VIRGINIA: H. J. 
Humm, York River at Gloucester Point. 

Fosliella lejolisii (Rosanoff) Howe: VIRGINIA: Marsh 
(1970), York River. 

Family SOLIERIACEAE (Harvey) Hauck 

Agardhiella tenera (J. Agardh) Schmitz: VIRGINIA: 
H. J. Humm, York River at Gloucester Point; K. M. S. 
Aziz, York River at Gloucester Point; P. E. Hargraves, 
York River at Gloucester Point; F. D. Ott, Hampton Roads 
at Lynnhaven. Zaneveld and Barnes (1965), lower Chesa- 
peake Bay; Zaneveld (1966-67); Mangum, Santos and 
Rhodes (1968), York River at Sandy Point; Mathieson and 
Fuller (1969), Gwynn Island 300 yards inshore of #1 
beacon; Marsh (1970), York River; Rhodes (1970), Bur- 
ton’s Bay near Wachapreague. MARYLAND: F. D. Ott, As- 
sateague Beach, abundant in wash. Zaneveld (1966-67). 

Family HYPNEACEAE J. Agardh 

Hypnea musciformis (Wulfen) Lamouroux: VIRGINIA: 
H. J. Humm, York River at Yorktown; F. D. Ott, Chesa- 
peake Bay at Cape Charles on stones of jetty. Zaneveld 
(1966-67). Rhodes (1970), Burton’s Bay near Wacha- 
preague. MARYLAND: F. D. Ott, Ocean City in marsh area 
just south of jetty. Zaneveld (1966-67). 

Family GRACILARIACEAE (Nägeli) Kylin 

Gracilaria foliifera (Forsskål) Borgesen: VIRGINIA: F. D. 
Ott, Burton's Bay near Wachapreague. Zaneveld and 
Barnes (1965), lower Chesapeake Bay; Zaneveld (1966- 


1973] Marine Algae — Ott 275 


67); Mangum, Santos and Rhodes (1968), York River at 
Sandy Point; Rhodes (1970), Burton’s Bay near Wacha- 
preague. MARYLAND: B. H. Robison and J. K. Lowry, 
jetty at Ocean City; F. D. Ott, Ocean City on shells in 
marsh area just south of jetty. Zaneveld (1966-67) ; Wulff 
et al (1968), jetty at Ocean City. 

Gracilaria verrucosa (Hudson) Papenfuss: VIRGINIA: 
F. D. Ott, Chincoteague Inlet at Chincoteague. Zaneveld 
and Barnes (1965), lower Chesapeake Bay; Zaneveld 
(1966-67) ; Mathieson and Fuller (1969), Mobjack Bay off 
Guinea Marsh, Gwynn Island 300 yards inshore of #1 
beacon; Rhodes (1970), Burton’s Bay near Wachapreague. 
MARYLAND: F. D. Ott, on stones in marsh area just south of 
jetty. Zaneveld (1966-67); Mathieson and Fuller (1969) 
beach midway between Cedar Point and Point No Point 
south of Patuxent River mouth, beach north of Cove Point, 
Holland Strait off South Marsh Island near #2 beacon, 
midway between #9 beacon and James Island Light off 
Crisfield, Great Rock Oyster Bar off James Island Light, 
Little Annemessex River at mouth of Battle Creek, Poco- 
moke Sound off Broad Creek, Shark Fin Shoal east of 
Bloodsworth Island, mouth of Patuxent River near Ob- 
struction Buoy west of Drum Point, between Poplar Island 
Narrows Light and Can #1, Crab Alley Bay between Can 
#1 and Bodkin Island, channel between James Island and 
Red Nun #2, east and west of Knapps Narrows, Kent 
Narrows and pilings at Piney Narrows Marina. 

Family GIGARTINACEAE Bory de Saint-Vincent orth. mut. 

Cohn 

Chondrus crispus Stackhouse: VIRGINIA: Zaneveld (1966- 
67). 

Family CHAMPIACEAE Kiitzing orth. mut. Bliding 

Champia parvula (C. Agardh) Harvey: VIRGINIA: K. M. 
S. Aziz, York River at Gloucester Point; M. Wass, York 
River at Gloucester Point; F. D. Ott, Chesapeake Bay at 
Cape Charles. Zaneveld and Barnes (1965), lower Chesa- 
peake Bay; Marsh (1970), York River; Rhodes (1970), 
Burton’s Bay near Wachapreague. MARYLAND: F. D. Ott, 


276 Rhodora [Vol. 75 


Ocean City on shells in marsh area just south of jetty. 
Mathieson and Fuller (1969), Great Rock Oyster Bar 
south of James Island Light, Little Annemessex River and 
Pocomoke Sound, Pocomoke Sound off Broad Creek, Shark 
Fin Shoal east of Bloodsworth Island. 

Lomentaria baileyana (Harvey) Farlow: VIRGINIA: F. D. 
Ott, Burton's Bay near Wachapreague. Rhodes (1970), 
Burton's Bay near Wachapreague. 

Family CERAMIACEAE (S. F. Gray) Harvey orth. mut. 

Rabenhorst 

Antithamnion cruciatum (C. Agardh) Nägeli: VIRGINIA: 
Rhodes (1970), Burton's Bay near Wachapreague. 

Callithamnion baileyi Arnott: VIRGINIA: B. L. Wulff, 
York River at Gloucester Point. Zaneveld (1966-67). 
MARYLAND: B. H. Robison, J. K. Lowry and B. L. Wulff, 
north jetty at Ocean City. Wulff et al (1968), jetty at 
Ocean City. 

Callithamnion byssoides Arnott: VIRGINIA: M. Lynch, 
jetty in Little Creek in Norfolk; H. J. Humm, Hummock 
Channel near Wachapreague; F. D. Ott, Burton’s Bay near 
Wachapreague. Zaneveld and Barnes (1965), lower Ches- 
apeake Bay; Wulff (1967), York River at Gloucester Point; 
Wulff and Webb (1969), York River at Gloucester Point; 
Marsh (1970), York River; Rhodes (1970), Burton’s Bay 
near Wachapreague. MARYLAND: F. D. Ott, Assateague 
Beach adrift in wash. 

Callithamnion corymbosum (Smith) C. Agardh: VIR- 
GINIA: F. D. Ott, Chincoteague Inlet at Chincoteague. 
Rhodes (1970), Burton’s Bay near Wachapreague. 

Ceramium diaphanum (Lightfoot) Roth: VIRGINIA: 
Zaneveld and Barnes (1965), lower Chesapeake Bay; Zane- 
veld (1966-67); Marsh (1970), York River. MARYLAND: 
Zaneveld (1966-67), sine locus. 

Ceramium fastigiatum (Roth) Harvey: VIRGINIA: M. 
Wass, mouth of Chesapeake Bay; F. D. Ott, Chesapeake 
Bay at Cape Charles, on stones of jetty. Zaneveld and 
Barnes (1965), lower Chesapeake Bay; Zaneveld (1966- 
67); Marsh (1970), York River; Rhodes (1970), Burton's 


1973] Marine Algae — Ott 277 


Bay near Wachapreague. MARYLAND: B. H. Robison, J. K. 
Lowry and B. L. Wulff, north jetty at Ocean City; F. D. 
Ott, Assateague Beach abundant in wash. Wulff et al 
(1968), jetty at Ocean City. 

Ceramium rubriforme Kylin: VIRGINIA: Marsh (1970), 
York River; Rhodes (1970), Burton’s Bay near Wacha- 
preague. 

Ceramium rubrum (Hudson) C. Agardh: VIRGINIA: B. L. 
Wulff, York River at Yorktown; P. E. Hargraves, adrift 
in the York River at Gloucester Point; F. D. Ott, Chesa- 
peake Bay at Cape Charles. Woodson (1959), mouth of 
James River; Zaneveld and Barnes (1965), lower Chesa- 
peake Bay; Zaneveld (1966-67) ; Wulff (1967), York River 
at Gloucester Point; Mangum, Santos and Rhodes (1968), 
York River at Sandy Point; Mathieson and Fuller (1969); 
Mobjack Bay off Guinea Marsh; Wulff and Webb (1969), 
York River at Gloucester Point; Marsh (1970), York 
River; Rhodes (1970), Burton's Bay near Wachapreague. 
MARYLAND: J. K. Lowry, B. L. Wulff and B. H. Robison, 
north jetty at Ocean City; F. D. Ott, Assateague Beach, 
abundant in the wash. Zaneveld (1966-67); Wulff et al 
(1968), jetty at Ocean City; Mathieson and Fuller (1969), 
beach on south shore of Patuxent River just west of en- 
trance to Naval Station seaplane harbor, rock retaining 
wall on Naval Base property south shore of Patuxent River, 
beach north of Cove Point, Holland Strait off South Marsh 
Island, midway between £9 beacon and James Island Light 
off Crisfield, Great Oyster Bar south of James Island Light, 
Little Annemessex River at mouth of Battle Creek, mid- 
channel in Broad Creek connecting Little Annemessex 
River and Pocomoke Sound, Shark Fin Shoal east of 
Bloodsworth Island, 1.5 miles east of Bloodsworth Island, 
mouth of Patuxent River near Obstruction Buoy west of 
Drum Point, between Poplar Island Narrows Light and 
Can £1, Crab Alley Bay between Can £1 and Bodkin 
Island, channel between James Island and Red Nun #2, 
Kent Narows and pilings at Piney Narrows Marina, Ches- 
ter River off Cedar Point, Patuxent River 200 yards west 
of Broomes Island. 


278 Rhodora [Vol. 75 


Ceramium strictum (Kiitzing) Harvey: VIRGINIA: M. 
Wass, York River at Gloucester Point; P. E. Hargraves, 
adrift in the York River at Gloucester Point; J. Vogel, 
eastern side of Gwynn Island near mouth of the Rappa- 
hannock River. Zaneveld and Barnes (1965), lower Chesa- 
peake Bay; Zaneveld (1966-67) ; Wulff (1967), York River 
at Gloucester Point; Mangum, Santos and Rhodes (1968), 
York River at Sandy Point; Mathieson and Fuller (1969), 
Mobjack Bay off Guinea Marsh; Wulff and Webb (1969), 
York River at Gloucester Point; Marsh (1970), York 
River. MARYLAND: B. L. Wulff, J. K. Lowry and B. H. 
Robison, north jetty at Ocean City. Zaneveld (1966-67) ; 
Wulff et al (1968), jetty at Ocean City; Mathieson and 
Fuller (1969), beach on shore of Patuxent River west of 
entrance to Naval Station seaplane harbor, rock retaining 
wall on Naval Base property south shore of Patuxent River, 
abandoned Cedar Point Light House, beach north of Cove 
Point, Holland Strait off South Marsh Island, midway be- 
tween #9 beacon and James Island Light off Crisfield, 
Great Rock Oyster Bar south of James Island Light, Little 
Anneniessex River at mouth of Battle Creek, mid-channel 
in Broad Creek connecting Little Annemessex River and 
Pocomoke Sound, Pocomoke Sound off Broad Creek. Shark 
Fin Shoal east of Bloodsworth Island, 1.5 miles east of 
Bloodsworth Island, mouth of Patuxent River near Ob- 
struetion Buoy west of Drum Point, mouth of Crab Alley 
Bay off Parson Island and Bodkin Island, channel between 
James Island and Red Nun #2, off Half Pone Point, 200 
yards west of Broomes Island. 

Griffithsia tenuis C. Agardh: VIRGINIA: H. J. Humm, 
York River at Gloucester Point. 

Spyridia filamentosa (Wulfen) Harvey: VIRGINIA: F. D. 
Ott, Chesapeake Bay at Cape Charles. Mangum, Santos 
and Rhodes (1968), York River at Sandy Point; Mathieson 
and Fuller (1969), Mobjack Bay off Guinea Marsh; Poco- 
moke Sound near Red Nun #6; Marsh (1970), York River; 
Rhodes (1970), Burton's Bay near Wachapreague. MARY- 
LAND: F. D. Ott, Ocean City on shells in marsh area just 


1973] Marine Algae — Ott 279 


south of jetty. Mathieson and Fuller (1969), Holland 
Strait off South Marsh Island, midway between #9 beacon 
and James Island Light off Crisfield, Great Rock Oyster 
Bar south of James Island Light, Little Annemessex River 
at mouth of Battle Creek, mid-channel in Broad Creek 
connecting Little Annemessex River and Pocomoke Sound, 
Pocomoke Sound off Broad Creek, between Red Nun #4 
and Hooper Strait Light, mouth of Patuxent River near 
Obstruction Buoy west of Drum Point. 

Trailliella intricata (J. Agardh) Batters: MARYLAND: 
Mathieson and Fuller (1969), Holland Strait off South 
Marsh Island, between #9 beacon and James Island Light 
off Crisfield, Great Rock Oyster Bar south of James Island 
Light, mid-channel in Broad Creek connecting Little An- 
nemessex River and Pocomoke Sound, Pocomoke Sound 
off Broad Creek, between Red Nun #4 and Hooper Strait 
Light. 

Family DASYACEAE Kiitzing orth. mut. Rosenberg 

Dasya pedicellata (C. Agardh) C. Agardh: VIRGINIA: 
K. M. S. Aziz and J. Vogel, Haven Beach, Mathews County; 
F. D. Ott, Burton’s Bay near Wachapreague. Zaneveld and 
Barnes (1965), lower Chesapeake Bay; Wulff (1967), 
York River at Gloucester Point; Wulff and Webb (1969), 
York River at Gloucester Point; Marsh (1970), York 
River; Rhodes (1970), Burton's Bay near Wachapreague. 
MARYLAND: Mathieson and Fuller (1969), beach north of 
Cove Point, 1300 yards east of Red Nun £6 off Sharps 
Island Light, between Poplar Island Narrows Light and 
Can #1, 900 yards east of Coaches Island, mouth of Crab 
Alley Bay off Parson Island and Bodkin Island, east of 
Knapps Narrows, off Half Pone Point. 

Family DELESSERICEAE Bory de Saint-Vincent orth. mut. 

Nageli 

Caloglossa leprieurii (Montagne) J. Agardh: VIRGINIA: 
F. D. Ott, York River at Sandy Point on Spartina stems. 
Post (1968), James River on shore of Mulberry Island 
above Nells Creek, R. Patrick; James River at Cobham's 
Wharf, J. C. Strickland. MARYLAND: Post (1968), at east- 


280 Rhodora [Vol. 75 


ern shore of mouth of St. Leonard Creek near Solomon's 
Island, G. Papenfuss. 

Grinnellia americana (C. Agardh) Harvey: VIRGINIA: 
H. J. Humm, adrift at Norfolk end of bridge in Hampton 
Roads; H. J. Humm, York River at Gloucester Point; J. 
Vogel and K. M. S. Aziz, Haven Beach in Mathews County ; 
F. D. Ott, Burton’s Bay near Wachapreague. Zaneveld and 
Barnes (1965), lower Chesapeake Bay; Marsh (1970), 
York River; Rhodes (1970), Burton’s Bay near Wacha- 
preague. MARYLAND: F. D. Ott, Ocean City in marsh area 
just south of jetty. 

Family RHODOMELACEAE (J. Agardh) Harvey 

Bostrychia radicans Montagne: VIRGINIA: F. D. Ott, 
James River at College Creek on old wooden retaining wall. 
Post (1968), James River on shore of Mulberry Island 
above Nells Creek, R. Patrick (as Bostrychia rivularis) ; 
on the sea wall ca. 14, mile above the York River Bridge at 
Yorktown, H. J. Humm; James River at Cobham’s Wharf, 
J. C. Strickland; Old Plantation Creek ca. two miles south 
of Cape Charles, H. G. Richards. MARYLAND: Post (1968), 
at eastern shore of mouth of St. Leonard Creek, near Solo- 
mon’s Island, G. Papenfuss (as Bostrychia radicans forma 
moniliforme) ; at Patuxent River opposite Solomon’s Is- 
land Bridge, G. Papenfuss. 

Bostrychia rivularis Harvey: VIRGINIA: H. J. Humm, 
York River in the Guinea Marshes. Rhodes (1970), Bur- 
ton’s Bay near Wachapreague. 

Chondria baileyana (Montagne) Harvey: VIRGINIA: H. J. 
Humm, York River in the Guinea Marshes; F. D. Ott, 
Burton’s Bay near Wachapreague. 

Chondria sedifolia Harvey: VIRGINIA: Rhodes (1970), 
Burton’s Bay near Wachapreague. 

Chondria tenuissima (Goodenough et Woodward) C. 
Agardh: VIRGINIA: H. J. Humm, jetty at Cape Charles. 
Zaneveld (1966-67); Rhodes (1970), Burton’s Bay near 
Wachapreague. MARYLAND: Zaneveld (1966-67), sine locus. 

Polysiphonia denudata (Dillwyn) Kiitzing: VIRGINIA: 
P. E. Hargraves, adrift in the York River at Gloucester 


1973] Marine Algae — Ott 281 


Point; H. J. Humm, York River at Gloucester Point; D. 
Kapram, York River at Gloucester Point; F. D. Ott, Chesa- 
peake Bay at Cape Charles. Zaneveld and Barnes (1965), 
lower Chesapeake Bay; Zaneveld (1966-67) ; Wulff (1967), 
York River at Gloucester Point; Rhodes (1970), Burton's 
Bay near Wachapreague. MARYLAND: B. L. Wulff, B. H. 
Robison and J. K. Lowry, intertidal region at Ocean City; 
J. K. Lowry, B. H. Robison and B. L. Wulff, north jetty at 
Ocean City; F. D. Ott, Assateague Beach, abundant in the 
wash. Zaneveld (1966-67); Wulff et al (1968), jetty at 
Ocean City. 

Polysiphonia harveyi Bailey: VIRGINIA: H. J. Humm, 
Hummock Channel near Wachapreague; M. Wass, York 
River near Gloucester Point; J. Vogel, east side of Gwynn 
Island near mouth of Rappahannock River; F. D. Ott, 
Hampton Roads at Lynnhaven Inlet. Zaneveld and Barnes 
(1965), lower Chesapeake Bay; Zaneveld (1966-67) ; Man- 
gum, Santos and Rhodes (1968), York River at Sandy 
Point; Mathieson and Fuller (1969), Mobjack Bay off 
Guinea Marsh; Marsh (1970), York River; Rhodes (1970), 
Burton’s Bay near Wachapreague. MARYLAND: B. H. Robi- 
son, J. K. Lowry and B. L. Wulff, north jetty at Ocean 
City; Zaneveld (1966-67); Wulff et al (1968), jetty at 
Ocean City; Mathieson and Fuller (1969), rock retaining 
wall on Naval Base property on south shore of Patuxent 
River, abandoned Cedar Point Light House, beach north 
of Cove Point, Holland Strait off South Marsh Island, mid- 
channel in Broad Creek connecting Little Annemessex 
River and Pocomoke Sound, Shark Fin Shoal east of 
Bloodsworth Island, 1.5 miles east of Bloodsworth Island, 
between Poplar Island Narrows Light and Can #1, 900 
yards east of Coaches Island, mouth of Crab Alley Bay off 
Parson Island and Bodkin Island, channel between James 
Island and Red Nun #2, east and west of Knapps Narrows, 
Kent Narrows and pilings at Piney Narrows Marina. 

Polysiphonia nigrescens (Hudson) Greville: VIRGINIA: 
B. L. Wulff, York River at Gloucester Point; F. D. Ott, 
Burtons Bay near Wachapreague. Zaneveld and Barnes 


282 Rhodora [Vol. 75 


(1965), lower Chesapeake Bay; Zaneveld (1966-67) ; Wulff 
(1967), York River at Gloucester Point; Wulff and Webb 
(1969), York River at Gloucester Point; Marsh (1970), 
York River; Rhodes (1970), Burton's Bay near Wacha- 
preague. MARYLAND: F. D. Ott, Ocean City on stones of 
jetty. Zaneveld (1966-67). 

Polysiphonia novae-angliae Taylor: MARYLAND: B. H. 
Robison, J. K. Lowry and B. L. Wulff, north jetty at Ocean 
City; J. K. Lowry, B. L. Wulff and B. H. Robison, north 
jetty at Ocean City. Wulff et al (1968), jetty at Ocean 
City. 

Polysiphonia subtilissima Montagne: VIRGINIA: J. Vogel, 
west side of Gwynn Island near mouth of the Rappahan- 
nock River; F. D. Ott, Chesapeake Bay at Cape Charles. 
Zaneveld (1966-67); Marsh (1970), York River; Rhodes 
(1970), Burton's Bay near Wachapreague. MARYLAND: 
Zaneveld (1966-67). 

Polysiphonia urceolata (Lightfoot) Greville: MARYLAND: 
J. K. Lowry and B. L. Wulff, north jetty at Ocean City. 
Wulff et al (1968), jetty at Ocean City. 


DIVISION CYANOPHYCOPHYTA 


Family CHROOCOCCACEAE Nägeli 

Agmenellum quadruplicatum (Meneghini) de Brébisson: 
GENERAL: Griffith (1961), Chesapeake Bay, R. E. Griffith; 
D. C. Morris. 

Agmenellum thermale (Kützing) Drouet et Daily: MARY- 
LAND: Drouet and Daily (1956), Potomac River at Plum- 
mers Island west of Cabin John, F. Drouet, E. P. Killip 
and D. Richards. 

Anacystis cyanea Drouet et Daily: VIRGINIA: B. L. Wulff 
bloom in the Rappahannock between Tappahannock and 
Port Royal. Drouet and Daily (1956), Queens Creek at 
Haw Tree Landing, A. F. Chestnut. 

Anacystis dimidiata (Kützing) Drouet et Daily: VIR- 
GINIA: Zaneveld (1966), Chincoteague Island in a ditch, 
L. C. Goldstein, E. S. Luttrell and J. C. Strickland. MARY- 


1973] Marine Algae — Ott 283 


LAND: Zaneveld (1966), Smith Island at Tylerton, P. W. 
Wolle. GENERAL: Griffith (1961), Chesapeake Bay, F. 
Drouet. 

Anacystis marina Drouet et Daily: VIRGINIA: E. T. and 
B. L. Wulff, York River at Gloucester Point; H. J. Humm, 
York River at Gloucester Point. GENERAL: Griffith (1961), 
Chesapeake Bay, F. Drouet. 

Anacystis montana (Lightfoot) Drouet et Daily forma 
montana (Lightfoot) Drouet et Daily: GENERAL: Griffith 
(1961), Chesapeake Bay. 

Anacystis montana (Lightfoot) Drouet et Daily forma 
minor (Wille) Drouet et Daily: GENERAL: Griffith (1961), 
Chesapeake Bay. 

Anacystis thermalis (Meneghini) Drouet et Daily forma 
thermalis (Meneghini) Drouet et Daily: GENERAL: Griffith 
(1961), Chesapeake Bay. 

Chroococcus turgidus (Kützing) Nägeli: MARYLAND: 
Drouet (1939), in brackish waters: marsh pool at Chance, 
marsh pool west of Ewell on Smiths Island, marsh pool at 
Tylerton on Smiths Island, P. W. Wolle. 

Chroococcus sp.: MARYLAND: Morse (1947), mouth of 
Patuxent River. 

Coccochloris elabens (de Brébisson) Drouet et Daily: 
GENERAL: Griffith (1961), Chesapeake Bay, F. Drouet. 

Coccochloris peniocystis (Kiitzing) Drouet et Daily: 
GENERAL: Griffith (1961), Chesapeake Bay. 

Coccochloris stagnina Sprengel: VIRGINIA: Drouet and 
Daily (1956), Virginia Beach in pool and on wet soil in 
the Back Bay National Wildlife Refuge, H. A. Bailey, E. S. 
Luttrell and J. C. Strickland; Zaneveld (1966), Virginia 
Beach in the Back Bay National Wildlife Refuge in pool 
and on wet soil under slightly brackish conditions, H. A. 
Bailey, E. S. Luttrell and J. C. Strickland. MARYLAND: 
Zaneveld (1966). GENERAL: Griffith (1961), Chesapeake 
Bay, F. Drouet. 

Gomphosphaeria aponina Kützing: MARYLAND: Drouet 
(1939), in brackish water: marsh pool between Chance 


284 Rhodora [Vol. 75 


and Dames Quarter. GENERAL: Griffith (1961), Chesapeake 
Bay, F. Drouet. 

Gomphosphaeria lacustris Chodat: GENERAL: Griffith 
(1961), Chesapeake Bay. 

Johannesbaptistia pellucida (Dickie) Taylor et Drouet: 
MARYLAND: Drouet (1939), brackish waters: marsh pool 
west of Ewell on Smiths Island, marsh pool between 
Chance and Dames Quarter, P. W. Wolle. GENERAL: Grif- 
fith (1961), Chesapeake Bay, F. Drouet. 

Merismopedia glauca (Ehrenburg) Nägeli: MARYLAND: 
Morse (1947), mouth of Patuxent River. 

Family ENTOPHYSALIDACEAE Geitler 

Entophysalis conferta Drouet et Daily: VIRGINIA: M. 
Wass, York River at Gloucester Point; Mangum, Santos 
and Rhodes (1968), York River at Sandy Point. MARY- 
LAND: Wulff et al (1968), jetty at Ocean City. 

Entophysalis deusta (Meneghini) Drouet et Daily: VIR- 
GINIA: Drouet and Daily (1956), York River at Yorktown, 
J. C. Strickland, York River at Yorktown, H. J. Humm; 
Zaneveld (1966), York River at Yorktown, J. C. Strickland, 
Zaneveld (1966); Wulff (1967), York River at Gloucester 
Point; Mangum, Santos and Rhodes (1968), York River 
at Sandy Point; Wulff and Webb (1969), York River at 
Gloucester Point. MARYLAND: B. H. Robison, J. K. Lowry 
and B. L. Wulff, north jetty at Ocean City. Drouet and 
Daily (1956), salt marsh flat at Chesapeake Beach, E. C. 
Leonard, in Patuxent River at Solomons Island, F. Drouet, 
E. P. Killip and F. R. Forsberg, on mud at Wenona, P. W. 
Wolle and F. Drouet, on hardened soil on the shore of Tan- 
gier Sound near Prickly Point, F. Drouet and P. W. Wolle; 
Zaneveld (1966). Wulff et al (1968), jetty at Ocean City. 
GENERAL: Griffith. (1961), Chesapeake Bay, F. Drouet. 

Family OSCILLATORIACEAE (S. F. Gray) 
Dumortier ex Kirchner 

Arthrospira brevis (Kützing) Drouet: VIRGINIA: Drouet 
(1968), culture from York River at Yorktown, J. C. Strick- 
land. MARYLAND: Drouet (1968), on mud and pilings at 


1973] Marine Algae — Ott 285 


Wenona, P. W. Wolle and F. Drouet; bank of Potomac 
River above Point of Rocks, M. H. Hohn. 

Hydrocoleum | holdenii Tilden: MARYLAND: Drouet 
(1939), in a salt marsh one mile south of Public Landing 
in Assateague Bay, P. W. Wolle; marsh ditch at Tylerton 
on Smiths Island, P. W. Wolle. GENERAL: Griffith. (1961), 
Chesapeake Bay, F. Drouet and F. K. Daily. 

Lyngbya  aerugineo-coerulea  (Kützing) Gomont: GEN- 
ERAL: Griffith (1961), Chesapeake Bay. 

Lyngbya aestuarii (Mertens) Lyngbye in Liebman ex 
Gomont: VIRGINIA: Strickland (1940), from a marine 
aquarium at West Point, J. W. Bailey and J. C. Strickland; 
Zaneveld (1966), Chincoteague Island, L. C. Goldstein, 
E. S. Luttrell and J. C. Strickland; coating on dead tree 
five miles upstream from the York River Bridge, at the 
old ferry landing at Gloucester Point, on border of York 
River near Gloucester Point, C. F. Rhyne; Zaneveld (1966- 
67). MARYLAND: Drouet (1939), subaerial in brackish and 
marine waters: salt marsh at Public Landing in Assa- 
teague Bay, P. W. Wolle, brackish ditch at Ewell; Zaneveld 
(1966), Rattlesnake Landing at Chincoteague Bay, J. Zane- 
veld, W. D. Barnes and H. W. West; Zaneveld (1966-67). 
GENERAL: Griffith (1961), Chesapeake Bay, F. Drouet. 

Lyngbya confervoides C. Agardh ex Gomont: VIRGINIA: 
Zaneveld (1966). MARYLAND: Drouet (1939), on rocks 
and wood in marine waters: on pilings of old wharf at 
Public Landing in Assateague Bay, P. W. Wolle, jetty on 
Upper Thoroughfare on Deal Island, on rocks at Wenona; 
Zaneveld (1966), Ocean City in harbor on wooden pilings, 
J. Zaneveld, W. D. Barnes and H. W. West; Zaneveld 
(1966). GENERAL: Griffith (1961), Chesapeake Bay, F. 
Drouet. 

Lyngbya gracilis (Meneghini) Rabenhorst: VIRGINIA: 
H. J. Humm, York River in the Guinea Marshes. 

Lyngbya lutea (C. Agardh) Gomont ex Gomont: VIR- 
GINIA: Strickland (1940), in brackish and salt water habi- 
tats: from pilings of West Norfolk Bridge in Portsmouth, 
on stump in James River at Cobham’s Wharf; Zaneveld 


286 Rhodora [Vol. 75 


(1966), on pilings of West Norfolk Bridge in Portsmouth, 
J. C. Strickland, on stump in James River at Cobham's 
Wharf, J. C. Strickland; York River at Seaford, R. W. 
Menzel and J. C. Strickland. MARYLAND: Drouet (1939), 
on rocks and wet soil on shores: on banks of Nanticoke 
River opposite Sand Hill Beach at Tyaskin. GENERAL: 
Griffith (1961), Chesapeake Bay, F. Drouet. 

Lyngbya semiplena (C. Agardh) J. Agardh ex Gomont : 
VIRGINIA: Zaneveld (1966), Chincoteague Island, L. C. 
Goldstein, E. S. Luttrell and J. C. Strickland, Wacha- 
preague in harbor on wooden pilings, J. Zaneveld, W. D. 
Barnes and H. W. West, Cape Charles city harbor on rocks 
of breakwater, C. F. Rhyne, Virginia Beach at Lynnhaven 
Inlet, W. D. Barnes, Virginia Beach at Lynnhaven Inlet 
forming a slimy coating on a bulkhead, W. D. Barnes, 
western side of Lynnhaven Bay on oyster shells, W. D. 
Barnes, Little Creek Inlet on the western breakwater, C. F. 
Rhyne, B. Phillips and J. Zaneveld, Back Bay, H. A. Bailey, 
E. S. Luttrell and J. C. Strickland, Chesapeake at west side 
of Great Bridge, C. M. Wilson, Norfolk on rocks, W. D. 
Barnes and H. W. West, in York River at Queens Creek, 
A. F. Chestnut and J. C. Strickland, York River five miles 
upstream from York River Bridge, along border of York 
River, C. F. Rhyne. MARYLAND: J. K. Lowry, B. H. Robison 
and B. L. Wulff, north jetty at Ocean City. Drouet (1939), 
in marine and brackish waters: in Chesapeake Bay, H. C. 
Bold; Zaneveld (1966) ; Wulff et al (1968), jetty at Ocean 
City. GENERAL: Griffith (1961), Chesapeake Bay, F. 
Drouet. 

Microcoleus chthonoplastes (Mertens) Zanardini ex 
Gomont: VIRGINIA: K. M. S. Aziz, Mobjack Bay; Zaneveld 
(1966), at the old ferry landing at Gloucester Point, on 
border of York River near Gloucester Point, C. F. Rhyne; 
Zaneveld (1966-67) ; Wulff (1967), York River at Glouces- 
ter Point; Wulff and Webb (1969), York River at Glouces- 
ter Point. MARYLAND: Drouet (1939), on mud flat between 
Ewell and Rhodes Point on Smiths Island, brackish ditch 
between Wenona and Deal Island, wet sand along Upper 


1973] Marine Algae — Ott 287 


Thoroughfare on Deal Island, on wet soil at White Haven 
Ferry at Mt. Vernon; Zaneveld (1966), Rattlesnake Land- 
ing in Chincoteague Bay, J. Zaneveld, W. D Barnes, and 
H. W. West; Zaneveld (1966-67). GENERAL: Griffith 
(1961), Chesapeake Bay, F. Drouet. 


Microcoleus lyngbyaceus (Kiitzing) Crouan: VIRGINIA: 
Wulff (1967), York River at Gloucester Point; Drouet 
(1968), Mulberry Island in James River, R. Patrick; 
Wulff and Webb (1969), York River at Gloucester Point. 
MARYLAND: Drouet (1968), Crisfield, F. Drouet and P. W. 
Wolle; Cabin John, F. Drouet and E. P. Killip, Chesapeake 
Bay, H. C. Bold, Chincoteague Bay, J. Zaneveld, W. D. 
Barnes and H. W. West. 

Microcoleus tenerrimus Gomont: VIRGINIA: K. M. S. Aziz, 
York River between York River Bridge and Nava] Weap- 
ons Station; Zaneveld (1966), western side of Lynnhaven 
Inlet on oyster shells, W. D. Barnes and H. W. West. 
MARYLAND: Drouet (1939), in brackish and marine waters: 
on pilings at Wenona. GENERAL: Griffith (1961), Chesa- 
peake Bay, F. Drouet. 


Microcoleus vaginatus (Vaucher) Gomont: VIRGINIA: 
Drouet (1968), James River downstream from J amestown, 
R. Patrick. MARYLAND: Drouet (1968), Patuxent River at 
Benedict, F. Drouet and C. W. Reimer, Plummers Island 
in Potomac River west of Cabin John, Drouet et al. 

Oscillatoria amphibia C. Agardh ex Gomont: VIRGINIA: 
Zaneveld (1966), west of Queen Sound, L. C. Goldstein, 
E. S. Luttrell, and J. C. Strickland. MARYLAND: Drouet 
(1939), in brackish and almost fresh water: marsh pool 
between Chance and Dames Quarter; Zaneveld (1966). 
GENERAL: Griffith (1961), Chesapeake Bay, F. Drouet. 

Oscillatoria brevis Kützing ex Gomont var. neapolitana 
(Kützing) Gomont: MARYLAND: Drouet (1939), in brack- 
ish and semi-marine waters, on pilings at Wenona, puddle 
at old wharf at Shelltown, P. W. Wolle. GENERAL: Griffith 
(1961), Chesapeake Bay, F. Drouet. 

Oscillatoria laetevirens Crouan ex Gomont: VIRGINIA: 


288 Rhodora [Vol. 75 


Zaneveld (1966), York River at Seaford, J. C. Strickland ; 
Zaneveld (1966-67). 

Oscillatoria lutea C. Agardh: VIRGINIA: Drouet (1968). 
on pilings at Seaford, J. C. Strickland, on shells in James 
River 4.5 miles downstream from Jamestown Island, R. 
Patrick, pilings of bridge at Portsmouth, J. C. Strickland 
et al. MARYLAND: Drouet (1968), bank of Nanticoke River 
opposite Sandy Hill Beach at Tyaskin, P. W. Wolle and 
F. Drouet. 

Oscillatoria margaritifera Kützing ex Gomont: VIRGINIA: 
Zaneveld (1966), west of Queens Sound, L. C. Goldstein, 
E. S. Luttrell and J. C. Strickland, York River at Seaford, 
L. C. Goldstein, E. S. Luttrell and J. C. Strickland. MARY- 
LAND: Drouet (1939), in brackish and semimarine waters: 
marsh ditch at Weona, P. W. Wolle, marsh pool at Chance. 
marsh ditch at Tylerton on Smiths Island, P. W. Wolle; 
Zaneveld (1966). 

Oscillatoria  nigro-viridis Thwaites in Harvey ex Go- 
mont: VIRGINIA: B. L. Wulff, York River at Gloucester 
Point. Strickland (1940), on pilings of the West Norfolk 
Bridge at Portsmouth; Zaneveld (1966), west of Queen 
Sound, L. C. Goldstein, E. S. Luttrell and J. C. Strickland, 
on pilings of West Norfolk bridge, J. C. Strickland. MARY- 
LAND: Drouet (1939), in marine and semimarine waters: 
on pilings at Wenona. 

Oscillatoria princeps Vaucher ex Gomont: VIRGINIA: 
Zaneveld (1966), northwest of Wachapreague, L. C. Gold- 
stein, E. S. Luttrell and J. C. Strickland, Portsmouth west 
of Hickory, C. M. Wilson. MARYLAND: Zaneveld (1966). 

Oscillatoria retzii C. Agardh: VIRGINIA: Drouet (1968), 
on logs in James River north of the Fort Eustis Dock, 
R. Patrick. 

Oscillatoria salinarum Collins, Holden and Setchell: 
VIRGINIA: Zaneveld (1966), west of Queen Sound, L. C. 
Goldstein, E. S. Luttrell and J. C. Strickland. MARYLAND: 
Drouet (1939), brackish ditch at Ewell on Smiths Island. 
marsh diteh near Wenona, P. W. Wolle, Tylerton, P. W. 
Wolle. 


1973] Marine Algae — Ott 289 


Oscillatoria splendida Greville ex Gomont: VIRGINIA: 
Zaneveld (1966), Portsmouth west of Great Bridge, C. M. 
Wilson; Zaneveld (1966-67). MARYLAND: Zaneveld (1966). 

Oscillatoria submembranaceae Ardissone et Strafforello: 
VIRGINIA : Drouet (1968), on barnacles on a pier at Seaford, 
J. C. Strickland. MARYLAND: Drouet (1968), on mud at 
Ewell on Smiths Island, P. W. Wolle and F. Drouet, on an 
alluvial flat on Plummers Island in the Potomac River west 
of Cabin John. 

Oscillatoria subuliformis Thwaites in Kützing ex Go- 
mont: VIRGINIA: M. Wass, on barnacles on a buoy in Sa- 
rah's Creek. Strickland (1940), from experimental jars in 
which oysters were active at West Point, J. W. Bailey, in 
Richmond from a marine aquarium at the University of 
Virginia, J. W. Bailey and J. C. Strickland. Zaneveld 
(1966), Yorktown, R. M. Moore and J. C. Strickland, south 
border of York River. 

Oscillatoria sp.: VIRGINIA: Marshall (1967), Elizabeth 
River. MARYLAND: Morse (1947), mouth of Patuxent 
River. 

Phormidium persicinum (Reinke) Gomont: VIRGINIA: 
Strickland (1940), from marine aquaria at West Point, 
J. W. Bailey and J. C. Strickland. 

Phormidium retzii (C. Agardh) Kützing ex Gomont: 
VIRGINIA: Strickland (1940), James River at Cobham's 
Wharf. 

Phormidium submembranaceum (Ardissone et Straffo- 
rello) Gomont: VIRGINIA: H. J. Humm, York River at 
Gloucester Point. 

Phormidium uncinatum (J. Agardh) Gomont: VIRGINIA: 
M. Wass, York River at Gloucester Point. 

Phormidium weissi Drouet: MARYLAND: Drouet (1939), 
in a brackish pool at Wenona, on Ruppia in a marsh pool 
between Ewell and Rhodes Point on Smiths Island, road 
puddle at old wharf at Shelltown, P. W. Wolle. GENERAL: 
Griffith (1961), Chesapeake Bay, F. Drouet. 

Porphyrosiphon notarisii (Meneghini) Kützing: VIR- 
GINIA: Wulff (1967), York River at Gloucester Point; 


290 Rhodora [Vol. 75 


Drouet (1968), Mulberry Island in the James River; Wulff 
and Webb (1969), York River at Gloucester Point. MARY- 
LAND: Drouet (1968), in a saltmarsh near Jenkins Creek 
at Crisfield, F. Drouet and P. W. Wolle. 

Porphyrosiphon splendidus (Greville) Drouet: VIRGINIA: 
Drouet (1968), Portsmouth west of Great Bridge, C. M. 
Wilson. MARYLAND: Drouet (1968), Potomac River three 
miles below mouth of Monocacy River, M. H. H ohn, Water- 
loo Marsh at Monie Creek, P. W. Wolle. 

Schizothrix arenaria (Berkeley) Gomont: VIRGINIA: 
Drouet (1968), James River north of Fort Eustis Dock, 
R. Patrick. MARYLAND: Drouet (1968), Plummers Island 
in Potomac River west of Cabin John, F. Drouet and E. P. 
Killip, marsh on shore of Patuxent River, F. Drouet and 
C. W. Reimer, salt marsh near Rattlesnake Landing at 
Chincoteague Bay, Moul. 

Schizothrix calcicola (C. Agardh) Gomont: VIRGINIA: 
E. T. and B. L. Wulff, York River at Gloucester Point; 
H. J. Humm, York River at Gloucester Point. Zaneveld 
(1966), Strickland in MS 1943; Zaneveld (1966-67) ; Wulff 
(1967), York River at Gloucester Point; Drouet (1968), 
James River at Mulberry Island, R. Patrick, York River 
at West Point, J. W. Bailey, in James River downstream 
from Jamestown Island; Mangum, Santos and Rhodes 
(1968), York River at Sandy Point; Wulff and Webb 
(1968), York River at Gloucester Point. MARYLAND: 
Drouet (1968), in Potomac River west of Cabin John, 
F. Drouet and E. P. Killip, salt marsh at Crisfield, F. 
Drouet and P. W. Wolle, in Patuxent River at Solomons 
Island, E. P. Killip, F. Drouet and F. R. Fosberg; Wulff 
et al (1968), jetty at Ocean City. 

Schizothrix tenerrima (Gomont) Drouet: VIRGINIA: 
Drouet (1968), James River at Mulberry Island, R. Pat- 
rick, James River at Cobham’s Wharf, J. C. Strickland. 
MARYLAND: Drouet (1968), Patuxent River at Solomons 
Island, F. Drouet, E. P. Killip and F. R. Fosberg, at 
Wenona, P. W. Wolle and F. Drouet. 

Spirulina major Kützing ex Gomont: MARYLAND: Drouet 


1973] Marine Algae — Ott 291 


(1939), brackish pool at Wenona, Cove Point, H. C. Bold. 
GENERAL: Griffith. (1961), Chesapeake Bay, F. Drouet. 

Spirulina subsalsa Oersted ex Gomont: VIRGINIA: Strick- 
land (1940), York River near West Point, J. W. Bailey, at 
West Point from jars in which oysters were active, J. W. 
Bailey, in Richmond at the University of Richmond in a 
marine aquarium; Zaneveld (1966), York River near York- 
town, J. C. Strickland; Zaneveld (1966-67) ; Wulff (1967), 
York River at Gloucester Point; Wulff and Webb (1969), 
York River at Gloucester Point. MARYLAND: Drouet 
(1939), in marine and brackish waters: in a marsh pool 
between Chance and Dames Quarter; Zaneveld (1966) ; 
Drouet (1968), Cove Point, H. C. Bold, Jerico Marshes 
west of Fairmount, F. Drouet and P. W. Wolle, between 
Ewell and Rhodes Point on Smiths Island, P. W. Wolle 
and F. Drouet, marsh south of Jenkins Creek near Cris- 
field, F. Drouet and P. W. Wolle. GENERAL: Griffith (1961), 
Chesapeake Bay, F. Drouet. 

Spirulina subsalsa Oersted forma oceanica (Crouan) 
Gomont: VIRGINIA: H. J. Humm, on barnacles from a buoy 
in Sarah’s Creek. 


Spirulina tenerrima Kiitzing ex Gomont: MARYLAND: 
Drouet (1939), in brackish and marine waters: marsh 
pool between Ewell and Rhodes Point on Smiths Island. 
GENERAL: Griffith (1961), Chesapeake Bay, F. Drouet. 

Spirulina sp.: MARYLAND: Morse (1947), mouth of Pa- 
tuxent River. 

Symploca atlantica Gomont: VIRGINIA: Strickland 
(1940), James River at Cobham’s Wharf; Zaneveld (1966), 
York River near Gloucester Point, C. F. Rhyne, James 
River at Cobham’s Wharf, J. C. Strickland. MARYLAND: 
Zaneveld (1966), on wooden harbor pilings at Snow Hill 
Public Landing, J. Zaneveld, W. D. Barnes, and H. W. 
West. 

Trichodesmium thiebautii Gomont: VIRGINIA: F. D. Ott, 
very abundant in the phytoplankton during August and 
September in the lower Chesapeake Bay, 1970. 


292 Rhodora [Vol. 75 


Family NOSTOCACEAE Dumortier ex Engler 

Anabaena inaequalis (Kützing) Bornet et Flahault: VIR- 
GINIA: H. J. Humm, York River at Yorktown. 

Anabaena sphaerica Bornet et Flahault: MARYLAND: 
Drouet (1939), in a brackish ditch between Wenona and 
Deal Island. GENERAL: Griffith (1961), Chesapeake Bay, 
F. Drouet. 

Anabaena torulosa (Carmichael) Lagerheim ex Bornet 
et Flahault: VIRGINIA: Zaneveld (1966), south border of 
York River near the York River Bridge. 

Family MICROCHAETACEAE Lemmermann 

Nodularia harveyana (Thwaites) Thuret: VIRGINIA: 
K. M. S. Aziz, York River at Yorktown. MARYLAND: 
Drouet (1939), floating in a brackish pond at Rhodes 
Point on Smiths Island. GENERAL: Griffith (1961), Chesa- 
peake Bay, F'. Drouet. 

Family RIVULARIACEAE Kiitzing ex Bornet et Flahault 

Dichothrix penicillata Zanardini ex Bornet et Flahault: 
VIRGINIA: Zaneveld (1966), York River 15 miles down- 
stream from the York River Bridge. 

Calothrix confervicola (Roth) C. Agardh: VIRGINIA: 
K. M. S. Aziz, York River between the York River Bridge 
and the Naval Weapons Station. MARYLAND: B. L. Wulff, 
B. H. Robison and J. K. Lowry, north jetty at Ocean City, 
J. K. Lowry, B. H. Robison and B. L. Wulff, north jetty 
at Ocean City. Wulff et al (1968), jetty at Ocean City. 

Calothrix crustaceae Schousboe et Thuret ex Bornet et 
Thuret: VIRGINIA: H. J. Humm, York River at Gloucester 
Point. Zaneveld (1966), on oyster shells on the west side 
of Lynnhaven Inlet, W. D. Barnes and H. W. West, on 
wooden pole of the old ferry landing at Gloucester Point, 
J. Zaneveld; Zaneveld (1966-67); Wulff (1967), York 
River at Gloucester Point; Wulff and Webb (1969), York 
River at Gloucester Point. MARYLAND: B. H. Robison, J. K. 
Lowry and B. L. Wulff, north jetty at Ocean City; J. K. 
Lowry, B. H. Robison and B. L. Wulff, north jetty at 
Ocean City. Zaneveld (1966), Snow Hill Publie Landing 
on concrete blocks in Chincoteague Bay, J. Zaneveld, W. D. 


1973] Marine Algae — Ott 293 


Barnes and H. W. West; Zaneveld (1966-67) ; Wulff et al 
(1968), jetty at Ocean City. 

Calothrix pulvinata (Mertens) C. Agardh: MARYLAND: 
Drouet (1939), in marine and brackish waters: on pilings 
in Nanticoke River at Sandy Hill Beach in Tyaskin, on 
pilings of old wharf at Shelltown, P. W. Wolle. GENERAL: 
Griffith (1961), Chesapeake Bay, F. Drouet. 

Rivularia nitida C. Agardh: MARYLAND: Drouet (1939), 
in wet brackish places: on cedar stumps at head of Poco- 
moke Sound below Shelltown, P. W. Wolle. GENERAL: Grif- 
fith (1961), Chesapeake Bay, F. Drouet. 

Family SCYTONEMATACEAE Kützing ex Bornet et Flahault 

Fremyella grisea (Bornet et Flahault) J. de Toni: MARY- 
LAND: Drouet (1939), in marine and brackish waters: on 
pilings of old wharf at Shelltown, P. W. Wolle. GENERAL: 
Griffith (1961), Chesapeake Bay, F. Drouet. 

Plectonema calothrichoides Gomont: VIRGINIA: K. M. S. 
Aziz, York River above the York River Bridge. 

Plectonema golenkinianum Gomont: MARYLAND: Drouet 
(1939), in marine and brackish waters: on pilings of old 
wharf at Shelltown, P. W. Wolle. GENERAL: Griffith 
(1961), Chesapeake Bay, F. Drouet. 

Plectonema terebrans Bornet et Flahault: VIRGINIA: 
K. M. S. Aziz, York River near Yorktown. 

Family STIGONEMATACEAE Kirchner 

Mastigocoleus testarum Bornet et Flahault: VIRGINIA: 
H. J. Humm, York River at Gloucester Point. 


SUM MARY 


In the preceding treatment 213 taxa have been recorded 
for the area under study. A number of these taxa, espe- 
cially in the Chlorophycophyta, are strictly freshwater 
algae and are included herein only because they have been 
reported as being present. A few other taxa of the Cyano- 
phycophyta may be entered in this treatment under two 
different names for the reasons discussed in the introduc- 
tion to this paper. 


294 Rhodora [Vol. 75 


These 213 taxa are distributed in the four algal divi- 
sions as follows: 


Division Number Percent 
Chlorophycophyta 55 25.8 
Phaeophycophyta 29 13.6 
Rhodophycophyta 54 25.4 
Cyanophycophyta 75 35.2 


It is interesting to note that the majority of the collect- 
ing sites within the area under study center about either 
1) the marine laboratories located at Solomons Island, 
Gloucester Point, Norfolk, and Wachapreague, or, 2) the 
stone jetties at Ocean City and in Norfolk. Little atten- 
tion has been given to the algal flora elsewhere including 
the thousands of acres of Spartina salt marshes, the oyster 
reefs, and the possible algal flora of the deeper waters off 
the Atlantic coast and in the Chesapeake Bay. As these 
particular areas are more thoroughly and more systemati- 
cally surveyed, it should be anticipated that additional taxa 
will be added to the marine algal flora of Maryland and 
Virginia. For each algal species reported, its known geo- 
graphical distribution in the area is given. 


ACKNOWLEDGMENTS 


I acknowledge with pleasure the use of the Virginia 
Institute of Marine Science herbarium. I am appreciative 
to Dr. Russell Rhodes and Dr. Marvin Wass for referring 
me to pertinent papers in the periodical literature which 
were unknown to me. Lastly, I express my sincere thanks 
to Dr. Marvin Wass and to my wife, Dr. Aleta Petrik-Ott, 
who proofread this paper. 


LITERATURE CITED 


Borp, H. C. 1967. Morphology of Plants, 2nd ed. Harper & Row, 
Publishers, New York. XXIX + 541 pp. 

Drovet, F. 1939. The Myxophyceae of Maryland. Field Mus. Nat. 
Hist., Bot. Ser. (Chicago) 20(1): 1-14. 


1973] Marine Algae — Ott 295 


1968. Revision of the Classification of the Oscilla- 
toriaceae. Monograph No. 15, The Academy of Natural Sciences 
of Philadelphia. Philadelphia. 370 pp., including 131 figs. 

DROUET, F. and W. A. DAILY. 1956. Revision of the Cocoid Myxo- 
phyceae. Butler Univ. Bot. Studies (Indianapolis) XII: 1-218. 

GRIFFITH, R. E. 1961. Phytoplankton of Chesapeake Bay, an Illus- 
trated Guide to the Genera. Contribution No. 172, Cheasapeake 
Biological Laboratory, Maryland Department of Research and 
Education, Solomons, Maryland. IV + 79 pp., including 123 figs. 

HUMM, H. J. 1963. Dictyota dichotoma in Virginia. Virginia. Jour. 
Sci. (Richmond) (N.S.) 14(3): 109-111. 

MANGUM, C. P., S. L. SANTOS, and W. R. RHopEs, JR. 1968. Distri- 
bution and feeding in the onuphid polychaete, Diopatra cuprea 
(Bose.) Mar. Biol. (Berlin) 2: 33-40. 

MansH, G. A. 1970. A Seasonal Study of Zostera Epibiota in the 
York River, Virginia. Doctor of Philosophy Dissertation, College 
of William and Mary, Faculty of the School of Marine Science, 
Gloucester Point, Virginia. XI + 156 pp. ` 

MARSHALL, H. G. 1967. Plankton in James River Estuary, Vir- 
ginia II. Phytoplankton in the Elizabeth River. Virginia Jour. 
Sci. (Richmond) (N.S.) 18(3): 105-109. 

MATHIESON, A. and S. W. FULLER. 1969. A preliminary investiga- 
tion of the benthonie marine algae of Chesapeake Bay region. 
Rhodora (Boston) 71: 524-534. 

MonsE, D. C. 1947. Some observations on seasonal variations in 
plankton population, Patuxent River, Maryland, 1943-1945. 
Chesapeake Biol. Lab. Publ. (Solomons, Maryland) 65: 1-31. 

PAPENFUSS, G. F. 1955. Classification of the Algae, pp. 115-224 
in R. C. Miller, E. L. Kessel and G. F. Papenfuss. A. Century 
of Progress in the Natural Sciences, 1853-1953. California 
Academy of Science, San Francisco. XII + 807 pp. 

PATTEN, B. C, R. A. MurFoRD and J. E. WARINNER. 1963. An 
annual phytoplankton cycle in the lower Chesapeake Bay. 
Chesapeake Sci. (Solomons, Maryland) 4(1): 1-20. 

Post, E. 1968. Zur  Verbreitungs-Ókologie des  Bostrychietum. 
Hydrobiologia (Den Haag) 31(2): 241-3106. 

RHODES, R. G. 1970. Seasonal occurrence of marine algae on an 
oyster reef in Burton's Bay, Virginia. Chesapeake Sci, (Solo- 
mons, Maryland) 11(1): 61-63. 

STRICKLAND, J. C. 1940. The Oscillatoriaceae of Virginia. Amer. 
Jour. Bot. (Lancaster) 27: 628-633. 

Woopson, B. E. 1959. A study of the Chlorophyta of the James 
River basin, Virginia. I. Collection points and species list. Vir- 
ginia Jour. Sci. (Richmond) (N.S.) 10(2): 70-82. 


296 Rhodora [Vol. 75 


WuLrrF, B. L. 1967. Intertidal Zonation of Marine Algae at Glou- 
cester Point, Virginia. Master of Arts Thesis, The College of 
William and Mary, Department of Marine Science, The Vir- 
ginia Institute of Marine Science, Gloucester Point, Virginia. 
VII 4- 49 pp. 

Wutrr, B. L., E. M. T. WULFF, B. H. RoBISoN, J. K. Lowry, and 
H. J. HuMM. 1968. Summer marine algae of the jetty at 
Ocean City, Maryland. Chesapeake Sci. (Solomons, Maryland) 
9(1): 56-60. 

Wutrr, B. L. and K. L. WEBB. 1969. Intertidal zonation of marine 
algae at Gloucester Point, Virginia. Chesapeake Sci. (Solomons, 
Maryland) 10(1): 29-35. 

ZANEVELD, J. 1966. The marine algae of the American coast be- 
tween Cape May, N. J. and Cape Hatteras, N. C. I. The Cyano- 
phyta. Bot. Mar. (Hamburg) IX(94): 101-128, including figs. 
1-7. 

ZANEVELD, J. S. and W. D. BARNES. 1965. Reproductive periodicities 
of some benthic algae in lower Chesapeake Bay. Chesapeake 
Sci. (Solomons, Maryland) 6(1): 17-32. 

ZANEVELD, J. S. 1966-67. The Benthic Marine Algae of Delaware: 
A Preliminary Checklist, Revised from 1966. Scientific Series, 
Publication No. 2, Old Dominion College, Norfolk, Virginia. 
36 pp. mimeographed. 


VIRGINIA INSTITUTE OF MARINE SCIENCE 
GLOUCESTER POINT, VIRGINIA 23062 


THE PRINGLE HERBARIUM 
FRANK C. SEYMOUR 


In the Pringle Herbarium of the University of Vermont 
are 335,000 specimens of plants ranging all the way from 
Fungi and Algae to Orchids and Compositae. While Ver- 
mont is the area by far the best represented, the specimens 
come from many far-scattered and remote parts of the 
earth. Between 5,000 and 6,000 specimens are being added 
each year. We believe that few botanists realize what 
extensive resources are available in this Herbarium for 
research and study. Visiting botanists have exclaimed re- 
peatedly over the wealth of material which they have found 
here. *I had no idea that you had so many significant 
specimens," one commented. Or, “If we had only known 
you had such rare numbers, we should have come sooner." 
*[ supposed this specimen was not to be seen nearer than 
Europe," another added. When there are such choice 
specimens in this herbarium, to make these resources 
known to the scientific world becomes almost a duty. 

What proportion of our 337,000 specimens come from 
our own state is difficult to estimate. In view of the many 
sheets from other parts of the world, it cannot be large, — 
pehaps 1 in 10 from the state of Vermont. A County List, 
now being prepared by the Vermont Botanical and Bird 
Club, will, when published, show how wide a representa- 
tion is from Vermont. That County List will include vari- 
ous herbaria, with symbols to indicate in which herbaria 
voucher specimens are to be found. 

Among collectors of Vermont plants, Cyrus Guernsey 
Pringle still leads the list. Beginning his collecting in 
1876, Vermont, especially his home town of Charlotte, was 
naturally the scene of his earliest as well as his latest 
endeavors. By his own labor and initiative, he built up the 
herbarium to 150,000 sheets. While the majority of these 
are from Mexico, he made extensive collections from his 
own state. 


297 


298 Rhodora [Vol. 75 


Mrs. Nellie F. Flynn assembled a massive herbarium, 
mostly by her own collecting, largely from the region of 
Burlington. Her entire herbarium is deposited with the 
Pringle Herbarium as a permanent loan. 

Ezra Brainerd, at one time President of Middlebury 
College, is best known for his published studies of Violets 
and Blackberries. Also in the genus Crataegus (Haw- 
thorns), he collected great quantities of specimens. His 
entire herbarium is now in the Pringle Herbarium. 

Dana S. Carpenter built up a large herbarium of notable 
importance, mostly from Vermont, but including also much 
material from distant lands. D. Lewis Dutton accomplished 
a similar work with specimens mostly from Vermont. Both 
of their herbaria are now parts of the Pringle Herbarium. 

A. J. Grout, most famous for his work with mosses, is 
another of the early collectors, thousands of whose speci- 
mens add greatly to our knowledge of the flora of Vermont. 

Among others who have contributed notably to this her- 
barium and to our knowledge of Vermont plants, the fol- 
lowing botanists should be mentioned. L. R. Jones, at one 
time Curator of this Herbarium, added many significant 
specimens. Sidney F. Blake, known throughout the world 
as an authority on the family, Compositae, contributed 
many specimens especially from the northwestern part of 
Vermont. George L. Kirk, by profession a journalist, by 
his keen observation added extensively to the known flora 
of the state. The late Leopold A. Charette contributed 
greatly to the knowledge of distribution records in Ver- 
mont and would have done much more if an untimely 
death had not cut short his botanical career. His earlier 
herbarium containing a large collection of plants of Japan 
as well as of Vermont was purchased by the Pringle 
Herbarium. The herbarium of Ernest L. Smith, with 
many choice plants of Vermont, was a gift of his daugh- 
ter, Marion L. Smith. 

From the other New England states, there are substan- 
tial numbers of specimens, with Maine probably leading. 
M. L. Fernald, John J. Cushman and Frank C. Seymour 


1973] Pringle Herbarium — Seymour 299 


are among the collectors there. J. W. Congdon's plants 
from Rhode Island include some items which probably are 
no longer to be found growing in that populous state. In 
Connecticut, collections by Arthur E. Blewitt, E. H. Eames 
and D. C. Eaton are notable. From Massachusetts, we have 
many specimens of F. F. Forbes and some of Thomas 
Morong, distinguished for his studies in aquatics. Mrs. 
Nellie F. Flynn's collections from Nantucket are remark- 
able not only for their quantity but for their variety. 
Among the large number of species are some which ap- 
parently have not been found there by any other botanist. 
Many specimens from New Hampshire were collected by 
A. S. Pease, author of “The Flora of Northern New 


Hampshire." 


Nearly every one of the fifty states in the United States 
is amply represented. Among the outstanding collectors 
are the following: New York, A. J. Grout; New Jersey, 
Levi J. Miller “Halsted’ American Weeds”; Pennsylvania, 
J. K. Small, A. A. Heller; North Carolina, A. H. Curtiss; 
Florida, A. H. Curtiss, Nellie F. Flynn; Michigan, Frank 
C. Gates; Alabama, G. R. Vasey and “Ea herb. C. Mohr, 
Mobile, Ala.," F. S. Earle, C. F. Baker; Gulf States, S. M. 
Tracy; Minnesota, J. M. Holzinger, G. B. Ownbey; Wiscon- 
sin, T. G. Hartley; Illinois, C. G. Pringle; Iowa, Jack L. 
Carter; Missouri, B. F. Bush, N. C. Henderson; Louisiana, 
Nellie F. Flynn; Texas, J. Reverchon, N. C. Henderson; 
Wyoming, Aven Nelson; Colorado, George W. Letterman, 
C. F. Baker, H. N. Patterson, C. C. Parry; Utah, Marcus 
E. Jones, S. L. Welch, G. Moore; Nevada, C. F Baker; Ari- 
zona, C. G. Pringle, Elinor Lehto; California, C. G. Pringle, 
Joseph P. Tracy, Lewis S. Rose, A. D. E. Elmer, S. B. and 
W. F. Parish; Washington, C. V. Piper, W. N. Suksdorf, 
A. D. E. Elmer; Pacific Coast, Thomas J. Howell. 


Many of the collectors are pioneers in their areas, so 
that their specimens are of special value, and often cited 
in literature. From Alaska, we have many specimens of 
Wiliam Chadsworth. Cynthia J. Bigelow has made us a 


300 Rhodora [Vol. 75 


gift of her summer's collection from the Yukon. Green- 
land's flora, most nearly related to North America, is 
represented by several hundred specimens distributed by 
the University of Copenhagen. 


Turning to areas south of the United States, from Mexico 
we have the very extensive collections of Pringle, himself, 
whom Asa Gray called “the Prince of Plant Collectors." 
While his specimens have been distributed far and wide 
to many other herbaria, it is not certain that all numbers 
were so distributed. We probably have many which are 
not to be found in any other herbarium. Since his time, 
our Mexican material has been augmented by specimens 
from other collectors such as Edward Palmer, E. Matuda, 
C. Conzatti, Forrest M. Shreve, D. E. Breedlove, Harde 
LeSueur, Elwood Molseed, Stephen S. White, and C. L. 
Smith. 


From Central America there are specimens collected by 
the following: Guatemala, H. von Türckheim and Charles 
C. Deam; British Honduras, P. H. Gentle; Honduras, C. 
Thieme; Costa Rica, C. W. Dodge, W. S. Thomas, Austin 
Smith; Panama, M. E. Davidson, S. Hayes, H. von Wedel, 
P. H. Allen, R. E. Woodson, Jr., R. W. Schery. 


As a result of recent expeditions to N icaragua, the 500 
sheets already mounted from the largest country of Central 
America are only a small part of those soon to be added 
from the large collections distributed by Frank C. Seymour. 
While many numbers are being distributed to other her- 
baria, the unicates are being deposited in this herbarium. 


From the West Indies, our largest collections are those 
from Guadeloupe by Pere Duss and others labeled “Herbier 
Capitaine.” Cuba is one of the many areas from which 
Pringle has provided us with much material. To his have 
been added specimens of A. H. Curtiss. 


South America is represented by important series from 
certain areas, such as the following: Brazil, Ex Museo 
Botanico Berolinensi, Sellow; “Flora von Mattogrosso,” 
“Dr. Pilger,” 1899; Plantae Chilensis, Otto Buchtein; Peru, 


1973] Pringle Herbarium — Seymour 301 


C. M. Belshaw; Colombia, Herbert H. Smith, H. W. Vogel- 
mann. 

In smaller numbers, there are specimens from nearly 
every other country of South America. 

For an American herbarium, we have a surprisingly 
large number of specimens from Europe. Here are listed 
some of the most numerous sets. Plantae Vasculares Dani- 
cae, Plantae Scandinavicae, Einar Sóderen, Jutland, Flora 
Exsiccata, Plantae Danicae, Alfred Hansen, Plantae Sue- 
cicae, Emil Almquist, Plantes de France, E. J. Neyraut, 
Plantes de France, Fr. Sennen, Plantae France, C. J. 
Pitard, Herbier, C. Copineau, Plantes des Hautes Alpes, 
Herbier, A. Faure, Silicia, Mary F. Spencer, Flora Ingieae 
(Russia), Flora Exsiccata Italica, Macedonia, Dimonie. 

At least a few thousand specimens have come from 
Africa including the following important series: “Plantae 
Schlechterianae Occidentali-Africanae", Ex Herbario Nata- 
lense, Legit J. Medley Wood and M. Franks, Cape Govern- 
ment Herbarium, Austro-Africanum, Cent. XVI, Austro- 
Africanum, Cent. XVII, MacOwan, Fragmenta Florae 
algeriensis exsiccata, Rec. par Ed. Lafrance, Plantes de 
Tunisie, C. J. Pitard, Plantes d'Algerie — 1898, Elisee 
Reverchon, “W. Schimper pl. Abyssin." 

The most interesting feature of the specimens from 
Africa and likewise from Europe is that a large number 
of them appear to have come from the Berlin Herbarium. 
Some of them are clearly so marked, as for instance nu- 
merous specimens from Kamerun, bearing the heading on 
the label: Museum Botanicum Berolinense. Thus the 
Pringle Herbarium may have the only extant duplicates 
of numbers which were destroyed when the Berlin Her- 
barium was bombed in March 1943. 

The part of Asia best represented in the Pringle Her- 
barium is Japan. The specimens from that country were 
obtained by purchase of the early herbarium of Leopold 
A. Charette who collected many of them himself and who 
evidently exchanged with Japanese herbaria, as many of 
his specimens were collected by various Japanese botanists. 


302 Rhodora [Vol. 75 


Many from India belong to the rare series bearing the 
label with the familiar heading, “Herb. Ind. Or. Hook. f. 
& Thomson" or “Flora of the Sikkim Himalaya" coll. G. A. 
Gammie; and others from «Herb. Hort. Bot. Calcuttensis" 
are significant. The Philippines are represented by a large 
series of the well known collections by E. D. Merrill and 
A. D. E. Elmer. 


Even from the Antipodes come large collections by well 
known botanists. That the Australian continent is well 
represented is especially remarkable. Among those sheets 
are many from the «National Herbarium of New South 
Wales," coll. J. L. Boorman, J. H. Maiden, J. B. Cleland, 
and J. H. Camfield. Those from Western Australia were 
collected by Max Koch. 


How many types are ineluded in this herbarium is un- 
known. Of those already known and recognized, there are 
about 200. Formerly unrecognized types are frequently dis- 
covered. Most of these are collections by Pringle in Mexico. 


A unique feature is the Pringle Index, compiled largely 
by Leopold A. Charette. It consists of about 15,000 cards, 
3 x 5 inches, numerically arranged by collection numbers, 
each containing a note on the specimens of Cyrus Guernsey 
Pringle in Mexico. They have been compiled (1) from 
specimens in the Pringle Herbarium; (2) from many 
periodicals and especially monographs; (3) from the 
“Catalogo de Plantas Mexicanas" by Manuel Urbina; (4) 
and from “Ferns and Fern Allies of Chihuahua" by I. W. 
Knobloch and D. S. Correll. The cards give references to 
citations, changes in identification and often indicate by 
international symbols the names of herbaria where dupli- 
cates are to be found. The code symbol used internationally 
to designate the Pringle Herbarium is VT, obviously se- 
lected because it is the state herbarium of Vermont. 


Support of the Pringle Herbarium comes from an endow- 
ment of $35,000. The one full-time worker is the curator. 
Part-time assistants, usually two at a time, are selected 
from among the students in the University. Altho the 


1973] Pringle Herbarium — Seymour 303 


turnover is rather frequent, students prove to be highly 
capable assistants. 

The first curator was none other than Dr. Pringle him- 
self who started his herbarium apparently in 1876. It 
remained his private herbarium until 1902 when it was 
given to the University of Vermont with him continuing 
as curator until the time of his death in 1911. For some 
years thereafter the herbarium was cared for by members 
of the Department of Botany without a designated curator. 
The second curator, E. J. Dole, who filled the office for 
several years, was succeeded by H. W. Vogelmann from 
1955 to 1971. The present incumbent, F. C. Seymour, be- 
came curator in 1971. 

The Pringle Herbarium is readily accessible at 601 Main 
Street, Burlington, Vermont. Its brick building, formerly 
a weather bureau station, is set back from the street with 
a driveway and parking space in the rear. Qualified bota- 
nists are cordially invited to visit the Pringle Herbarium 
and to make use of these rich resources. 


THE PRINGLE HERBARIUM 
UNIVERSITY OF VERMONT 
BURLINGTON, VERMONT 05401 


A RANGE EXTENSION AND A NEW SYNONYM 
FOR ASTRAGALUS POLARIS BENTH. 


R. C. BARNEBY 


In a discussion of Astragalus sect. Polares (Rydb.) 
Barneby I have mentioned (1964, p. 370) sect. Vesicularia 
Gontsch. (in Komarov, 1946, p. 51, 877) as a possible 
synonym, but as the sectional typus, A. atlasovi Kom., was 
known to me only from descriptions I could make no posi- 
tive statement. Now through the kindness of Dr. I. T. 
Vassilezenko I have been privileged to borrow an authentic 
example of A. atlasovi: basin of Anadyr’ river, 27. VII. 
1933, fl., fr, Chashevy 1393, LE, perhaps the basis of 
Gontscharov's record (1. c.) of the species from Anadyrskiy 
Kray. The plant matches in fine detail some Alaskan speci- 
mens of A. polaris Benth., and leaves little doubt that 
A. atlasovi (Komarov in Feddes Repert. 13: 229. 1914. — 
“Legi 7. VII. 1909 in peninsulae Kamtshatka valli centrali 
in alpibus Kaschchan non procul a pago Pustshina.") and 
A. polaris, described from plants collected by Berthold 
Seeman in 1849 at Eschscholtz Bay, Alaska (BM!) are 
taxonomic synonyms in a strict sense, although the Siberian 
plant represents the phase of the species with relatively 
narrow, fusiform pods which was segregated in Alaska as 
Homalobus amblyodon Rydb. 

The range of A. polaris, known up to the present only 
from western coastal and interior montane Alaska (Bar- 
neby, 1964, map 39), is now seen to be amphi-Beringian. 
This is, of course, a familiar pattern of dispersal, but has 
particular interest in Astragalus, for A. polaris is the first 
member of the genus with the chromosome number x = 12 
to be recorded from Asia, where the basic number 8 is 
virtually ubiquitous. Asiatic groups have contributed a 
number of species to the astragaline flora of North Amer- 
ica, but traffic in the reverse direction through Beringia 
has not been recognized hitherto. 

Starting from the assumption that the plant from 


304 


1973] Astragalus Polaris — Barneby 305 


Anadyr' is indeed conspecific with Kamtchatkan A. atla- 
sovi, an assumption fully supported by the protologue of 
the latter, the monotypic sect. Vesicularia Gontsch. (1946) 
automatically falls into the synonymy of sect. Polares 
(Rydb.) Barneby, based on Phaca sect Polares Rydb. 
(1929). 


LITERATURE CITED 


BARNEBY, R. C. 1964. Atlas of N. American Astragalus. Mem. 
N. Y. Bot. Gard. vol. 13. 
KoMaRov, V. L. (ed.) 1946. Flora S. S. S. R. vol. 12. 


NEW YORK BOTANICAL GARDEN, 
BRONX, NEW YORK 10458 


THE STATUS OF RAMALINA SUBAMPLIATA 
(NYL.) FINK. IN NORTH AMERICA 


PETER A. BOWLER and PHILLIP W. RUNDEL 


Ramalina subampliata (Nyl.) Fink is a poorly under- 
stood lichen species reported from North America by many 
authors. At least two distinct North American taxa, an 
esorediate and a sorediate form, have been attributed to 
R. subampliata. The esorediate form, discussed by Howe 
(1914) as R. fastigiata var. subampliata, has been reported 
from New Mexico (Bouly de Lesdain, 1942; Egan, 1972), 
although Howe (1914) strangely gave its distribution as 
“uncommon in the Upper Austral and Transition zones 
from New Hampshire to North Dakota south to Louisiana 
and Florida.” We have examined specimens from Arizona 
and New Mexico. The sorediate form has been collected in 
Alberta, Canada. This paper is the result of a study made 
to correctly determine the proper identity of R. subampli- 
ata in North America. 


The new combination made by Fink (1935) of R. sub- 
ampliata from R. calicaris var. subampliata Nyl. included 
an orthographic error, the specific epithet being misspelled 
as subamplicata. Although some subsequent authors have 
corrected this error (Bouly de Lesdain, 1942; Follman and 
Huneck, 1968), most Americans have perpetuated it (Hale 
and Culberson, 1970; Egan, 1972). 


An examination of syntypes in the Nylander Herbarium, 
Botanical Museum, Helsinki, annotated as R. calicaris var. 
subampliata by Nylander himself, revealed seven collec- 
tions. Fragments of each of these collections were analyzed 
chemically by thin-layer chromatography using Merck 
silica gel F-254 TLC plates in a benezene :dioxane : acetic 
acid solvent system (90:25:4 v/v/v), developed with 10% 
H.SO, and heat (Culberson and Kristinsson, 1970). 


hb. Nyl. 37286. France; Lenormand. Unknown acid A. 


306 


1973] Ramalina — Bowler and Rundel 307 


hb. Nyl 37284. Portugal; Welwitsch. Unknown acid A. 
(as forma pyrifera). 

hb. Nyl. 37283. India, Nilgherrensibus; Perrotet. Seki- 
kaic acid, unknown acid B. (as forma subpapillosa). 

hb. Nyl. 37290. India, Northwest Himalaya, Pangi. Col- 
lector unknown; usnic acid. 

hb. Nyl. 37319. Scotland, Barcaldine; Crombie 1880. No 
medullary substances. 

hb. Nyl. 37282. Cersica, Bonifacio; Norrlin 1877. Un- 
known acid C, usnic acid. (annotated R. canariensis 
Steiner by Magnusson). 

hb. Nyl. 37285. France, Pyrenees-Orientales, Forca Real. 
Collector unknown, 1872. Evernic acid (?). 


The problems involved in selecting a lectotype for R. 
subampliata are complicated by the fact that distinct 
morphological differences occur among these seven collec- 
tions. Clearly, more than a single species is included. The 
observed variations in thallus chemistry reinforce this 
view. 

Howe (1914) selected a lectotype for his new combina- 
tion R. fastigiata var. subampliata based on hb Nyl. 37282 
from Corsica. His description and photograph are of this 
collection. Unfortunately, the collection date on this packet 
is 1877, seven years after the original description was pub- 
lished. Of the seven collections listed above, the latter 
three (37319, 37282, 37285) all postdate the original de- 
scription of the taxa, and thus cannot be considered as 
syntypes. 

Nylander’s original description of R. calicaris var. sub- 
ampliata is as follows: 

"thalli laciniis latioribus (latit. circiter 6-12 millim., 
altit. 5-10 centim.) et facie jam Ramalinae fraxineae 
jamve fastigiatae, sporis vero rectis mox distincta. 

Occurrit haecce in Gallia occidentali (etiam ad Parisios), 
in Lusitania (Welwitsch), in-Algeria et in India orientali 
(montibus Nilgherrensibus, Himalaya). 

In Lusitania adest forma sporis saepius pyriformibus 


308 Rhodora [Vol. 75 


(vel quidem infra subcaudatis), quae dici possit pyrifera. 
Forma subpapillosa thallo magis rugoso, marginibus (et 
quoque lateribus) papillis minutis apice saepe albo-sore- 
diellis exasperato, obvenit in India orientali, Nilgherris 
( Perrottet ) ." 

From this description, three of the remaining four col- 
lections appear to be mentioned by Nylander. The Wel- 
witsch (37284) and Perrotet (37283) collections are 
clearly the packets he examined and designated f. pyrifera 
and f. subpapillosa, respectively. The Lenormand collection 
(37286) from “Gallia occidentali" must be considered the 
logical choice as the letcotype for this taxa. Although the 
Welwitsch collection of f. pyrifera is fragmentary, it ap- 
pears to morphologically be similar to the lectotype in its 
cortical and apothecial characteristics as well as being 
chemically identical. The Perrotet collection of f. subpapil- 
losa is both morphologically and chemically distinct. 

The remaining collection on the above list (37290) from 
India is not mentioned in the original description. It is 
both morphologically and chemically distinct from any of 
the three collections mentioned in Nylander’s description. 


The single specimen (Fig. 1) in the Lenormand collec- 
tion (37286) is 7 cm. long, sparsely branched with 
branches up to 0.7 em wide. Other characteristics are as 
follows: rigid, strongly flattened; surfaces indistinguish- 
able with longitudinally oriented furrows in the shiny and 
smooth (though occasionally bumpy), virescent cortex; 
branch tips narowing; no isidia or soredia; medulla white; 
apothecia marginally disposed the length of the branches; 
the disk more or less flattened, pale, up to 4 mm in diame- 
ter. 

The packet of f. subpapillosa (37283) consists of two 
morphologically and chemically similar plants, the largest 
being 6 cm long. Other characteristics are: sparsely 
branching, the branches up to 0.75 cm wide, rigid and 
flattened; surfaces indistinguishable and white-papillate 
with longitudinal and transverse furrows; the branch tips 


1973] Ramalina — Bowler and Rundel 309 


Figure 1. Lectotype of Ramalina subampliata (Nyl.) Fink; scale 
in mm. (Photograph by B. Gassner). 


narrowing; the lacunose surface of cortex dull, apothecia 
marginal and laminal, subterminal, disk pale, up to 3 mm 
in diameter, flat to convex, morphologically and chemically 
it differs distinctly from the leucotype. 

There is no question that North American reports of R. 
subampliata are incorrect. Neither the specimens figured 
by Howe in 1914 (hb. Nyl. 37282) from Corsica nor by 
Fink (1935) from New Mexico are conspecific with the 
lectotype of R. subampliata (hb. Nyl. 37286). It seems 
probable that chemical reports of usnic acid in Himalayan 
specimens of R. subampliata (Follmann and Huneck, 1968; 
Sarma and Huneck, 1968), are based not on this species, 
but on a taxa conspecific with Himalayan material men- 
tioned earlier in this paper (hb. Nyl. 37290). More re- 
search will be necessary to properly assess the taxonomic 
position of North American taxa formerly reported as 
R. subampliata. 


310 Rhodora [Vol. 75 


LITERATURE CITED 


BouLY DE LESDAIN, M. 1942. Lichens de l'Etat de New Mexico 
(U.S.A.) recueilis par le Frere G Arsene Brouard, Rev. Bryol. 
Lichen. 12: 44-66. 

CULBERSON, C. F. 1970. Supplement to “Chemical and Botanical 
Guide to Lichen Products." Bryologist 73: 177-377. 

CULBERSON, C. F. and H. KRISTINSSON. 1970. A standardized 
method for the identification of lichen products. Jour. Chroma- 
togr. 46: 85-93. 

EGAN, R. S. 1972. Catalog of the Lichens of New Mexico. Bryolo- 
gist 75: 7-35. 

Fink, B. 1935. The Lichen Flora of North America. Univ. of 
Mich. Press, Ann Arbor. 

FOLLMAN, G. and S. HuNECK. 1968. Mitteilung uber Flechtenin- 
haltsstoffe. LXI. Zur Chemotaxonomie der Flechtenfamilie 
Ramalinaceae. Willdenowia 5: 181-216. 

HALE, M. E., JR. and W. L. CULBERSON. 1970. A Fourth Checklist 
of the Lichens of the Continental United States and Canada. 
Bryologist 73: 499-543. 

Howe, R. H. 1914. North American species of the genus Ramalina. 
Part IV. Bryologist 17: 17-27. 

NYLANDER, W. 1870. Recognitio Monographica Ramalinarum. Bull. 
Soc. Linn. Normand. II, 4: 108-180. 

SARMA, K. G. and S. HuNECK. 1968. Uber Flechteninhaltsstoffe. 
52. Mitteilung: Uber die Inhaltsstoffe einiger Flechten aus dem 
Himalaja. Die Pharmazie 23: 583-584. 


DEPARTMENT OF POPULATION AND 
ENVIRONMENTAL BIOLOGY, 
UNIVERSITY OF CALIFORNIA, 
IRVINE, CALIFORNIA 92664. 


THE VASCULAR FLORA OF KENT ISLAND, 
GRAND MANAN, NEW BRUNSWICK* : 


JOHN MCCAIN, R. B. PIKE and A. R. HopGDON 


In the Grand Manan cluster of islands at the entrance of 
the Bay of Fundy there are three neighboring isles which 
are the Kent Island group. Kent, the largest, has 150 acres, 
Hay 50 and Sheep Island only 20. These somewhat remote 
islands are set off by themselves as the outermost of the 
Grand Manan Archipelago. Kent Island lies about 15 miles 
east of Cutler, Maine, the nearest point on the mainland. 

Since Kent Island is an important nesting place for 
Herring gulls, Black guillemots, Leach's petrels and other 
pelagic birds it holds great interest for ornithologists and 
has been maintained as a biological station by Bowdoin 
College since 1934. 

During the second and third seasons of the Kent Island 
station, Dr. David Potter of Clark University compiled lists 
of the vascular species of Kent and the two neighboring 
islands, the results of which were published in the first and 
third annual reports of the Bowdoin Scientific Station 
(1936) (1937). Unfortunately these lists did not receive 
wide circulation and only came to the attention of the sec- 
ond and third authors of this paper in 1966 through Dr. 
Philip Sawyer of the Zoology Department at the University 
of New Hampshire. 

When Professor Charles A. Weatherby and John Adams 
published their excellent flora of Grand Manan (1945) 
there was no mention of Potter's lists nor of his collections 
which had been placed in the herbarium at Clark Univer- 
sity. This is not surprising in view of the obscure publica- 
tion of the lists, but it is much more difficult to explain the 


*This paper is published with the approval of the Director of the 
Bowdoin Scientific Station as Contribution No. 40. 

‘Published with the approval of the Director of the New Hamp- 
shire Agricultural Experiment Station as Scientific Contribution 
No. 668. 


311 


312 Rhodora [Vol. 75 


failure of communication between Professor Weatherby 
and Dr. Potter who knew each other and who must have 
met often at the Gray Herbarium and at meetings of the 
New England Botanical Club. Furthermore, Weatherby 
and Adams’ paper made eighteen specific citations of plants 
growing on Kent or Hay Island. 


In 1970 Pike and Hodgdon with Potter's lists in hand 
studied his collections at Clark University. Funds had been 
granted in support of this study and a subsequent field 
survey of the Kent Island flora. At about the same time 
they received, as editors of Rhodora, a letter from McCain, 
expressing interest in submitting a paper on the flora and 
vegetation of Kent Island. The present paper brings to- 
gether all contributions to our knowledge of the Kent 
Island flora. 

The first author (McCain) made extensive collections in 
1968 and 1969 on Kent, Hay and Sheep Islands. His speci- 
mens are now deposited in the herbaria of the University 
of New Hampshire and Albion College, Albion, Michigan. 
The second and third authors visited Kent Island for two 
days in 1970 and were able to add a few additional taxa in 
groups of special interest to them. Their collections also 
have been placed in the herbarium of the University of 
New Hampshire. 

Each of the four independent studies made of Kent and 
associated islands has added appreciably to the flora. The 
work of the senior author in 1968-69 resulted in 61 addi- 
tions, and the most recent visit by Pike and Hodgdon sup- 
plied six more from Kent Island alone. It is doubtful, how- 
ever, that further gleaning would be as profitable. 


The composite list of species of vascular plants from 
Kent, Hay and Sheep now includes 142 genera with 240 
species and 10 additional varieties representing 54 families. 
Any taxa not found on Kent Island are indicated in the 
following list as coming from Hay or Sheep, thus permit- 
ting an assessment of the tota] vascular flora of Kent Island 
itself. It should be noted also that twenty-eight species in 


1973] Kent Island — McCain, Pike & Hodgdor 313 


the flora have not been previously listed for Grand Manan. 
Most of these species are either halophytic or boreal. 

The nomenclature is based essentially on Seymour 
(1969), the most up-to-date major treatment of vascular 
plants of the general region. However, the authors prefer 
not to follow Seymour in splitting off other genera from 
Pyrus and Vaccinium as these groups are conservatively 
understood. The blueberries and the Rosaceae, therefore, 
are listed according to the 8th Edition of Gray’s Manual. 

We wish to thank Dr. R. F. Nunnemacher for his special 
kindnesses to Pike and Hodgdon on the occasion of two 
pleasant visits to Clark University. All of us also were 
granted the utmost in hospitality by Dr. Charles E. Hunt- 
ington, the director of the Bowdoin Science Station on Kent 
Island, during the course of our work. A grant from the 
University of New Hampshire Central University Research 
Fund (cuRF 391) supported travel to Clark University and 
to Kent Island by Pike and Hodgdon. McCain's work on 
Kent Island was supported by National Science Foundation 
Undergraduate Research Participation Grants Gy-4343 in 
1968 and Gy-6095 in 1969. He acknowledges the taxonomic 
advice of Dr. E. A. Stowell of Albion College and Dr. E. G. 
Voss and Dr. W. H. Wagner of the University of Michigan, 
and the support and encouragement of Dr. Clara L. Dixon 
of Albion College. 


List of vascular plants found growing on Kent Island. 


EQUISETACEAE 
Equisetum arvense L. var. boreale (Bong.) Ledeb. 
E. sylvaticum L. var. pauciramosum Milde. 
LYCOPODIACEAE 
Lycopodium annotinum L. var. acrifolium Fern. 
L. obscurum L. forma dendroideum (Michx.) Blom- 
quist & Correll. (Hay Island only) 
OSMUNDACEAE 
Osmunda cinnamomea L. 
POLYPODIACEAE 
Onoclea sensibilis L. (Hay Island only) 


314 Rhodora | [Vol. 75 


* Thelypteris palustris (Salisb.) Schott var. pubescens 
(Lawson) Fern. 
Dryopteris spinulosa (O. F. Mueller) Watt. var. ameri- 
cana, (Fischer) Fern. 
D. cristata (L.) Gray 
* Dennstaedtia punctilobula (Michx.) Moore 
Athyrium filix-femina (L.) Roth. var. michauxii 
(Sprengel) Farw. forma rubellum (Gilbert) 
Farw. 
PINACEAE 
Abies balsamea (L.) Miller 
Picea glauca (Moench.) Voss. 
P. rubens Sarg. 
P. mariana (Miller) BSP. 
Larix laricina (DuRoi) K. Koch 
* Juniperus communis L. var. depressa Pursh. 
TYPHACEAE 
* Typha latifolia L. 
» SPARGANIACEAE 
* Sparganium americanum Nutt. 
ZOSTERACEAE 
Zostera marina L. var. stenophylla Ascherson & 
Graebner. 
J UNCAGINACEAE 
Triglochin maritimum L. 
T. palustre L. 
GRAMINEAE 
Festuca rubra L. var. rubra 
Puccinellia paupercula (Holm.) Fern. & Weath. var. 
alaskana (Scribner & Merr.) Fern. & Weath. 
Poa annua L. 
* P. compressa L. 
P. pratensis L. 
Poa palustris L. 
Agropyron repens (L.) Beauv. forma aristatum 
(Schum.) Holmb. | 
A. repens (L.) Beauv. forma trichorrachis Rohlena. 
A. repens (L.) Beauv. forma pilosum (Scribner) Fern. 
Hordeum jubatum L. 


1973] Kent Island — McCain, Pike & Hodgdon 315 


Elymus arenarius L. var. villosus Meyer 
* E. virginicus L. 
Deschampsia flexuosa (L.) Trin. 
Danthonia spicata (L.) Beauv. 
Calamagrostis canadensis (Michx.) Nutt. var. cana- 
densis 
Ammophila breviligulata Fern. 
* Agrostis stolonifera L. var. major (Gaud.) Farw. 
A. stolonifera L. var. palustris (Hudson) Farw. 
A. tenuis Sibth, forma aristata (Sinclair) Wiegand 
* A. scabra Willd. 
* A. perennans (Walter) Tuck. 
* Cinna arundinacea L. 
+ C. latifolia (Trev.) Griseb. 
Phleum pratense L. 
Alopecurus pratensis L. 
Spartina alternifiora Loisel. 
Anthoxanthum odoratum L. 
CY PERCEAE 
* Eleocharis smallii Britton var. major (Sonder) Sey- 
mour 
E. halophila Fern. & Brackett. 
E. elliptica Kunth. 
Scirpus americanus Pers. 
* S. atrovirens Willd. 
* S. cyperinus (L.) Kunth. forma pelius Fern. 
S. atrocinctus Fern. (Hay Island only) 
Eriophorum spissum Fern. 
E. angustifolium Honckeny 
E. virginicum L. 
Carex stipata Muhl. (Hay Island) 
trisperma Dewey 
canescens L. var. disjuncta Fern. 
canescens L. var. subloliacea Laestad. 
brunnescens (Pers.) Poiret. var. sphaerostachya 
(Tuck.) Kukenthal. 
cephalantha (Bailey) Bickn. 
tribuloides Wahlenb. 
hormathodes Fern. 


onn agana 


318 Rhodora [Vol. 75 


RANUNCULACEAE 
Ranunculus cymbalaria Pursh. 
R. acris L. 
Thalictrum polygamum Muhl. (Hay Island only) 
Coptis groenlandica (Oeder) Fern. 
CRUCIFERAE 
Capsella bursa-pastoris (L.) Medicus 
Cakile edentula (Bigelow) Hooker 
Rorippa islandica (Oeder) Borbas var. hispida (Desv.) 
Butters & Abbe. 
* R. islandica (Oeder) Borbas var. fernaldiana Butters 
& Abbe. 
SARRACENIACEAE 
Sarracenia purpurea L. 
DROSERACEAE 
Drosera rotundifolia L. 
CRASSULACEAE 
* Sedum telephium L. forma purpureum (L.) Seymour 
S. rosea (L.) Scop. 
SAXIFRAGACEAE 
Ribes hirtellum Michx. 
R. lacustre (Pers.) Poiret. 
R. glandulosum Grauer. 
ROSACEAE 
Spiraea latifolia (Ait.) Borkh. 
Pyrus malus L. 
Pyrus floribunda Lindl. 
Pyrus americana (Marsh) DC. 
Pyrus decora (Sarg.) Hyland 
Fragaria virginiana Duchesne 
Potentilla norvegica L. 
Potentilla anserina L. 
Filipendula ulmaria (L.) Maxim. (Hay Island only) 
Rubus chamaemorus L. 
Rubus idaeus L. var. canadensis Richardson 
Rubus hispidus L. 
Rosa rugosa Thunb. 
LEGUMINOSAE 
Trifolium pratense L. var. sativum (Miller) Schreber 
(Hay Island) 


1973] Kent Island — McCain, Pike & Hodgdon 319 


T. repens L. 

T. agrarium L. (Hay Island only) 

Vicia cracca L. (Hay Island only) 

Lathyrus japonicus Willd. var. pellitus Fern. 
OXALIDACEAE 

Oxalis montana Raf. 
CALLITRICHACEAE 

Callitriche verna L. (Hay Island only) 
EMPETRACEAE 

Empetrum nigrum L. 
BALSAMINACEAE 

Impatiens capensis Meerb. forma immaculata. (Weath.) 

Fern. & Schub. 

GUTTIFERAE 

Hypericum virginicum L. var. fraseri (Spach.) Fern. 
VIOLACEAE 

Viola cucullata Aiton 

* V. septentrionalis Greene 

V. pallens (Banks) Brainerd 
ONAGRACEAE 

Epilobium angustifolium L. 

* E. palustre L. 
E. glandulosum Lehm. var. adenocaulon (Haussk.) 
Fern. 

Oenothera perennis L. 

Circaea alpina L. 
HIPPURIDACEAE 

Hippuris vulgaris L. (Hay Island only) 
ARALIACEAE 

Aralia hispida Vent. 
UMBELLIFERAE 

Carum carvi L. 

Ligusticum scothicum L. 

Coelopleurum lucidum (L.) Fern. 

Heracleum maximum Bartr. (Hay Island only) 
CORNACEAE 

Cornus canadensis L. 
PYROLACEAE 

Moneses uniflora (L.) Gray 

* Monotropa uniflora L. 


320 Rhodora [Vol. 75 


+ Monotropa hypopithys L. 
ERICACEAE 
Ledum groenlandicum Oeder 
* Rhododendron canadense (L.) Torrey 
Kalmia angustifolia L. 
* Cassandra calyculata (L.) D. Don. var. angustifolia 
(Aiton) Seymour 
* Chiogenes hispidula (L.) T. & G. 
Vaccinium angustifolium Aiton. var. laevifolium House 
V. vitis-idaea L. var. minus Lodd. 
V. oxycoccus L. 
ti V. oxycoccus L. var. ovalifolium Michx. 
PRIMULACEAE 
Lysimachia terrestris (L.) BSP. 
Trientalis borealis Raf. 
Glaux maritima L. 
PLUMBAGINACEAE 
Limonium nashii Small 
GENTIANACEAE 
Menyanthes trifoliata L. var. minor Raf. 
CONVOLVULACEAE 
Convolvulus sepium L. 
Convolvulus sepium L. forma malachophyllus Fern. 
BORAGINACEAE 
Mertensia maritima S.F. Gray 
LABIATAE 
Scutellaria epilobiifolia A. Hamilton 
Galeopsis tetrahit L. var. bifida (Boenn.) Lej. & Court. 
* Lycopus virginicus L. 
Lycopus uniflorus Michx. 
SCROPHULARIACEAE 
Euphrasia randii B. L. Robinson (Sheep Island only) 
Euphrasia americana Wettst. 
Rhinanthus crista-galli L. var. fallax (Wimmer & 
Grab.) Druce 
PLANTAGINACEAE 
* Plantago major L. 
Plantago juncoides Lam. var. decipiens (Barneoud) 
Fern. 


1973] Kent Island — McCain, Pike & Hodgdon 321 


* Plantago juncoides Lam. var. glauca (Hornem.) Fern. 
+ Plantago oliganthos R. & S. 
RUBIACEAE 
Galium trifidum L. 
CAPRIFOLIACEAE 
Lonicera canadensis Bartram 
Linnaea borealis L. var. americana (Forbes) Rehder. 
COMPOSITAE 
* Solidago sempervirens L. 
Solidago rugosa Miller var. villosa (Pursh.) Fern. 
Solidago graminifolia (L.) Salisb. var. nuttallii 
(Greene) Fern. 
t Solidago macrophylla Pursh. 
Aster foliaceus L. var. arcuans Fern. 
* Aster johannensis Fern. 
Aster acuminatus Michx. 

* Aster nemoralis Aiton 

t Aster X Blakei (Porter) House 
Aster umbellatus Miller 
Anaphalis margaritacea (L.) B.&H. var. subalpina 

Gray. 
* Anaphalis margaritacea (L.) B.&H. var. intercedens 
Hara. 

* Gnaphalium uliginosum L. 

Ambrosia artimisiifolia L. var. elatior (L.) Descortils 

* Bidens cernua L. 

Bidens frondosa L. 
Achillea millefolium L. 
i Matricaria maritima L. var. agrestis (Knaf.) Wilmott. 
Matricaria matricarioides (Less.) Porter 
Chrysanthemum leucanthemum L. var. pinnatifidum 
Lecoq. & Lamotte 

Tanacetum vulgare L. (Hay Island only) 

t Senecio vulgaris L. 

Senecio sylvaticus L. 

Cirsium vulgare (Savi) Tenore 

Cirsium arvense (L.) Scop. 

Leontodon autumnalis L. var. pratensis (Link.) W. D. 
J. Koch 


322 Rhodora [Vol. 75 


Taraxacum officinale Weber 

Sonchus asper (L.) Hill. 

Hieracium aurantiacum L. 

Hieracium floribundum Wimm. & Grab. 
Hieracium pratense Tausch. 

Hieracium pilosella L. 


STATISTICAL SUMMARY 
54 Families 
142 Genera 
240 Species (or 239 spp. and 1 hybrid) 
10 Varieties 


Unless otherwise noted, species were collected and iden- 
tified by John McCain in 1968 and 1969. 
* Species collected by Dr. David Potter 1935-1936. 
T Species listed by C. A. Weatherby and J. Adams, 1945. 
i Species collected by Dr. Albion Hodgdon and Radcliffe 
Pike in 1970. 


LITERATURE CITED 


PoTTER, DAvip. 1936. List of Kent's Island Flowering Plants 1935. 
Bull. Bowdoin Sci. Sta. 1: 10-12. 

. 1937. Plants found growing on Kent’s Island 1935- 
1936. Bull. Bowdoin Sci. Sta. 3: 19-26. 

WEATHERBY, C. A. and J. ApAMs. 1945. A list of the vascular 
plants of Grand Manan, Charlotte County, New Brunswick, 
Canada. Contrib. Gray Herbarium 158: 96 pp. 


DEPARTMENT OF BIOLOGY 
ALBION COLLEGE 
ALBION, MICHIGAN 49224 


DEPARTMENTS OF PLANT SCIENCE 
AND BOTANY 

UNIVERSITY OF NEW HAMPSHIRE 
DURHAM, N. H. 03824 


Correction in Book Review, Flora of the Galápagos Islands 


The meaning of the sentence on lines 14-17 on page 144 
of Rhodora for March 1973 is quite different from that 
intended. The sentence should read as follows: The 
presence of the endemic Miconia and Cyathea would seem 
to depend on reasonably moist conditions having obtained 
for some time in the areas where they are now growing 
or in places not far away. 

Librarians and others who are interested in this review 
are advised to substitute the word now for not. 


— A. R. HODGDON 


323 


324 Rhodora [Vol. 75. 


INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA 


Manuscripts must be double-spaced or preferably triple- 
spaced (not on corrasable 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 indicate the style of type through the use of capitals 
or underscoring, partieularly in the citations of specimens, 
except that the names of species and genera may be under- 
lined 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 System- 
atics and Ecology of Poison-Ivy and the Poison-Oaks," 
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu- 
larly with reference to the indentation of keys and syno- 
nyms. Papers of a floristic nature should follow, as far as 
possible, the format of “Contribution to the Fungus Flora 
of Northeastern North America. V.,” H. E. Bigelow & M. E. 
Barr, Rhodora 71: 177-203. 1969. For bibliographic cita- 
tions, a recommended list of standard journal abbreviations 
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. 
Club 85: 277-300. 1958. 


Volume 75, No. 802, including pages 169-324, was issued June 29, 1973. 


CONTENTS: — continued 


Palynological Evidence for the Late Glacial Occurrence of 
Pringlea and Lyallia on Kerguelen Islands 
Steven B. Young and Eileen K. Schofield ........................ 239 


Phytogeography of the Carices of Virginia 
A. M. Harvill, Tre ciccccccccccccccsssscssssccscssesesssesvssasesssecssssesseseesseess 248 


The Marine Algae of Virginia and Maryland Including the 
Chesapeake Bay Area 
Franklyn D. Ott ................. J... UU... a. teta nana 258 


The Pringle Herbarium 
Frank C. Seymowr ................................................................. 297 


A Range Extension and a New Synonym for Astragalus 
polaris Benth. 
B. C. HOPED Leere rere enn reno enne menn enii std 304 


The Status of Ramalina subampliata (Nyl) Fink in North 
America 
Peter A. Bowler and Phillip W. Rundel ............................ 306 


The Vascular Flora of Kent Island, Grand Manan, New 
Brunswick 
John McCain, R. B. Pike and A. F. Hodgdon ................ 311 


Correction in Book Review, Flora of the Galápagos Islands 
A. R. Hodgdom wisccccccccccsssccsssececssssccssseccseseseseeeecseeeessseeeseeeasesseees 323 


Instructions for Contributors ................. eere eene 324 


Dovdova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
ALBION REED HODGDON, Editor-in-Chief 


^ 


ROLLA MILTON TRYON 
RADCLIFFE BARNES PIKE 
STEPHEN ALAN SPONGBERG ' Associate Editors 
GERALD JOSEPH GASTONY 
ALFRED LINN BOGLE 


Vol. 75 September, 1973 No. 803 
CONTENTS: 


Biosystematic Study of Chamaesaracha (Solanaceae) 
PAVE S Aapa K, 1A Lau n n E E s 825 


Some Notes on the Flora of the Southern States, Particu- 
larly Alabama and Middle Tennessee 
ZR PO A E supricbóoutidistezt Dad s UD UNEFE el UP ETE DEI: 366 


Additions to the Bahama Flora since Britton and Mills- 
paugh 
William T. Gillis, Richard A. Howard, and George R. 
PEIUS S damit ez m ua E 411 


Revision of the Genus Pontederia L. 
Richard M. Lowden u u] u ua s 0 426 


(Continued on Inside Back Cover) 


Che New England Botanical Club, Ine. 
Botanical Museum, Oxford St., Cambridge, Mass. 02138 


RHODORA. — A quarterly journal of botany, devoted primarily to the 
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Rhodora 


JOURNAL OF THE 
NEW ENGLAND BOTANICAL CLUB 


Vol. 75 September, 1973 No. 803 


BIOSYSTEMATIC STUDY OF 
CHAMAESARACHA (SOLANACEAE): 


JOHN E. AVERETT? 


Chamaesaracha was established as a genus by A. Gray 
in 1876, having previously been recognized as a section 
of Saracha. The genus has been treated in part by Gray 
(1878) and by Rydberg (1896). However, an inclusive 
revision of Chamaesaracha has not been forthcoming. 

As herein conceived, Chamaesaracha is an exclusively 
North American genus with its greatest concentration of 
species in the Chihuahuan desert. A total of 18 species 
has been attributed to the genus, but only 7 are recognized 
in the present treatment. The remaining 11 have been 
relegated to synonymy or have been transferred to other 
genera (see excluded species). 


CHROMOSOME STUDIES 


Meiotic chromosomes were observed best at anaphase I, 
because of clumping and poor staining of the chromosomes 
in diakinesis and metaphase I. The latter is particularly 
notable in polyploid cells. Except for several triploid indi- 
viduals, meiosis appeared to be regular with equal separa- 


"Part of a doctoral dissertation presented to the faculty of the 
Graduate School of The University of Texas at Austin. 

"Department of Biology, University of Missouri — St. Louis 63121 
and Missouri Botanical Garden. 


325 


326 Rhodora [Vol. 75 


tion of chromosomes at anaphase I. All chromosome num- 
bers reported here are from meiotic material. 

Several of the earlier chromosome number reports for 
Chamaesaracha (Powell and Averett, 1967) require clarifi- 
cation in keeping with the results of my present systematic 
account of the genus. In the 1967 paper, counts were listed 
for the following: C. coronopus (n = 24); and C. cf. 
sordida (n = 12, 18, 36). With the exception of C. sordida 
and C. cf. sordida, all of the 1967 reports were applied to 
taxa that were correctly identified. (C. cf. crenata is C. 
crenata). The following corrections must be noted in 
regard to the 1967 reports attributed to C. sordida and 
C. cf. sordida: The n = 12 and n = 18 collections identi- 
fied as C. cf. sordida are now recognized as C. sordida. 
The » — 24 populations referred to C. sordida are now 
regarded as C. coniodes, a species then regarded as con- 
specific with C. sordida. Chamaesaracha cf. sordida (n = 
36), except for Averett 155, is now recognized as C. coro- 
nopus. Collection 155 is C. pallida, a newly described 
taxon (Averett, 1971). Listed in Table 1 are additional 
counts for the taxa first reported in 1967, as well as reports 
for the remaining species of Chamaesaracha. 

Polyploidy is found in five of the seven species of Cha- 
maesaracha, four being typically polyploid. Although there 
are areas lacking convincing data some comment may be 
made regarding the origin of selected polyploids, Because 
each is somewhat unique, they are considered individually 
in the following discussion. 


Chamaesaracha sordida. There is evidence for an auto- 
polyploid origin of the triploid (n = 18) individuals in 
C. sordida. In a regional study of the taxon, a triploid plant 
was collected from a population in northern Culberson 
County, Texas. Meiosis in this individual was irregular. 
Diakinetic and metaphase chromosomes were not clearly 
observed, but unequal segregation of 36 chromosomes could 
be observed in anaphase I. From 14 to 22 chromosomes 
could be found going to one side of the diad in about 100 
cells, but 18 choromsomes could be observed in one side 


1973] Chamaesaracha — Averett 327 


or the other of an equivalent number of cells, The irregu- 
larity was first attributed to hybridization between a 
diploid and a tetraploid. However, subsequent resampling 
of the area from which the triploid individual was collected 
indicated that only diploid individuals occurred in the 
population concerned. (Chromosome numbers were ascer- 
tained for about 15 individuals from the general area; all 
were diploid with n = 12). 

Two additional triploid individuals have been found in 
widely disjunct populations of the same species. These 
have not been carefully studied, but in one population in 
Yoakum County, Texas, chromosome numbers from three 
individuals were determined (Averett & Tomb 320A, B, C). 
One of these was triploid and the other two were diploid. 
The only other triploid found in this species came from a 
population in Lincoln County, New Mexico. 

The morphology and flavonoid chemistry of the triploids 
referred to above are nearly identical to that of the diploids. 
Considering the sporadic occurrence of the triploids, the 
plants do not seem to have a common origin nor do they 
appear to be the result of the interspecific hybridization. 
Rather, each appears to have had an independent origin 
through the chance fertilization of an unreduced gamete 
with a normal haploid gamete. The additional set of 
chromosomes would account for the observed unequal 
chromosomal segregation found in the triploid populations. 


Chamaesaracha coronopus. This species is typically 
tetraploid with n — 24, but occasional diploids (n — 12) 
and hexaploids (n — 36) are found in western and south- 
western Texas and also in southeastern Arizona, Unlike 
the triploids of C. sordida, the hexaploids are quite fertile. 
Meiosis appears to be regular, at least in anaphase I, and 
unequal segregation has not been observed. Additionally, 
seed set and germination is approximately equivalent to 
that of the tetraploids. Because of their apparent fertility, 
an autoploid origin such as proposed in C. sordida does 
not seem likely. However, as in the triploid C. sordida, 
there is no evidence that another taxon has contributed 


328 Rhodora [Vol. 75 


to the genome of the hexaploids in question, even though 
C. sordida (n = 12) does occur sympatrically in some 
areas with C. coronopus. Rather, the hexaploids concerned 
appear to be derived from hybridization between the 
diploid and tetraploid races of C. coronopus. Chemical 
and morphological data are currently being accumulated 
and preliminary results appear to confirm this suggestion. 

Evidence is not available to suggest the origin of the 
tetraploid race of C. coronopus and the remaining poly- 
ploid species of Chamaesaracha. However, two species, 
C. pallida (n = 36) and C. edwardsiana (n = 24) are 
believed to be closely related to, and derived from, C. coro- 
nopus or its ancestors. Chamaesaracha coniodes is not 
particularly close to any of the former species and is likely 
independently derived. Autoploidy or segmental alloploidy 
may account for the polyploidy in both C. coniodes and 
C. coronopus n = 24. 


Table I. Chromosome numbers and number of plants 
counted of Chamaesaracha.’ 


Taxon Chromosome Number of plants 
number (n) counted 
C. coronopus* 12 3 
C. coronopus 24 26 
C. coronopus 36 5 
C. coniodes 24 68 
C. crenata 12 8 
C. edwardsiana 24 11 
C. pallida 36 6 
C. sordida 12 53 
C. sordida 18 3 
C. villosa 12 8 


*Data from Powell & Averett (1967) and Averett (1970). 
*Not previously reported. Discussion & localities in text. 


FLAVONOID STUDIES 


Populations were sampled for flavonoid studies so as 


1973] Chamaesaracha — Averett 329 


to represent as much of the total range, as well as intra- 
specifie variability, of each of the species as time and 
opportunity permitted. One to ten individuals from the 
various populations were analyzed to determine the extent 
and significance of inter- and intrapopulational variation. 
Voucher specimens for the populations studied are de- 
posited in the Herbarium of the University of Texas at 
Austin. Duplicate specimens will be distribued at a later 
date. ! 


Flavonoids were extracted overnight from leaf material 
with 85% methanol and separated by two-dimensional 
paper chromatography. Solvent systems of t-butanol, 
glacial acetic acid, and water (3:1:1 v/v) and 15% glacial 
acetic acid were employed. The compounds detected were 
then characterized structurally by ultraviolet spectroscopy 
and appropriate acid and enzyme hydrolyses. The tech- 
niques utilized for chromatographic and spectral analysis 
follow those presented by Mabry, Markham and Thomas 
(1971). 

All of the compounds extracted from Chamaesaracha 
are based on either quercetin or kaempferol skeletons. 
Differences among the several compounds primarily reflect 
glycosidic substitutions, but a few methoxylated compounds 
also are present. Except for the methoxylated compounds, 
the flavonoids of Chamaesaracha are the same as those 
reported for Leucophysalis by Averett and Mabry (1971), 
but greater interspecific variability is found in Chamae- 
saracha. 


Flavonoid data, in general, were found to substantiate 
conclusions based upon morphological data. Two-dimen- 
sional chromatographic patterns were, for the most part, 
species-specific, showing relatively little variation, either 
within or between populations of the same species. A 
notable exception was found in Chamaesaracha coniodes 
in which significant variations in the flavonoid patterns 
were observed. This species is also quite variable morpho- 
logically and much of the flavonoid variation probably 
reflects genetic heterogeneity from population to popula- 


330 Rhodora [Vol. 75 


tion. Within the more discrete populations, however, 
flavonoid patterns were essentially uniform (this is also 
true of morphological variation). 

Closely related taxa were found to have similar flavo- 
noids; consequently, chemical data have been utilized for 
many of the taxonomic conclusions reached in the present 
study, especially in situations where morphological and 
chromosomal distinctions were not decisive. 


BIOECOLOGY 


Chamaesaracha, a closely knit genus of perennial herbs, 
is largely restricted to the desert regions of the south- 
western United States and northern Mexico. The plants 
are fairly common along roadsides in early and later sum- 
mer, but populations become depauperate in the hot, dry, 
midsummer period, being found with some difficulty, except 
in areas of accumulated moisture. 

The plants often occur in closely clustered populations of 
25.30 plants. Such populations likely originated from one 
or a few seeds, and are believed to be largely the result 
of vegetative reproduction from spreading rhizomes. In 
garden plots, individual plants form colonies 8-10 feet in 
diameter within a single growing season. Intrapopulational 
variation, both chemical and morphological, within these 
clusters is essentially absent, suggesting a single source 
for most of the individuals within a given population. 
Vegetative reproduction, along with perenniality of the 
rootstocks, must have much to do with the variation pat- 
terns, or lack of them, in the natural populations. 

The role of vegetative reproduction and perennial habit 
becomes even more apparent when seed germination is 
considered. Although seed set is more than adequate, the 
percentage of seed germination is very low. In germina- 
tion experiments, seeds were allowed to imbibe for about 
seven days. After imbibition, various exposures to light 
and dark were given, but the latter seemed to have little 
effect on germination. Typically fewer than 5% of the 
seeds germinated. Germination in diploid species was con- 


1973] Chamaesaracha — Averett 331 


siderably lower than in polyploid species. It is presumed 
that seed germination is equally poor in nature, though this 
may not be a valid assumption. 

Since fruit with relatively high seed set are found in 
nature, the various insects that visit the flowers must be 
fairly effective pollinators. No consistent pollen vectors 
have been observed, but bee flies and several types of small 
beetles are commonly found on the flowers of Chamaesara- 
cha. Evidence for self-sterility has not been obtained for 
any of the species, but in isolated plants in garden plots, 
fruit was observed only on C. crenata. Even in this spe- 
cies, however, seed set was reduced some 80% from that 
found in nature. In the field, probably both self and out 
pollination occur, with outcrossing predominating. 


GENERIC RELATIONSHIPS 


Chamaesaracha is included in the tribe Solaneae (Wett- 
stein, 1895) and has generally been associated with Phy- 
salis and Leucophysalis, largely because of their similar 
corolla forms. It is probably most closely related to Leu- 
cophysalis, differing in characters of the berry; whereas 
Physalis is separated primarily by its inflated fruiting 
calyx, and to some extent by its flower. The Asian genus 
Physaliastrum is related to Chamaesaracha in much the 
same way that the latter is related to Leucophysalis. Phy- 
saliastrum is, however, believed to be more closely related 
to Leucophysalis; indeed, they may well be congenic (Ave- 
rett, 1971). All of the species that now comprise Leu- 
cophysalis and Physaliastrum have, at one time or another, 
been relegated to Chamaesaracha. In fact, workers are 
not generally in accord as to the preferred treatment of 
these several genera of Solaneae. For example, Waterfall 
(1958) suggested that Physalis and Chamaesaracha might 
be combined and treated as subgeneric categories. If these 
were the only two genera involved, the merger would not 
be unreasonable. However, under such a treatment, the 
status of Margaranthus would be affected since it is hardly 
more distinct from Physalis than is Chamaesaracha, differ- 


332 Rhodora [Vol. 75 


ing only in the color and urceolate form of its corolla. 
While the sinking of Margaranthus with Physalis may also 
be reasonable, it is difficult, at least for me, to accept the 
merger of Chamaesaracha and Margaranthus. Other gen- 
era are interrelated in a similar manner, and for that 
reason it seems worthwhile to consider them in more detail. 

Leucophysalis was established as a monotypic genus by 
Rydberg (1896) to accommodate L. grandiflora, formerly 
included in Physalis by Hooker (1928). Fernald (1949) 
compared L. grandiflora with certain Asian species of what 
he considered to be Chamaesaracha, and as a consequence 
transferred L. grandiflora to the latter genus. However, I 
have recognized Leucophysalis as distinct from Chamae- 
saracha, and have added an additional two species (Ave- 
rett, 1971). The distinctive morphological features of the 
two taxa are comparable to those that differentiate most 
of the other genera of the Solaneae. Leucophysalis is sim- 
ilar to Chamaesaracha with respect to characters of the 
flower and the fruiting calyx, but differs in having a fleshy 
berry with a placenta extending the entire length of the 
berry. In Chamaesaracha the berry is dry, and the placenta 
is mostly basal (not extending the entire length of the 
berry). The species of Chamaesaracha are extremely uni- 
form in floral, fruit, and seed morphology and as such, 
departure of the fruiting character seems significant. In 
addition, the two taxa have quite distinct habitat prefer- 
ences and occupy different geographical regions. In con- 
trast to the arid desert habitat of Chamaesaracha, Leuco- 
physalis is found in more mesic habitats, e.g., the Sierra 
Nevada of California and the Great Lakes Region. 

Physaliastrum is a small Asian genus of two species 
distinguished from Leucophysalis by the form of its coro!la 
lobes, which are not plicate (as they are in Leucophysalis). 
The berry is like that of Leucophysalis, further differenti- 
ating the genus from Chamaesaracha. 

Physalis differs from Chamaesaracha, as well as from 
Leucophysalis and Physaliastrum, in having a ribbed, in- 
flated fruiting calyx. The only genus with a structure of 
this type is Margaranthus, which, as mentioned above, 


1973] Chamaesaracha — Averett 333 


differs from Physalis in corolla shape. Physalis may be 
differentiated from Chamaesaracha, at least to some extent, 
on characters of the perianth. The corolla of Physalis is 
campanulate, and the flowering calyx is truncate. How- 
ever, the floral morphology of Chamaesaracha is at least 
approached in a few species of Physalis (e.g., P. lobata 
and P. wrightii). Physalis is probably more closely related 
to Margaranthus on the basis of the fruiting calyx, and on 
other characters it is more closely related to Leucophysalis 
and Physaliastrum than it is to Chamaesaracha. 


Nicandra is native of Peru, but is widly naturalized in 
the United States being found in wastelands and around 
old gardens. Nicandra is closely related to and often mis- 
taken for Physalis, but differs in having 5 carpels and dis- 
tinct, sagittate sepals. 


Saracha is primarily a tropical genus of about 30 species 
extending from Peru through Central America north into 
northern Mexico and the southwestern United States.* 
Chamaesaracha and Saracha were once considered con- 
generic, but the two genera may be distinguished on fruit- 
ing characters. In Saracha the fruiting calyx is rotate and 
expanding at the base of a fleshy berry; in Chamaesaracha 
the fruiting calyx is tightly investing a dry berry. Saracha 
is probably not as closely related to Chamaesaracha as the 
genera mentioned above. 


Oryctes is a monotypic genus of western Nevada and 
southeastern California, resembling Chamaesaracha in 
habit and somewhat in fruit. It is, however, easily dis- 
tinguished by its small tubular flower and flat, orbicular 
seeds. 

The few remaining genera of the Solaneae are not con- 
sidered here since they are not so closely related to, and 
rarely confused with, Chamaesaracha. 


*Recent studies by W. G. D’Arcy and Johnny Gentry indicate that 
the Mexican and Central American species are properly treated as 
Bellinia R. & S. Earlier suggestions of affinity between Saracha and 
Chamaesaracha were likely based upon these species. 


334 Rhodora [Vol. 75 


KEY TO CHAMAESARACHA AND RELATED GENERA 


Flowers tubular, seeds orbicular ............... Oryctes 
Flowers campanulate to repand-rotate or urceolate, seed 
reniform, 
Fruiting calyx expanding under and not enclosing 
the berry ................................. Saracha 
Fruiting calyx loosely or tightly investing the berry ... 
Calyx inflated around the berry. 


Flowers urceolate ................ Margaranthus 
Flowers campanulate to repand-rotate. 
2 carpelate, calyx of united sepals ...... Physalis 
5 carpelate, calyx of distinct, 
sagittate-based sepals ............... Nicandra 


Calyx tightly investing the berry. 
Berry dry, placentation mostly basal, seeds 


rugose-reticulate ................ Chamaesaracha 

Berry fleshy, placentation axile, seeds punctate. 
Flowers repand-rotate .......... Leucophysalis 
Flowers campanulate ........... Physaliastrum 


PHYLOGENETIC CONSIDERATIONS 


Chamaesaracha is largely restricted to the arid regions 
of the southwestern United States and northern Mexico. 
What are believed to be the more primitive, diploid species, 
C. villosa, C. crenata and C. sordida, have their centers of 
distribution in the Chihuahuan desert of northern Mexico. 
Of these, C. sordida is the only species to have any appre- 
ciable distribution in the United States. Still, it has a 
distinctly southern distribution as compared to the poly- 
ploid species, C. coronopus and C. coniodes. Chamaesaracha 
coronopus is the most widespread species, extending essen- 
tially over the entire range of the genus ; C. coniodes is also 
widespread, but does not extend so far west as C. corono- 
pus. The remaining polyploid species are fairly restricted 
in distribution, with C. edwardsiana occurring primarily on 
the Edwards Plateau of south-central Texas and adjacent 
Mexico and C. pallida extending from the Edwards Plateau 
into western Texas and northern Mexico. 


1973] Chamaesaracha — Averett 335 


From the distributional data, it seems reasonable to sug- 
gest that Chamaesaracha had its origin in the highlands of 
northern Mexico, the ancestral species of which probably 
belonged to the more xeric members of the Madro-Tertiary 
Geoflora (cf. Axelrod, 1958). With continued adaptation 
to xeric conditions, expansion from this area was likely 
coneurrent with the spread of an arid environment to the 
north and east. In that the polyploid and derived species 
are marginal] to the Chihuahuan desert proper, it is diffi- 
cult to imagine an opposing direction of migration and 
speciation. 

The interpretation of evolutionary trends in Chamae- 
saracha is based for the most part on chromosomal and 
distributional data in combination with correlated morpho- 
logical features. Some of the characters that I consider to 
be primitive and derived, at least within Chamaesaracha, 
are listed below : 


Primitive Derived 
1. woody rootstock subligneous root 
2. large, ovate, entire, small, lanceolate, lobed, and 
and thin leaves thickened or leathery leaves 
3. large flowers small flowers 
4. O-methylation of absence of O-methylation 
flavonoids of flavonoids 
5. increased glycosylation decreased glycosylation 
of flavonoids of flavoids 


The assessment of morphological characters generally 
follows classical consideration for those features (i.e., 
perenniality is generally considered primitive, as are large 
flowers). Polyploidy is considered to be derived in Chamae- 
saracha, quite apart from numerical considerations, be- 
cause the polyploids possess fewer of the characters con- 
sidered primitive in the listing above. Hence, the fairly 
widesperad occurrence of polyploidy offers a sound basis 
upon which to establish evolutionary trends within the 
genus. 

Considering the above, it is possible to construct a sche- 
matic representation of the proposed origin and evolution 


336 Rhodora [Vol. 75 


TOPHY SALIS 


edwordsiona 


coronopus 


PHYSALIASTRUM 


CHAMAESARACHA 


ANCESTRAL 


COMPLEX 


Figure 1. Hypothetical phyletie relationships within Chamaesara- 
cha. 


1973] Chamaesaracha — Averett 337 


of the species of Chamaesaracha (Figure 1). While the 
diagram is highly conjectural from a phyletic point of 
view, it nonetheless expresses visually my views on the 
relationships among the taxa. 

The flavonoid chemistry of the various species of the 
genus supports the relationships suggested in Figure 1. 
In what are considered to be advanced taxa, there is a 
general decrease of O-glycosylation and O-methylation, 
presumably reflecting the loss of enzymes necessary for 
these substitutions. In addition, the more closely related 
taxa (as I have grouped them) do have a greater similarity 
in their chromatographic profiles. 


ACKNOWLEDGMENTS 


A significant portion of this work, particularly the field 
studies, was made possible by NSF Traineeship grants for 
Environmental and Systematic Study while the author was 
a graduate student at The University of Texas at Austin. 
The latter part of the study was supported by a Summer 
research fellowship from The University of Missouri — 
St. Louis. 

I am indebted to Drs. M. C. Johnston, T. J. Mabry, A. M. 
Powell, and B. L. Turner for their interest and assistance 
in this study. I would also like to acknowledge those herb- 
aria from which some 2,000 specimens were borrowed.. 


TAXONOMIC TREATMENT 
Chamaesaracha Gray 


Chamaesaracha A. Gray, Bot. Calif. 1:540, 1876. TYPE 
SPECIES: Chamaesaracha coronopus (Dunal) Gray. 

Saracha Ruiz & Pav. sec. Chamaesaracha Gray, Proc. 
Amer. Acad. 10:62. 1874. 


Rhizomatous perennial herbs; stems erect, ascending, 
or prostrate, 10-50 em long, much branched, round striate 
or angular; glabrous to variously pubescent; leaves simple, 
broadly ovate, ovate, rhomic, lanceolate, oblanceolate, or 
linear-lanceolate, entire to pinnately or irregularly lobed, 


338 Rhodora [Vol. 75 


subsessile to distinctly petiolate, base attenuate to trun- 
cate, apex acute, rounded, or obtuse with veins prominent 
or obscure; flowers 1-5 in axils, pedicels slender, elongating 
and becoming curved or twisted in fruit; calyx campanu- 
late, rounded at the base, connate 14-834 its length, pu- 
bescent; corolla repand-rotate, white, ochroleuceous, or 
yellow-green, often tinged with purple on the outer surface 
of the lobes, with white tomentose appendages in the 
throat; stamens inserted near the base of the corolla; 
filaments slender, ca 3 mm long, anthers yellow, ca. 1.5 mm 
long, dehiscing longitudinally; styles slender and somewhat 
curved; stigmas blunt, or minutely 2-cleft; fruit a globose 
berry, tightly invested, but not enclosed, by the accrescent 
calyx; seeds flattened, reniform, rugose-reticulate, brown- 
ing with maturity; x = 12. 


KEY TO THE SPECIES 


a. Leaves 1.5-3.5 cm wide, ovate to broadly ovate-lanceo- 
late, entire to crenate; flowering calyces exceeding 5mm 
long. .... ee ee f b. 
b. Herbage villous with. long forked and simple tri- 

chomes; leaves subsessile, blades ca 14 as wide as 
long. D] f 4. C. villosa. 
b. Herbage basically glandular- pubescent, but with 
some longer simple trichomes; leaves petiolate, 


blades ca as wide as long. .... . . 5. C. crenata. 
a. Leaves less than 2.0 em wide; flowering calyces usually 
less than 4 mm long. C. 


c. Herbage predominately glabrous o or - pubescent with 
branched or stellate hairs; leaves linear, linear- 
lanceolate, or rhombic, entire irregular, or few- 
lobed; stems usually purplish or gray at base.. . d. 
d. Leaves linear to linear-lanceolate, usually few- 

lobed; sparsely pubescent with scurfy, white, 
stellate or branched hairs. e. 
e. Leaves sparsely pubescent with stellate hairs; 
pedicels without glandular hairs. ........ 
......................... 1. C. coronopus. 


1973] Chamaesaracha — Averett 339 


€. Leaves glabrous; pedicels with glandular 


hairs. . 2s. s. 2 C. edwardsiana. 

d. Leaves broadly NINE to rhombic, subentire 
or entire, pubescent with stellate hairs. .. f. 

f. Leaves densely pubescent with stellate hairs. 
ae ee fae . . €. C. pallida. 

f. Leaves glabrous. |... ... 2. C. edwardsiana. 


c. Herbage  predominately  glandular-pubescent or 
glandular-pubescent mixed with simple, longer tri- 
chomes; leaves broadly lanceolate or rhombic, en- 
tire to pinnately or irregularly lobed; stems green. 


Pe aes ae gies eae ne g. 
g. pus See of short @landular NaS S 

20 G PE 

g. P chos of trichomes mixed with glandular 

Husa) 5 cid A Mp 7. C. coniodes. 


1. Chamaesaracha coronopus (Dunal) Gray, Bot. Calif. 
1:540. 1876. Holotype (G!), Mexico, ca Laredo: [Texas, 
Webb Co. near Laredo] Berlandier 1494. Isotypes: not 
seen. Fig. 2. 

Solanum coronopus Dunal, in DC. Prod. 13:54. 1852. 
Saracha, coronopus (Dunal) Gray, Proc. Amer. Acad. 
10:62. 1874. 


Rhizomes slender and subligneous, 1.5-3.5 mm in diam- 
eter; stems prostrate or ascending, 10-50 cm long, ca 1.5 
mm in diameter, usually purplish or grey below; herbage 
glabrous to sparsely pubescent with scurfy, short, white 
stellate hairs; leaves linear to linear-lanceolate, 2.0-6.5 cm 
long, 1-10 (-15) mm wide, gradually narrowing to an at- 
tenuate, subsessile base, apex of blade acute, rounded, or 
obtuse, midrib prominent, sparsely pubescent with stellate 
hairs, margin subentire to deeply lobed; flowers 1 or 2 in 
axils; calyx 2.5-4.0 mm long, rather densely pubescent; 
corolla 6-10 (-15) mm wide; fruit 4-8 mm in diameter; 
chromosome number, n = 12, 24, 36. 

A rather widespread, weedy species occurring through- 
out most of the range of the genus (Fig. 3). At various 


340 Rhodora [Vol. 75 


perm 1970 


Figure 2. Habit sketch of Chamaesaracha coronopus, X 1; 
A, Leaf, X ca. 1; B, Stem, X 10. 


1973] Chamaesaracha — Averett 341 


elevations from sea level to 7000 ft., but more frequently 
in grassland areas with Larrea, Prosopis, and Juniperus, 
the species does not seem to be restricted to particular soil 
types and has even been collected on gypseous soils. Flow- 
ering from March to October, primarily in the early and 
again in the later summer, depending on rains. 

Chamaesaracha coronopus is most closely related to C. 
edwardsiana. The plants from south Texas differ some- 
what from those found in west Texas, New Mexico, and 
Arizona in having broader, more lobed leaves. In leaf 
shape, the plants from south Texas superficially resemble 
C. coniodes, but the two species are not believed to be 
closely related. 


REPRESENTATIVE SPECIMENS: Arizona: APACHE CO. Win- 
dow Rock, 19 Aug 1962. Bohrer 737 (uc). COCHISE co. Ft. Lowell, 
31 Jul 1903, Thomber 245 (ARIZ, NY, UC). COCONINO CO. Black Point, 
near Cameron, 18 May 1961, Demaree 44084 (ARIZ, SMU, UC). GILA 
CO. 16 mi S of Safford, 21 Oct 1964, Gould 3989 (ARIZ, UC). MOHAVE 
co. 5 mi NE of Tweep, Parker et al. 6256 (ARIZ, US). NAVAJO CO. 
Holbrook, 27 Jun 1896, Zuck s.n. (GH, NY, US). PIMA CO. Vail, 22 
Oct 1926, Peebles et al. 3306 (ARIZ, MICH). PINAL CO, near Sacaton, 
2 May 1926, Harrison 1779 (ARIZ). LAVAPAI CO. near Montezuma 
Castle Natl Monument, 21 May 1936, Nelson & Nelson 2050 (GH, 
NY, UC). YUMA co. Antelope, May-Oct 1902, Purpus 95 (UC). 

Colorado: FREEMONT CO. Canyon City, 1872, Brandegee 391 (GH, 
NY, UC, US). LAS ANIMAS CO. 10 mi N of Bronson, 16 Jul 1968, 
Averett & Tomb 348 (SMU, MO, TEX). LINCOLN CO. 7 mi S of Forder 
School, 6 Jul 1937, Ownbey 1327 (GH, NY, UC). PUEBLO CO. mesas 
near Pueblo, 15 May 1900, Rydberg & Vreeland 5666 (NY). 

Oklahoma: CIMARRON CO. 3.5 mi N of Kenton, 7 Oct 1950, Waterfall 
9713 (ARIZ, OKLA, SMU). 

New Mexico: BERNALILLO CO. Albuquerque, 4 Sep 1884, Jones 4147 
(ARIZ, GH, NY, UC, US). COLFAX CO. vicinity of Raton, 21-22 Jun 1911, 
Standley 6365 (US). DONA ANA CO. plains E of Organ mts. 28 Aug 
1897, Wooton 406 (GH, NY, UC, US). GRANT CO. Mangas Springs, 18 
mi NW of Silver City, 31 May 1903, Metcalfe 97 (ARIZ, NY, UC, US). 
HIDALGO CO. 5 mi N of Animas, 9 Sep 1950, Castetter 3030 A & B 
(UNM). LINCOLN CO. above Tularosa, 24 Aug 1897, Wooton s.n. 
(ARIZ, US). LUNA CO. foothills, Tres Hermanas Mts, W of Columbus, 
12 Sep 1942, Clark s.n. (UNM). MCKINLEY CO. Gallup, 28 Jul 1933, 
Degener 4834 (NY). OTERO CO. White Mts, divide above Mescalero, 
1 Aug 1897, Wooton s.m. (NY, US). SANDOVAL CO. 19 mi W of Santa 
Fe on Rio Grande River, 31 May 1897, Heller & Heller 3633 (GH, 


342 Rhodora [Vol. 75 


Figure 3. Distribution of Chamaesaracha coronopus. 


1973] Chamaesaracha — Averett 343 


NY, US). SAN JUAN CO. Sheep Springs, 18 Aug 1880, Rusby 302 
(MICH, NY). SAN MIGUEL CO. near Pecos, 18 Aug 1908, Standley 
(GH, NY, US). SANTA FE CO. S of Santa Fe near hwy 85, 14 Jul 1966, 
Plowman AP195 (ARIZ). SOCORRO co. Ojo Caliente, 28 Jul 1904, 
Wooton 2683 (US). UNION CO. 2 mi W of Oklahoma line, NW of 
Kenton, 11 Jun 1948, Rogers 5973 (MICH, US). VALENCIA CO. 5 mi W 
of Correo, 15 Jun 1959, Martin 3273 (UNM). 

Texas: BEXAR CO. San Antonio, 23 Apr 1917, Palmer 11626 (TEX). 
BREWSTER CO. Alpine, 7 Jun 1940, Warnock 21586 (TEX). BROWN CO. 
Brownwood, 31 Mar 1917, Palmer 11439 (TEX). BURNET CO. Marble 
Falls, 1 Apr 1913, Young s.n. (TEX). CAMERON CO. Boca Chica rd, 
2 mi E of Palmito battleground monument, 29 Apr 1959, Travers 
1160 (SMU, TEX). CHILDRESS CO. Spr 1931, Biology class sm. (TEX). 
CULBERSON CO. railroad near Kent, 1 May 1902, Tracy & Earle 407 
(NY, TEX, US). DUVAL CO. San Diego, 1885, Croft 5994 (US). EL PASO 
co. vicinity of El Paso, 1911, Stearns 39 (US). ERATH CO. 21 Mar 
1921, Gough s.n. (TEX). HIDALGO CO. 6 mi E of Sulivan City, 8 Mar 
1959, Turner 4493 (TEX). JEFF DAVIS CO. 9 mi NE of Ft. Davis in 
Limpia Canyon, 27 Apr 1947, Cory 53554 (SMU). KARNES CO. 0.5 mi 
NW of Karnes City, 14 Mar 1953, Johnson 1140 (SMU, TEX). KLE- 
BERG CO. Kingsville, 11 Apr 1905, Tracy 9225 (GH, NY, TEX, US). LA 
SALLE CO. 10 mi N of Encinal, 30 Mar 1935, Wiegand 2065 (GH). 
MAVERICK CO. on the Rio Grande River, Eagle Passs, 7 Mar 1852, 
Schott s.n. (NY, US). MCKINNEY CO. 1881, Garrett s.n. (NY). MITCH- 
ELL CO. b mi SE of Colorado City, 29 Jul 1945, Pohl 5113 (SMU). 
NUECES CO. Corpus Christi 21 Mar 1907, York s.n. (TEX). OLDHAM CO. 
Tascosa, 16 Jun 1932, Reed 3510 (US). PECOS CO. 3 mi N of Ft. Stock- 
ton, 22 Jun 1952, Warnock 10490 (SMU). PRESIDIO CO. Marfa, Jun 
1936, Hinckley 658 (ARIZ, GH, SMU, TEX). SAN PATRICIO CO. 8 mi E of 
Taff, 20 Oct 1951, Jones 652 (SMU). SCURRY CO. 26 mi SW of Snyder, 
17 Apr 1960, Waterfall 15875 (OKLA). STARR CO. 4 mi W of Ft 
Ringold, Tharp & York 51-250 (TEX). TAYLOR CO. 8 mi S of Abilene, 
8 Apr 1960, McCoy 3727 (OKLA). TRAVIS co. Austin, 18 Mar 1940, 
Warnock 88 (TEX). UVALDE co. NW of Uvalde, 3 Apr 1942, Lundell 
& Lundell 10974 (SMU, US). VAL VERDE Co. Del Rio, 7 Dec. 1891, 
Plank s.m. (NY). WEBB CO. Laredo, 1-20 Aug 1879, Palmer 920 (GH, 
NY). WICHITA CO. 215 mi W 1 mi S of Iowa Park, 8 Jun 1960, 
Mahler 1168 (TEX). WILLACY CO. 3 mi S of Raymondville, 6 Nov 
1953, Shinners 17108 (SMU). WILSON CO. 4 mi S of Stockdale, 25 
Mar 1955, Johnston & McCart 5082 (TEX). 

Utah: KANE CO. 8 mi E of Kanab, 7 Jun 1957, Anderson 760 (NY, 
UC). SAN JUAN CO. along the San Juan River, 20-25 Aug 1911, 
Rydberg & Garrett 9970 (Ny, US). 

Mexico: CHIHUAHUA, 6 mi S of Camargo, 29 Jun 1937, Shreve 
8070 (ARIZ, GH, US). COAHUILA, Saltillo and vicinity, 1898, Palmer 
145 (GH, NY, UC, US). DURANGO, 85 mi NE of Durango, 11 Aug 1959, 


344 M Rhodora [Vol. 75 


Figure 4. Habit sketch of Chamaesaracha edwardsiana, X 1. 


1973] Chamaesaracha — Averett 345 


Waterfall 15399 (OKLA). NUEVO LEON, 16 mi N of Matehuala, 21 
Aug 1959, Waterfall 15749 (OKLA). SAN LUIS POTOSI, San Luis 
Potosi, 1879, Shaffner 404 (Ny, US). SONORA, mesa de las Carreras, 
W of Colonia Morelos, 15 Sep-4 Oct 1941, White 4547 (ARIZ, GH, 
MICH). TAMAULIPAS, 10 mi S of Nuevo Leon, 26 Feb 1944, Barkley 
14326 (GH, NY, TEX). 


2. Chamaesaracha edwardsiana Averett, Sida 5: 48-49. 
1972. Holotype (TEX): TEXAS, Travis Co.: 44 mi E 
of the Pedernales River on hwy 620. 28 Jun 1968. 
Averett 289. Isotypes: GH, MO, SMU. (Fig. 4). 


Rhizomes slender and subligneous, 1.5-3.5 (-7) mm in 
diameter; stems erect or ascending, 7-30 cm long, ca 1.5 mm 
in diameter, usually purplish or grey below; herbage 
glabrous to pubescent with stellate hairs, or occasionally 
with simple and branched trichomes (especially in the 
upper portions); leaves linear-lanceolate to rhombic, typi- 
cally undulate, but occasionally few-lobed, 2.5-7.0 em wide, 
base attenuate, subsessile to slightly petiolate, midrib 
prominent, but secondary veination usually present, es- 
sentially glabrous; flowers 1 or 2 in axils, flowering pedicel 
pubescent with trichomes and glandular hairs; calyx 4-5 
mm long, pubescent with stellate, branched and simple 
trichomes; corolla 1.0-1.5 cm wide; fruit 5-8 mm in diam- 
eter; chromosome number, n = 24. 

Primarily on limestone soils in the Edwards Plateau of 
Texas (Fig. 6). Flowering from March to October, but 
mostly in the early and late summer, depending on rains. 

Chamaesaracha edwardsiana is probably most closely re- 
lated to C. coronopus, differing from the latter in having 
glabrous, entire leaves. It is also related to C. pallida, dif- 
fering in the narrow, linear-lanceolate, glabrous leaves. 


REPRESENTATIVE SPECIMENS: Texas: BEXAR CO. near 
Bracken, 13 Jul 1903, Groth 73 (GH, US). BLANCO CO. ca 1 mi W of 
Johnson City, 5 Apr 1969, Tomb & Tomb 386 (TEX). BRAZOS CO. 
15 mi from A & M Campus, College Station, 26 Apr 1967, Lonard 
1899, (TEX). coMaL co. Comanche Springs, New Braunfels, May, 
1850, Lindheimer 1938 (ARIZ, GH, NY, SMU, TEX, UC, US). CROCKETT CO. 
McKenzie’s Well, May 188?, Reverchon 1569 (NY, SMU, US). ED- 
WARDS CO. Ranch Expt. Sta., 2 Nov 1929, Cory 3469 (GH). HAYS CO. 


346 Rhodora [Vol. 75 


Figure 5. Chamaesaracha pallida: A, Habit sketch, X 1; B, Seed, 
X ca. 5; C, Leaf X 2; D, Stem, X ca. 10; E, Pubescence, X 50. 


1973] Chamaesaracha — Averett 347 


E of Henly, 20 Apr 1941, Lundell & Lundell 10279 (MICH, US). 
LAMPASAS CO. 8 mi E of Lampasas, 15 Jun 1929, Wolff 940 (us). 
LLANO CO. near Llano, 4 Aug 1931, Wolff 3080 (US). KENDALL CO. 
5 mi N of Boerne, 10 May 1969, Averett 454 (SMU, MO, TEX). KERR 
co. Kerville, 19-25 Apr 1894, Heller 1647 (ARIZ, GH, MICH, NY, SMU, 
UC, US). MENARD CO. 5 mi NE of Menard, 3 Jun, 1968, Shinners 
32274 (SMU). PECOS CO. 15 mi W of Sanderson, 22 Apr 1948, White- 
house 19729 (MICH, NY, OKLA, SMU). TRAVIS CO. Austin, 17 May 1940, 
Lundell & Lundell 8957 (GH, MICH, NY, SMU, UC, US). UVALDE CO. 
Montell, 15 Oct 1917, Palmer 13006 (TEX, UC). WEBB CO. Laredo 
Junior College Pasture, 14 Apr 1963, Abrego s.n. (TEX). WILSON CO. 
Sutherland Springs, Aug 1879, Palmer s.m. (GH). WILLIAMSON CO. 
SW of Georgetown, 16 May 1942, Wolcott 118 (TEX). 

Mexico: COAHUILA, Sierra del Pinto, 26 Aug 1940, Johnston & 
Muller 745 (GH, MICH). NUEVO LEON, 10 mi S of Roberto Junction 
on hwy 56, 22 Jul 1969, Bierner & Turner 90 (TEX). SAN LUIS 
POTOSI, Charcas, Jul-Aug 1934, Lundell 5065 & 5065A (ARIZ, MICH, 
OKLA, TEX, US). 


3. Chamaesaracha pallida Averett, Sida 5: 48-49. 1972. 
Holotype (TEX): TEXAS, Presidio Co.: 35 mi SW of 
Marfa on Pinto Canyon Road. 15 Jul 1966. Averett 
155. Isotypes: GH, MO, SMU, US. Fig. 5. 


Rhizomes slender and subligneous, occasionally woody; 
stems suberect to ascending, 7-15 (-20) cm long, 1.0-1.5 
em wide (ca 1% the length), narrowly rhombic, broadly 
lanceolate, or oblanceolate, entire to deeply undulate, apex 
acute, obtuse, or rounded, base attenuate, subsessile or 
short-petiolate, pubescent with stellate or branched hairs; 
flowers 1 or 2 in axils, pedicels slender 1-2 cm long; calyx 
3-4 mm long, pubescent; corolla usually 1-1.5 cm wide; 
fruit 5-7 mm in diameter; chromosome number, n = 36. 


A rather rare species of Trans-Pecos, Texas and north- 
ern Mexico (Figure 6). Flowering from March to October, 
primarily in the early and late summer depending on rains. 
The species appears to be mostly confined to limestone soils, 
although it has been reported from igneous soils. 

Chamaesaracha pallida has its closest affinities with C. 
edwardsiana, and to some extent, C. sordida, However, a 
dense stellate pubescence is typically found in C. pallida. 


348 Rhodora [Vol. 75 


^ 


Figure 6. Distribution of Chamaesaracha edwardsiana (open cir- 
cles) and Chamaesaracha pallida (closed circles). 


1973] Chamaesaracha — Averett 349 


Some specimens of C. edwardsiana and C. coronopus may 
have considerable pubescence, but not so much as C. pallida. 


REPRESENTATIVE SPECIMENS: New Mexico: EDDY CO. Sitting 
Bull Falls, 24 May 1944, Hershey 3098 (GH). 

Texas: BREWSTER CO. 30 mi S of Marathon, 2 Apr 1938, Warnock 
T283 (TEX, US). CULBERSON CO. S of McKittrick Canyon, Guadalupe 
Mts, 3 Aug 1952, Warnock 10941 (SMU). EL PASO CO. Hueco Pass, 
29 May 1932, Whitehouse s.n. (TEX). HUDSPETH CO. Calcareous Can- 
yon, Malone Mts, 21 Aug 1946, Waterfall 6663 (GH, NY). IRION CO. 
1 mi S of Mertzon, 24 Apr 1948, Warnock 7700 (TEX). KIMBLE CO. 
16 Apr 1930, Wolff 1601 (Us). KINNEY CO. Anacacho Mts, ca 30 mi 
SE of Brackettville, 15 May 1965, Strother 253 (TEX, SMU). PECOS 
co. ca 15 mi W of Sanderson, 22 Apr 1948, Whitehouse 19729 (SMU). 
PRESIDIO CO. 8 mi SW of Shafter, 30 Jul 1945, Muller 8471 (NY, 
OKLA, SMU). REGAN CO. 714 mi E of Bit Lake, 24 Apr 1947, Cory 
53440 (NY, SMU). TERRELL CO. Blackstone Ranch, ca 15 mi S of 
Sheffield, 15 Jun 1949, Webster 269 (MICH, TEX). TOM GREEN CO. 7% 
mi S of Christoval, 28 Oct 1945, Cory 50565 (GH, NY, SMU, UC, US). 
UVALDE CO. 4 mi E of Kline, 2 Nov 1963, Moore et al. 8845 (SMU). 
VAL VERDE CO. 30 (airline mi N of Del Rio, 25 Apr 1943, Cory 
41528 (GH). 

Mexico: COAHUILA, 26 mi SW of Monterrey, 1 Dec 1945, Barkley & 
Warnock 14700M (GH, NY, TEX). DURANGO, 9 mi N of La Zarca on 
hwy 45, Sikes & Patterson 398A,B,C, (SMU, SRSC, TEX). NUEVO LEON, 
Villa de Garcia, 19 Sep 1959, Fuentes 19 (OKLA). 


4. Chamaesaracha villosa Rydb., Mem. Torr. Bot. Club 4: 
368. 1896. Holotype (US!), MEXICO, Coahuila: moun- 
tains 24 mi NE of Monclova. Sept 1880. Palmer 924. 
Isotypes: GH!, K!, NY!, US!, vr!. Fig. 7. 


Plants strongly perennial; rhizomes woody, stems sub- 
erect to ascending, 15-50 em long, 1.5-3.5 mm in diameter, 
herbaceous to somewhat woody, densely pubescent with 
long, villous, forked trichomes, occasionally somewhat 
glandular; leaves 4-7 cm long, 1.5-3.0 cm wide, ovate to 
ovate-lanceolate, entire to undulate-crenate, apex acute to 
obtuse, base attenuate, petioles less than !/4 the total 
length of the leaf if present; flowers 1 to few in axils; 
pedicels 1-3 em long; calyx ca 5 mm long, pubescent; 
corolla 0.5-3.0 cm wide; fruit 5-8 mm in diameter; chromo- 
some number, 7-12. 


350 Rhodora [Vol. 75 


Figure 7. Chamaesaracha villosa: A, Habit sketch, X 1; B, Leaf, 
X 1; C, Stem, X 10; D, Pubescence, X 40. 


1973] Chamaesaracha — Averett 351 


Creosote bush dominated desert areas of northern Mexico 
(Durango, Chihuahua, and Coahuila) and Trans-Pecos 
Texas along the Rio Grande River (Fig. 9), usually occur- 
ring in sandy or gravelly washes or along the roadside 
in disturbed areas. Flowering primarily in early spring 
(March-April) and fall (September-October). 


Considerable data indicate that Chamaesaracha villosa 
is most closely related to C. crenata. Further discussion of 
the two species is found following treatment of the latter. 


REPRESENTATIVE SPECIMENS: Texas: BREWSTER CO. near 
Comanche Springs between Terlingua and Lajitas, 17 Jul 1937, 
Warnock 20701 (TEX). CULBERSON CO. N of Van Horn, 10 Jun 1905, 
Reverchon sm. (SMU). HUDSPETH CO. Indian Hot Springs, 20 Apr 
1932, Whitehouse 8500 (TEX). PRESIDIO CO. b mi N of Ruidosa, 26 
May 1964, Seudday 372 (OKLA, SRSC). 


Mexico: CHIHUAHUA, 31 mi N of Jimenez, 1 Aug 1939, White 2171 
(ARIZ, GH, MICH). COAHUILA, Mohano, Jun 1910, Purpus 4550 (GH, 
UC, US). DURANGO, 2 mi W of Bermejillo, 18 Sep 1938, Shreve 8783 
(ARIZ, MICH, UC). 


5. Chamaesaracha crenata Rydb., Mem. Torr, Bot. Club 4: 
368. 1896. Holotype (US!), MEXICO, Coahuila: Par- 
ras, 111 mi W of Saltillo. Jun 1880. Palmer 923. Iso- 
types GH! K!, NY!, US! vr!. Fig. 8. 


Plants strongly perennial; rhizomes woody; stems 15-40 
cm long, 1.5-3.5 mm in diameter, herbaceous or somewhat 
woody, much branched, densely pubescent with glandular 
hairs mixed with longer trichomes; leaves 4-6 cm long, 
1.5-3.5 em wide, ovate to broadly rhombic, entire-undulate 
to crenate, apex obtuse, base cuneate or somewhat attenu- 
ate: petioles 2-3 cm long (ca 1/3 the total length of the 
leaf); flowers 1-5 in axils: pedicels 1-3 cm long; calyx 
(3-) 5-7 mm long, densely pubescent, primarily with long 
trichomes; corolla 1-2 cm wide; fruit 8-10 mm in diameter; 
chromosome number, n = 12. 


Creosote-desert shrub areas of northern Mexico and 
along the Rio Grande River in Texas (Fig. 9). Usually 
found in sandy or gravelly washes or along the roadside in 


352 Rhodora [Vol. 75 


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`. 


E 


a 
MV 


Chamaesaracha crenata: A, Habit sketch, X 1; B, 


Figure 8. 
Stem X 10; C, Pubescence, X 40. 


1973] Chamaesaracha — Averett 353 


disturbed areas. Flowering primarily in early spring and 
in the fall, depending on rains. 


Chamaesaracha crenata is most closely related to C. 
villosa. The two species are largely allopatric, but their 
ranges overlap in Coahuila and in the Trans-Pecos area of 
Texas (Figure 9). In the latter area, particularly in Big 
Bend National] Park and around Lajitas (Brewster and 
Presidio Co), the two species are often difficult to dis- 
tinguish and hybridization is suspected. At the extremities 
of their ranges, however, the two species are fairly clear 
and they can be distinguished by the characters given in 
the key to the species. 

Rydberg, in distinguishing the two species, used differ- 
ences in flower size, fruit size, seed number, and calyx 
shape. The type specimens do exhibit such differences; 
however, they are not particularly useful in distinguishing 
between the taxa since these characters are quite variable 
both within and among populations. 


I have maintained the species as distinct for biological 
reasons although it might be equally sound to treat the 
taxa as varieties of but a single species. The characters 
that distinguish the two are largely of the same kind that 
distinguish other species in the genus (viz., leaf shape and 
pubescence), although, perhaps because of gene exchange 
in regions of overlap, the differences are often not so easily 
seen. The decision for varietal or specific status is some- 
what arbitrary in this instance, so I have chosen to follow 
Rydberg's treatment in recognizing allopatric species show- 
ing minimal gene exchange in regions of sympatry. 


REPRESENTATIVE SPECIMENS: Texas: BREWSTER CO. Glenn 
Springs, Big Bend Nat] Park, 25 May 1955, Warnock & Wallmo 
12300 (SRSC, TEX). STARR CO. 3 mi N of Roma, 31 Jan 1954, Shinners 
17714 (SMU). TERRELL CO. Feodora, 26 Apr 1928, Palmer 33582 (GH, 
NY, US). VAL VERDE CO. Shumla, 10 Oct 1936, Tharp, s.m. (GH, NY, 
SMU, UC, US). WEBB CO. Laredo, 15 Mar 1917, Palmer 11285 (vc). 

Mexico: CHIHUAHUA, 5 km W of Santa Fe, 15 Sep 1942, Stewart 
2604 (GH). COAHUILA, Parras, Mar 1905, Purpus 1081 (GH, NY, UC). 


6. Chamaesaracha sordida (Dunal) Gray, Syn. Fl. N. 


354 Rhodora [Vol. 75 


max 


Figure 9. Distribution of Chamaesaracha villosa (open circles) 
and Chamaesaracha crenata (closed circles). 


1973] Chamaesaracha — Averett 355 


Amer. 1: 232. 1876. Holotype (G!), tracing (GH!) : 
TEXAS: Webb Co. near Laredo, Aug 1829. Berlandier 
2076 (—666). Isotypes: GH!, NY(2)!. Fig. 10. 
Withania sordida Dunal. in D.C. Prod. 13: 456. 1852. 


Rhizomes subligneous to woody; stems suberect to as- 
cending, 9-30 cm long (typically ca 15 em), 1.5-2.0 mm in 
diameter, pubescent, herbage densely glandular pubescent, 
occasionally mixed with simple trichomes; leaves rhombic, 
entire or subentire to only slightly lobed, 1.5-4.0 cm long, 
4-8 (-10) mm wide, acute to somewhat rounded at the apex, 
gradually narrowing to a subsessile base; flowers 1-2 in 
axils; pedicels 1.5-3.0 em long; calyx ca 3 mm long, rather 
densely pubescent; corolla typically 1 cm or less wide; 
fruit 4-8 mm in diameter. 


Southwestern United States and adjacent Mexico (Fig- 
ure 11). Fairly widespread in this area, commonly occur- 
ring in disturbed soils along the roadside. Usually in asso- 
ciation with Prosopis or Larrea. Principal flowering dates 
from March to June and August to October, depending on 
rains. 

Chamaesaracha, sordida is related to C. pallida and C. 
coniodes, probably more closely to the former. Further 
consideration of this species is found in the discussion of 
C. coniodes. 


REPRESENTATIVE SPECIMENS: Arizona: COCHISE CO. 6 mi 
NW of Chiricahua, 18 May 1945, Gould & Pultz 3157 (ARIZ, GH, NY, 
UC, US). PIMA CO. 26 mi E of Tucson, 10 May 1935, Maguire 11214 
(GH, NY). PINAL CO. 4 mi N of Christmas, Anderson et al. 1047 
(ARIZ). 

New Mexico: DE BACA CO. 7 mi W of Ft. Sumner, hwy 60, 15 Jun 
1961, Mahler 1814 (OKLA). DONA ANA CO. Organ Mts, 15 Jul 1897, 
Wooton 156 (GH, NY, UC, US). EDDY CO. Carlsbad Caverns 24 May 
1931, Nelson 11403 (GH, US). GRANT CO. 3 mi E of Red Rock, 20 May 
1935, Maguire et al.; R 14. W., 12 Aug 1954, Gordon & Buchanan 
74 (UNM). LEA CO. b mi NW of Hobbs, 7 Apr 1967, Pearce 2666 
(ARIZ). LINCOLN CO. 5 mi W of Lincoln, 26 Jul 1938, Hitchcock et al. 
4277 (uc). OTERO co. Road to Rinconado, 15 Aug 1938, Humphrey 
57 (ARIZ, UNM). SIERRA CO. Kingston, 10 Sep 1904, Metcalfe 1335 (GH, 
NY, UC, UNM, US). SOCORRO CO. 6.5 mi N of Old Bingham, and 2 mi 
E, 1 Aug 1949, Dunn 6067 (SMU). 


356 Rhodora [Vol. 75 


A.CANO 'CG'] 


Figure 10. Chamaesaracha sordida: A, Habit sketch, X 1; B, 
Leaf, X 2; C, Stem, X 10. 


1973] Chamaesaracha — Averett 357 


Texas: BREWSTER CO. foothills of Chisos Mts, 19 Jun 1931, Tharp 
sin. (GH, MICH, TEX, UC). BRISCOE CO. caprock W of Quitaque on 
Silverton road, 18 Jun 1945, Whitehouse 10001 (SMU). CRANE CO. 
6 mi S of Crane, 11 Jul 1941, Tharp s.n. (GH, SMU). CROCKETT CO. 
roadside park over Pecos River Canyon, 26 Sep 1964, Turner 5109 
(TEX). CULBERSON CO. below McKittrick Canyon, 23 Jul 1931, Moore 
& Steyermark (GH, MICH, NY, UC, US). DAWSON CO. 7 mi SE of 
Lamesa, 2 May 1967, Shinners 31814 (SMU). SIMMITT CO. 3 mi NW 
of Carrizo Springs, 2 May 1954, Johnston et al. 3543 (OKLA, TEX). EL 
PASO co. El Paso, 16 Apr 1884 Jones 3704 (ARIZ, UC). FLOYD CO. 
7 mi S of Floydad, 19 Jun 1964, Delso 57 (OKLA). GLASSCOCK CO. 
10 mi W of Garden City, 26 Apr 1964, Turner 5015 (TEX). HOWARD 
CO. Scenie Mt, Big Spring, 14 Apr 1963, Fields 53 (OKLA). HUDSPETH 
co. ca 30 mi E of El Paso, hwy 180, 7 Sep 1955, Warnock 13812 
(TEX). LOVING CO. ca 10 mi E of Mentone, 3 Jun 1949, Turner 966 
(SMU). LUBBOCK CO. Buffalo Springs, Lubbock, 29 May 1982, Reed 
3494 (US). MAVERICK CO. Rio Grande River, Eagle Pass, 7 Mar 1852, 
Schott s.n. (NY). NOLAN CO. oilfield W of Sweetwater, 9 Jul 1941, 
Tharp s.m. (GH, TEX). PECOS CO. Ft. Stockton, 30 Jun 1938, Warnock 
T407 (ARIZ, GH, TEX, US). PRESIDIO CO. ca 20 mi SW of Marfa on 
Ruidoso road, 24 Jul 1966, Averett 153 (SMU, MO, TEX). RANDALL CO. 
floodplains, Red River, Palo Duro Canyon, 2 Sep 1907, Ball 1249 
(NY, UC, US). REEVES CO. 25 mi N of Pecos, 4 Aug 1943, Waterfall 
5491 (GH, NY). SCURRY CO. 12.6 mi E of Snyder, 16 Sep 1966, Shin- 
ners 81572 (SMU). STARR CO. 10 mi NW of Roma, 15 Mar 1964, Pena 
21 (sMU). STONEWALL CO. 8.4 mi NW of Aspermont, 29 May 1963, 
Shinners 30036 (SMU). SUTTON CO. 10 mi W of Sonora, 30 Apr 1950, 
Cory 50054 (SMU). TAYLOR CO. ca 1 mi S of Wylie, 29 Mar 1963, 
Henderson 63-102 (SMU, TEX). TERRELL CO. 2-10 mi N of Sanderson, 
1 Jun 1957, Warnock & McBryde 14868 (TEX). TOM GREEN CO. San 
Angelo, 18-19 May 1920, Pennell 10497 (NY). UPTON CO. 14 mi N 
of Rankin, 18 Apr 1933, Cory 5589 (GH). VAL VERDE CO. 5 mi NE of 
Juno, 7 Jun 1957, Warnock & McBryde 15078 (TEX). WEBB CO, 11 mi 
S of Laredo, 23 Mar 1963, Robles 16 (SMU). WINKLER Co. 10 mi N of 
Kermit, 8 Dee 1965, Collins 1088 (SMU). ZAPATA CO. 22 mi S of 
Laredo, 4 Apr 1965, Guerra et al. 609 (ARIZ, TEX). 

Mexico: CHIHUAHUA, 3.5 mi S of Trincheras, 10-12 Aug 1941, 
Johnston 8061 (GH). COAHUILA, Muzquiz, Apr 1938, Marsh 1179 
(OKLA, SMU, TEX). NUEVO LEON, 23 mi N of Sabinas Hidalgo, 26 
Mar 1944, Webster & Barkley 14580 (Ny, TEX, UC). 


7. Chamaesaracha coniodes (Moric. ex Dunal) Britt., Mem. 
Torr. Bot. Club 5: 287. 1895. Lectotype (G) not seen: 
MEXICO: Canada Verde, between Lardo and Bexar 
[San Antonio], [TEXAS: McMullen Co] Mar 1828. 


[Vol. 75 


Rhodora 


358 


"DpWpA4os mw90408$2D14717) JO uomnmnqrasqstq 


‘TT ean3rq 


` 
` 


1973] Chamaesaracha — Averett 359 


Berlandier 1494 (234). Isotype GH! Berlandier 1463 
(—203) was also cited in the original description, but 
I have selected the other collection since the specimen 
at GH is the only one I have seen. Fig. 12. 

Solanum coniodes Moric. ex Dunal, in D.C. Prod. 13(1) : 
456. 1852. 


Rhizomes subligneous to woody; stems prostrate or as- 
cending, 9-30 em long, 2.0-3.0 mm in diameter, pubescent ; 
herbage pubescent with simple trichomes, occasionally 
mixed with fewer glandular hairs, usually somewhat vil- 
lous; leaves broadly lanceolate, 2.5-5.5 cm long, 1.0-2.0 cm 
wide, crenate to deeply and pinnately lobed; apex acute to 
rounded, base attenuate, subsessile; flowers 1-2 in axils; 
pedicel 2.0-3.5 em long; calyx ca 4.0 mm high; corolla 
1.0-1.5 em wide; fruit ca 7 mm in diameter, chromosome 
number, n — 24, 36. 


Southwestern United States and adjacent Mexico (Fig- 
ure 13). Fairly widespread, commonly occurring in dis- 
turbed soils along the roadside, usually in association with 
Prosopis or Larrea. Principal flowering dates, March to 
June and August to October, depending on rains. 

Chamaesaracha coniodes was merged with C. sordida by 
Gray (1876). However, the two species may be distin- 
guished by the characters given in the key to the Species. 
Chemical and chromosomal differences also serve to dis- 
tinguish the two species. In areas of sympatry both taxa 
retain their diagnostic features, even in mixed populations 
and, hence, appear to be good biological species. 


REPRESENTATIVE SPECIES: Colorado: BACA CO. breaks of 
Carrizo Creek, 5 mi SW of Kirkwell, 29 Aug 1949, Weber 5120 (SMU, 
UC). LAS ANIMAS Co. Tecolate mesa, near Troy, 15 Jul 1947, Rogers 
4848 (MICH, UC). 

Kansas: CLARK CO. 9 mi N of Ashland, 28 Apr 1939, Horr E249 
(OKLA, SMU, TEX, UC, US). ELLIS CO. Smoky Hill River, 22 Jun 1935, 
Bondy 121 (ARIZ, OKLA, US). FINLEY CO. 4 mi W of Kalvesta, 2 Jun 
1967, Stephens 11300 (OKLA, SMU, UC). HAMILTON CO. Syracuse, 5 
Jul 1893, (OKLA, SMU, UC); Syracuse, 5 Jul 1898, Thompson 190 
(NY, UC). MORTON CO. 9 mi N, 3 mi W of Elkhart, 7 Jun 1961, 


360 Rhodora [Vol. 75 


Qudruy Ü ot 970 


Figure 12. Chamaesaracha coniodes: A, Habit sketch, X 1; B, 
Stem, X 15; C, Leaf, X ca. 1. 


1973] Chamaesaracha — Averett 361 


Richards 2559 (sMU). SEWARD CO. 12 mi N of Liberal, 8 Jul 1960, 
McGregor 15778 (SMU). ULYSSES GRANT CO. cellar excavations, 26 
Jun 1893, Thompson 29 (GH, NY, UC, US). 

New Mexico: CURRY CO. 5.7 mi W of Melrose, 5 Aug 1945, Shinners 
20905 (SMU). DE BACA co. 2 mi N of Ft. Sumner, 3 Jun 1969, Tomb 
497 (TEX). LINCOLN CO. Gray, 6 Jun 1898, Skehan 31 (GH, NY, US). 
quay co. S of San Juan, 26 Aug 1945, Waterfall 6123 (GH, NY, OKLA, 
SMU). SAN MIGUEL co. 5 mi SW of Las Vegas, 11 Aug 1949, Gordon 
& Norris 495 (UNM). SIERRA CO. Elephant Butte Dam, 30 Apr 1933, 
Nelson s.m. (UNM). UNION CO. 3 mi W of Oklahoma line W of 
Kenton, 11 Jun 1948, Rogers 5960 (MICH, US). 

Oklahoma: BEAVER CO. 2 mi W of Turpin, 16 Aug 1960, Hindman 
236 (OKLA). BECKHAM CO. 2 mi E of Sayne, 9 Aug 1927, Stratton 
354 (SMU). CIMARRON CO. 3.5 mi N of Kenton, 7 Oct 1950, Waterfall 
9714 (ARIZ, OKLA, SMU). CUSTER CO. 1 mi W, 1 mi S of Weatherford, 
11 May 1952, Waterfall 10716 (OKLA, SMU). ELLIS CO. 3.8 mi S of 
Shattuck, 25 Sep 1964, Shinners 30415 (SMU). GREER CO. near Gran- 
ite, Stevens 1010 (GH, OKLA, US). HARMON CO. floodplain of Elm 
Fork, 14 mi S of Erick, 25 May 1940, Waterfall 2002 (GH, NY, OKLA). 
JACKSON CO. 6 mi SE of Duke, 29 May 1957, Waterfall 18105 (ARIZ, 
OKLA, US). KIOWA CO. 1.7 mi N of Snyder, 23 Apr 1938, Fry 186 
(OKLA). MAJOR CO. near Waynoka, 23 May 1913, Stevens 599 (GH, 
NY, OKLA). OKFUSKEE CO. 2 mi E of Paden, 7 Jun 1938, Ray 94 
(OKLA). SWANSON CO. near Mountain Peak, 23 Jun 1913, Stevens 
1239 (GH, OKLA). TILLMAN co. 2 mi S of Frederick, 2 Apr 1961, 
Myrick 49 (OKLA). woops co. 8 mi NW of Alva, 8 May 1932, Strat- 
ton 3014 (OKLA). 

Texas: BAILEY CO. Muleshoe Nat’l Wildlife Refuge, 20 mi S of 
Muleshoe, 18 Jun 1964, Rosson 576 (OKLA, UC). BAYLOR CO. 4.2 mi 
ENE of Mabelle, 24 Apr 1954, Shinners 18584 (SMU). BELL CO. near 
Sparta, 4 May 1931, Wolff 2885 (US). BORDEN CO. red bed lands 
Colorado to Gail, 25 May 1899, Bray s.n. (TEX). BREWSTER CO. hwy 
90 12 mi E of Alpine, 8 Sep 1966, Averett & Watson 161 (SMU, MO, 
TEX). BROWN CO. Brownwood, 31 Mar 1917, Palmer 11439 (vc). 
CALLAHAN CO. hwy 183, 4.2 mi S of Baird, 29 Sep 1950, Cory 58391 
(SMU). CASTRO CO. Nazareth Community, 3 May 1946, Rachaner 32 
(TEX). CHILDRESS CO. Childress, 21 Apr 1968, Shinners 32119 (SMU). 
COLEMAN CO. 1 mi SE of Santa Anna, 9 Jun 1957, Shinners 26386 
(SMU). coTTLE CO. S Pease River, 12 mi NE of Paducah, 29 Apr 
1963, Rowell 8072 (OKLA). COMAL CO. Comanche Springs New Braun- 
fels, ete., Jun 1851, Lindheimer 1037 (GH, NY, SMU, TEX, UC, US). 
COMANCHE co. 8 mi N of Comanche, 6 Apr 1968, Seale 32 (OKLA). 
DALLAM co. Rita Blanca Creek, N of Dalhart, Aug 1949, York & 
Rodgers 252 (OKLA, SMU, TEX). DICKENS CO. 2 mi N of Dickens, 28 
Jun 1964, Rowell 10301 (OKLA). DIMMIT CO. Carrizo Springs, 30 
Apr 1930, Hoaglund s.n. (TEX). DUVAL CO. 6 mi W of Freer, 20 Mar 


362 Rhodora [Vol. 75 


Figure 13. Distribution of Chamaesaracha coniodes. 


1973] Chamaesaracha — Averett 363 


1960, McCart 7188 (OKLA). EASTLAND CO. Rising Star, 11 Dec 1931, 
Potter s.m. (TEX). ECTOR CO. NE Odessa, 28 Mar 1967, Averett 
227 (SMU, MO, TEX). EDWARDS CO. 28 mi W of Rockspring, 18 Apr 
1929, Cory 689 (GH). EL PASO CO. Loyal Valley (Frontera), 22 May 
1852, Buckley 7837 (SMU). ERIO CO. hwy 81, 6 mi N of Dilley, 21 
Mar 1964, Vasquez 50 (SMU, TEX). GARZA CO. Post, 4 Jun 1925, Ruth 
1270 (MICH, US). HALE CO. 4 mi N of Hale Center, 17 Jul 1968, 
Averett & Tomb 354 (SMU, MO, TEX). HALL CO. ca 4 mi from hwy 
287, E of Estelline, 6 Sep 1945, Whitehouse 10736 (MICH, SMU, US). 
HAMILTON CO. 5 mi ESE of Hamilton, 14 Apr 1965, Shinners 30952 
(SMU). HARDEMAN CO. 2 mi W of Quanah, 5 May 1963, Stokes 65 
(OKLA). HARTLEY CO. junct. of hwy 87 & 385 near Hartley, 16 Jul 
1968, Averett & Tomb 351 (SMU, MO, TEX). HEMPHILL CO. 7 mi NE 
of Canadian, 20 Jun 1964, Brown 76 (OKLA). HIDALGO CO. 2 mi S 
of Alamo, 7 Dec 1933, Clover 1482 (MICH, NY). HOCKLEY CO. 10 mi 
E of Levelland, 20 Apr 1964, Forhand s.m. (OKLA). HUTCHINSON CO. 
Canadian River breaks, 4 May 1965, Drake 45 (TEX). JACK CO. 3 mi 
W of Jacksboro, 14 Apr 1946, Whitehouse 15308 (SMU). JEFF DAVIS 
co. hwy 166, 1 mi S of Ft Davis, 26 Jun 1966, Averett 136 (SMU, 
MO, TEX). JIM HOGG CO. 5 mi E of Hebbronville, 16 Mar 1963, Areola 
et al. 119 (SMU, TEX). LIVE OAK CO. hwy 59, 15 mi SW of George 
West, 13 Mar 1965, Rodriquez 17 (ARIZ, TEX). LUBBOCK CO. ca 5 mi 
N of Slayton, 17 Jul 1968, Averett & Tomb 356 (SMU, MO, TEX). 
LYNN CO. 11 mi W of Tahoka, 25 Apr 1954, Shinners 18621 (SMU). 
MAVERICK CO. 30 mi SE of Eagle Pass, 14 Mar 1964, Bruni 13 (OKLA, 
SMU, TEX). MC MULLEN Co. 15 mi N of Freer, 28 Apr 1958, Thomp- 
son & Graham 80 (TEX). MIDLAND CO. 15.6 mi ESE of Midland, 30 
May 1963, Shinners 30101 (SMU). MILLS CO. 2 mi N of Priddy, 26 
Sep 1950, Cory 58175 (OKLA, SMU). MOTLEY CO. 3.3 mi W of Matador, 
28 Apr 1954, Shinners 18693 (SMU). NOLAN CO. Sweetwater, 27 May 
1918, Palmer 13709 (us). OCHILTREE CO. Canadian River, 27 mi S 
of Perryton, 19 Sep 1958, Wallis 7820 (SMU, TEX). OLDHAM CO. Tas- 
cosa, 16 Jun 1932, Reed 3515 (US). RANDALL Co. entrance to Palo 
Duro State Park, 17 Jul 1968, Averett & Tomb 353 (SMU, MO, TEX). 
REEVES CO. S of Pecos, 31 Mar 1942, Nelson & Nelson 5004 (GH). 
ROBERTS CO. Canadian River Valley, 27.5 mi S of Perryton, 6 Jun 
1958, Wallis 7172 (OKLA, TEX). PALO PINTO CO. 9 mi ENE of Gordon, 
6 May 1951, Shinners 13230 (SMU). PARKER CO. 16.5 mi SW of 
Weatherford, Brazos River bottoms, 4 Apr 1948, Shinners 9971 
(SMU). PECOS CO. 1 mi E of Ft Stockton, 19 Apr 1946, Warnock 
46151 (TEX, UC). PRESIDIO CO. Spur Ranch, S of Marfa, 10 Aug 1941, 
‘Warnock 21370 (TEX). SAN PATRICIO CO. near Mathis, 5 Apr 1931, 
MC KELVEY 1724 (GH, US). SAN SABA CO. 20 mi W of Cherokee, 4 
Apr 1966, Henderson 8 (TEX). SHACKELFORD CO. !$ mi N of Callahan 
Co line on hwy 283, 24 Apr 1963, Henderson 63-516 (SMU, TEX). 
STARR CO. 9 mi N of Roma, 13 Apr 1962, Alvarez et al. 8043 (TEX). 


364 Rhodora [Vol. 75 


TAYLOR CO. Camp Barkeley, 25 Oct 1942, Tolstead 5779 (NY, SMU). 
TERRELL CO. 1 mi E of Sanderson, 30 Apr 1949, Warnock & Turner, 
599 (SMU). TERRY CO. 5 mi E of Brownfield, 25 Apr 1954, Shinners 
18618 (sMU). THROCKMORTON CO. 8.3 mi E of Throckmorton, 29 Apr 
1954, Shinners 18806 (SMU). TOM GREEN CO. San Angelo, 3 Jul 1917, 
Palmer 12379 (uc). TRAVIS CO. Onion Creek, near Austin, 17 May 
1918, Young s.n. (TEX). UVALDE CO. Garner State Park, 5 Apr 1969, 
Tomb 391 (TEX). VAL VERDE CO. E city limits, Del Rio, 5 Apr 1969, 
Tomb 401 (TEX). WEBB CO. Laredo, 15 Mar 1917, Palmer 11286 (vc). 
WICHITA CO. Red River 7.4 mi N of Electra, 30 Mar 1945, Whitehouse 
9590 (MICH, SMU, US). YOUNG CO. 3 mi W of New Castle, 2 Apr 1958, 
McCart 873 (SMU). ZAPATA CO. 1.5 mi S of San Ygnacio, 31 Jan 
1954, Shinners 17690 (SMU). ZAVALA CO. near Batesville, 25 Mar 
1961, Stewart 58 (OKLA). 

Mexico: CHIHUAHUA, Mts NW of Chihuahua, 1 Aug 1936, Le Sueur 
47 (OKLA, TEX UC). COAHUILA, 24 mi W of Cd Acuna, 20 Aug 1966, 
Powell et al. 1412 (SRSC, TEX). NUEVO LEON, 12 mi E of Monterrey, 
27 Feb 1944, Barkley 14351 (GH, TEX, UC). 


EXCLUDED SPECIES 


C. potosina Rob. & Greenm. 

— Saracha potosina (Rob. & Greenm.) Averett 
C. boliviensis Dammer 

— Solanum chamaesarachidium (Dammer) Bitt. 
C. grandiflora (Hook.) Fernald 

— Leucophysalis grandiflora (Hook). Rydb. 
C. nana, Gray 

— Leucophysalis nana (Gray) Averett 
C. heterophylla Hemsl. 

— Leucophysalis heterophylla (Hemsl.) Averett 
C. japonica Fr. & Sav. 

— Physaliastrum japonicum (Fr. & Sav.) Honda 
C. echinata Yatabe 

— Physaliastrum japonicum (Fr. & Sav.) Honda 
C. savatieri Makino 

— Physaliastrum savatieri (Makino) Makino 
C. watanabei Y atabe 

— Physaliastrum savatieri (Makino) Makino 
C. sinensis Hemsl. 

— Physalis sinensis (Hemsl.) Averett 
C. physaloides Green 

= Physalis labata Torr. fide Waterfall (1958). 


1973] Chamaesaracha — Averett 365 


LITERATURE CITED 


AVERETT, J. E. 1970. Systematics of Chamaesaracha (Solanaceae) : 
A chemosystematie and cytotaxonomic study. Ph.D. Dissertation. 
University of Texas at Austin. 

1971. New Combinations in the Solaneae (Solana- 
ceae) and comments regarding the taxonomy of Leucophysalis. 
Ann. Mo. Bot. Gard. 57: 380-382. 

AVERETT, J. E. & T. J. MABRY. 1971. Flavonoids of the North 
American Species of Leucophysalis (Solanaceae). Phytochem- 
istry 10: 2199-2200. 

AVERETT, J. E. 1972. Two New Species of Chamaesaracha (Sol- 
anaceae) from Texas. Sida 5: 48-49. 

AXELROD, D. I. 1958. Evolution of the Madro-Tertiary Geoflora. 
Bot. Rev. 24: 433-509. 

FERNALD, M. L. 1949. Contributions from the Gray Herbarium of 
Harvard University. No. CLXIX. Part II. Studies of Eastern 
American Plants. Rhodora 51: 82-83. 

Gray, A. 1876. Synopsis of the North American species of Phys- 
alis. Proc. Amer. Acad. 10: 62. 

1876. Bot. Calif. 1.540. 
1878. Syn. Fl. N. Amer. 2:232. 

Hooker, W. J. 1841. Flora Boreali-American. Vol. 2. 

Lewis, H. 1967. The taxonomic significance of autopolyploidy. 
Taxon 16: 267-271. 

Masry, T. J., K. MARKHAM, & M. B. THoMas. 1971. The system- 
atic identification of Flavonoids. Springer-Verlag, New York. 

MAKINO, T. 1914. Observations on the flora of Japan. The Botani- 

ca] Magazine 28: 20-22. 

PowELL, A. M. & J. AVERETT. 1967. Chromosome numbers of 
Chamaesaracha, (Solanaceae) in Trans-Pecos Texas and adja- 
cent regions. Sida 3: 156-162. 

RYDBERG, P. A. 1896. The North American species of Physalis and 
related genera. Mem. Torr. Bot. Club 4: 279-374. 

TURNER, B. L. & M. C. JoHNSTON. 1961. Chromosome numbers in 
the Compositae III. Certain Mexican species. Brittonia 13: 
64-69. 

WATERFALL, U. T. 1958. A taxonomic study of the genus Physalis 
in North America north of Mexico. Rhodora 60: 113-114. 

WETTSTEIN, R. 1895. Solanaceae in Engler & Prantl, Die Natur- 
lichen Pflanzenfamilien IV (36): 4-39. 


DEPT. OF BIOLOGY 
UNIVERSITY OF MISSOURI 
ST. LOUIS, MO. 63121 


368 Rhodora [Vol. 75 


Isoétes melanopoda Gay. and Dur. 

ALABAMA. CHEROKEE COUNTY: locally abundant 
on clay of low flat places in savanna-pasture by Alabama 9, 
ca. 6 miles south Centerville, 5 June 1969, Kral 35013. 

Not hitherto recorded for Alabama. According to Pfeif- 
fer (Ann. Mo. Bot. Garden 9: 151-152, 1922) it is ex- 
tremely close taxonomically to E. butleri, and intergrades 
with it. It has the same ephemeral habit. On the Alabama 
site, it has collected along with Schoenolirion croceum, 
Fimbristylis puberula, and the very rare Marshallia mohrit. 


Lycopodium alopecuroides L. 

TENNESSEE. COFFEE COUNTY: sandy clay of 
sphagnous area by pond, by Tennessee 55, ca. 5.2 miles 
north of Manchester, 7 August, 1970, Kral 40215. 

This lycopod, here in association with an abundance of 
Rhynchospora perplexa, has previously been collected in 
Tennessee from Coffee and Fentress counties by Shaver. 
It is reported by Wherry (Southern Fern Guide: 288, 
1964) as abundant, though local, on sandy seepage and in 
bogs from Florida north to New York and west in the 
coastal plain into Mississippi. As Wherry states, L. alo- 
pecuroides and L. prostratum Harper are similar enough 
to be treated by some as varieties of the same species. 
However the case, my own. material shows the arching 
stems so characteristic of the former and in sharp contrast 
to the prostrate, rooting habit of the latter. Material 
agreeing with L. prostratum has also been collected from 
Coffee County, Tennessee, by Shaver. 


Sagittaria isoetiformis T. G. Sm. 

ALABAMA. COVINGTON COUNTY: sandy shallows 
of Blue Pond, Blue Springs Wildlife Management Area, 
Conecuh National Forest, southwest of Andalusia, 1 June 
1970, Kral 39506. HOUSTON COUNTY: Indigo Pond, 9 
June 1971, Kral 43119. 

Previously reported for Alabama by Dr. Godfrey (Sida 1 
[5], 1964), and by others, but still infrequent enough to 
deserve mention. This species appears only around lime- 


1973] Southern States Flora — Kral 369 


sink lakes and ponds, and in Alabama is in association with 
the same kinds of rarities (namely Xyris longisepala, 
Rhezia salicifolia, Eriocaulon lineare, Lachnocaulon minus) 
as are found in similar sites in northwest Florida. 


Festuca myuros L. 

ALABAMA. CHEROKEE COUNTY: sandy empty lot, 
Centre, 6 June 1969, Kral 35039. PICKENS COUNTY: 
sandy clearing in oak-pine, 1.5 miles north Carrolltown, 
5 June 1971, Kral 43026. 

An extension of known range westward from Georgia or 
eastward through the gulf south. 


Eragrostis amabilis (L.) Wight and Arn. ex Nees. 

ALABAMA. ESCAMBIA COUNTY: weed on sandy 
grounds of Southland Motel, east side of Atmore, 20 Octo- 
ber 1969, Kral 38071. GENEVA COUNTY: sandy grounds 
of Eunola Motel, east side of Geneva, 13 October 1970, 
Kral 41737. 

In Hitchcock’s Manual of the Grasses of the United 
States, ed. 2 (1950) this grass is not reported for Alabama. 
It is of frequent occurrence in Georgia and Florida, prob- 
ably as an introduction from the tropics. 


Diarrhena americana Beauv. 

Diarina festucoides Raf. 

Not previously reported for Alabama, though reported 
in Hitchcock from Tennessee and the Virginias. 

ALABAMA. JACKSON COUNTY: rich limestone 
woods by upper Paint Rock Creek and Alabama 65, ca. 
7 miles north Princeton, 30 September, 1971, Kral 44597. 


Chasmanthium nitidum (Baldwin in Ell.) Yates 

Uniola nitida Baldwin in Ell. 

ALABAMA. HOUSTON COUNTY: Cowart’s Creek 
bottoms, rich woods, by Alabama 53, ca. 5 miles southeast 
Cottonwood, 16 July, 1970, Kral 40003. 

Reported by Yates (Southwestern Naturalist 11 [4]. 
1966) as occurring from South Carolina through south 
Georgia and Florida, but not reported by him for Alabama. 


310 Rhodora (Vol. 75 


Spartina spartinae (' Trin.) Merr. 

ALABAMA. MOBILE COUNTY: on brackish sands by 
causeway just north of Dauphin Island, 22 September 1969. 
Kral 36368. 

This cordgrass occurs in salt marshes along the gulf 
coast from Florida west to Texas, but no previous reports 
from Alabama have been made. 


Leptochloa fascicularis (Lam.) Gray 

ALABAMA. MOBILE COUNTY: Battleship Park, by 
causeway US 90-98, east of Mobile on the Bay; moist sands, 
20 August 1968, Kral 32754. 

This grass, though previously reported from Mobile 
County by Mohr (Plant Life of Alabama, 1900) was for 
some reason not mapped for Alabama in Hitchcock, where 
it is reported from Mississippi and Florida. 


Leptochloa uninervia (Presl.) Hitchcock and Chase 

ALABAMA. MOBILE COUNTY: sands of large clear- 
ing by Battleship Park, along US 90-98 causeway over 
Mobile Bay, 16 April, 1967, Kral 28449. MORGAN 
COUNTY: exposed drying silt of Flint Creek bottoms, just 
south of Flint on US 31, 22 September 1970, Kral 41193. 

Reported in Hitchcock (op. cit.) as from Mexico, north 
to Oregon in the west; introduced locally in Maine, Massa- 
chusetts and New Jersey, also in the coastal plain from 
Louisiana into Texas. 


Chloris floridana (Chapman) Vasey 

ALABAMA. COFFEE COUNTY: sandy bluffs above 
Pea River, 9.5 miles east Opp, 12 October 1970, Kral 41678. 
ESCAMBIA COUNTY: Sandy Conecuh River bottoms 
6.0 miles east of Boykin off US 29, 4 October 1971, Kral 
44832. GENEVA COUNTY: sandy live oak-longleafpine 
banks of Pea River just west of Samson on Alabama 52, 
19 October 1969, Kral 38045; sandy field at edge of gum 
swamp, ca. 7 miles southeast of Samson on Alabama 87, 
19 October 1970, Kral 38060. 

This Chloris has previously been found only in Florida 
and Georgia, essentially along the sandy banks of coastal 


1973] Southern States Flora — Kral 371 


plain rivers, very frequently in association with live oak. 
There are no prior reports of it from Alabama. 


Deschampsia flexuosa (L.) Trin. 

ALABAMA. BLOUNT COUNTY: mesic woods of N 
facing bluffs above Mulberry Fork of Warrior River where 
crossed by US 20, 13 miles south of Cullman, 10 May 1966, 
Kral 26631. JACKSON COUNTY: arenaceous, open out- 
crops, Flat Rock, 4 June 1969, Kral 34944. 

Reported by Mohr (op. cit.) from Dekalb County in 
Alabama, but not reported for Alabama in Hitchcock (op. 
cit.). A wide ranging hairgrass, essentially throughout 
the Appalachians and from the old mountains of Arkansas 
and Oklahoma. In Alabama and elsewhere it is on arena- 
ceous rock, or the sands derived. 


Melica nitens Nutt. ex Scribn. 

ALABAMA. FRANKLIN COUNTY: open limestone 
glade ca. 5 miles south of Russellville, 28 May 1970, Kral 
39120. BIBB COUNTY: rocky bluffs above Schultz Creek 
1.8 miles north of Centreville off Alabama 5, 15 April 19770, 
Kral 38502. 

An extension of known range into Alabama from Ten- 
nessee and North Carolina. Frequent in the cedar glades 
of middle Tennessee (Marshall, Maury counties). Appears 
to have an affinity for calcareous barrens and bluffs. In 
the Bibb County locality, it is in association with Croton 
alabamensis. 


Aristida tuberculosa Nutt. 

ALABAMA. GENEVA COUNTY: sandy high banks of 
Choctawhatchee River between Geneva and Eunola beside 
Alabama 52, 18 September 1971, Kral 44354, 

Here reported as new for Alabama, though already re- 
ported in Hitchcock (op. cit.) for Georgia and Mississippi. 
A grass of dry sandy areas along the Atlantic and Gulf 
coasts, also inland in the Great Lakes region. 


Muhlenbergia glabriflora Scribn. 
TENNESSEE. MAURY COUNTY: banks of Duck 


312 Rhodora [Vol. 75 


River, rocky bluffs, by I-65, 11.7 miles north northeast 
Lewisburg, 14 August 1971, Kral 43551. 

Here reported as new for Tennessee. Similar to M. 
frondosa, differing from it in the scaberulous nodal areas 
and in the awnless glumes. 


Paspalum minus Fourn. 

ALABAMA. MOBILE COUNTY : sandy peat at edge of 
hillside pineland bog, by US 45, 3.8 miles southeast of 
Chunchula, 3 June 1970, Kral 39624; sandy lot, west side 
of Mobile near junction International Drive and Interstate 
10, 2 September 1970, Kral 41021. 

A species of central America and the West Indies but 
already reported from eastern Texas. A first record how- 
ever for Alabama. 


Reimarochloa oligostachya (Munro) Hitchcock. 
ALABAMA. MOBILE COUNTY: Battleship Park, 
brackish sands by US 90-98 causeway over Bay, 23 Sep- 
tember 1969, Kral 37405. 
Previously reported only from Florida, Cuba. Super- 
ficially similar to Paspalum vaginatum Sw. 


Brachiaria platyphylla (Griseb.) Nash. 

B. extensa Chase 

ALABAMA. PICKENS COUNTY: ca. 2.5 miles north 
of Aliceville; common on sandy clay of banks of Alabama 
17, 3 August 1971, Kral 48486. WILCOX COUNTY: ca. 
4 miles north of Camden on Alabama 41; sandy clay of 
highway bank, 21 August 1968, Kral 32788. 

Previously reported for the state but scarce enough in 
Alabama to be noteworthy. No record of it for Alabama 
appears in Hitchcock (op. cit.). 


Setaria faberi Herrm. 

ALABAMA. LAWRENCE COUNTY: sandy clearing 
in oak-pine by Alabama 33, 8.8 miles south of Moulton, 
22 September 1970, Kral 41250. 

A widespread, aggressive weed, rapidly spreading south- 
ward into Tennessee and North Carolina, just recently 


1973] Southern States Flora — Kral 878 


reported from Georgia, but hitherto not reported for Ala- 
bama. 


Panicum adspersum Trin. 

ALABAMA. COVINGTON COUNTY: sandy road 
shoulder by Pond Creek, just west of Florala, 29 October 
1969, Kral 38151. 

Frequent on coral rock in southern Florida and showing 
up at a variety of U.S. ports, including Mobile, as reported 
by Hitchcock. However, this indication of its establishment 
so far inland is noteworthy. 


Panicum amarum Ell. 

ALABAMA. MOBILE COUNTY: sand dunes, Dauphin 
Island, 3 September 1970, Kral 41041. 

A species of coastal dunes in Florida and Mississippi, 
west into Texas; also on similar situations north into New 
England. Not reported for Alabama by Hitchcock and 
Chase, though reported from Cedar Point in Mobile County 
by Mohr. 


Panicum linearifolium Scribn. 

ALABAMA. LAUDERDALE COUNTY: sandy clay of 
oak flats by US 43, ca. 0.5 miles west of Green Hill, 4 June 
1968, Kral 31052. 

This entity reportedly intergrades with P. werneri 
Scribn. Whatever the taxonomic status, there are no prior 
reports of it from Alabama. 


Panicum malacophyllum Nash. 

ALABAMA. BIBB COUNTY: sandy oak-pine with 
some longleaf, along Alabama 5, ca. 5 miles south of Brae- 
head, 9 May 1966, Kral 26575. 

An extension of known range southward from Tennessee. 


Panicum webberianum Nash. 

ALABAMA. COVINGTON COUNTY: 4.3 miles west of 
Lockhart by county 4 in sandy bulldozed longleaf pineland, 
18 April 1970, Kral, 38757. 

While reported from Mobile County by Mohr, the spe- 
cies is not credited to Alabama by Hitchcock. The species 


374 Rhodora [Vol. 75 


is distinctive within the sect. “Lancearia” in its rather 
broad leafblades together with its papillose fertile lemmas. 


Panicum werneri Scribn. 

ALABAMA. LAUDERDALE COUNTY : sandy clay of 
oak flats by US 43, ca. 0.5 miles west of Green Hill, 4 June 
1968, Kral 31066. 

Probably but a form of P. linearifolium; in any case not 
previously reported for Alabama. 


Amphicarpon muhlenbergianum (Schult.) Hitchcock. 

A. floridanum Chapm. 

ALABAMA. COVINGTON COUNTY: upper sandy 
shores of Open Pond, in Conecuh National Forest, south- 
west of Andalusia, 5 October 1971, Kral 44731. 

This genus produces dimorphic fruit, only the under- 
ground spikelets seemingly developing viable seeds. A first 
report for Alabama for the genus; the species was pre- 
viously known to occur from Florida to South Carolina. 


Manisuris tuberculosa Nash 

ALABAMA. COVINGTON COUNTY: sandy peat of 
longleaf pine savanna 9.5 mi. s. Opp, 5 October 1968, Kral 
33681. GENEVA COUNTY: cypress swamp 5.8 miles 
south of Hartford, 2 September 1969, Kral 36722. HOUS- 
TON COUNTY: sandy peat of longleaf pine flatwoods ca. 
1 mile southeast Cottonwood, 12 October 1970, Kral 41659. 

Close taxonomically to M. rugosa (Nutt.) Kuntze, sep- 
arable only in degree of rugosity of the first glume and in 
the comparative straightness of its rachis joints, and per- 
haps therefore but a variant. Previously known only from 
peninsular Florida. 


Andropogon arctatus Chapm. 

ALABAMA. COVINGTON COUNTY: sandy savanna, 
ca. 12 miles south of Andalusia, 3 October 1971, Kral 
44749. 

Reported in Hitchcock as only from Florida; similar to 
A. ternarius but racemes duller, more grayish, and less 
obscured by shorter hairs. 


1973] Southern States Flora — Kral 375 


Andropogon divergens (Hack.) Anderss. ex Hitchcock 
ALABAMA. ESCAMBIA COUNTY: pitcher plant bog 
southwest corner of county, Perdido, 5 October 1971, Kral 
44892. MOBILE COUNTY: 10.7 miles south Theodore on 
Alabama 59, wet flatwoods 8 October 1967, Kral 29734. 
An extension of known range eastward from the low 
pinelands of Texas and Mississippi. 


Andropogon capillipes Nash 

ALABAMA. BALDWIN COUNTY: sandy pine flats due 
east of Orange Beach by Alabama 180, 22 October 1969, 
Kral 38250. GENEVA COUNTY: ca. 5 miles south south- 
east of Hartford on Alabama 167, sandy savanna, 13 Octo- 
ber 1970, Kral 41714. 

Very distinctive because of its dullish, short racemes, 
and its highly glaucous foliage. Reported in Small to occur 
from Florida to Mississippi and North Carolina, but not 
reported for Alabama by Hitchcock. 


Andropogon hirtiflorus (Nees) Kunth 

ALABAMA. BALDWIN COUNTY: west junction Ala- 
bama 59 and Alabama 180 near Gulf Shores Golf Course, 
along Alabama 180; sandscrub, 9 October 1967, Kral 29851. 
GENEVA COUNTY: savanna by junction Alabama 153 
and county 4, south of Ganer and southwest of Samson, 
18 September 1971, Kral 44517. 

This plant of the pine flatwoods of southern Georgia and 
of Florida is here recorded as new for Alabama. 


Andropogon littoralis Nash 

ALABAMA. BALDWIN COUNTY: sandhills, Romar 
Beach, 22 October 1969, Kral 38218. Reported by Hitch- 
cock and Chase as from sandy coasts along the Atlantic 
and intermittently from the Great Lakes, also from south- 
eastern Texas. However, fairly common on coastal] dunes 
of Baldwin County. 


Andropogon niveus Swallen 
GEORGIA. LOWNDES COUNTY: ca. 4 miles north of 
Alapaha on River bottoms, 27 July 1967, Kral 28672. 
Reported previously only from central Florida. 


376 Rhodora [Vol. 75 


Andropogon perangustatus Nash 

ALABAMA. BALDWIN COUNTY: sandy splash pine 
flats due east of Orange Beach by Alabama 180, 22 Octo- 
ber 1969, Kral 38228. 

Reported in Hitchcock (op. cit.) as from bogs and moist 
pine woods of Florida and Mississippi but not from Ala- 
bama. 


Scirpus purshianus Fernald 

S. debilis Pursh 

ALABAMA. CHEROKEE COUNTY: sandy, coldwater 
slough by campgrounds, Canyon Mouth State Park, by 
Little River, 3 September 1971, Kral 43817. MISSISSIPPI 
ITAWAMBA COUNTY: 0.5 miles east of Fulton City 
limits by US 278; wet clay along pond edge, 28 July 1966, 
Kral 27269. 

Previously not reported from either Alabama or Missis- 
sippi. 

Fuirena pumila Torr. 

ALABAMA. COVINGTON COUNTY: sandy moist 
shores of Open Pond, Conecuh National Forest, southwest 
of Andalusia, 5 October 1971, Kral 44735. TEXAS. HAR- 
RIS-LIBERTY COUNTY: sandy shores of Lake Houston 
where crossed by causeway of county farm road 1960, 
2 September 1965, Kral 25804. 

Seemingly first records for this small, annual Fuirena 
for Alabama and Texas; the species is particularly abun- 
dant on moist sandy disturbed situations in the lake areas 
of Florida but ranges northward throughout the Atlantic 
coastal plain. 


Eleocharis melanocarpa Torr. 

ALABAMA. COVINGTON COUNTY: sandy shores of 
Blue Pond, Blue Springs Management Area, Conecuh Na- 
tional Forest, southwest of Andalusia, 18 April 1970, Kral 
38728; 1 June 1970, Kral 39500. 

A first report for Alabama, though already reported 
from states east and west in the coastal plain. 


1973] Southern States Flora — Kral 377 


Eleocharis olivacea Torr. 

ALABAMA. BALDWIN COUNTY: almost entirely sub- 
mersed in swift flowing sandy creek through gum swamp 
by I-65, 4 miles northeast junction Alabama 59 southwest 
of Atmore, 19 September 1971, Kral 44397. ESCAMBIA 
COUNTY: seepage slope, west side of Brewton, 4 October 
1971, Kral 44778. 

According to Svenson (Rhodora 31: 231-232, 1929) this 
species ranges from N. S. to Ontario, southward to Florida, 
western Pennsylvania, Ohio and Michigan but chiefly on 
the Atlantic Coastal Plain. This is a first report of it from 
Alabama. A specimen that is probably this species and 
gotten from Mobile area is at the Mohr Herbarium but 
Mohr did not identify it, so did not include it in his Flora. 
Some authors, the latest being Correll and Johnston (Man- 
ual of the Vascular Plants of Texas, p. 274, 1970) place it 
in the synonymy of E. flavescens (Poir.) Urban. It is how- 
ever distinguishable from E. flavescens on the basis of its 
greenish akenes, the cusp-like prolongation of the tubercle 
of the akene, and by the blunter, greenish-ribbed spikelet 
bracts. The Baldwin County material shows a very inter- 
esting trailing, lax submersed habit, though material] gotten 
from wet sandy shores is similar to E. flavescens super- 
ficially. 


Eleocharis robbinsii Oakes 

ALABAMA. GENEVA COUNTY: shallows of savanna 
pond, by Alabama 4 ca. 10 miles east of Florala, 2 August 
1971, Kral 43441. 

Previously reported from Florida and Georgia and inter- 
mittently northward and westward, but not for Alabama. 


Eleocharis tricostata Torr. 

ALABAMA. COVINGTON COUNTY: sandy peat of 
flatwoods by gum bay, 13 miles south Opp, 1 June 1970, 
Kral 39447. GENEVA COUNTY: sandy peat of savanna 
ditch ca. 3.5 miles west of Geneva, 11 June 1971, Kral 
43288. HOUSTON COUNTY: sandy bands of Indigo 
Pond, southeast Cottonwood, 9 June 1971, Kral 43125. 


378 Rhodora [Vol. 75 


A first report for Alabama, though the species is listed 
from both east and west of Alabama within the coastal 
plain. 


Eleocharis wolfii Gray 

ALABAMA. FRANKLIN COUNTY: moist swales in 
calcareous barrens ca. 5 miles east of Russellville beside 
Alabama 24, 4 June 1971, Kral 42932. 

Essentially a northern plant, according to Fernald 
(Gray’s Manual, 8th ed., 1950, p. 253) sporadically in Ten- 
nessee, Louisiana. A first report for Alabama. 


Bulbostylis capillaris (L.) Clark 

Stenophyllus capillaris (L.) Britton 

FLORIDA. JACKSON COUNTY: Just south of Florida 
line north of Malone on Florida 71-Alabama 53; sandy 
disturbed pocosin edge, 1 August 1971, Kral 43420. 

A second authentic report of this widespread sedge for 
Florida, the earliest being made from a Chapman specimen 
gotten from near Apalachicola. 


Fimbristylis perpusilla Harper 
GEORGIA. SEMINOLE COUNTY: silty margin of 
pond in slash pine savanna-pasture beside US 84, ca. 1.5 
miles east of Donaldsonville, 1 August 1971, Kral 43421. 
Noteworthy primarily for its sporadic occurrence even 
at the same locality. Its history of collection is peculiar. 
Harper collected the type in 1903 in Sumter County, Geor- 
gia, where it was very abundant on the drying margins of 
a flatwoods pond very similar to the one in Seminole 
County. He made several trips to the same locality in sub- 
sequent years but never again found the plant. My dis- 
covery of it in Seminole county in July 1962 turned out to 
be the second authentic record (Kral 15486). The follow- 
ing year Dr. Godfrey visited the Seminole county area and 
found a few plants (Godfrey 63204). However, in spite of 
several trips to that place made subsequently either by Dr. 
Godfrey or myself, the plants were not in evidence. Yet 
in 1971, seven years later, it was again in great abundance 
in the same locality. As was true in 1962, there were sev- 


1973] Southern States Flora — Kral 379 


eral other small ponds in the same area, any of these seem- 
ing to have the same conditions for growth, but none actu- 
ally having the plants. As I stated in my treatment of 
Fimbristylis (Sida 4: 126, 1971), F. perpusilla appears to 
have such a refined niche that conditions for successful 
germination of its seed are but rarely met. It also becomes 
evident from this one case that its seed may remain dor- 
mant in large quantity for at least 8 years, and with no 
apparent loss of viability. This is but one of many peculiar 
examples of the sort of complex niches occupied by aquatic 
and marsh plants, the study of which should be immensely 
rewarding to ecologists and biosystematists. 


Cyperus dentatus Torr. 

ALABAMA. COVINGTON COUNTY: shallow water at 
sandy edge of Open Pond, Conecuh National Forest, south- 
west of Andalusia, 3 October 1971, Kral 44736. 

This sedge is essentially northern, in the Atlantic coastal 
plain occurring south into South Carolina. Mohr (loc. cit., 
p. 391) reported it from Mobile County as an adventive 
from the northern states, but subsequent workers have not 
cited it from Alabama. 


Cyperus difformis L. 

ALABAMA. LIMESTONE COUNTY: abundant on al- 
luvial flats, bottoms of Tennessee River, north of Dekatur, 
Wheeler National Wildlife Refuge, 22 September 1970, Kral 
41221. 

A sedge of the Old World tropics reported by Gleason 
and Cronquist (Manual of Vascular Plants, p. 125, 1963) 
as adventive in southeastern Virginia; reported from 
Louisiana (Delahoussaye 71) and from California, but a 
first record for Alabama. 


Dichromena colorata (L.) Hitchcock 
ALABAMA. BIBB COUNTY: swale beside dolomite 
outcrop, also in pockets of dolomite, by Alabama 5, 3.6 
miles north of Centerville, 29 May 1970, Kral 39316. 
Noteworthy because of its occurrence so far from the 
present coast, evidence to the fact that some coastal plain 


380 Rhodora [Vol. 75 


species still persist along a geologically very old coast. 
This species is ordinarily not found in Alabama save in the 
very coastal counties (Mobile, Baldwin) and there is con- 
fined usually to the inland sides of brackish marsh areas. 
The only other species in my area is D. latifolia, a species 
of more acid situations inland in the lower coastal plain, 
but with one very interesting disjunction into Tennessee 
(A. Gattinger, Flora of Tennessee, 1900). 


Rhynchospora fernaldii Gale 

ALABAMA. BALDWIN COUNTY : sandy peat of swale 
in sand pine hills, Romar Beach, 22 October 1970, Kral 
38192. 

Already reported from Alabama (Buckley 35) but with 
no definite locality by Shirley Gale (Rhodora 46: 1944). 
Here reported simply to establish a definite locality. The 
species is essentially in Florida. 


Rhynochospora pleiantha (Kuk.) Gale 

ALABAMA. COVINGTON COUNTY: sandy beaches, 
Blue Pond, by recreation area, Blue Springs Wildlife Man- 
agement area, Conecuh National Forest, southwest of An- 
dalusia, 1 June 1970, Kral 39499. 

Reported by Gale (loc. cit.) as from southeast North 
Carolina south to Florida and Cuba but not by her from 
Alabama. 


Carex communis Bailey 

TENNESSEE. DAVIDSON COUNTY: 7.6 miles north- 
east junction Tennessee 12 along Little Marrowbone Creek; 
shaley bluffs, 16 May 1971, Kral 42602. 

Already reported from east Tennessee by Sharp et al. 
(Preliminary Checklist of Monocots in Tennessee, 1960) 
but a first report from middle Tennessee. 


Carex austro-caroliniana Bailey 

ALABAMA. JACKSON COUNTY: shaded sandy loam 
of Pisgah Gorge, south side of Pisgah, 24 April 1969, Kral 
34406. 
_A first report for Alabama of this southern Appalachian 
Carez. 


1973] Southern States Flora — Kral 381 


Carex microdonta Torr. 

ALABAMA. WILCOX COUNTY: chald prairie ca. 2.5 
miles north of Furman, 15 April 1970, Kral 38542. TEN- 
NESSEE. WILSON COUNTY: Cedars of Lebanon State 
Park, west of US 231, 15 miles north of Murphreesboro, 
J. Baskin and C. Caudle 230. 

An extension of known range eastward from Mississippi 
and probably including material of C. crawei Dewey re- 
ported from Alabama and Tennessee. A plant of calcareous 
prairies, hence probably much more common in the lime- 
stone or chalk openings of these two states than present 
collection records indicate. 


Carex tenax Chapman 

C. validior Mackenzie 

ALABAMA. AUTAUGA CO.: sandy longleaf pineland 
by US 82, 21.5 mi. n.w. Prattville above Little Mulberry 
Creek, 30 May 1970, Kral 39365. CHOCTAW COUNTY: 
longleaf pine sand hills by Alabama 17, 4.6 miles south of 
Butler, 4 June 1970, Kral 39660. ESCAMBIA COUNTY: 
sandy longleaf pineland by US 29, ca. 7 miles east of Brew- 
ton, 7 June 1968, Kral 31253. HOUSTON COUNTY: sand- 
hills above Little Choctawhatchee River by Alabama 92, 
1.7 miles west junction US 84, 9 June 1971, Kral 43151. 

This Carex, in section *Triquetrae", which is comprised 
of but two species, is here listed tentatively as a record for 
Alabama. The other species of the complex, C. dasycarpa 
Muhl, is in Alabama, but in very different situations, 
namely moister sands of the beech-magnolia forests. 


Xyris difformis Chapman var. floridana Kral 

ALABAMA. HOUSTON COUNTY: sandy peat of low 
place in savanna, southeast side of Cottonwood, 16 July 
1970, Kral 39993. 

A first report of this variety from Alabama, though it is 
known to range west to Louisiana and north in the Atlantic 
coastal plain to North Carolina. 


Xyris fimbriata Ell. 
TENNESSEE. COFFEE COUNTY: peaty shores and 


382 Rhodora [Vol. 75 


shallows of Goose Pond, just east of Arnold Center, 17 
August 1970, Kral 40682. 

This is probably a record station inland for X. fimbriata, 
though it does get out of the coastal plain in Georgia and 
Alabama. A definite first record for Tennessee. In this 
particular location growing mixed with Xyris iridifolia 
and Lachnanthes caroliniana. 


Xyris jupicai Ell. 

TENNESSEE. PUTNAM COUNTY: boggy places in 
hardwood bottom, 6.9 miles south of Cookeville on Tennes- 
see 42, 14 August 1970, Kral 40589. 

A first record for Tennessee, probably an extension 
northward from northern Alabama. Primarily a wetlands 
weed of the coastal plain, Florida north to New Jersey and 
west to Texas. Widespread in the tropics of the New 
World. 


Xyris longisepala Kral 

ALABAMA, COVINGTON COUNTY: sandy shores and 
shallows of Blue Pond, Blue Springs Wildlife Management 
Area, Conecuh National Forest, southwest of Andalusia, 
31 August 1970, Kral 40935. 

A first Alabama record for this rare endemic, previously 
thought (Sida 2: 246, 1966) to occur only in northwest 
Florida. Very similar to X. smalliana, and at this locality 
mixed with it, but with distinctively different seed char- 
acter and blooming earlier in the day. 


Xyris platylepis Chapman 

TEXAS. JASPER COUNTY: 9 miles south of Brooke- 
land on US 96; seepage area in longleaf pineland, 28 
August 1967, Kral 29058. 

Probably the first authentic record from Texas of this 
species whose range previously was known to extend from 
Florida north to Virginia, and west into Louisiana (adven- 
tive in Hawaii). 


Eriocaulon lineare Small 
ALABAMA. BALDWIN COUNTY: peaty edge of hy- 
pericum pond just north junction I-10 on Alabama 59 and 


1973] Southern States Flora — Kral 383 


6 miles south of Stapleton, 8 June 1971, Kral 43805. 
COVINGTON COUNTY: wet sands and shallows of north 
shore Lake Jackson, Florala, 3 September 1969, Kral 
36821; sandy beaches, Blue Pond, southwest of Andalusia, 
1 June 1970, Kral 39494. HOUSTON COUNTY: Indigo 
Pond, southeast Cottonwood, 9 June 1971, Kral 43124. 

This species, while already reported for Alabama by 
Harper, is rare there. It is part of a group of species that 
frequents sandy shores of sinkhole lakes and ponds, in 
contrast to its nearest relative taxonomically, E. texense, 
which frequents acid pineland bogs from Texas to Georgia 
and which has usually died back by the time E. lineare 
comes into bloom and seed. 


Lachnocaulon digynum Korn. 

ALABAMA. CONECUH COUNTY: sandy longleaf pine- 
land bog 3 miles west of Castleberry, 1 September 1970, 
Kral 40960. ESCAMBIA COUNTY: 1.3 miles east Waw- 
beek on US 31; sphagnous swale in longleaf pine sandhills, 
7 October 1968, Kral 33835. MOBILE COUNTY: pine flat- 
woods bog between Fowl River and Delchamps on co. 59, 
15 July 1969, Kral 35642. WASHINGTON COUNTY: 
boggy bottom on county 28, ca. 4 miles southwest of St. 
Stephens 14 July 1969, Kral 35556. 

This diminutive species, while already reported for Ala- 
bama by Moldenke (Eriocaulaceae. N. Amer. 19 (1), 
1937) is scarce enough in that state to be noteworthy. It 
ranges in the lower terraces of the coastal plain from 
Florida west to southern Mississippi, but is nowhere 
abundant. 


Lachnocaulon engleri Ruhl. 

ALABAMA. BALDWIN COUNTY: peaty swale in slash 
pine-sand pine woods near estuary, Gulf Shores, 19 August 
1968, Kral 32646. 

This species was until now known only for Florida, 
where it is locally abundant on sandy lakeshores, particu- 
larly in areas of karst topography. 


384 Rhodora [Vol. 75 


Lachnocaulon minus (Chapman) Small 

ALABAMA. COVINGTON COUNTY: sandy shores of 
Blue Pond, southwest of Andalusia in Conecuh National 
Forest, 1 June 1970, Kral 39503. HOUSTON COUNTY: 
sandy edges of Indigo Pond, southeast of Cottonwood, 9 
June 1971, Kral 43120. 

A first report for Alabama for this species whose range 
has been thought to be in the lower coastal plain from 
Florida north to North Carolina. 


Juncus brachycephalus (Engelm.) Buchenau 

TENNESSEE. LEWIS COUNTY: seepage bog in oak- 
hickory, by Tennessee 99, 3.5 miles southwest of Hamp- 
shire, 26 August 1969, Kral 36371. MAURY COUNTY: 
seepage area on bluffs above Big Bigby Creek just south- 
west of Rockdale on US 43, 6 October 1967, Kral 29393. 
WAYNE COUNTY: 5.5 miles east of Waynesboro off US 
64; gravelly streambank in oak-hickory hills, 4 September 
1971, Kral 43947. 

This essentially northern rush has not been reported 
previously from further south than Indiana and Illinois. 
In the abovementioned Tennessee localities it is extremely 
abundant, often in association with masses of Parnassia. 


Juncus nodatus Coville 

ALABAMA. PICKENS COUNTY: backwater slough, 
bases submersed, 4.6 miles south of Aliceville off Alabama 
17, 5 June 1971, Kral 43022. 

This is an essentially western gulf coastal plain Juncus, 
previously known from no further east than Mississippi, 
with stations inland from Indiana west to Kansas. 


Allium stellatum Ker 

TENNESSEE. DAVIDSON COUNTY : cedar glade, east 
Nashville, across from Mt. View School, Baskin and Caudle 
5-b, September 25, 1965; RUTHERFORD COUNTY : cedar 
glade 4 miles northeast of La Vergne, 28 September 1946, 
E. Quarterman 1874; 23 September 1950, E. Quarterman 
4227. 

This species, an extension southward or eastward from 


1973] Southern States Flora — Kral 385 


the prairie provinces, distinguishable from A. cernuum, 
on the basis of its shorter bulbs, erect flowering scapes (in 
contrast to distal nodding in A. cernuum), and its narrower 
and more spreading perianth segments, is locally abundant 
in the limestone barrens of Middle Tennessee, where its 
height of flowering is usually from mid-September well into 
October. Putative intermediates between it and A. cernum 
are abundant in the same area. 


Lachnanthes caroliniana, (Lam.) Dandy 

Gyrotheca tinctoria (Walt.) Salisb. 

TENNESSEE. COFFEE COUNTY: peat and muck 
along edge of Goose Pond, just east of Arnold Center, 17 
August 1970, Kral 40679. 

This species so typical of bogs and flatwoods of the lower 
Atlantic and Gulf coastal plain, has not been reported pre- 
viously from Tennessee. Indeed this record might consti- 
tute the furthest one inland for the species, further proof 
of the strong coastal plain floristic affinities of the “oak 
barren” country of middle Tennessee. 


Hypoxis juncea J. E. Sm. 

ALABAMA. GENEVA COUNTY: sandy peat of pine- 
land bog edge, between Geneva and Samson on Alabama 52, 
27 April 1969, Kral 34551. 

An extension of known range westward from Florida 
and Georgia. 


Canna flaccida Salisbury 

ALABAMA. MOBILE COUNTY: inland edge of salt 
marsh, in slash pine, Bayou La Batre, 7 June 1971, Kral 
43053. 

A first report for Alabama of this distinctive, yellow- 
flowered Canna that has been known previously from Flor- 
ida north to South Carolina and also from coastal Missis- 
sippi. 

Salix rigida Michx. 
ALABAMA. PIKE COUNTY: banks of Whitewater 


Creek 5 mi. e.s.e. Spring Hill on Ala. 6, 30 Mr. 1968, Kral 
29967. 


386 Rhodora [Vol. 75 


Sufficiently rare in Alabama (though reported for the 
state by Small, 1933) as to be worth mention here. 


Boehmeria nivea (L.) Gaudich. 

Ramium niveum (L.) Small 

ALABAMA. COVINGTON COUNTY: weedy subshrub 
of sandy empty lot by US 331, s. side Opp, 12 October 1970, 
Kral 41696. ESCAMBIA COUNTY: weedy subshrub, At- 
more, 19 September 1971, Kral 44413. 

Reported by Mohr as an occasional escape from cultiva- 
tion in southern Alabama, but infrequent enough to deserve 
mention here. 


Polygonum erectum L. 

ALABAMA. JACKSON COUNTY : rich limestone woods 
by upper Paint Rock Creek & Ala. 65, ca. 7 mi. n. Princeton, 
30 September 1971, Kral 44588. 

A first report for Alabama of this more northerly weed. 


Polygonella fimbriata (Ell.) Horton 

Thysanella fimbriata (Ell.) Gray 

ALABAMA. GENEVA COUNTY: sandy, live oak for- 
ested west banks of Choctawhatchcee River between Geneva 
and Eunola beside Ala. 52, 18 September 1971, Kral 44348. 

Reported by Small for Alabama, but not by Horton for 
the state in his revision of Polygonella (Brittonia 15: 177- 
203, 1963). The Small citation is based on a collection used 
by Mohr, made by Dr. E. A. Smith from Geneva County, 
perhaps from the same locality. 


Persicaria bicornis (Raf.) Nieuwl. 

Polygonum longistylum Small 

ALABAMA. LIMESTONE COUNTY: gravelly creek 
bottoms by US 31, 1 mi. n. Athens and ject. Ala. 127, 
3 October 1968, Kral 33471. 

Range extended eastward from Mississippi. 


Antigonon leptopus H. & A. 
Corculum leptopus (H. & A.) Stuntz. 
ALABAMA. MOBILE COUNTY: naturalized along rail- 


1973] Southern States Flora — Kral 387 


way, s. side Mobile in old part of town near Bay, 23 Sep- 
tember 1969, Kral 37385. 

This showy species, called “Coral Vine" in Florida, is 
frequently cultivated and escaped there; it is often found 
in cultivation in the gulf coastal plain toward the actual 
coast. In the Mobile area, it persists around old homesites 
and in the negro parts of the old city, but was not reported 
from there by Mohr. 


Silene gallica L. 


ALABAMA. MOBILE COUNTY: sands of large clear- 
ing by Battleship Park, along US 90-98 causeway over 
Mobile Bay, 16 April 1967, Kral 28441. 

Range extended southward from Kentucky and Tennessee 
or over from Georgia. 


Tunica prolifera (L.) Scop. 

Dianthus prolifer L. 

ALABAMA. MARION COUNTY: bluffs on North Fork 
Creek by Ala. 17, n. of Hamilton; sandy roadbank of Ala. 
17, 20 July 1970, Kral 40202. 

A first report for Alabama of this European weed which 
appears (according to Fernald, op. cit.) to be naturalizing 
in the northeast. 


Thalictrum mirabile Small 


TENNESSEE. OVERTON COUNTY: bluffs above West 
Fork of Obey River by Tenn. 52, 3.4 mi. e. Alpine, 31 April 
1971, Kral 42321. 

A first report for Tennessee of this rare species, previ- 
ously found only in Kentucky and Alabama. 


Thalictrum debile Buckl. 


ALABAMA. GREENE COUNTY: blackland woods off 
US 43, 2.7 mi. s. Eutaw, 1 April 1968, Kral 30006. LAW- 
RENCE COUNTY: silty clay of Town Creek bottoms n.w. 
Hatton by where crossed by Ala. 157, 10 April 1969, Kral 
34151. 

This unique little species of heavy clay soils, usually in 
prairie border woods or alluvial woods, has its type locality 


388 Rhodora [Vol. 75 


in Wilcox County. It is similar in habit to T. dioicum, but 
more delicate, and its rootstock consists of a dense fascicle 
of inch-long, fusiform tubers (very similar to those of 
Anemonella). It flowers in March and April, and is usually 
gone back to rootstock by late May. Taxonomically it is 
closest to T. texanum (gray) Small. This complex, which 
includes some Mexican species as well, should be appraised 
further. 


Thalictrum macrostylum Small & Heller 

ALABAMA, JACKSON COUNTY: low sandy woods by 
Ala. 71, ca. 2 mi. s. Pisgah, 4 June 1969, Kral 34979. 

An extension of known range southward from Tennessee. 


Cardamine douglassii (Torr.) Britt. 

ALABAMA. COLBERT COUNTY: clay loam of bluffs 
above Tennessee River at Muscle Shoals, ca. 0.5 mi. w. of 
US 43 bridge crossing, 9 April 1969, Kral 34091. JACK- 
SON COUNTY: rich wooded slopes by Ala. 65, 5.1-7.1 mi. 
s. Huntland, 18 April 1971, Kral 42095. LAWRENCE 
COUNTY: clay of wooded slope above tributary stream 
above Tennessee River near paved county rd., ca. 15 mi. e. 
Joe Wheeler Dam, 30 March 1971, Kral 41498. MARENGO 
COUNTY : woods, n. facing slope, 1 mi. n. Linden by US 43, 
1 April 1968, Kral 30025. 

Reported here as new to the Alabama flora. Mr. Richard 
Simmers of Ithaca, N.Y., an excellent field biologist now 
active in Alabama, has records for the species from Clarke 
and Madison counties. The pubescence character that is 
fairly consistent in populations of this species in middle 
Tennessee varies greatly in the northern Alabama popula- 
tions, so that some individuals of a given population are 
usually completely glabrous. Petal color also ranges more, 
from pure white to the deep fine pink that is typical. It 
would appear that there is intergradation with Cardamine 
bulbosa (L.) B.S.P.; the complex should be reappraised. 


Diplotaxis muralis (L.) DC 
ALABAMA. MARENGO COUNTY: 2.3 mi. s. Demop- 
olis on US 43, chalk prairie, 6 June 1968, Kral 31217. 


1973] Southern States Flora — Kral 389 


HALE COUNTY: chalk glade, 4.2 mi. w. Faunsdale, 9 
September 1968, Kral 33087. | 

A European weed, heretofore not reported from Alabama, 
and rarely from the Small's Manual range. The flowers, a 
bright yellow, are quite showy against the white of the 
chalk barrens it frequents. 


Warea amplexifolia (Nutt.) Nutt. 

W. sessilifolia Nash 

ALABAMA. PIKE COUNTY: sandy clearings amongst 
longleaf pine, Big Pocosin, s.e. Troy, n.e. Fairgrounds, 10 
September 1968, Kral 33165. 

According to Small, the only species of Warea known to 
occur in Alabama is W. cuneifolia, a species whose leaves 
broaden upward. The Pike county material has distinctive, 
sessile, ovate blades, and agrees in all other respects with 
material of W. amplexifolia I have from western Florida. 


Polanisia dodecandra (L.) DC 

TENNESSEE. HICKMAN COUNTY: gravelly banks 
of Beaverdam Creek s. of Centerville on Tenn. 100, 20 Sep- 
tember 1968, Kral 33430. HOUSTON COUNTY: banks of 
Yellow Creek 7 mi. w. Van Leer, 2 August 1969, Kral 
35860. HUMPHRIES COUNTY: banks of Big Richland 
Creek by Tenn. 13, 6 mi. n. Waverly, 4 August 1969, Kral 
35076. 

Iltis (Brittonia 10:46, 1958) shows in his excellent mono- 
graph but two counties in western Tennessee for this spe- 
cies. It appears to be quite abundant in middle Tennessee 
in that additional records of it from Cheatham, Davidson 
and Lewis counties, but not yet processed, have been gotten. 
In the area of this study, the species is almost always on 
gravel bars of streams. 


Fothergilla major Lood. 

ALABAMA. CHEROKEE COUNTY: boulder strewn, 
sandy, wooded banks of Little River Canyon, Canyon 
Mouth branch of De Soto State Park, 12 April 1969, Kral 
34269. 

Reported long previously from Alabama by Small (op. 


392 Rhodora [Vol. 75 


A first report for Alabama, but appears already well 
established in Mississippi. Another clover of Old World 
origin that is being experimented with by agronomists here. 


Geranium robertianum L. 


TENNESSEE. SMITH COUNTY: shaded limestone 
cliffs above Cumberland River, 3.8 mi, n. Chestnut Mound 
off Tenn. 53, 30 April 1971, Kral 43249. 


A first report for Tennessee of this essentially northern 
species. According to Fernald the species is introduced 
from Eurasia, so that its appearance in the rugged, to all 
appearances undisturbed, Smith County location is sur- 
prising. 


Linum westii Rogers 


FLORIDA. FRANKLIN COUNTY: peaty shallows of 
ditch in slash pine-saw palmetto flatwoods near tidal estu- 
ary, 8.2 mi. n. jct. Fla. 65 and US 98, 9 July 1970, Kral 
39896. 


C. M. Rogers (Brittonia 15: 1963) reports this as one 
of the rare plants of the south and indicates that most of 
the collections examined in his study were gotten in Florida 
by Chapman. Only one of these had definite locality, 
"Apalachicola", which seems to have meant a large part 
of Franklin County to Chapman. The type of the species 
is from the only other Florida collection, this made from 
far to the east in Baker County by West and Arnold in 
1946. In addition Rogers cites one collection from Rome, 
in Floyd County, Georgia, but with some reservations. In 
that this particular specimen is again from Chapman, and 
in that flatwoods habitat of the sort L. westii frequents is 
so far removed from Rome, Georgia, one might suspect a 
label mix-up. 

I discovered this Linum whilst travelling north toward 
Telogia along the lonely stretches of Fla. 65, and was at 
once struck by the large patches of handsome, pale yellow 
corollas, these fully open at ca. 6:00 p.m. As Rogers sus- 
pected and mentioned in his description, it is a perennial. 


1973] Southern States Flora — Kral 393 


It spreads by basal offshoots, but tends also to form one 
rootstock several ascending, strict, flowering shoots. 


Acalypha setosa Richard 
ALABAMA. ESCAMBIA COUNTY: lot weed, sandy 
ground, Atmore, 19 September 1971, Kral 44404. LEE 
COUNTY: lawn weed, Opelika, 16 July 1971, Kral 43370. 
Already reported by L. A. W. Miller (Thesis, 1970) for 
Alabama, but shown only for Mobile County. 


Vitis munsoniana Simpson 

ALABAMA. ESCAMBIA COUNTY: sandy bottoms of 
Escambia Creek where crossed by I-65, n.e. Atmore, 27 July 
1969, Kral 35636. 

Extension of known range westward from Florida and 
Georgia. 


Sida elliottii T. & G. 

ALABAMA. BIBB COUNTY: calcareous rock outcrop 
area ca. 5 mi. n. Centerville on Ala. 5, 5 September 1970, 
Kral 41092. 

This has been reported from Alabama since Mohr, but 
it is not at all a common species there or anywhere else 
within its range. I am of the opinion that narrow-leaved 
material of other species is often referred erroneously to 
this taxon. To those like myself who have had experience 
primarily with the commoner southeastern sidas, a first 
experience with this one is startling. The plants of the 
Bibb Co. locale had their flowers fully open at ca. 4:00 p.m. 
The corollas are particularly striking, being of an intense 
orange-yellow (far more orange than other eastern species 
I have seen). Most of the plants had several strong, but 
slender and decumbent shoots from a central, stout, very 
large caudex. The overall effect of these particular plants 
was quite harmonious, and were the species a heavier 
bloomer, it would make a very good horticultural risk. The 
open corollas are fully 2.5 cm. broad, much larger than 
those of other eastern species. 


Cuphaea carthaginensis (Jacq.) MacBr. 
ALABAMA. MOBILE COUNTY: between Fow] River 


394 Rhodora [Vol. 75 


and Delchamps on Ala. 59; sandy roadbank, 5 October 
1971, Kral 44869. 

A first report for Alabama; fairly common as a weed of 
moist sandy ground in southern Georgia, northwestern 
Florida, and southern Mississippi. 


Rhexia salicifolia Kral & Bostick 

ALABAMA. COVINGTON COUNTY: sandy shores of 
Open Pond, Blue Spring Mgmt. Area, Conecuh Nat. Forest, 
s.w. Andalusia, 31 August 1970, Kral 40928; Blue Pond, 
s.w. Andalusia in Coneeuh Nat. Forest, 31 August 1970, 
Kral 40932. HOUSTON COUNTY : sandy shores and shal- 
lows of Indigo Pond, s.e. Cottonwood, 9 June 1971, Kral 
43118. 

An extension northward from northwestern Florida, 
where it is not rare on the shores of limesink lakes and 
ponds. In Alabama, as in Florida, it often is in association 
with Rhexia mariana var. mariana and sometimes produces 
hybrids with it (Kral 40929, 40937). These usually form 
few seeds, the capsules more often falling off before ripen- 
ing. In this they are similar to hybrids between R. mariana 
and R. virginica. 


Epilobium ciliatum Raf. 

TENNESSEE. GILES COUNTY : by Tenn. 11, 3 mi. n.e. 
Minor Hill; wet meadow in shaley hills, 17 September 1969, 
Kral 36940. 

An extension southward from Kentucky. 


Ludwigia spathulata T. & G. 

Isnardia spathulata (T. & G.) Small 

ALABAMA. BALDWIN COUNTY: just s. Perdido on 
county 47 on drying bottoms of sandy pond, 21 October 
1969, Kral 38179. BARBOUR COUNTY: : sandy open seep- 
age area by Lake Eufaula, ca. 5 mi. s. Eufaula off US 431, 
11 Sept. 1968, Kral 33226. COVINGTON COUNTY: sandy 
north shore of Lake Jackson, Florala, 3 September 1969, 
Kral 36818. GENEVA COUNTY: 1 mi. s.w. Hartford on 
edge of pond cypress lake, 12 August 1966, S. McDaniel 
7646. 


1973] Southern States Flora — Kral 395 


An extension westward from Georgia or northward from 
s.w. Florida. First discovery of it in Alabama is credited 
to Dr. McDaniel. 


Ludwigia sphaerocarpa Ell. 

TEXAS. MONTGOMERY COUNTY: peaty bottoms, 12 
mi. w. Cleveland, 24 July 1964, Kral 21041. 

Dr. Raven, who made the identification in 1967, anno- 
tates: “extremely pubescent. Apparently new to Texas.” 


Ludwigia suffruticosa Walt. 

ALABAMA. HOUSTON COUNTY: sandy upper banks 
of Indigo Pond, s.w. Cottonwood, 1 August 1971, Kral 
43405. 

Extension of range westward from Georgia or north- 
ward from west Florida. 


Myriophllym exalbescens Fern. 

ALABAMA. JACKSON COUNTY: ca. 4 mi. e. Scotts- 
bora; edge of lake, formerly Coosa River, 14 September 
1968, Kral & Blum 33311. 

Of general distribution as an aquatic, but apparently 
not of frequent occurrence in the southeast; new for Ala- 
bama. 


Spermolepis inermis (Nutt.) M. & C. 

S. patens (Nutt.) B. L. Robins. 

ALABAMA. BALDWIN COUNTY: ca. 3 mi. w. jet. Ala. 
59 and Ala. 180 toward Ft. Morgan; sand pine-magnolia, 
16 April 1967, Kral 28397. GENEVA COUNTY: sandy 
areas above Pea River 2 mi. w. Samson, 17 April 1970, 
Kral 38680. 

More western, from prior reports, save for isolated rec- 
ords in North Carolina and Tennessee; a first record for 
Alabama. 


Ptilimnium costatum (Ell.) C. & R. 

ALABAMA. ST. CLAIR COUNTY: ca. 2 mi. s. Ashe- 
ville on US 231; prairie-like area by road, 15 August 1968, 
Kral 32348. 


396 Rhodora [Vol. 75 


Reported already for Alabama (Radford et al, Guide to 
the Vascular Flora of the Carolinas, p. 252, 1964; Manual 
of the Vascular Flora of the Carolinas, p. 784, 1968), but 
this essentially western gulf coastal plain umbellifer is in- 
frequent in the state. 


Cornus amomum Mill. ssp. microcarpa (Nash) Wilson 

Svida microcarpa (Nash) Small 

ALABAMA. ESCAMBIA COUNTY: ca. 10 mi. e. Brew- 
ton on co. 4, by Conecuh River, 8 June 1969, Kral 35154. 
GENEVA COUNTY : sandhills above Pea River ca. 6 mi. s. 
jct. Ala. 52-153, 7 June 1969, Kral 35044. 

A first record for Alabama. 


Cornus purpusii Koehne 

TENNESSEE. MAURY COUNTY : just above Rockdale 
on US 43, rocky shaley bluffs above small tributary of Big 
Bigby Creek, 26 June 1970, Kral 39738. 

An extension southward from Kentucky. 


Rhododendron austrinum (Small) Rehder 

Azalea austrina Small 

ALABAMA. COVINGTON COUNTY : sandy bottoms of 
5 Runs Creek by Stanley Community off Ala. 55, 16 April 
1970, Kral 38596; sandy creekbank 4.5 mi. e. Wing on Co. 
4, 18 April 1970, Kral 38739. COFFEE COUNTY: sandy 
Pea River bottoms off Ala. 134, e. of Opp and Perry, 17 
April 1970, Kral 38652. ESCAMBIA COUNTY: sandy 
bottoms Conecuh River, e. of Brewton on US 29, by hwy. 
bridges, 18 April 1970, Kral 38724. GENEVA COUNTY: 
sandy woods by Flat Creek, by Ala. 54, w. of Samson, 16 
April 1970, Kral 38619; sandy bluffs above Pea River 2 mi. 
w. Samson, 17 April 1970, Kral 38672; sandy bluffs above 
branch, 4 mi. w. Geneva on Ala. 52, then ca. 3 mi. n.e. by 
county rd., 17 April 1970, Kral 38688. 


Already reported for Alabama, though not mentioned 
for the state by Small. Dr. Clark (thesis, 1969) lists 9 
counties, one of which is Appalachian. This species equals 
R. calendulaceum (Michx.) Torr. in flamboyance, and is 


1973] Southern States Flora — Kral 397 


hardly distinguishable save for the pubescence on the backs 
of the bud scales and the glandular hairs of the twigs. The 
difference sometime utilized, namely that flowers emerge 
after the leaves have unfolded, as has already been noted 
by others, is inconstant. R. austrinum will hybridize with 
R. canescens (Michx.) Sweet, which almost always is in the 
same areas with it, though usually R. canescens is in bloom 
first. Both have the same “‘woodsy” fragrance, to me very 
similar to that of Phlox divaricata L. 


Lysimachia gramineum (Greene) Handel-Mazzetti 
Steironema gramineum Greene 
ALABAMA. CHEROKEE COUNTY: ca. 2 mi. s. Cen- 
tre; sandy swale by Ala. 9, spreading by slender rhizomes, 
22 July 1968, Kral 31898. 


To quote from J. S. Ray (Ill. Biol. Monogr. 24: 39) : 
“Lysimachia gramineum Greene, known only from 
specimens from the type locality, Northern Ala- 
bama,” is included here in 1878 by G. R. Vasey, 
probably near Gadsden, the type material, repre- 
sented by two sheets in the U.S. Nationa] Herb- 
arium and a fragmentary specimen in the Gray 
Herbarium, is perhaps not even typical of its 
kind. The stem, either cut or grazed near its base, 
consists of several attenuated lateral shoots. Nar- 
rowly linear leaves, small flowers, and capsules 
all range in size below those of ssp. lanceolata, 
yet there is not sufficient evidence for a distinct 
taxon. E. L. Greene emphasized the long filiform 
filaments which greatly exceed the short oblong- 
oval anthers. However, these are not unlike others 
of the southeastern populations." 


My own samples were from an enormous population, 
probably an acre in extent, but strikingly uniform and with 
fruits uniformly smaller than those of L. lanceolata. Fur- 
ther field work in northeastern Alabama ought to reveal 
still more of this, now known to be in at least two counties. 


398 Rhodora [Vol. 75 


Lysimachia quadriflora Sims 

Steironema quadriflorum (Sims) Hitchc. 

ALABAMA. CHEROKEE COUNTY: peaty seepage 
area by Ala. 9, 6 mi. s. Centre; forming large stand by 
rhizomes, 21 July 1968, Kral 37142. 

According to Ray (op. cit.) This essentially northern 
species does extend south into Arkansas and Georgia, but 
not into Alabama. It has very large, handsome yellow 
flowers similar to those of L. lanceolata. 


Sabatia difformis (L.) Druce 

ALABAMA. GENEVA COUNTY: sandy peat of sa- 
vanna-bog by Ala. 4, ca. 10 mi. e. Florala, 2 August 1971, 
Kral 43437. 


Extension of range probably northward from north- 
western Florida. 


Sabatia quadrangula Wilbur 

S. paniculata (Michx.) Pursh 

ALABAMA. HOUSTON COUNTY: e. side Ashford on 
sandy peaty swale by railroad, 5 June 1964, Kral 20523; 
16 July 1970, Kral 40019. 

Extension of range westward from Georgia or northward 
from Florida. 


Amsonia rigida Shuttlw. 

ALABAMA. GENEVA COUNTY: 5.3 mi. s. Samson on 
Ala. 87; edge of gum bottom; cor. near white, fls. faintly 
fragrant, 26 April 1969, Kral 34502. 

Extension of known range westward from Georgia or 
northward from Florida. 


Asclepias hirtella (Pennell) Woodson 

Acerates hirtella Pennell 

TENNESSEE. BLEDSOE COUNTY: upper edge of 
marsh meadow by Tenn. 30, 8 mi. e. entrance to Bledsoe 
Forest, 8 August 1970, Kral 40297. LEWIS COUNTY: 
sandy oak-hickory woods, clearing, ca. 2 mi. s. Meriwether 
Lewis entrance at Tenn. 20 and by road to Metal Ford, 


1973] Southern States Flora — Kral 399 


6 August 1969, Kral & Demaree 36192; 4.2 mi. e. Hohen- 
wald on sandy clay of high pasture, 27 August 1969, Kral 
36404. ALABAMA. ST. CLAIR COUNTY : prairie, s. side 
Asheville, 16 July 1971, Kral 43388. 

Some slight mystery as to range of this species exists. 
Small reports it from Alabama, thence “to Okla., Ill., and 
Mich." ; however Woodson (Ann. Mo. Bot. Gard. 41: 170- 
171, 1954) does not report it for Alabama and shows but 
one station for it in Tennessee (Franklin Co.). Therefore, 
the species is mentioned here simply to add locality in- 
formation. 


Asclepias tomentosa Ell. 

A. aceratoides M. A. Curtis 

FLORIDA. FRANKLIN COUNTY: sandy areas by 
coast, Alligator Point, 9 July 1970, Kral 39898. 

Some restatement as to distribution of this species is 
perhaps in order. Small reported A. aceratoides from 
North Carolina and South Carolina and A. tomentosa for 
Florida and Georgia; Woodson (op cit.), considering the 
two as one species, shows what appears to be an interesting 
disjunction, namely that the species is in peninsular and 
northeastern Florida, then in North and South Carolina 
(as indicated by Small). Then he shows it in Texas, in- 
dicating Polk County in his text, but indicates several 
counties in Texas for it on his map. The disjunction be- 
comes much less if the several collections by Godfrey (and 
others) from northwestern Florida are added to the in- 
formation. It appears there primarily in sandhills along 
the coast. 


Ipomoea muricata (L.) Jacq. 

ALABAMA. ESCAMBIA COUNTY: weedy along rail- 
road, w. side Atmore; fls. in evening, 19 September 1971, 
Kral 44416. 

Very distinctive with its bristly muricate stems and nar- 
rowly funnelform, lavender-tinted, corollas that open 
toward evening; a first report for Alabama. Previously 
reported from Florida and Texas. 


400 Rhodora [Vol. 75 


Phlox pilosa L. var. ozarkana Wherry 

TENNESSEE. MAURY COUNTY: shaley, w. facing 
oak-hickory slopes by US 43, just above Rockwood, on rim, 
3 June 1971, Kral 42847. 

Wherry indicates (The Genus Phlox, Morris Arboretum 
Monographs III, pp. 44-45, 1955) two disjunct areas, one 
essentially ozarkian and the other primarily in central and 
northern Alabama, also extreme western Georgia. He does 
not report it for Tennessee. 


Verbena bipinnatifida Nutt. 

Glandularia drummondii (Lindl.) Small 

ALABAMA. GREENE COUNTY: chalk prairie 6.5 mi. 
s. Eutaw on US 43, 5 June 1968, Kral 31082. MARENGO 
COUNTY: 2.3 mi. s. Demopolis on US 43; chalk prairie, 
6 June 1968, Kral 31215. MONTGOMERY COUNTY: 
roadside weed s. side Montgomery, 21 May 1968, Kral 
30953. SUMTER COUNTY: chalk outcrop 3 mi. n.n.e. 
Livingston, Kral 23786; patch of chalk prairie, by Ala. 17, 
8 mi. s. Dancy, 4 June 1970, Kral 39643. 

Very abundant and showy in late spring and into the 
summer on the black earths and outcrops of the chalk 
prairies in Alabama; probably in every black belt county. 
However, not reported by Small from e. of La. A common 
verbena of the prairie provinces of the west and midwest; 
already reported for Alabama by Harper (Ala. Geol. Surv. 
Bull. 53, 1944) and L. M. Perry (Annals Mo. Bot. Gard. 20: 
323, 1933). 


Collinsonia canadensis L. 
ALABAMA. CRENSHAW COUNTY: longleaf pine 
flatwoods 3.5 mi. n. Brantley, 5 October 1968. Kral 33711. 
According to Shinners (Sida 1, p. 80) this species is rare 
in the coastal plain; he cites but one collection, this by 
Buckley and without definite locality. 


Hedeoma drummondii Benth. 
ALABAMA. MARENGO COUNTY: 2.3 mi. s. Demopo- 
lis on US 43; chalk prairie, 6 June 1968, Kral 31210. SUM- 


1973] Southern States Flora — Kral 401 


TER COUNTY: north of Livingston on hwy. 11, prairie 
area, K. E. Rogers, 29 May 1970. 

This species of the prairie districts of the western U.S. 
is here recorded as a rarity; it has already been reported 
for Alabama by Harper (op. cit.). 


Stachys floridana Shuttlw. 

ALABAMA. BALDWIN COUNTY: e. end of US 90-98 
causeway over Bay, e. of Mobile; moist sandy cleared area 
by highway, 19 April 1970, Kral 38799. 

While Small reported this only from Florida, Dr. Sam 
Jones and others have collected and reported this from 
Alabama. It is still rare enough there to be noteworthy, 
and very distinctive because of the slender, elongate stolons 
terminating in pale, fusiform tubers. Very weedy where 
found, and likly to expand its range accordingly. 


Stachys hispida Pursh 

TENNESSEE. HOUSTON COUNTY: sandy gravelly 
wooded banks of Yellow Creek, near where crossed by 
Tenn. 49, w. of Van Leer, 16 June 1969, Kral 35276. 

An extension of known range southward from Kentucky. 


Scutellaria racemosa Pers. 

ALABAMA. BALDWIN COUNTY: sandy-gravelly rail- 
road embankment through Hypericum pond just n. of ject. 
1-10 off Ala. 59 and 6 mi. s. Stapelton, 8 June 1971, Kral 
43084. 

A first report for this species for the U.S.A. There are 
specimens of it at the U.S. National Museum from Argen- 
tina, Bolivia, Chile, Paraguay, Uruguay and Central Amer- 
ica. It is distinctive in its very small flowers (ca. 3 mm. 
long), and in its peculiar rootstock which consists of nu- 
merous, pale, quite elongate stolons from which branch 
fleshy, fusiform-cylindrica] tubers. 


Mosla dianthera (Buch.) Maxim. 

TENNESSEE. MARION COUNTY: sandy pockets in 
sandstone glades 8.5 mi. s. Grundy Co. line and Tracy City, 
29 September 1971, Kral 44514; sandstone outcrop area by 


402 Rhodora [Vol. 75 


Fish Branch, by Tenn. 108, 3 mi. n. Whitwell, 29 September 
1971, Kral 44522. 

An extension southward from Kentucky. Very like 
Perilla. 


Monarda clinopodia L. 

ALABAMA. MADISON COUNTY: rich loamy banks of 
Mountain Fork Creek 4.5 mi. n.e. New Market, 15 July 
1971, Kral 43341. 

An extension southward from Tennessee. 


Hyptis mutabilis (Rich.) Briq. 

ALABAMA. BALDWIN COUNTY: edge of longleaf 
pine sandridge by US 98, 3.5 mi. n. Montrose, 9 October 
1967, Kral 29806. COVINGTON COUNTY: sandy road- 
bank, in pine flatwoods ca. 13.5 mi. s. opp on US 331, 17 
August 1968, Kral 32504. RUSSELL COUNTY: weed in 
empty sandy lot, Hurtsboro, 12 September 1968, Kral 
33281. I have other, yet unprocessed records from Escam- 
bia and Mobile counties. 

This native of South America is credited in the United 
States only to Florida by Small. 


Physalis longifolia Nutt. 

P. sulglabrata Mackenzie & Bush 

ALABAMA. MONTGOMERY COUNTY: clay loam of 
cultivated area, 4 mi. s. of Loop Road, s. side Montgomery, 
1 June 1970, Kral 39479. PICKENS COUNTY: just s. of 
Cochrane; woods at border of chalk prairie, 5 June 1971, 
Kral 42997. 

An extension of range southward from Tennessee; prob- 
ably more abundant in the black belt than present collec- 
tions indicate. 


Veronica anagallis-aquatica L. 

ALABAMA. MADISON COUNTY: banks of Mountain 
Fork Creek, 1.3 mi. ne. New Market off Old Mountain 
Fork Road, 15 July 1971, Kral 43359. 

An extension southward from Tennessee. 


1973] Southern States Flora — Kral 403 


Lindernia crustacea (L.) F. von Muell. 

ALABAMA. ESCAMBIA COUNTY: wet sands of 
nursery lot, Atmore, 19 September 1971, Kral 44407. 

A first report for Alabama; already reported from Flor- 
ida and Mississippi. 


Utricularia resupinata, Greene 

Lecticula resupinata (Greene) Barnh. 

ALABAMA. COVINGTON COUNTY: sandy shallows 
of north shore Lake Jackson at Florala, 3 September 1969, 
Kral 36008. 

An extension northward from northwest Florida. 


Diodia hirsuta Pursh. 

ALABAMA. WASHINGTON COUNTY: 5 mi s. 
Chatom in pineland bog, 14 July 1969, Kral 35534. 

This entity, considered part of D. virginiana L. by some 
authors, (i.e., Radford et al., op. cit.), is certainly distinc- 
tive, and has not been reported previously save from the 
Atlantic coastal plain from Florida to North Carolina. 


Plantago hookeriana Fisch. & Mey. 

P. wrightiana Decaisne 

ALABAMA. BALDWIN COUNTY: railroad gravels 
above Hypericum pond just n. jet. I-10 on Ala. 59 and 6 mi. 
s. Stapelton, 8 June 1971, Kral 43083. GENEVA COUNTY: 
sandy edge of field between Geneva and Samson on Ala. 52, 
27 April 1969, Kral 34565; sandy railroad right of way, 
0.5 mi. w.n.w. Geneva on Ala. 52, 17 April 1969, Kral 38703. 
HOUSTON COUNTY: sandy railbanks above Little Choc- 
tawhatchee Creek on Ala. 92, 1.7 mi. w. jct. US 94, 9 June 
1971, Kral 43156. LEE COUNTY: dry sandy oak-pine 
woods, clearing, by Ala. 169, 4 mi. n. Crawford, 9 May 
1968, Kral 30713. MONTGOMERY COUNTY: NW side 
Montgomery on RR gravels by US 31-jet. bypass, 5 June 
1969, Kral 34989. 

Shinners (op. cit.) treated this part of the P. patagonica 
complex as P. wrightiana rather than as a variety of P. 
hookeriana as did Poe. My own records of it in Alabama 
appear to be new records for the state, but this essentially 


404 Rhodora [Vol. 75 


western weed has been collected from the coastal plain of 
Georgia and North Carolina (Radford, et al). 


Valeriana pauciflora Michx. 

ALABAMA. MADISON COUNTY: rich loamy woods 
along Mountain Fork Creek, n.e. New Market, 4 May 1971, 
Kral 32485. 


An extension southward from Tennessee. 


Liatris chapmanii T. & G. 

Laciniaria chapmanii (T. & G. Kuntze 

ALABAMA. BALDWIN COUNTY: sandy banks and 
flats of scrub above River Styx, by where crossed by co. rd., 
ca. 5 mi. n.n.e. Elsanor, 2 September 1970, Kral 41029. 
ESCAMBIA COUNTY: 2.7 mi. s.w. Flomaton along US 
31; sandy clay roadbank in longleaf pine-turkey oak hills, 
6 September 1965, Kral 25950. 

An extension northward from northwest Florida, where 
it locally abounds in sandhills. 


Xanthocephalum dracunculoides (DC) Shinners 

Gutierrezia dracunculoides (DC) Blake 

TENNESSEE. DAVIDSON COUNTY: ca. 1 mi. w. 
Scottsboro on Tenn. 12, 9 September 1957, H. F. L. Rock 
887. RUTHERFORD COUNTY: edge of wet limestone 
glade, ca. 2 mi. s.s.e. Murfreesboro by US 41, 6 October 
1969, Kral 37551; limestone glade by US 708, 3.8 mi. e. 
Murfreesboro, 9 August 1970, Kral 40326; limestone glades 
3 mi. s. Murfreesboro by US 231, and 8.1 mi. n. Deason, 
27 August 1970, Kral 40755. 

This broomweed is extremely abundant in open limestone 
glades in Rutherford and Davidson counties but evidently 
has not been in the area very long, for while it is men- 
tioned for Alabama by Fernald (op. cit), there is no refer- 
ence to it for Tennessee prior to the Rock collection and 
other contemporary collections made through the Univer- 
sity of Tennessee. Common westward in Missouri, Okla- 
homa, Arkansas, and Texas in essentially calcareous dis- 
tricts; more local northward in the interior lowlands. 


1973] Southern States Flora — Kral 405 


Solidago elliottii T. & G. 

ALABAMA. GENEVA COUNTY: 0.3 mi. n. jet. Ala. 
153 and Ala. 52 on Ala. 52; sandy peat of longleaf pine 
savanna-bog, 6 October 1968, Kral 33756. 

An extension of known range westward from the south- 
ern Atlantic coastal plain. 


Solidago ulignosa Nutt. 

ALABAMA. JACKSON COUNTY: sandy clay summit, 
seepage, of Sand Mt., off Ala. 117, 5.2 mi. n. Flat Rock, 
7 October 1969, Kral 37590. 

This essentially northeastern bog species has been re- 
ported previously from no further south than North Caro- 
lina. 


Aster chapmanüi T. & G. 

ALABAMA. GENEVA COUNTY: sandy peat of long- 
leaf pineland savanna by county 4, 11 mi. e. Black, 28 
October 1969, Kral 37951. HOUSTON COUNTY: sandy 
peat of longleaf pine flatwoods savanna ca. 1 mi. s.e. Cot- 
tonwood, 12 October 1970, Kral 41656. 

Extension of range northward from northwestern Flor- 
ida. The Alabama localities are both threatened by either 
real estate development or farmers, who convert the rich 
black peats to use for row crops. 


Aster eryngiifolius T. & G. 

ALABAMA. COVINGTON COUNTY: longleaf pine- 
land sandhills bog by US 331, ca. 3 mi. n. Florala, 26 July 
1968, Kral 32096. GENEVA COUNTY: by county 4, ca. 
10 mi. s. intersection US 231, in pineland savanna, 8 June 
1968, Kral 31350. 

An extension of known range northward from north- 
western Florida. 


Aster spectabilis Ait. 

ALABAMA. DEKALB COUNTY: swale in sandrock 
hills, oak-pine, on county 89, ca. 4 mi. s. DeSoto State Park 
Hq., 29 August 1969, Kral 36491; sandy loam around sand- 
rock, and under oak-pine, ca. 4 mi. n.e. Ft. Payne, 8 Octo- 


406 Rhodora [Vol. 75 


ber 1969, Kral 37644; sandy powerline clearing DeSoto 
Parkway just n. of Ala. 35, e. of Ft. Payne, 28 August 1970, 
Kral 40793. 

This very showy aster, a worthy subject for cultivation 
on sandy soils, appears not to have been reported previously 
for Alabama. An extension southward and eastward from 
western North Carolina. 


Conyza bonariensis (L.) Cronquist 

Leptilon bonariense (L.) Small 

ALABAMA. ESCAMBIA COUNTY: sandy longleaf 
pine woods above Perdido Creek, ca. 7 mi. w.s.w. Atmore, 
18 August 1968, Kral 32576. MOBILE COUNTY: Battle- 
ship Park, by causeway e. of Mobile on the Bay, 20 August 
1968, Kral 32756; sandy empty lot, w. side Mobile near jet. 
International Drive and Interstate Hwy. 10, 2 September 
1970, Kral 41012. 

Reported in Small as only from Florida but since col- 
lected by Demaree in Mississippi and probably more abun- 
dant in Alabama than present records indicate. It is always 
on sandy disturbed grounds, and therefore much in evi- 
dence along the newer highways toward the coast. 


Erigeron myrionactis Small 

E. repens A. Gray, 1884; not (HBK) Weddell, 1857. 

MISSISSIPPI. JACKSON COUNTY: trailing over 
sandy road shoulder, Old US Rt. 90, ca. 5 mi. e. Ocean 
Springs, R. B. Channell 3561, 12 July 1955. 

Here cited as new for Mississippi. Reported in Small 
(Flora, 1903) as a weed of the coast of Texas and Louisi- 
ana. 


Iva microcephala Nutt. 

ALABAMA. BARBOUR COUNTY: sandy pineland 
above W. F. George Reservoir (Lake Eufaula), s. of Eu- 
faula off Ala. 95, 18 October 1969, Kral 38023. HOUSTON 
COUNTY: sandy bluff above Cedar Creek where crossed 
by Ala. 95, s. of Columbia, 18 October 1969, Kral 37962. 


An extension westward from Florida, Georgia. 


1973] Southern States Flora — Kral 407 


Iva annua L. 

I. ciliata Willd. 

TENNESSEE. RUTHERFORD COUNTY: edge of 
marshy area, heavy soil, s. side Murphreesboro on US 231, 
27 August 1970, Kral 40764. 

This essentially western weed is of scattered distribution 
east of the Mississippi, save in the black belt where it is 
common on disturbed heavy soils. This Tennessee report 
is probably an extension from Alabama. 


Ambrosia bidentata Michx. 

TENNESSEE. DAVIDSON COUNTY: field next to 
open limestone glades by Mountain View Road, n. of 
Murphreesboro Rd., toward Percy Priest Lake, 14 August 
1968, Kral 32342. RUTHERFORD COUNTY: gravelly, 
calcareous road shoulders beside I-24 jct. Buchanan Road, 
7.7 mi. n.w. Beech Grove, 19 August 1971, Kral 43641. 

This ragweed hitherto reported for w. Tennessee by Gat- 
tinger (Flora of Tennessee, 1900) and from east Tennessee 
recently (Sharp et al, Preliminary Checklist of Dicots in 
Tennessee, p. 105, 1960) but not subsequently in the man- 
uals, must be an extension from the calcareous districts of 
the mid-west. Evidently spreading rapidly in Tennessee. 


Rudbeckia mollis Ell. 

ALABAMA. HENRY COUNTY: sand hills 6.5 mi. s. 
Abbeville on US 431, 24 July 1968, Kral 31951; sandy oak- 
pine woods 1.5 mi. s. Abbeville on US 431, 24 July 1968, 
Kral 31966. 

Perdue (Rhodora 59: 1957), reports this species as from 
southeastern Georgia and northeastern Florida, but not 
from Alabama. In the field quite similar from a distance 
to R. hirta, but paler green because of the soft, long, 
clammy, spreading pubescence of stem and leaves, and 
because of the paler yellow rays. 


Ratibida columnifera (Nutt.) Woot. & Standl. 

R. columnaris (Sims) D. Don 

ALABAMA. LOWNDES COUNTY: chalk prairie by 
US 80, 3.2 mi. w. Lowndesboro, 28 June 1970, Kral 39807. 


408 Rhodora [Vol. 75 


Appearing to be a recent introduction to Alabama. At 
this locality it was extremely abundant, the population 
showing a great range of ligule pigmentation from deep 
maroon to yellow or orange. It appears well established, in 
a locality strikingly similar to the western calcareous 
prairies where this ratibida is sometimes weedy. This part 
of Alabama is cattle country, so that akenes of the species 
could have been brought in either with shipments of stock 
or feed. 


Bidens cernua L. 

ALABAMA. LIMESTONE COUNTY : 5 mi. n. Dekatur 
by US 31; abundant on wet gravelly creekbank, 16 October 
1969, Kral 37860. 

An extension southward from Tennessee. 


Bidens tripartita L. 

ALABAMA. LIMESTONE COUNTY: north of Dekatur 
across reservoir of Tennessee River at Wheeler Wildlife 
Refuge; silt of drying bottoms, 22 September 1970, Kral 
41207. 

An extension southward from Tennessee. 


Viguirea porteri (A. Gray) Blake 

ALABAMA. CHAMBERS COUNTY: granite outcrop 
area 1.9 mi. n.w. Penton, 17 Oct. 1969, Kral 37889. 

This representative of a southern and southwestern 
north American genus has already been reported for Ala- 
bama by Harper and later by McVaugh (Ecol. Monogr. 13, 
no. 2, 1943). The Chambers County locality is however an 
additional county record in Alabama for this rather re- 
stricted endemic. 


Marshallia mohrii Beadle & Boynt. 

ALABAMA. CHEROKEE COUNTY: peaty seepage 
area by Ala. 9, 6 mi. s. Centre, 21 July 1968, Kral 31739; 
sandy clay of savanna pasture swale, ca. 6 mi. s. Centre by 
Ala. 9, 5 June 1969, Kral 35018. 

The above two collections represent flowering and fruit- 
ing material from a very large population. The find is here 


1973] Southern States Flora — Kral 409 


considered noteworthy in that Dr. Channel] (Contrib. Gray 
Herb. CLXXXI, 1955) mentions in his revision that the 
last known collection of this rare species was made at the 
type locality in Cullman, Alabama in 1941 and that no cur- 
rent stations for it were presently known to him. 


Artemisia ludivociana Nutt. 

TENNESSEE. RUTHERFORD COUNTY: clearings in 
cedar glades by Percy Priest Lake, n. of Luvergne, 3 Octo- 
ber 1969, Kral 37511. 

Reported by Sharp et al. (op. cit.) from the Great Smoky 
Mountain National Park Headquarters and as a probable 
escape in Hardeman County. However appearing well 
naturalized in the cedar glades. An extension eastward 
from the western prairie districts, or southward from 
Illinois. 


Cacalia diversifolia T. & G. 

Mesadenia diversifolia (T. & G.) Greene 

ALABAMA. HOUSTON COUNTY: 5.5 mi. s.e. Cotton- 
wood, near creek on floodplain of Cowarts Creek, 24 May 
1967, McDaniel 9059; ca. 2 mi. s.w. Cottonwood in creek 
bottom, 8 June 1968, Kral 31321; Cowart’s Creek bottoms 
ca. 6 mi. s.e. Cottonwood, 8 June 1968, Kral 31317. 

Proper credit for the find must go to Dr. McDaniel who 
told me of the locality. Formerly, the species was thought 
to occur only at a few isolated points in Florida. The Chi- 
pola River of Florida along which it is locally abundant, 
heads up in Alabama partly as Cowart's Creek. 


Hypochaeris glabra L. 

ALABAMA. MONTGOMERY COUNTY: calcareous 
pasture and adjacent railroad embankment n.w. side Mont- 
gomery by US 31, 28 April 1969, Kral 34572; heavy soil 
of motel lawn, by Loop, s. side Montgomery, 1 June 1970, 
Kral 39584. 

This representative of a mainly South American genus, 
has been reported, in the southeast, from North and South 
Carolina; it appears to be abundant locally in the cal- 


410 Rhodora [Vol. 75 


careous areas about Montgomery and is to be looked for 
throughout the black belt. Well established in the Pacific 
states. 


Lactuca saligna L. 

TENNESSEE. CLAY COUNTY: shaley banks by 
Tenn. 52, ca. 2 mi. s. Celina, 11 August 1970, Kral 40430. 
MARION COUNTY: steep limestone roadbank by US 41, 
ca. 4 mi. s.s.e. Monteagle, 7 October 1969, Kral 37630. 
WILLIAMSON COUNTY: elm dotted calcareous meadow 
by 1-65, 0.5 mi. n. jet. Tenn. 96 (Franklin turnoff), 3 
October 1968, Kral 33530. 

An extension southward from Kentucky for this weed. 


Taraxacum laevigatum (Willd.) DC 

T. erythrospermum Andrz. ex. Besser 

ALABAMA. DALLAS COUNTY: 2 mi. s.e. US 80 cross- 
ing of Alabama River at Selma; banks of highway above 
wet ditch, 15 April 1970, Kral 38575. 

Not previously reported for Alabama; at scattered locali- 
in the southeastern United States. 


DEPARTMENT OF BIOLOGY 
VANDERBILT UNIVERSITY 
NASHVILLE, TENNESSEE 37203 


ADDITIONS TO THE BAHAMA FLORA 
SINCE BRITTON AND MILLSPAUGH — I. 


WILLIAM T. GILLIS, RICHARD A. HOWARD, 
AND GEORGE R. PROCTOR 


Additional taxa may be found in a region covered by a 
flora if (1) they were overlooked by the original flora 
writers, (2) they were described since the flora was com- 
pleted, (8) they have been segregated from other taxa in 
the meantime, or (4) they have been introduced since the 
flora was completed. We compile here 81 taxa of vascular 
plants which are added to the Bahama flora since the pub- 
lication of Britton and Millspaugh’s work (1920). Some 
are reported for the first time; others are cited as having 
been gleaned from the diffuse taxonomic literature as it 
relates to the Bahama flora. Three previous works are of 
principal concern. A check-list of the plants of Bimini was 
published in connection with vegetational studies there 
(Howard, 1950). Additions to the flora of Inagua, the 
southernmost island in the archipelago, were noted by 
Howard and Dunbar (1964). Still other plants, not pre- 
viously recorded by Britton and Millspaugh, were pointed 
out for Grand Bahama Island and Eleuthera (Lewis, 1971). 


We present here, in the process of preparing a new vas- 
cular flora of the Bahama Islands, a compilation of addi- 
tional vascular plants, growing without cultivation, in the 
Bahama Islands including the Turks and Caicos group. 
We include the Turks and Caicos Islands in the Bahamas, 
as did Britton and Millspaugh, because they all comprise 
the same geographical unit, despite the fact that the Turks 
and Caicos Islands are a separate Crown Colony from the 
Commonwealth of the Bahama Islands. A second paper 
on additional species is in preparation, the result of a 
recent study in the southern islands by Gillis and Proctor. 
Another paper will follow, summarizing nomenclatural 
changes from usage in Britton and Millspaugh. Together, 


411 


412 Rhodora [Vol. 75 


these papers should represent the indigenous Bahamian 
flora as it is now understood. 

Voucher specimens for plants reported here for the first 
time are deposited in the herbaria of the Fairchild Tropical 
Garden (for specimens of Gillis and John Popenoe), of the 
New York Botanical Garden (for specimens of Howard), 
and of the Institute of Jamaica (for specimens of Proctor), 
with duplicates at the Arnold Arboretum. 


POLYPODIACEAE 

Trismeria trifoliata (L.) Diels. This species was found 
on the cay Lubbers Quarters off Marsh Harbour, Great 
Abaco (Gillis 7328A and Popenoe 131). It occurs on marl 
fill over an old Conocarpus swamp at the south end of the 
island, possibly introduced with the fill. 

Pteris vittata L. This species was also found in the same 
marl fill as the Trismeria above (Gillis 7323 and Popenoe 
132), on a road cut near Marsh Harbour (Proctor 30554), 
and on Grand Bahama Island near the former town site 
of Freeport, abandoned about 1954 (Gillis 7841). 


NAJADACEAE 

Najas marina L. This species is not reported by Clausen 
(1936) as occurring in the Bahamas. Its having been 
found in the Duck Pond on South Bimini, the island near- 
est to the Florida mainland, is however not surprising. It 
most likely has been brought in from Florida and may now 
spread by birds to other islands in the chain. This species 
represents a new family of flowering plants for the Ba- 
hamas (Gillis 11289). 


GRAMINEAE 

Cenchrus gracillimus Nash. This species is reported 
from Fortune Island (now known as Long Cay) in the 
monograph by DeLisle (1936), represented at the Mis- 
souri Botanical Garden by an old Hitchcock collection. 

Eragrostis ciliaris var. laxa Kuntze. Hitchcock (1936) 
from Grand Turk and South Caicos. 

Eragrostis excelsa Griseb. A pan-Caribbean species re- 
ported by Howard (1950) from a cemetery in Bimini. 


1973] Bahama Flora — Gillis, Howard and Proctor 413 


Eragrostis urbaniana Hitchc. in Urb. Hitchcock (1936) 
from South Caicos, Long Cay (Caicos), and Grand Turk. 
This species was included in E. ciliaris in Britton and 
Millspaugh. It is represented by a recent collection (Proc- 
tor 9194) from Grand Turk. 

Monanthochloé littoralis Engelm. Howard (1950) from 
Mosquito Point, North Bimini. 

Panicum echinulatum Mez. Proctor 8955 from South 
Caicos. Hitchcock (1936) indicates that the species has 
been introduced to the West Indies from Paraguay and 
Bolivia. 

Paspalum bakeri Hack. Collected on Abaco from the 
Eastern Shore District (Proctor 30734), this species was 
also reported from Inagua by Chase (1929). 

Paspalum laxum Lam. Howard (1950) from old fields 
on Bimini. 

Paspalum molle Poir. Proctor 30564 from Marsh Har- 
bour, Abaco. 

Paspalum propinquum Nash. Chase (1929) from Inagua. 

Paspalum urvillei Steud. Proctor 30621 from seven to 
eight miles southwest of Marsh Harbour, Abaco. 

Rhyncheletrum repens (Willd.) Hubb. Reported as R. 
roseum from Eleuthera by Lewis (1971), and found by 
Gillis on New Providence, Abaco, and Exuma. This weed 
of the southern United States, a native of Africa, has be- 
come common in the southeastern United States within 
the last 50 years. It surely would not have been overlooked 
by Britton and Millspaugh if it had been present prior to 
1920. 

Setaria geniculata (Lam.) Beauv. Howard (1950) from 
old fields on Bimini. 

Sorghum halepense (L.) Pers. Proctor 30624 from 
Abaco, where it is extensively naturalized. 


CYPERACEAE 
Cyperus aristata Rottb. Both Horvat (1941) and Mc- 
Laughlin (1944) cite specimens of this species from the 
Bahamas in their revisions of certain groups of Cyperus. 


414 Rhodora [Vol. 75 


Cyperus globulosus Aubl. Reported (as C. globosus) by 
Howard (1950) from old fields on Bimini. 

Eleocharis geniculata (L.) R. et S. Stimson (1967) 
from Bimini. 

Fimbristylis annua (All. R. et S. Kral (1971) records 
this species from Andros and New Providence. 

Fimbristylis autumnalis (L.) R. et S. This species was 
cited by Kral (1971) as inhabiting the “Caribbean Islands" 
with no specific reference to the Bahamas. Proctor 30811 
confirms its presence at Morgan’s Bluff, North Andros. 


PALMAE 

Coccothrinax inaguensis Read. This palm was described 
as new by Read (1966) as a segregate from C. argentata. 
It was described initially from Inagua but has since been 
found on San Salvador, along the shore near Sandy Point 
(Gillis 8796 and Popenoe s.n.). Presumably it is on other 
southern islands in the chain as well. 

Phoenix dactylifera L. Proctor has seen this escaping 
from cultivation in the Caicos group. 


JUNCACEAE 
Juncus roemerianus Scheele. Presence of this plant adds 
another plant family to the Bahama flora. Specimens have 
been collected by Peter Garrett near Red Bays, North 
Andros, and sent to the Fairchild Tropical Garden herbar- 
ium for identification. Proctor 30515 records its presence 
on Abaco, about 8 miles southeast of Marsh Harbour. 


LILIACEAE 

Smilax domingensis Willd. Proctor 9082 from North 
Caicos. Britton and Millspaugh state that Hitchcock’s 
material determined as this species actually represents S. 
havanensis. The present assignment of the North Caicos 
material to this species must, therefore, be viewed with 
doubt, although the Proctor specimen resembles S. domin- 
gensis much more than S. havanensis. 


DIOSCOREACEAE 
Dioscorea bulbifera L. Found as a weed on the north 
side of New Providence Island in a coppice (Gillis 7520). 


1973] Bahama Flora — Gillis, Howard and Proctor 415 


IRIDACEAE 
Sisyrinchium exile Bicknell. Reported from Inagua by 
Howard and Dunbar (1964), a presumed waif. Daniel B. 
Ward (personal communication) suggests that this spe- 
cies may actually be the same as S. micranthum Cav. 


PORTULACACEAE 
Talinum triangulare (Jacq.) Willd. Determined by 
Howard from material sent in from the Exuma keys where 
it had some use in bush medicine. Reported as introduced 
and naturalized at Red Bays on North Andros (Proctor 
30887) and at the Current on Eleuthera (Proctor 30952). 


NYCTAGINACEAE 
Bougainvillea glabra Choisy. Howard (1950) reported 
this species as persistent after cultivation on Bimini. It is 
not known to produce seed in the Bahamas, and therefore 
probably does not escape from cultivation. 


PHYTOLACCACEAE 
Phytolacca purpurascens A.Br. et Bouché. Lewis (1971) 
reported this species from Grand Bahama Island. Gillis 
7385 is from bulldozed marl about 15 miles north of Marsh 
Harbour, Abaco. (The Gillis specimen determined by 
Jonathan Sauer whose assistance is gratefully acknowl- 
edged.) 


AIZOACEAE 
Sesuvium microphyllum Willd. This West Indian species 
was collected on Grand Turk (Proctor 8808). More re- 
cently, it has also been confirmed from a dry pond near 
Matthew Town, Inagua (Gillis 11707). 


PAPAVERACEAE 

Bocconia frutescens L. This species was first noted on 
Abaco by a local naturalist, Jack Patterson. It was col- 
lected and identified for inclusion in the Bahama planting 
at the Fairchild Tropical Garden where it has not been 
successful. (Gillis 7395 and Proctor 30599 from the same 
locality.) The occurrence of the plant is very local, extend- 
ing along the road and into the pine woods for about a mile, 


416 Rhodora [Vol. 75 


15 miles north of Marsh Harbour. It is obviously doing 
well and becoming locally abundant. So far as is known, 
this is its only station in the Bahamas. 


CRUCIFERAE 
Lepidium filicaule C.L. Hitchc. Described as new by 
Hitchcock (1945) from specimens determined by Britton 
and Millspaugh as L. virginicum, presumably depauperate 
ones. It is known from rocks on the east shore of Amber- 
gris Cay in the Caicos group, and from South Caicos. 


LEGUMINOSAE 

Acacia tortuosa (L.) Willd. Represented by a specimen 
(Lewis, s.n.) at the Institute of Jamaica. It is interesting 
that this species should be reported as new to the Bahamas 
inasmuch as it has only recently been reported as new to 
South Florida (Alexander, 1968). 

Caesalpinia divergens Urb. Howard (1950) reports this 
plant (as Gwilandina divergens) from Bimini where it 
inhabits coastal thickets. 

Lonchocarpus sericeus Kunth. This plant was collected 
by John Popenoe at South Beach, New Providence in a 
seemingly wild state even though it is difficult to suspect 
any place so close to Nassau as not having been in cultiva- 
tion. Seed from this population has grown to plants of 
mature, bearing age at the Fairchild Tropical Garden. 
Specimens from this cultivated population have been de- 
termined by Mr. Mario Sousa at Harvard University. This 
is a West Indian species which could easily be in the 
Bahama flora; its having been found only once in the 
middle of the island chain is, however, puzzling. 

Melilotus alba Desr. Naturalized in old fields seven to 
eight miles southwest of Marsh Harbour, Abaco (Proctor 
30609). 

Stylosanthes tuberculata Blake. In his revision of the 
genus Stylosanthes, Mohlenbrock (1957) reports this spe- 
cies from Southwest Landing on New Providence. 

Stylosanthes calcicola Small. Mohlenbrock (ibid.) also 
reports this species from New Providence where it has 
been collected at West Bay. 


1973] Bahama Flora — Gillis, Howard and Proctor 417 


RUTACEAE 

Citrus aurantium L. Howard (1950) records this plant 
as escaped from cultivation on Bimini. 

Zanthoxylum bifoliolatum Leonard. Discovered on San 
Salvador by John Popenoe. It has been found at the south- 
west corner of the island near the Sandy Point House 
(erroneously called *Watling's Castle") and from the dirt 
road that leads from near Rocky Point to Great Lake 
(Gillis 8811). Indigenous to Hispaniola, its occurrence on 
the eastern fringe of the Bahama archipelago 500 miles 
from Hispaniola is puzzling, but it is not the only species 
to have this distribution (see Croton discolor). One of the 
authors (GRP) prefers use of the generic name Fagara, 
but the other two have chosen to follow Brizicky (1962). 

Spathelia bahamensis Marie-Victorin. Described by 
Marie-Victorin (1948) as a new species, endemic to the 
Bahamas. The type came from a point northwest of Set- 
tlement, Cat Island. Proctor 30973 confirms its occurrence 
on Eleuthera, south of Rock Sound and 1 mile north of the 
entrance to the Cotton Bay Club. This species was included 
within S. vernicosa by Britton and Millspaugh. 


MELIACEAE 
Cedrela odorata L. Known from the wild in a limited 
area on North Eleuthera near the settlements of Bluff and 
Upper Bogue (Jack Patterson, s.n.). 


EUPHORBIACEAE 
Croton discolor Willd. Gillis 8729 from a ridge on the 
southwest corner of San Salvador where it was growing 
with C. linearis which it resembles. Their differences were 
evident when seen together. Gillis 6205 (deposited at 
Michigan State University) is from near Graham’s Har- 
bour, also on San Salvador. This species was under con- 
sideration by Britton and Millspaugh, but all the material 
which they saw in this complex was considered to be C. 
linearis. These San Salvador collections confirm the pres- 

ence of this species in the Bahamas. 
Poinsettia cyathophora (Murr.) Kl. et Gke. This plant 


418 Rhodora [Vol. 75 


may possibly be the species referred to by Britton and 
Millspaugh as P. heterophylla. Burch (1966) has pointed 
out that these two species have long been confused, but 
that P. cyathophora has red markings on the bracts and 
P. heterophylla has white markings. In any event, this 
ruderal plant is found on various islands, but true P. 
heterophylla has been collected thus far only from Bimini 


(Gillis 11308). 


ANACARDIACEAE 
Spondias purpurea L. Howard (1950) reports this spe- 
cies as escaped from cultivation on Bimini. 


CELASTRACEAE 
Maytenus phyllanthoides Benth. J ack Patterson of 
Abaco sent this species for determination. It appears to be 
a new record for the Bahamas of a shrub indigenous to the 
Florida Keys. It is probably more common than indicated, 
having been mistaken for M. buxifolia which is already 
known for the flora. 


RHAMNACEAE 

Colubrina asiatica (L.) Brongn. var. asiatica. Johnston 
(1971) shows Andros Island as having this species accord- 
ing to his dot map, but does not specifically cite specimens 
from the Bahamas. A recent collection (Gillis 11680) 
has been made from the strand on the southwest shore of 
Inagua. This Asiatic species has been known from the 
West Indies since 1860. Because of its habitat as a strand 
plant and because of its viability in sea water, it can be 
expected to appear more frequently in the Bahamas. 


MALVACEAE 

Cienfuegosia yucatanensis Millsp. Regarded as weedy 
in Cuba and Puerto Rico, Howard and Dunbar (1964) 
reported it first in the Bahamas from Inagua as C. hetero- 
phylla. Fryxell (1969) has recognized this to be C. yuca- 
tanensis. 

Hibiscus acetosella Welwit. ex Hiern. This species was 
found growing without cultivation near a garbage dump 
on the east shore of the lake near Lake City, Abaco. It 


1973] Bahama Flora — Gillis, Howard and Proctor 419 


may not persist, considering where it was found, but it 
nevertheless was healthy at the time of collection (1969), 
and appeared to be spreading (Gillis 7450). We are herein 
using the nomenclature of Bates (1965). 


TURNERACEAE 

Piriqueta caroliniana (Walt.) Urb. This species is com- 
mon on some of the islands, but can become exceedingly 
difficult to find if a flush of flowering is past. It was 
abundant one day and nearly invisible the next day at the 
same site, once the bright yellow flowers had closed and/or 
fallen. Now known from Abaco (Gillis 7426) in a sandy 
swale south of the Marsh Harbour airport, Great Exuma 
(Gillis 9383) from roadsides north of Georgetown, and on 
the former townsite of Freeport, Grand Bahama Island 
(Gillis 7848). Ornduff (1970) had previously reported it 
as absent from the Lesser Antilles or Bahamas. 

Piriqueta tomentosa Kunth. Found four miles west- 
northwest of Marsh Harbour, Abaco (Proctor 30434-A). 
In the same place, the glabrous form also was found (Proc- 
tor 30434, 30448). The latter is treated by some as P. 
viridis Small, but by others as only a glabrous form of 
P. tomentosa (Ornduff, 1970). 


CACTACEAE 

Harrisia undata (Pfeiff.) Britt. Proctor has collected 
this species on Grand Turk. His specimen fits the descrip- 
tion of this Cuban species fairly well, with the exception 
that the hairs of the bud-scales are longer and more 
abundant, and the spines on the average are somewhat 
fewer than those of the Cuban plant. Britton's type speci- 
men, however, was from cultivated material, and the pro- 
cess of cultivation may have produced modifications of 
these epidermal characters. On the other hand, the Grand 
Turk plant may represent a geographical race differing in 
these characters. 


COMBRETACEAE 
Terminalia muelleri Benth. This Australian native has 
been planted occasionally in Florida and the West Indies. 


120 Rhodora [Vol. 75 


It has been found, apparently escaped from cultivation, on 
Eleuthera (Proctor 30913). 


UMBELLIFERAE 

Oxopolis filiformis (Walt.) Britt. From Abaco, 8 miles 
southwest of Marsh Harbour (Proctor 30463). 

Hydrocotyle umbellata L. From Morgan's Bluff, North 
Andros (Proctor 30812). The old record of Dolley was 
disallowed by Britton and Millspaugh, and the species was 
thus excluded from the flora. The species, however, is now 
definitely confirmed from the Bahamas. 


OLEACEAE 
Jasminum fluminense Vell. Howard and Dunbar (1964) 
record this plant from Inagua. Native to Brazil, it is now 
weedy in the West Indies. 


GENTIANACEAE 
Sabatia stellaris Pursh. Reported by Lewis (1971) from 
Eleuthera. In his treatment of the North American spe- 
cies of this genus, Perry (1971) has also indicated that 
the species is known from Grand Bahama Island and 
Abaco as well. 


ASCLEPIADACEAE 
Cynanchum graminifolium (Griseb.) Alain. Reported 
by Howard (1950) as Metastelma graminifolium from 
Bimini. 
CONVOLULACEAE 
Merremia tuberosa (L.) Rendle. Gillis 7350 from Abaco, 
on the west shore of the lake near Lake City in a garbage 
dump. Gillis 7860 from near the Colonial Research Insti- 
tute compound on Grand Bahama Island in the vicinity of 
Freeport. 
Ipomoea nil (L.) Roth. Proctor 30783 from south of the 
San Andros airstrip on North Andros. 


VERBENACEAE 
Callicarpa americana L. Popenoe 129 from Abaco, 6 
miles north of the Hole-in-the-Wall at the south end of the 
island in a pineland. Proctor 30769 from thickets between 


1973] Bahama Flora — Gillis, Howard and Proctor 421 


cultivated fields south of the San Andros airstrip on North 
Andros. 

Clerodendrum speciosissimum Van Geert. This plant is 
known to escape from cultivation on roadsides near the 
Current, Eleuthera (Lewis, 1971). 


LABIATAE 

Micromeria bahamensis Shinners. Shinners (1962) felt 
that true Micromeria brownei (as treated in Bahama 
Flora) did not occur in the Bahamas. He set up three 
varieties of M. brownei, one from Jamaica, one from Cuba 
and Mexico, and one from the southeastern United States 
and Mexico. For plants with narrower leaf blades (4-6 mm 
vs. 5-20 mm) and short calyx teeth, he erected this new 
species, endemic to the Bahamas. He added that a later 
worker mght choose to make this species another variety 
of M. brownei. For the moment, we follow Shinners’s 
treatment. 

Satureja rigida Bartr. Proctor 30456 from Abaco, eight 
miles southwest of Marsh Harbour. 


SOLANACEAE 
Solanum eleagnifolium Cav. Proctor 30960 from Hatchet 
Bay Farms, Eleuthera. This attractive plant may well be 
in other of the western islands of the archipelago, since its 
native range includes the southern Florida Keys. 


PLANTAGINACEAE 
Plantago virginica L. Howard and Dunbar (1964) report 
this species from the flat land beyond Conch Shel] Point on 
Inagua. More recently, Gillis and Proctor have found it at 
Smith’s Thatch Pond on the southern coast of Inagua. 


RUBIACEAE 
Hedyotis nigricans var. filifolia (Chapm.) Shinners. 
Gillis 7838 from a ruderal site of Grand Bahama Island. 
Lewis (1971) reported the species from a similar locality. 
Proctor 30583 from Abaco, five miles west-northwest of 
Marsh Harbour. 
Morinda citrifolia L. This East Indies native is now 


422 Rhodora [Vol. 75 


widely naturalized in the West Indies. Proctor 8772 came 
from a comparatively remote sea-beach at the eastern 
shore of Grand Turk and thus believed to be a natural 
sea-borne introduction. 

Spermacoce aspera Aubl. Lewis (1971) tentatively re- 
ported this species as present at the Current, Eleuthera. 
The stems were upright to 0.3 m, and the corollas white. 
A specialist might segregate a microspecies from Lewis’s 
specimen, but for now, it is recognized as this species. 


LYTHRACEAE 
Rotala ramosior (L.) Koehne. Reported by Lewis 
(1971) from the edge of a sandy beach at Dead Man’s 
Reef, 9 miles east-southeast of West End, Grand Bahama 
Island. 


ACANTHACEAE 
Stenandrium carolinae Leonard et Proctor ex Leonard. 
See Leonard, 1960. This plant is an apparent endemic to 
North Caicos. It is known only from the type collection 
(Proctor 9094). 


COMPOSITAE 

Ageratum conyzoides L. Johnson (1971), in his treat- 
ment of the genus Ageratum, reported two subspecies of 
A. conyzoides from the Bahamas, both widely scattered 
among the islands, but neither appearing on the same is- 
lands together. Subspecies latifolium is the Ageratum 
latifolium of Britton and Millspaugh, but the typical sub- 
species conyzoides is herein reported as new to the Ba- 
hamas. 

Gaillardia pulchella Foug. This common species of south- 
ern United States was picked up by Howard (1950) as an 
escape on Bimini. 

Flaveria bidentis (L.) Kuntze. The record from Inagua 
by Howard and Dunbar (1964) was the first of this South 
American species from the West Indies but specimens from 
the Institute of Jamaica indicate that it has since been 
collected on Puerto Rico as well. 

Flaveria trinervia (Spreng.) C. Mohr. Proctor 30502 


1973] Bahama Flora — Gillis, Howard and Proctor 423 


from Abaco, about 10 miles south of March Harbour; 
Proctor 30784 from Morgan’s Bluff on North Andros. More 
recently, Gillis and Proctor have found it on Mayaguana. 

Senecio confusus Britten. Gillis 6372 from a cemetery 
at Calabash Bay, Andros where it escapes from cultivation. 
It does so as well on the south side of New Providence and 
probably elsewhere near habitations. 

Emilia javanica (Burm.) C. B. Robinson. It is not sur- 
prising that this weed in South Florida has turned up in 
the Bahamas. It has been collected on New Providence 
(Gillis 7303), Grand Bahama (Gillis 7827), Eleuthera 
(Lewis 7239), and Abaco (Proctor 30630). Its lavender- 
flowered relative, E. sonchifolia, more rare in South Flor- 
ida, was reported by Britton and Millspaugh from the 
Bahamas. 

Solidago sempervirens L. Howard (1950) reported this 
species from Bimini; Lewis (1971) from Grand Bahama. 
A population from Lake City, Abaco (Gillis 7351) has 
been determined by Dr. Gary Morton to be S. sempervirens 
var. mexicana (L.) Fern. 

Youngia japonica (L.) DC. Proctor 30759 found as a 
weed at Marsh Harbour, Abaco. We suspect that this plant 
has been introduced since the publication of Bahama Flora. 
We herein are following the nomenclature of Vuilleumier, 
1973. 


LITERATURE CITED 


ALEXANDER, TAYLOR R. 1968. Acacia choriophylla, a tree new to 
Florida. Quart. J. Florida Acad. Sci. 31: 197-98. 

Bates, Davip M. 1965. Notes on the cultivated Malvaceae. I. Hi- 
biscus. Baileya 13: 56-130. 

BRITTON, NATHANIEL LORD and CHARLES FREDERICK MILLSPAUGH. 
1920. The Bahama Flora. New York Botanical Garden. New 
York, N.Y. (Reprinted 1962, Hafner Publishing Co., New York, 
N.Y.). 

Brizicky, GEoRGE K. 1962. Taxonomic and nomenclatural notes on 
Zanthoxylum and Glycosmis (Rutaceae). J. Arnold Arbor. 43: 
80-93. 


424 Rhodora [Vol. 75 


CHASE, AGNES. 1929. The North American species of Paspalum. 
Contr. U.S. Natl. Herb. 28: 1-310 + xvii. 


CLAUSEN, R. T. 1936. Studies in the genus Najas in the northern 
United States. Rhodora 38: 333-45. 


DELISLE, DONALD G. 1963. Taxonomy and distribution of the genus 
Cenchrus. lowa State Univ. J. Sci. 37: 239-351. 


FRYXELL, PAUL A. 1969. The genus Cienfuegosia Cav. (Malvaceae). 
Ann. Missouri Bot. Gard. 56: 179-250. 


HiTCHCOCK, A. S. 1936. Manual of the grasses of the West Indies. 
USDA Misc. Publ. 243: 1-439. 


HITCHCOCK, C. LEO. 1945. The Mexican, Central American, and 
West Indian Lepidia. Madrofio 8: 118-43. 


HORVAT, SR. Mary Licourr. 1941. A revision of the subgenus 
Mariscus found in the United States. Contr. Biol. Lab. Catholic 
Univ. Amer. 33: 1-147 + x. 


HowaRp, RICHARD A. 1950. Vegetation of the Bimini Island group, 
Bahamas, B.W.I. Ecol. Monog. 20: 317-49. 


HowaRD, RICHARD A. and HENRY F. DUNBAR. 1964. Additions to 
the flora of Inagua, the Bahamas. Rhodora 66: 6-15. 


JOHNSON, MILES F. 1971. A monograph of the genus Ageratum L. 
(Compositae — Eupatorieae). Ann. Missouri Bot. Gard. 58: 6-88. 

JOHNSTON, MARSHALL C. 1971. Revision of Colubrina (Rhamna- 
ceae). Brittonia 23: 2-53. 

KRAL, ROBERT. 1971. A treatment of Abildgaardia, Bulbostylis, and 
Fimbristylis for North America. Sida 4: 57-227. 

LEONARD, EMERY C. 1960. Acanthaceae americanae novae vel criti- 
cae. Wrightia 2: 75-82. 

LEWIS, WALTER H. 1971. Additions to the flora of the Bahama 
Islands. Rhodora 73: 46-50. 

MARIE-VICTORIN, FR. 1948. Le genre Spathelia (Rutacées) avec 


description de cinq espéces et d'une variété nouvelles, Contr. 
Inst. Bot. Univ. Montréal 63: 14-48. Références et Notes, p. 76. 


MCLAUGHLIN, ANNE DESMOND. 1944. The genus Cyperus in the 
West Indies. Catholic Univ. Amer. Biol. Studies No. 5, pp. 1-108 
+ viii. 

MOHLENBROCK, ROBERT H. 1957 (appeared 1958). A revision of the 
genus Stylosanthes. Ann. Missouri Bot. Gard. 44: 299-355. 
PERRY, JAMES D. 1971. Biosystematic studies in the North Ameri- 

can genus Sabatia (Gentianaceae). Rhodora 73: 309-69. 


1973] Bahama Flora — Gillis, Howard and Proctor 425 


READ, RoBERT W. 1966. Coccothrinax inaguensis — a new species 
from the Bahamas. Principes 10: 29-35. 

SHINNERS, LLoyp H. 1962. Micromeria brownei and its allies (La- 
biatae). Sida 1: 94-7. 

STIMSON, WILLIAM R. 1967. Additions to the flora of the Bimini 
Island group, Bahama Islands. Rhodora 69: 60. 


VUILLEUMIER, BERYL SIMPSON. 1973. The genera of Lactuceae 
(Compositae) in the Southeastern United States. J. Arnold 


Arbor. 54: 42-98. 


W. T. GILLIS AND R. A. HOWARD 
ARNOLD ARBORETUM, HARVARD UNIVERSITY 
CAMBRIDGE, MASS. 02138 


G. R. PROCTOR, SCIENCE MUSEUM 
INSTITUTE OF JAMAICA 
KINGSTON, JAMAICA, B.W.I. 


REVISION OF THE GENUS PONTEDERIA L. 
RICHARD M. LOWDEN 


This small group of emersed aquatics is comprised of 
five species which are distributed throughout the tropical, 
subtropical and temperate regions of the New World. The 
center of diversity of Pontederia occurs in Middle America 
where the two recognized subgenera perhaps originated. 
A similar morphology, geography, cytology and chemistry 
suggest that Reussia Endl. does not warrant generic status 
apart from Pontederia. Subgeneric status is selected for 
Reussia since the species comprise a natural but distinct 
phyletie unit within Pontederia 

Pontederia is quite variable in habit and leaf shape. 
Individual species, i.e. Pontederia cordata L., have many 
leaf forms which have contributed to the creation of many 
unwarranted taxa and combinations in the past. This is 
partly attributed to the limited geographical scopes of 
previous taxonomic studies of the genus. It has been im- 
portant to recognize that distinct species have a develop- 
ment of similar leaf polymorphisms throughout a wide geo- 
graphical range. Also where it has been possible, floral 
and fruit characters are selected to elucidate the natural 
phyletie units that comprise Pontederia. 


NOMENCLATURAL HISTORY 


Over two centuries ago Linnaeus (1737a, b, 1754) de- 
scribed the genus Pontederia. More than a few attempts 
have been made to clarify the typification of the genus. 
In order to elucidate the past (and consequently the 
present) status of Pontederia it is necessary to examine 
the following chronology of historical events: 

Linnaeus (1737a) — Synonymy of Morison as communi- 
cated by Gronovius is cited along with the generic 
description of Pontederia, translated as follows: 
calyx: Spathe common, oblong, laterally dehiscing. 
corolla: Connate, biparted, gaping. Upper lip erect, flat, 


426 


1973] Pontederia — Lowden 427 


of 3 equal parts. Lower lip reflexed, 3 parted, segments 
equal. 

stamens: Filaments 6, awl-shaped, of which [1 stamen] 
is situated on each segment of the lower lip of the 
fused corolla, the 3 remaining [stamens] are closed 
under the upper lip. 

pistil: Ovary not completely rounded. Style simple, 
short. Stigma undivided. 

perigone: Capsule [little box or encasement] ovate [in- 
side], 3 compartments, 3-valved, 3-angled, 3-furrowed. 
seeds: Not completely rounded, many. 

Linnaeus (1737b) — Dedication of the genus to J ulio Pon- 
tedera, with special reference to the American Pickerel- 
weed, Maryland and Virginia, of Morison, Petiver and 
Plukenet. The genus Carimgolo of Malabar was men- 
tioned. 

Linnaeus (1753) — Three species of Pontederia were enu- 
merated in the class Hexandria: (1) P. ovata, with 
ovate foliage, flowers in heads, Narukila of Rheed. mal., 
habitat Malabar; (2) P. cordata, with cordate foliage, 
flowers in spikes, with identical reference made of the 
Hortus Cliffortianus, habitat Virginia; (3) P. hastata, 
with foliage hastate, flowers in umbels, Carimgola of 
Rheed. mal, habitat India. 

Linnaeus (1754) — Generic description is the same as that 
in the first edition (1737) except for the following 
additions (translated) : 
corolla: tubular. 
stamens: inserted on corolla, of which 3 are awl-shaped, 
long, and inserted on the throat of the corolla tube; 
3 remaining stamens attached at the base of the same 
tube. Anthers pointed inwards. 
pistil: Ovary oblong, above receptacle. Style simple, 
very short, curved downwards. 
perigone: Capsule [encasement] fleshy, conie, apex wide 
and bent inwards. 

Adanson (1763) — Narukila was listed with reference to 
Pontederia L. [Narukila is a nomen confusum.] 


428 Rhodora [Vol. 75 


Rafinesque (1808, 1830) — Established the genus Unisema 
from that element of Pontederia described by Linnaeus 
(1753) as P. cordata. *. .. L. [Linnaeus] positively says 
that the fruit of it [Pontederia] is 3 locular and many 
seeded [Linnaeus; 1737a, 1754]. I observed . . . the 
singular one seeded fruit [of P. cordata] and established 
the genus [Unisema] ..." 

Farwell (1924, 1928) — “As defined by Linnaeus in the 
Genera [1754], this genus is restricted to his P. hastata, 
the only species named by him in the Species Plantarum 
that had a many-seeded, 3-celled capsule, as was pointed 
out by Rafinesque . . ." 

Sprague (1924) — Carimgola (Linnaeus, 1737b) was not 
definitely cited as a synonym of Pontederia since locality, 
the Maryland and Virginia habitat, and synonymy refer- 
ences (Linnaeus, 1737 a,b) are of P. cordata. “Thus it 
is evident that in 1737 he [Linnaeus] gave the new 
generic name Pontederia to the species subsequently 
named by him P. cordata, and that he was uncertain 
whether Carimgola (P. hastata) was congeneric or not." 

Fernald (1925) — “The description of the genus Pontederia 
in the 5th edition of the Genera Plantarum (1754) was a 
mixture based upon the 2nd and 3rd species of the Species 
Plantarum; but in general the name has been maintained 
by post-Linnaean botanists for the American Pontederia 
cordata; Linnaeus’s 1st species, P. ovata [‘a plant with 
1 stamen'], clearly not belonging in the class Hexandria, 
being excluded as a member of the family Marantaceae, 
and the 3rd species, P. hastata, separated off as Mono- 
choria Presl.” *'*. , . in the 1st edition of the Genera, 
Linnaeus [1737a] gave the same mixed description as 
in the 5th [1754], the capsules 3-valved and many-seeded, 
but stated that the plant was communicated by Gronovius 
(from Virginia)." 

It is apparent from his three species that Linnaeus 
(1753) accidentally assembled dissimilar taxa under Pon- 
tederia. Farwell (1924) and Fernald (1925) point out 
that P. ovata having 1 stamen obviously did not belong 


1973] Pontederia — Lowden 429 


in the class Hexandria which includes P. cordata and P. 
hastata. Taxonomists must examine more critically the 
1st and 5th editions of the Genera Plantarum (Linnaeus; 
1737a, 1754) to know if Linnaeus (1753) actually con- 
sidered P. cordata or P. hastata as the “type” of the genus. 

Rafinesque (1808, 1830) insisted that the genus Ponte- 
deria as described by Linnaeus (1737a, 1754) called for 
fruits “3 locular and many seeded.” Thus, Rafinesque 
established his genus Unisema from the Linnaean P. cor- 
data having 1-seeded fruits. Farwell (1924, 1928) adopted 
Rafinesque’s decision. 

Sprague (1924) and Fernald (1925) defended P. cor- 
data as the intended “type” of the Linnaean Pontederia 
based on locality (the Maryland and Virginia habitat) and 
synonomy (Morison, Petiver, Plukenet and Gronovius) as 
presented by Linnaeus (1737 a,b, 1753, 1754). Fernald 
(1925) also appealed to established custom which until 
today treats P. hastata as the Asian Monochoria and P. 
cordata as the American Pontederia, This appeal has been 
enforced by “principal systematic botanists” such as, Pursh 
(1814), Nuttall (1818), Torrey (1824), Solms-Laubach 
(1883), Britton and Brown (1913), Sprague (1924), Fer- 
nald (1925), Schwartz (1927), Hitchcock and Green 
(1929), and Castellanos (1951). All have retained and 
reinstated Pontederia cordata L. as the type of the Lin- 
naean Pontederia. 

The interpretation by Rafinesque (1808, 1830) of the 
Linnaean generic description of Pontederia (173a, 1754) 
was accepted beyond doubt by Sprague (1924) and Fernald 
(1925). Fernald openly admitted Rafinesque’s interpreta- 
tion that the Genera Plantarum (1737a, 1754) called for 
“capsules 3-valved and many-seeded” and therefore Lin- 
naeus based his generic description on a mixture of the 
2nd (P. cordata) and 3rd (P. hastata) species of the 
Species Plantarum (1753). Both Sprague and Fernald 
sought valid evidence (locality and synonomy) in the 
Genera Plantarum (1737a, 1754) for support of P. 
cordata as “type” ignoring the generic description of the 


430 Rhodora [Vol. 75 


Figure 1. Hardened Perigone Bases of Pontederia. A. Tooth- 
ridged perigone of P. cordata var. cordata, U.S., Ohio (Lowden, 36, 
37), 8 mm long. B. Smooth-ridged perigone of P. sagittata, Mexico 
(Lowden 6), 8 mm long. C. Tooth-ridged perigone of P. parviflora, 
Panama (Lowden 21), 8 mm long. D. Spinulose-ridged perigone of 
P. rotundifolia, Nicaragua (Lowden 17, 18), 10 mm long. 


1973] Pontederia — Lowden 431 


Linnaean protologue and the fact that it was misinterpreted 
by Rafinesque. 

Rafinesque (1808, 1830) confused the modern termi- 
nology of “capsules 3-loculed and many seeded" (which 
actually applies to P. hastata) with the Linnaean choice 
of terminology in the Genera Plantarum. Linnaeus (1737, 
1754) used “capsula” to mean a “small box" or “encase- 
ment" and it was properly identified by him as part of the 
perigone (perianth). In general appearance the mature 
perigone of P. cordata (Fig. 1A) is triangular in all aspects 
and is rounded inside.. Linnaeus never stated as Rafinesque 
insisted that the fruits were very many seeded, he only 
stated there were many seeds which implies without as- 
sumption each plant or inflorescence bears many seeds. This 
as does the rest of the generic description of the Genera 
Plantarum (1737a, 1754) fits perfectly P. cordata and can 
not be confused with the Asian P. hastata (Monochoria 
hastata). Pontederia hastata, with flowers actinomorphie, 
petals nearly free to the base (corolla not tubular), sta- 
mens 6 of which 1 is longer and seeds round, can mot 
possibly be confused with the Linnaean P. cordata, with 
flowers zygomorphic, perianth 2-lipped, petals connate 
(tubular), stamens 6 of which 3 are longer and seeds not 
completely rounded. The additional wording of the 5th 
edition of the Genera Plantarum (1754) reveals that Lin- 
naeus was describing the short style floral form (Fig. 3A, 
3 stamens longer and a very short style) of tristylous P. 
cordata. The perigone encasement (capsule) was accurately 
described as fleshy, conic, with apex (Fig. 1A, mature 
stamens and upper portion of perigone) wide and bent 
inwards, 

The Linnaean generic description of Pontederia (1737a, 
1754) is exclusively that of P. cordata and not P. hastata. 
Linnaeus (1753) was sure of the morphological structure 
of P. cordata as it was explicitly expressed by him in the 
Genera Plantarum. Linnaeus accidentally included P. ovata 
and P. hastata in the Species Plantarum (1753) as he 
accidentally mentioned Carimgolo (— P. hastata) in his 


432 Rhodora [Vol. 75 


Figure 2. Fruits and Seeds of Pontederia. A. Utricle of P. parvi- 
flora, Panama (Lowden 21), 8.5 mm long. B. Portion of copious 
endosperm surrounding embryo of P. parviflora, Panama (Lowden 
21), 4 mm long. C. Utricle of P. rotundifolia, Nicaragua (Lowden 
17, 18), 8.5 mm long. D. Ovoid seed of P. rotundifolia, Nicaragua 
(Lowden 17, 18), 3 mm long. 


1973] Pontederia — Lowden 433 


Hortus Cliffortiamus (1737b). It is difficult to understand 
why Linnaeus mentioned these taxa (perhaps they ap- 
peared superficially similar), since he made no mention 
of them in the generic description of the Genera Planta- 
rum. This revision adopts Pontederia cordata L. as the 
lectotype of the Linnaean Pontederia. 

In Solms-Laubach’s treatment (1883) of the Ponte- 
deriaceae, Pontederia L. was recognized as being generally 
distinct from Reussia Endl. Both have one-seeded inde- 
hiscent fruits and two-lipped perigones (perianth). 
Pontederia was characterized by having 3 lobes in each 
lip, whereas, Reussia was characterized by having 5 lobes 
in the upper lip and 1 lobe in the lower lip. This was 
followed by Schwartz (1927, 1930), Schulz (1942) and 
Castellanos (1951, 1958). In accordance with this dis- 
tinction, Castellanos (1951) transferred Pontederia ro- 
tundifolia L. f. to Reussia based on his reported observa- 
tion of 5 lobes in the upper lip of the perigone. 

My observations indicate that Pontederia species in- 
cluding P. rotundifolia have 3 lobes in each lip, even 
though the deepest incisions of the perigone occur on both 
sides of the smallest lobe of the lower lip. Contrary to 
later opinions, Endlicher (1836) described Reussia as 
having 4 lobes in the upper lip and 2 lobes in the lower 
lip. Certainly, the number of lobes in each lip is a con- 
fused and weak distinction for separating Pontederia and 
Reussia. Additional evidence obtained in the present in- 
vestigation indicates that Pontederia and Reussia are so 
morphologically, cytologically and chemically similar as 
to warrant the treatment of Reussia (nom. cons.) as a 
subgenus of Pontederia in accordance with the International 
Code (1972, Appendix III, p. 284 and Article 14). 


DISPERSAL MECHANISMS 


Pontederia inhabits inland fresh waters and marshes 
transitional to salt waters along the coasts. It is primarily 
a genus of tropical and subtropical America, whose habi- 
tats are probably held in check by latitude (temperature) 


434 Rhodora [Vol. 75 


and altitude (elevation). Mountain ranges, as exemplified 
by the Appalachian (Core, 1966) and Rocky Mountains 
in North America, the Sierras Madre Oriental and Occi- 
dental in Mexico, Sierras del Mico and Minas in Guatemala, 
Mayan Mountains in Guatemala and British Honduras, 
Cordillera Isabelia in Honduras, Cordillera de Talamanca 
in Costa Rica and the Andes in South America have been 
natural physiographic barriers promoting the speciation 
of Pontederia in the Americas. Geographical isolation is 
greatest among taxa within Middle America such as, 
P. sagittata (Fig. 16) and P. parviflora (Fig. 19). 
Dispersal by water of Pontederia utricles is thought to 
be the main mode of long range dispersal, with vegetative 
reproduction a prominent factor in population establish- 
ment. The utricle is buoyant and consequently its dis- 
tribution is due to the light aeriferous tissue of the perigone 
base (Fig. 1) surrounding it. Schulz (1942) reports a 
flotaion period longer than fifteen days for seeds enclosed 
in perigone bases which is long enough for the fruit to 
travel a considerable distance. The presence of fleshy 
perigone bases surrounding the utricles and the copious 
endosperm contained inside (Fig. 2B) greatly increase 
survival expectancy over long distances traveled. 


Long distant dispersal of utricles (Fig. 2A, C) by avian 
and terrestrial animals (Sculthorpe, 1967) is considered 
to be of less frequent occurrence. Field observations of 
Pontederia rotundifolia at the Laguna Zapotitan, Dept. 
La Liberatad, El Salvador and in the fields of Lago Arenal, 
Province Guanacaste, Costa Rica suggest limited local ter- 
restrial dispersal by animals. Both areas are inhabited 
by livestock on which the spinulose perigone bases (Fig. 
1D) are easily anchored. 

Vegetative reproduction by trailing prostrate branched 
stems and by rhizomes is of frequent occurrence throughout 
populations. Clones are recognized by clumping of plants 
within populations (Fig. 9). Rhizome fragmentation 
results in the establishment of new clones within close 
proximity. Field observations indicate that the prostrate 


1973] Pontederia — Lowden 435 


branched stems of P. rotundifolia with adventitious roots 
at each node may easily become severed and rooted. Trail- 
ing stems and rhizomes are able to survive harsh environ- 
mental pressures as organs of food accumulation during 
the climatie fluctuations of temperate and tropical regions. 


TRISTYLY 


Tristyly, as the term relates to Pontederia, is the type 
of heterostyly in which three kinds of plants occur in a 
species. These are (1) plants with flowers having a SHORT 
STYLE, 3 medium stamens and 3 long stamens (Sml) ; (2) 
plants with flowers having 3 short stamens, a MEDIUM 
STYLE and 3 long stamens (sMI); and (3) plants with 
flowers having 3 short stamens, 3 medium stamens and a 
LONG STYLE (smL. The Sml, sMl and smL floral forms 
(Fig. 3A-C) were found in Pontederia cordata (including 
varieties), P. sagittata, P. rotundifolia and P. subovata. 
Only P. parviflora has plants with 3 short stamens, 3 long 
stamens and a LONG STYLE (slL). The sIL floral form 
(Fig. 3D) is an excellent diagnostie character for dis- 
tinguishing this homostylous taxon from its closely related 
tristylous counterparts of subgenus Pontederia. 

Legitimate pollinations (Fig. 4, pollen transfer from a 
stamen to the stigmatic surface of a pistil of equivalent 
length) are more productive of seeds than illegitimate 
pollinations (Fig. 4, pollen transfer from a stamen to the 
stigmatic surface of a pistil not the equivalent length). 
Ornduff (1966) demonstrated with Pontederia cordata, 
that self-incompatibility (self and own-form pollinations 
are illegitimate) is “considerably stronger" in the smL 
and Sml floral forms than in the sMI floral form. Regard- 
ing legitimate pollinations the sMI floral form had a 
greater percentage seed production than the smL and Sml 
floral forms. 

An indieation of this breeding system of individual 
populations may be extrapolated through the study of the 
frequency of floral forms within populations of different 
Pontederia taxa (Table 1). In populations where all three 


436 


Table 1. 


Rhodora 


Herbarium Studies’ 


Taxon 


Populations & 


[Vol. 75 


Floral Form Frequency in Pontederia based on Population and 


# (96) Inflorescences 


Vouchers Sml 


sMI smL slL 


P. cordata 
var. 
cordata 


var. 
lancif olia, 


var. 
ovalis 


U.S. Ohio, Ottawa 8 (38.8) 
Co., Winous Pt. SW 
of Port Clinton, 
Lowden 36 & 37. 
British Honduras, 
Belize District: 9 
miles N of Belize 
City, Lowden 24; 

29 miles N of 
Belize City, 
Cowhead Creek, 
Lowden 26; 

32 miles N of 
Belize City, 
Lowden 27; 
Hattieville 

Burrell Boom 
Road, 514 miles 
from Burrell 

Boom, Lowden 28. 


5 (62.5) 


0(0) 


2 (66.7) 


0(0) 


U.S. Florida, + 
Holmes Co., 0.2 


miles E of Florida 


81 in Ponce de 
Leon, Stone 2589 
Sml, 2590 smL & 
2591 sMI. 


Brazil. Mato + 
Grosso, 300 km. 

past Cuiaba in 

route to Goiania, 

Magwire et al 

56931 Sml; Brasilia, 


5(20.8)  11(45.8) — 


2 (25.0) 1(12.5) — 


0(0) 3 (100) — 


1(33.3) 0 (0) — 


2 (40.0) 3(60.0) — 


'Code to floral forms: (Sml) SHORT STYLE, 3 medium stamens, 3 long stamens; 
(sM1) 3 short stamens, MEDIUM STYLE, 3 long stamens; (smL) 3 short stamens, 
3 medium stamens, LONG STYLE; (slL) 3 short stamens, 3 long stamens, LONG 
STYLE. A plus sign (+) indicates the occurrence of the floral form, whereas a 
dash (—) indicates the floral form is unknown in the taxon. 


1973] 


Table 1. 


Taxon 


Pontederia — Lowden 


(cont.) 


Populations & 
Vouchers 


# (96) Inflorescences 


437 


Sml 


sMI smL 


slL 


P. 
sagittata 


P: 
parviflora 


Federal District, 
Prance & Silva 59082 


smL. Colombia, Dept. 


Boyaca, near Orocue, 
Haught 2716 sMI. 
Mexico. State 
Vera Cruz: Vera 
Cruz, El Coyol, 
Lowden 6; 

El Puente de 
Teculapilla, 
Carretera Nacional 
180, 5 km. NW of 
Lerdo, Lowden 7; 
Laguna Catemaco, 
Arroyo Agrio, 
Lowden 9. 


Guatemala. Dept. 
Izabal: Quirigua, 
Lowden 10; 
Puerto Barrios, 
Lowden 11 


Honduras. Dept. 
Cortes: Puerto 
Cortes, Lowden 13; 
13.5 miles inland 
from Puerto Cortes, 
Lowden 14. 

Dept. Atlantida: 
Tela, La Curva, 
Lowden 15; 
Santiago, 11 

miles W of Tela, 
Lowden 16. 


Panama. Province 
Herrera, Los 
Llanos de Santa 
Maria, Lowden 20; 


46 (33.8) 


5 (55.6) 


10 (76.9) 


2(9.1) 


10 (82.3) 


0(0) 


16 (47.1) 


15 (34.1) 


21 (55.3) 


52(38.2) | 38(27.9) 


LOLLL) 3 (33.3) 


3 (23.0) 0(0) 


12(54.5) 8(36.4) 


14(45.2) 7 (22.2) 


6(37.5)  10(62.5) 


7(20.6)  11(32.3) 
16(36.4)  13(29.5) 


12(31.6) 5(13.2) 


89 (100) 


438 


Table 1. 


Taxon 


Rhodora 


(cont. ) 


Populations & 


[Vol. 75 


# (%) Inflorescences 


Vouchers Sml 


sMI smL 


P. 
rotundifolia 


P. subovata 


Province Panama, -— 
between Pacora 

and Chepo, Lowden 

21; 

17 herbarium — 
specimens cited 

under specimens 

examined. 


El Salvador. 
Dept. La 
Libertad, 

Laguna Zapotitan, 
Lowden 23. 


0 (0) 


Honduras. Dept. 
Comayagua, Lago 
Yojoa, Pito 

Solo, Lowden 12. 


0 (0) 


Nicaragua. Dept. 
Granada, near 
Granada, Lowden 
17, 18. 


0 (0) 


Costa Rica. 

Province 
Guanacaste, 1 km. 
from Arenal, 2 

km. from Trenadora, 
Lowden 19. 


184 (100) 


Paraguay, prope RE 
Concepcion, 
Hassler 1352 Sml; 
Brazil, Estado 
Minas, Mun. 
Ituiutaba, Loc. 

S. Terezuiha, 
Marcedo 3168 sMI; 
Argentina, Prov. 
Chaco, Loc. Colonia 
Benitez, Meyer 3864 
smL. 


0 (0) 29 (100) 


30 (81.1) 7 (18.9) 


1?(0.7?) 


145(99.3) 


0 (0) 0 (0) 


83 


1973] Pontederia — Lowden 439 


Figure 3. Floral Forms of Pontederia. A. SHORT STYLE, 3 
medium stamens, 3 long stamens (Sml); P. rotundifolia, Costa Rica 
(Lowden 19), 10 mm long. B. 3 short stamens, MEDIUM STYLE, 
3 long stamens (sMl1); P. rotundifolia, Honduras (Lowden 12), 13 
mm long. C. 3 short stamens, 3 medium stamens, LONG STYLE 
(smL); P. rotundifolia, Nicaragua (Lowden 17, 18), 11 mm long. 
D. 3 short stamens, 3 long stamens, LONG STYLE (slL); P. parvi- 
flora, Panama (Lowden 21), mm long. 


440 Rhodora [Vol. 75 


LEGITIMATE 


Figure 4. Legitimate and illegitimate pollinations in Pontederia 
floral forms (arrows represent the transfer of pollen from anthers 
to stigmas). See Fegure 3 for code to floral forms. 


1973] Pontederia — Lowden 441 


floral forms are present, it is difficult, only on the basis 
of frequency of occurrence, to know if any one floral form 
has greater breeding value compared with another floral 
form in the same population. However, in Pontederia 
rotundifolia three out of four populations examined 
(Table 1) had the frequent occurrence of one floral form 
in the absence of the other two (e.g., the smL floral form 
was found at Lago Nicaragua, Nicaragua and is absent at 
Lago Arenal, Costa Rica where the Sm1 floral form was 
found). These populations perhaps represent clonal estab- 
lishment of a single floral form in which a certain amount 
of inbreeding must be occurring through illegitimate pol- 
linations (selfing and own-form). Even though an in- 
compatibility system is operative there is not total incom- 
patibility in populations containing a single floral form. 

The homostylous Pontederia parviflora has most likely 
been derived from a tristylous floral form. This is perhaps 
a step towards increased self-compatibility in one member 
of the genus. Field observations (in Panama) indicate a 
compatibility system due to the high seed set of infructes- 
cences. 

In Pontederia a reproductive system has evolved which 
is highly tolerant to different pressures of the environment. 
Tristyly functions in juxtaposition with an effective mode 
of vegetative reproduction. Subgenus Reussia is charac- 
terized by having few flowered inflorescences (less seed 
production) and long trailing stems. Stem fragmentation, 
especially in populations having one floral form (P. ro- 
tundifolia), has greater immediate population survival 
value than reproduction through a tristylous breeding 
system. In subgenus Pontederia, inflorescences are many 
flowered (greater seed production), and the erect, above 
ground stems are much shorter. In this subgenus, tristyly 
perhaps has greater long range survival value and homo- 
styly (P. parviflora) appears to be derived from a tri- 
stylous ancestor. 


CYTOLOGICAL STUDIES 
Smith (1898) recorded 1—8 and 2n—16 for Pontederia 


442 Rhodora [Vol. 75 


C D 


Figure 5. Chromosome Configurations in Microsporocytes of Pon- 
tederia. A. Eight bivalents (diakinesis) of P. cordata var. cordata, 
U.S., Ohio (Lowden 36, 37). B. Secondary association of six chromo- 
somes (arrow points to three bivalents) of P. cordata var. cordata, 
U.S., Ohio (Lowden 36, 37). C. Sixteen bivalents of P. rotundifolia, 
Nicaragua (Lowden 17, 18). D. Diagramatic representation of C, 
showing sixteen bivalents of P. rotundifolia. 


1973] Pontederia — Lowden 443 


cordata. The cytological investigations of Bowden (1945) 
verified Smith's counts. This study reconfirms these earlier 
reports and elucidates counts (Table 2) for other Ponte- 
deria taxa. 

Young Pontederia inflorescences still enclosed by the 
spathe and the petiole base of the flower stalk leaf were 
collected for microsporocytes undergoing meiosis. Inflor- 
escences were placed in 100% ethanol and glacial acetic 
acid (3:1 respectively). After 24 hours, they were stored 
in 70% ethanol until anthers were stained with iron aceto- 
carmine. 

Microsporogenisis in taxa examined of subgenera Ponte- 
deria and Reussia is normal except for the occasional 
occurrence of secondary associations of either 4 or 6 
chromosomes (Fig. 5B). Haploid counts (Table 2) during 
diakinesis (Fig. 5) indicate a base number of z=8 in 
Pontederia, Subgenera of Pontederia might be distin- 
guished cytologically by the numbers, n=8 in subg. Ponte- 
deria and n=16 in subg. Reussia. The polyploid P. ro- 
tundifolia and possibly P. subovata (not examined) most 
likely evolved from an n=8 ancestor. 


CHEMOTAXONOMIC STUDIES 


Phenolic compounds, especially flavonoids, have recently 
been the subject of a large number of chemosystematic 
studies. These studies (Alston, 1967) have shown that 
color and position of phenolic compounds on chromato- 
grams can be used to show systematic relationships with- 
out necessitating chemical characterization. Considering 
the validity of this empirical chromatographic data the 
following chemical studies were initiated in order to assess 
the degree of variation in phenolic patterns within and 
between taxa of Pontederia, to see if phenolic patterns of 
Pontederia taxa correlate with that indicated by mor- 
phology and geography, and to note if similarity in phenolic 
patterns exists between taxa of Pontederia, Eichhornia 
and Heteranthera (related genera). 

The methods employed are essentially those outlined by 


K 


444 Rhodora [Vol. 75 


Mabry, Markham and Thomas (1970). Phenolic compounds 
were extracted from dried flower stalk leaf material in 
50% aqueous-methanol for at least 24 hours before 4 ml 
of individual leaf extract were spotted on Whatman 3MM 
chromatographic paper. The two-dimensional solvent 
systems were: (1) 3 parts TBA : 1 part glacial acetic 
acid : 1 part distilled water and (2) 15% HoAc. Chromat- 
ographic patterns were read using no reagent in visible 
light, then in Uv light with and without NH.. 

Twenty-six principal spots (Fig. 6) were discernible 
by position and color on chromatograms. Color charac- 
terizations (Table 3) were uniform throughout chromato- 
grams, however, for spots with mixed color indications, 
it is possible that more than one phenolic compound is 
present. Spot numbers 11 and 15 (Fig. 6) were subdivided 
because color and close proximity of spots prevented con- 
trasting delineations. 

Summary phenolic patterns (Table 4) for each taxon 
were compiled using the total number of spots found 
in each taxon. Infrageneric and  intergeneric com- 
parisons of summary phenolic patterns were based on 
positive matches (a spot occurring in two taxa compared) 
and differences (a spot occurring in one of the taxa com- 
pared but not in the other). The degree of relationship 
between pairs of taxa was calculated using the coefficient 


of similarity — "where p represents the positive matches 
and d the differences. Coefficients vary from 0 (no resem- 
blance) to 1 (identity). 

Chromatographic patterns were subject to interpreta- 
tion, although delimitations of infraspecific and interspecific 
taxa of Pontederia were not based strictly on phenolic 
patterns. Chromatograms were initially separated as to 
summary phenolic patterns (Table 4) in accordance with 
the morphological concept of taxa (see Taxonomy sec- 
tion). This morphological alignment of taxa was reinforced 
by the high coefficients of similarity (Fig. 7) obtained 
among taxa. The nine spots shared in common by all taxa 
of Pontederia examined (Table 4) are perhaps chemical 


1973] Pontederia — Lowden 445 


HOAc (2) 


TBA (1) 


Figure 6. Summary two-dimensional paper chromatographic pro- 
file of phenolic compounds found in Pontederia and related genera. 


indications of a similar operative genetics clearly visible 
by the close morphological identity among principal taxa. 

Infrageneric comparisons (Fig. 7) indicate higher 
identity among typical Pontederia cordata, P. sagittata, 
P. parviflora and P. rotundifolia than among these taxa 
with P. cordata varieties lancifolia and ovalis. There was 
a stronger affinity of these two varieties with P. rotundi- 
folia and P. parviflora than with typical P. cordata. This 
affinity should be tested further considering the very small 
sample sizes of varieties lancifolia and ovalis (Table 4). 

High phenolic identity, greater variation in phenolic 
patterns, greater morphological similarity and like 
chromosome complement were found between typical P. 
cordata and P. sagittata, each containing an isolated spot 
(spot 15b and 22 respectively). Perhaps these single iso- 
lated spots are the development of metabolic pathways in 
fairly restricted geographical regions (Figs. 10-12, 15-16). 
The highest phenolic similarity was between P. parviflora 
and P. rotundifolia. Partial and fluctuating geographical 
isolation in Central and South America (Figs. 19, 22, 25) 


446 Rhodora [Vol. 75 


82^ N59 
Pa f / \ YS. 
4^ /^8 MPO 
N 
va. in — / N 


Sp. 
CORDATA ROTUNDIFOLIA 


var. 
LANCIFOLIA 


Sp. 
PARVIFLORA 


Sp. 
SAGITTATA 


Figure 7. Graphic representation comparing two dimensional pa- 
per chromatographic profiles of Pontederia based on coefficients of 
similarity. 


1973] Pontederia — Lowden 447 


could account for a parallel and independent development 
or loss of metabolic systems in these two morphologically 
and cytologically distinct species. 

Intergeneric comparisons of phenolic patterns and 
morphology reveal a greater difference between Pontederia, 
Eichhornia and Heteranthera than among taxa of Ponte- 
deria. Seven out of the nine spots common to Pontederia 
taxa were shared equally with Eichhornia and Heteranthera 
taxa examined. No spots (Table 4) were added to the 
total number of spots found in Pontederia by the inclusion 
of Eichhornia and Heteranthera taxa. Undoubtedly similar 
metabolie systems have evolved in the Neotropical Pon- 
tederiaceae, perhaps divergently from an ancestral system 
comparable with that found in typical P. cordata and P. 
sagittata. 


EVOLUTIONARY CONSIDERATIONS 


A speculative origin of Pontederia is attempted based 
on the present studies. The ancestral stock is hypothesized 
as aquatic, tropical in origin, inflorescence a many-flowered 
spike, fruit a one-seeded utricle, ovule pendulous, tristylous, 
six stamens, flowers zygomorphic and _ perianth parts 
basally connate. Cytological evidence indicates a haploid 
base chromosome number of x—8. Chromatographic pat- 
terns of phenolic compounds suggest that similar metabolic 
systems have evolved. 

By the Lower Eocene, Pontederia had a restricted tropi- 
cal range from South America to North America via the 
Middle American landbridge. On several occasions from 
the Upper Eocene to the Lower Pliocene (Schuchert, 1935) 
portions of Middle America were inundated. Speciation 
occurred in isolated areas as this isthmian landbridge rose 
and fell. Changes in environmental conditions (Baker, 
1970), due in part to climatic instability and mountain 
building, were accompanied by floristic changes. A com- 
ponent of this change was Pontederia which persists. today 
throughout Middle America. 

Pontederia spread from Middle America into North 


448 Rhodora [Vol. 75 


America where it had perhaps at one time a more western 
distribution as indicated by the Green River Formation of 
Eocene Age. Pontederites (Knowlton, 1922) was described 
from the lake margin fossil flora of the Green River 
Formation in northwestern Colorado and the Gosiute Lake 
in southwestern Wyoming. The fossil material is an upper 
portion of a leaf fragment which looks like the charac- 
teristic venation of living Pontederia. The Green River 
Formation was bordered on the north and west by the 
Rocky Mountains. The barriers were less containing to 
the east and south towards the Gulf Coastal Plain which 
was covered with water during the Eocene. As the 
water receded Pontederia was redistributed to the warmer 
subtropical climate of the Gulf Coastal Plain where it 
started to inhabit the Atlantic Coastal Plain of North 
America. The Appalachian Mountains (Core, 1966) were 
then and still are the main obstacle (Fig. 11) to its spread 
into the interior of the eastern United States. 

The advent of the Pleistocene brought deleterious climatic 
changes resulting in a restricted southerly distribution of 
Pontederia toward the warmer climate of the Gulf Coast. 
As the climate fluctuated during glacial and interglacial 
periods, fluctuations also occurred in the distribution of 
Pontederia along the Atlantic Coast. Glacial withdrawal 
(Sculthorpe, 1967) brought an increasing warmth accom- 
panied by an optimum post glacial maximum warmth, 
culminating in today’s climatic conditions. The retreat 
of the last ice-sheets left behind numerous lakes, bogs and 
ponds. Since the Wisconsin glacier, Pontederia has spread 
and thrived throughout these aquatic glacial remnants of 
eastern North America reaching its present northern and 
northwestern limits within relatively recent times. 


TAXONOMY OF PONTEDERIA L, 


Pontederia L., Sp. Pl. 288. 1753. Gen. Pl. ed. 5, 140. 1754. 
Lectotype: Pontederia cordata. L. 

Umsema Raf. [Unisema Raf., Med. Fl. 2: 105. 1830.], 
Med. Repos. II 5: 352. 1808. 


1973] Pontederia — Lowden 449 


Reussia Endl., Gen. Pl. 1: 139, 1836. Type: Reussia triflora 
Endl. ex Seub. = Pontederia subovata (Seub.) Lowden. 


Perennial emersed herbs. Rhizomes branched, aerial 
stems erect or prostrate. Leaves obtuse; margins entire 
with parallel outwardly curved venation; blades (phyl- 
lodes) lanceolate, ovate, cordate, sagittate, hastate or reni- 
form; foliage dark green to grayish, often containing 
anthocyanin. Petiole of floral shoot leaf short; petioles of 
other leaves long; bases sheathed and clasping, sheaths 
ligulate. Inflorescence a few to many flowered spike en- 
closed (early in floral development) by a spathe (the most 
terminal phyllode of the floral shoot) and petiole base of 
the flower stalk leaf; peduncles longest with maturation 
of infructescences and subtended by the spathe; sheath 
(expanded phyllode base) in axil of floral shoot. Flowers 
bisporangiate, zygomorphic (bilaterally symmetrical), bi- 
labiate, blue to purple, white or white-green, largest 
perianth lobe with a bilobed yellow spot; perigone (peri- 
anth) of 6 basally connate parts; stamens 6 unequal, in two 
groups of 3 each, adnate to the perigone tube; anthers 
introrse, dorsifixed, versatile, dehiscing longitudinally; 
filaments and perigone tube covered with hairs terminated 
in pinheadlike glands; stigmas 3-lobed, each lobe bifid; tri- 
stylous or homostylous species; carpels originally 3, united, 
2 aborted and 1 fertile, unilocular, 1-seeded; ovules pen- 
dulous, anatropous, placentation parietal. Fruit a utricle, 
l-seeded, indehiscent; enclosed by a hardened perigone 
base, toothed, smooth or spinulose ridged; pericarp high 
in glutanous content. Seeds reniform or ovoid; endosperm 
white, copious, surrounding a cylindrical embryo. 


Key To The Taxa 


a. Hardened perigones toothed or smooth ridged (Fig. 
1A-C); floral bearing shoots erect (subgenus Ponte- 
demna) ia a cae ee b. 


b. Style length unequal to stamen length (Fig. 3A-C) 


150 Rhodora [Vol. 75 


c. Hardened perigones toothed ridged (Fig. 1A) 


LX» **Ps92329**»T9T*F*V*»922»XT223-«V*99-X2»1 d. 
d. Leaves cordate, sagittate (unauricled), hastate 
or reniform .... la. P. cordata var. cordata 


d. Leaves narrow to broadly lanceolate (leaf 
base not lobed) ......................... 
............. lb. P. cordata var. lancifolia 

d. Leaves ovate to ovate-lanceolate (leaf base 
not lobed) ; peduncles hairy .............. 
nan le. P. cordata var. ovalis 

c. Hardened perigones smooth ridged (Fig. 1B); 
leaves sagittate (usually deeply auricled) 

TM 2. P. sagittata 

b. Style as long as the three longest stamens (Fig. 3D) ; 


leaves subcordate .............. 3. P. parviflora 
a. Hardened perigones spinulose ridged (Fig. 1D) ; floral 
bearing shoots prostrate (subgenus Rewssia) ...... e. 


e. Flowers long lasting (more than 8 flowers per in- 
florescence) ; leaves reniform, sagittate or cordate 
TEM 4. P. rotundifolia 

€. Flowers ephemeral (usually fewer than 12 flowers 
per inflorescence) ; leaves subovate, sublanceolate or 
elliptical ............... lessen 5. P. subovata 


I. Pontederia subg. Pontederia (Lectotype:  Pontederia 
cordata, L.) 

Hardened perigone bases toothed or smooth ridged (Fig. 
1A-C) ; floral bearing shoots erect; aerial stems with short 
internodes; underground rhizomes long (mature plants). 

Subgenus Pontederia is found 50° North to 35° South of 
the equator in the Americas. 


1. Pontederia cordata L., Sp. Pl. 288. 1753. 


Plants 9-13 dm tall. Leaves cordate, sagittate (un- 
auricled), ovate or lanceolate; blades 0.4-21 em wide; spathe 
(terminal phyllode of floral shoot) 2-7.4 cm long; petiole 
of floral shoot leaf (includes sheath base) 4-31 cm long; 
petiole of other leaves 28.7-59.5 cm long, ligule of petiole 
sheath 4-10.5 em long; sheath in axil of floral shoots 22.3- 


1973] Pontederia — Lowden 451 


Figure 8. Inflorescence, clasping subtending spathe and flowering 
stalk leaf of Pontederia cordata var. cordata, U.S., Ohio, Ottawa 


County (Lowden photo, Jul 1966) ; upper figure. 

Figure 9. Clones of Pontederia cordata war. cordata in marsh 
habitat, U.S., Ohio, Ottawa County (Lowden photo, Jul 1966) ; lower 
figure. 


452 Rhodora [Vol. 75 


28 cm long. Inflorescence 2-16 cm long; peduncle (in- 
florescence base to floral shoot leaf base) 5-33 cm long. 
Perigone blue to white; hardened perigone bases (Fig. 1A) 
toothed ridged; anthers blue. Tristylous. Fruits and seeds 


reniform. 


la. Pontederia cordata L. var. cordata. Type missing: 

United States, Virginia and Maryland, communicated to 

Linnaeus by Gronovius. Neotype chosen: [Savage, 407.4] 

P. Kalm (LINN). 

Umsema obtusifolia Raf., Med. Repos. II, 5: 352. 1808. 
Based on the type of Pontederia cordata L. 

Umsema mucronata Raf., Med. Repos. II, 5: 352. 1808. 
Based on the type of Pontederia cordata L. 

Pontederia angustifolia Pursh, Fl. Am. Sept. 224. 1814. 
Type: United States: New York to Carolina (Holotype 
OXF?). 

?Unisema sagitata Raf., Fl. Ludov. 18. 1817. Type not 
seen. 

Pontederia cordata L. var. angustifolia (Pursh) Torrey, 
Fl. U.S. 1: 343. 1824. 

Unisema deltifolia Raf., Med. Fl. 2: 105, 107. 1830. Based 
on the type of Pontederia cordata L. 

Unisema purshiana Raf., Med. Fl. 2: 107. 1830. Based on 
the type of Pontederia angustifolia Pursh. 

Unisema media Raf., Med. Fl. 2: 107. 1830. Type: United 
States: *New York to Carolina" (Holotype NY!). 

Unisema media Raf. var. albiflora Raf., Med. Fl. 2: 107. 
1830. Type not seen. 

Unisema media Raf. var. angustifolia Raf., Med. Fl. 2: 107. 
1830. Type not seen. 

Unisema obliquata Raf., Med. Fl. 2: 107. 1830. Type: 
United States: “New Jersey and Virginia". Type not 
seen. 

Unisema latifolia Raf., Med. Fl. 2: 107. 1830. Type: United 
States: "Southern States". Type not seen. 

Unisema latifolia Raf. var. elatior Raf., Med. Fl. 2: 107. 
1830. Type not seen. 


1973] Pontederia — Lowden 453 


Unisema latifolia Raf. var. undulata Raf., Med. Fl. 2: 107. 
1830. Type not seen. 

Unisema, latifolia Raf. var. albiflora Raf., Med. Fl. 2: 107. 
1830. Type not seen. 

Unisema latifolia Raf. var. pallida Raf., Med. Fl. 2: 107. 
1830. Type not seen. 

Unisema acutifolia Raf., Med. Fl. 2: 107. 1830. Type: 
United States: Carolina. Type not seen. 

Unisema heterophylla Raf., Med. Fl. 2: 108. 1830. Type: 
United States: “New York to Louisiana", Type not seen. 

Unisema heterophylla Raf. var. lanceolata Raf., Med. Fl. 2: 
108. 1830. Type not seen. 

Unisema heterophylla Raf. var. stenocardia Raf., Med, FI. 
2: 108. 1830. Type not seen. 

Unisema rotundifolia Raf., Med. Fl. 2: 108. 1830. Type: 
United States: “Western States" [Rafinesque (1837); 
“Kentucky and Illinois"]. Type not seen. 

Pontederia cordata L. var. albiflora Short, Transylvania J. 
Med. Assoc. Sci. 32: 3, 7. 1835. Type: United States: 
Kentucky, marshes around Louisville, C. W. Short (Holo- 
type PH!; isotype NY!). 

Pontederia caerulea Maund, Bot. Gard. 6: 551. 1836. Type: 
Great Britain, plant under cultivation, introduced 1830. 
Type not seen. 

Unisema peduncularis Raf., New Fl. 2: 75. 1837. Based 
on the type of Pontederia anyustifolia Pursh. 

Unisema peduncularis Raf. var. parvifolia Raf., New FI. 
2: 75. 1837. Type not seen. 

Pontederia nymphaeifolia Kunth, Enum. Pl. 4: 126. 1848. 
Type: Brazil: Sello 235 (Holotype B?, photograph of 
holotype NY!; isotype PH!). 

Pontederia cordata L. var. typica, Solms in DC. Monog. 
Phan. 4: 532, in part. 1883. [that part based on the types 
of Pontederia cordata L., P. angustifolia Pursh and P. 
cordata L. forma brasiliensis Solms in Dc.] 

Pontederia cordata L. forma angustifolia (Pursh) Solms 
in DC., Monog. Phan. 4: 532, in part. 1883. [that part 
based on the type of Pontederia angustifolia Pursh] 


154 Rhodora [Vol. 75 


Pontederia cordata L. forma brasiliensis Solms in DC., 
Monog. Phan. 4: 533. 1883. Type not seen. 

Pontederia rotundifolia L. f. var. nymphaeifolia (Kunth) 
Solms in DC. Monog. Phan. 4: 534. 1883. 

Pontederia cordata L. [var.] lancifolia (Muhl.) Morong. 
Mem. Torrey Bot. Club 5: 105, in part. 1894. [that part 
based on types of Pontederia cordata L. and P. angusti- 
folia Pursh] 

Narukila cordata (L.) Nieuwland, Amer. Mid]. Naturalist 
3: 101. 1913. 

Narukila cordata (L.) Nieuwland var, lancifolia (Muhl.) 
Nieuwland, Amer. Midl. Naturalist 3: 101, in part. 1913. 
[that part based on the types of Pontederia cordata L. 
and P. angustifolia Pursh | 

Pontederia cordata L. forma angustifolia (Pursh) House, 
New York State Mus. Bull. 243-244: 62. 1923. 

Pontederia cordata L. forma latifolia (Raf.) House, New 
York State Mus. Bull. 243-244: 62. 1923. 

Pontederia cordata L. forma albiflora (Raf.) House, New 
York State Mus. Bull. 243-244: 62. 1923. Based on the 
type of Unisema media Raf. var. albiflora Raf. 

Unisema cordata (L.) Farwell, Pap. Michigan Acad. Sci. 
3: 91. 1924. 

Unisema cordata (L.) Farwell forma angustifolia (Pursh) 
Farwell, Pap. Michigan Acad. Sci. 3: 92. 1924. 

Unisema cordata (L.) Farwell forma latifolia Farwell, 
Pap. Michigan Acad. Sci. 3: 92. 1924. Type: United 
States: New Jersey: Secancus, 15 Sept. 1890, H. H. 
Rusby (Lectotype MICH !); Franklin, Aug. 1873, H. H. 
Rusby (Syntype MICH !). 

Pontederia lanceolata Nutt. forma trullifolia Fernald, 
Rhodora 27: 81. 1925. Type: United States: Florida, 
Okeechobee region, Brevard Co., 3 August 1903, A. Fred- 
holm 5927 (Holotype GH!). 

Pontederia lanceolata Nutt. forma brasiliensis (Solms) 
Fernald, Rhodora 27: 81. 1925. 

Unisema lancifolia (Muhl.) Farwell forma trullifolia (Fer- 
nald) Farwell, Amer. Midl. Naturalist 11: 73. 1928. 


1973] Pontederia — Lowden 455 


Pontederia cordata L. forma taenia Fassett, Rhodora 39: 
274. 1937. Type: United States: Maine, Lincoln Co. 
Damariscotta Lake, Jefferson, shallow mucky cove, 28 
August 1936, N. C. Fassett 16067 (Holotype WIS!; iso- 
types GH !, MO!). 

Pontederia cordata L. forma bernardi Lepage, Naturaliste 
Canad. 82: 101. 1955. Type: Canada: Quebec, Nomi- 
ningue, zone tourbeuse autour du lae Violon, 12 aout 
1951, Jean-Paul Bernard 386 (Holotype CAN!). 


Plants up to 11 dm tall (Fig. 9). Leaves cordate 
(Fig. 8), rarely auricled, or sagittate (unauricled), blades 
2.2-21 cm wide; spathe (terminal phyllode of floral shoot) 
2-7.4 cm long; petiole of floral shoot leaf (includes sheath 
base) 4-31 cm long; petiole of other leaves 28.7-59.5 cm 
long, ligule of petiole sheath up to 10.5 em long; sheath in 
axil of floral shoots up to 28 em long. Inflorescence (Fig. 8) 
2-16 cm long; peduncle (inflorescence base to floral shoot 
leaf base) 5-33 em long. Perigone blue, purple or white. 

In North America restricted to the eastern provinces of 
Canada (Fig. 10; Ontario, Quebec, New Brunswick, Prince 
Edward Island and Nova Scotia) and the eastern to mid- 
western United States (Fig. 11) with greatest concentra- 
tion along the Atlantic Coastal Plain and the Great Lakes 
Region as compared with the Mississippi embayment. 
In Middle America known only from British Honduras 
(Fig. 12). In South America (Fig. 15) found in Argen- 
tina, Brazil, Colombia, Paraguay and Uruguay. 


Representative Specimens: CANADA. Province New Brunswick. 
Norton, 20 Aug 1876, Hay (CAN). Province Nova Scotia. COLCHESTER 
co.: Earltown Lake, 29 Jul 1954, Smith, Webster & Bentley 11742 
(CAN). CAPE BRETON ISLAND: Louisburg, 19 Aug 1898, Macoun (CAN). 
Province Ontario. Vermilion River between Lake Superior and Lake 
Nipissing near Whitefish, 13 Aug 1936, Grass! 7494 (MICH). PELEE 
ISLAND: Essex Co., lagoon N end, 17 Aug 1967, Stuckey 5161 (os). 
GEORGIAN BAY AREA: Go Home, O.G.U. (CAN). Province Quebec. 
Bell’s Lake near Wakefield, 24 Jul 1903, Macoun (cAN). UNITED 
STATES. Alabama. BALDWIN co.: Battleship Parkway, 4 mi E of 
Mobile, Routes 31-90-38, Mobile Bay Causeway, 15 Jul 1970, Lowden 
5 (os). Arkansas. CRAIGHEAD CO.: Lake City, Demaree 7075 (US). 


456 Rhodora [Vol. 75 


Figure 10. Distribution of Pontederia cordata var. cordata in 
Ontario, Quebec, New Brunswick, Prince Edward Island and Nova 
Scotia Provinces of Canada. 


Connecticut. FAIRFIELD CO.: Green Pond, Sherman, 4 Sep 1916, Pen- 
nell 8592 (PH). Delaware. NEW CASTLE CO.: Wilmington, 6 Jul 1843, 
Brakely (PH). District of Columbia. Potomac bank, Washington, 
28 Jun 1891, Blanchard (Mo-2). Florida. ALACHUA CO.: 2 mi S of 
Gainesville, 17 May 1940, Martin 1457 (DUKE). Georgia. LONG CO.: 
Altamaha River Swamp, 4% mi SW of Ludowici, 10 Jun 1950, 
Duncan 11089 (GH). Illinois. cook co.: 131St between Wolf Rd and 
Will-Cook Rd W of Palos Park Pond, 16 Aug 1941, Steyermark 40917 
(F). Indiana. DEKALB CO.: Diamond Lake, 35 mi N of Auburn, 30 
Jul 1933, Shoop (F). Iowa. BENTON CO.: Vinton, 1879, Davis (WIS). 
Kentucky. BALLARD CO.: Wickliffe, Swan Pond, 18 Aug 1923, McFar- 
land & Anderson 167 (MO). Louisiana. BEAUREGARD PARISH: Near 
Longville, 29 Apr 1955, Cooley & Brass 4072 (GH). Maine. LINCOLN 
co.: 8 mi S of Newcastle Center off Route 1, 3 Sep 1969, Lowden 31 
(0S). SAGADAHOC CO.: Cathance River, 1 mi from Topsham, 1% mi 
from Bowdoinham, 3 Sep 1969, Lowden 29 (os); Foreside Road near 
Topsham off Route 24, 3 Sep 1969, Lowden 30 (os). YORK CO.: 
Bonnie Bay Pond, North Berwick, W of Route 4, 2 Jul 1970, Lowden 
1 (os); Kennebunk River, boundary between Kennebunk and Arun- 
del, Route 95, Jul 1970, Lowden 4 (os). Maryland. CECIL co.: Chesa- 
peake City, 7-11 1923, Tidestrom 11402 (GH). Massachusetts. BARN- 
STABLE CO.: Hyannisport, 22 Aug 1888, Churchill (Mo). Michigan. 
BARRY CO.: Long Lake, 14 Aug 1936, Woodbury 317 (MiCH). Minne- 


1973] Pontederia — Lowden 457 


Figure 11. Distribution of Pontederia cordata var. cordata (dots) 
and var. lancifolia (triangles) in the United States. 


458 Rhodora [Vol. 75 


mw 


— 2 N 


— 
MILES 


£c 
— 


Figure 12. Distribution of Pontederia cordata var. cordata (dots) 
in British Honduras (solid lines represent highways), C.A., and var. 
lancifolia (triangles) in Cuba, West Indies. 


I 
89*w 


1973] Pontederia — Lowden 459 


sota. AITKEN CO.: Rice Lake, Aug 1921, Kubichek 143 (Mo). Missis- 
sippi. HANCOCK CO.: Jordan River S of Kiln, 29 Apr 1967, Jones 
11846 (rsU). Missouri. BATES CO.: Prairie Lake, Cygnes River, 1!4 
mi SW of Rapinsville, 1 Oct 1938, Steyermark 9975 (F, MO). New 
Hampshire. ROCKINGHAM Co.: Exeter River, Philips Exeter Academy, 
Exeter, 8 Sep 1969, Lowden 32 (0S). STRAFFORD CO.: Bellamy River, 
Exit 7 Spaulding Turnpike, 4 Jul 1970, Lowden 2 (os). New Jersey. 
ATLANTIC CO.: Pleasant Mills, 4 Aug 1907, Bartram (PH). New York. 
ESSEX CO.: Lake Harris, Newcomb, 1 Aug 1921, House 8431 (CAN). 
North Carolina. BEAUFORT CO.: Cockold Creek, near Bellhaven, 25 
Jun 1935, Correll 1703 (DUKE). Ohio. DEFIANCE CO.: Little Pond off 
Route 49, 22 Sep 1969, Lowden 34 (0s); Big Pond off Route 49, 
22 Sep 1969, Lowden 35 (os). LICKING CO.: Cranberry Bog, Buckeye 
Lake, 12 Oct 1969, Lowden 33 (os). OTTAWA co.: Winous Pt., 3% 
mi SW of Port Clinton off Route 53, 10 Aug 1969, Lowden 36 (os); 
Winous Pt., 27 Jun 1969, Lowden 37 (os). Oklahoma. CHEROKEE CO.: 
Tahlequah, 21 Jun 1936, Gruchy 135 (GH, NY). Pennsylvania. BUCKS 
co.: Neshaminy Creek, Bridge Point [Edison], 22 Jul 1876, Meredith 
(PH). Rhode Island. NEWPORT Co.: Tiverton, Sawdy Pond, 16 Jul 
1932, Sanford (NEBC). South Carolina. ANDERSON CO.: Piedmont, 17 
Jul 1919, Davis (BKL, MO). Tennessee. GRUNDY CO.: Goose Pond, near 
Pelham, 27 Apr 1936, Svenson 7610 (BKL). Texas. BEXAR CO.: San 
Antonio, 3 Oct 1900, Bush 1258 (Mo). Vermont. ESSEX CO.: Bow 
Pond, Guildhall, 30 Aug 1940, Pease 28364 (NEBC). Virginia. FAIRFAX 
co.: Near Great Falls, 7 Aug 1910, Dowell 6435 (GH). Wisconsin. 
ASHLAND CO.: Torrey L., Morse, 16 Jul 1936, Knowlton 97 (wis). 
CENTRAL AMERICA. British Honduras. BELIZE DISTRICT: Northern 
River, Nov 1933, Gentle 944 (GH, MICH, MO, NY); 9 mi N of Belize 
City, Northern Highway, 24 Dec 1969, Lowden 24 (0S); 29 mi N of 
Belize City, Cowhead Creek, Northern Highway, 30 Dec 1969, Lowden 
26 (os); 32 mi N of Belize City, Northern Highway, 30 Dec 1969, 
Lowden 27 (os); Hattieville-Burrell Boom Road, 5% mi from Bur- 
rell Boom, 30 Dec 1969, Lowden 28 (os). COROZAL DISTRICT: 1931- 
1932, Gentle 388 (F, MICH, US). EL CAYO DISTRICT: 41 mi Sect., Belize- 
Cayo Road, 1 Apr 1958, Gentle 9708 (DUKE, F, MICH, NY, US). 
ORANGE WALK DISTRICT: Hillbank Lagoon, Feb 1933, Pelly 43, 44 (F). 
STANN CREEK DISTRICT: Cockscomb vic. 18 Jun [1930], Schipp 8119 
(F); 23 mi, Stann Creek Valley, 11 Mar 1932, Schipp 955 (F, GH, 
MICH, MO, NY). TOLEDO DISTRICT: 15 mi, San Antonio-Punta Gorda 
Road, 28 Nov 1951, Gentle 7531 (F, NY, US). SOUTH AMERICA. 
Argentina. PROVINCIA BUENAS AIRES: Depto. Campana, Campana, 
27 Nov 1938, Eyerdam & Beetle 23070 (GH). PROVINCIA CHACO: Dept. 
Resistencia, Colonia Benitez, 12 Dec 1928, Venturi 7907 (GH, US). 
PROVINCIA CORRIENTES: Mercedes, 12 leguas al N, XI 1936, Rodrigo 
750 (NY). PROVINCIA MISIONES: Dept. Apostoles, Tres Galpones, 8 
II 1947, Huidobro 4817 (MO). TERRITORY FORMOSA: Formosa, 5-1918, 


460 Rhodora [Vol. 75 


Jorgensen 3005 (Mo). Brazil PARANA: Rio Pequeno, auto estude 
Curitiba-Paranagua, XI 1960, Brage & Vesreira 320 (US). PROV. 
MINAS GERAIS: Brasilia (PH). RIO DE JANEIRO: Brasilia, pr. Lorena, 
L. Riedel, iter Brasiliensis 1821-36 (NY). RIO GRANDE DO SUL: Logoa 
dos Barros, pr. Osorio, 24 Nov 1949, Rambo 44574 (F). SANTA 
CATARINA: Mun. Chapeco, Fazenda Campo Sao Vicente, 24 km W of 
Campo Ere, Smith, Reitz & Sufridini 9468, 9482 (US). SAO PAULO: 
Butantan, 17 Oct 1917, Hoehne 726 (NY). Colombia. COMMISSARIA 
MAGALENA: Near Valencia, 12 Oct 1944, Haught 4406 (F, US). COM- 
MISSARIA VALLE [DEL CAUCA]: Costa del Pacifico, rio Cajambre, Que- 
brada de Guapecito, 16 May 1944, Cuatrecasas 17703 (F). Paraguay. 
Paraguaria Centralis, lacus Ypacaray, Nov 1913, Hassler 12683 (GH, 
MO, NY, US); Dep. Paraguari, Paraguari, Cerro Hu, 25 Nov 1950, 
Vervoorst 564 (Mo). Uruguay. DEPARTMENT CANELONES: Loc. Dict. 
Toledo, Nov 1926, Herter 522 (F, GH, MO, NY, WIS). DEPARTMENT 
MALDONADO: Lau [San] Carlos, M 1 1941 Descole 104 (GH). 


Since none of the original material was located the speci- 
men of Kalm in the Linnaean Herbarium by 1751 was 
selected as neotype. The numerous names of Rafinesque 
are placed in synonomy based on their leaf descriptions 
and geographical locations. A complete list of specimens 
examined is included in my original dissertation. 

The specimens from British Honduras are of particular 
interest in considering the puzzling disjunct distributions 
of both Pontederia cordata varieties cordata and lancifolia 
in North and South America. Actually these specimens 
are somewhat intermediate between typical P. cordata and 
P. sagittata. They have tooth ridged hardened perigone 
bases like P. cordata (Fig. 1A) with the overall perigone 
shape of P. sagittata (Fig. 1B). The leaves have deep 
sinuses giving the appearance of being slightly auricled. 
A hybrid origin is suggestive; however, an overlap in 
geographical ranges between P. cordata var. cordata and 
P. sagittata is not evident. The specimens in general are 
more characteristic of P. cordata var. cordata. 


lb. Pontederia cordata L. var. lancifolia (Muhl.) Torrey, 

Fl. U.S. 1: 343. 1824. 

Pontederia lancifolia Muhl., Cat. 34. 1813. Type: United 
States: Car. [Carolina] (Lectotype chosen, PH !), Figure 
13. 


1973] Pontederia — Lowden 461 


— Carers! Sareen ai PERE . 
l obo eases: jiTEO ATER MATIQNAL MUSEUM 


Figure 13. Pontederia cordata var. lancifolia. Lectotype of P. 
lancifolia Muhl. in the Muhlenberg Herbarium (PH), showing a 


narrow lanceolate flowering stalk leaf; at left. 
Figure 14. Pontederia cordata var. ovalis. Holotype of P. ovalis 


Mart. in Roemer & Schultes, showing ovate leaves and hairy pe- 
duncles; at right. 


462 Rhodora [Vol. 75 


Pontederia lanceolata Nuttall, Gen. 1: 216. 1818. Type: 
United States: Savannah, Georgia [W. Baldwyn] (Holo- 
type PH!). 

Unisema lancifolia Raf., Alt. Jour. 178. 1833. Based on the 
type of Pontederia lancifolia Muhl. 

Pontederia cordata L. var. lanceolata (Nutt.) Grisebach, 
Cat. Pl. Cub. 252. 1866. 

Pontederia cordata L. var. typica, Solms in DC. Monog. 
Phan. 4: 532, in part. 1883. [that part based on the type 
of Pontederia lanceolata Nutt.] 

Pontederia cordata L. forma angustifolia (Pursh) Solms 
in DC. Monog. Phan. 4: 533, in part. 1883. [that part 
based on the type of Pontederia lanceolata Nutt.] 

Pontederia cordata L. lancifolia (Muhl.) Morong. Mem. 
Torrey Bot. Club 5: 105, in part. 1894. [that part based 
on the type of Pontederia lancifolia Muhl.] 

Narukila cordata (L.) Nieuwland var. lancifolia (Muhl.) 
Nieuwland, Amer. Midl. Naturalist 3: 101, in part, 1913. 
[that part based on the type of Pontederia lancifolia 
Muhl.] 

Pontederia heterantherimorpha (K. Schum. ex Schwartz) 
Sehwartz in Engler, Bot. Jahrb. 61 (no. 139) : 41. 1927. 
Type: Brazil: “Im Staate Goyaz, A. Glaziow 22228. 
Type not seen. 

Unisema lancifolia (Muhl.) Farwell, Am, Midl. Nat. 11: 
73. 1928. 

?Pontederia oblonga Larranaga, Pub. Inst. Hist. Geog. 
Uruguay 2: 134. 1930. Type not seen. 


Plants up to 13 dm tall. Leaves narrow lanceolate (Fig. 
13) to broadly ovate-lanceolate, blades 0.4-8.3 cm wide; 
spathe (terminal phyllode of floral shoot) 2-6.5 em long.; 
petiole of floral shoot leaf (includes sheath base) 4-21 cm 
long; petiole of other leaves 29-59.5 cm long, ligule of 
petiole sheath up to 10.5 em long; sheath in axil of floral 
shoot up to 28 cm long. Inflorescence 2-15 cm long; 
peduncle (inflorescence base to floral shoot leaf base) 5-33 
cm long. Perigone blue to blue purple. 

In the United States (Fig. 11) concentrated along the 


1973] Pontederia — Lowden 463 


Gulf Coastal Plain of the southeast extending along the 
coasts of the northeastern Atlantic seaboard states. Along 
the southwestern coast of Cuba (Fig. 12), West Indies. 
In South America (Fig. 15) known from Argentina, 
Brazil, Colombia, Paraguay and Uruguay, 

Representative Specimens. UNITED STATES. Alabama. GENEVA 
co.: 8 mi S of Samson, 9 May 1967, McDaniel 8916 (FSU). Connec- 
ticut. NEW LONDON co.: E Lyme, Dodge’s Pond Niautic, 19 Aug 1913, 
Harger 6328 (NEBC). WINDHAM CO.: Thompson, Long Pond, 19 Aug 
1918, Weatherby 4373 (NEBC). District of Columbia. Potomac River, 
Jun 1896, Morris 3046 (BKL). Florida. BAY CO.: Panama City, Mine 
Defense Lab vicinity, 1 May 1959, Jones (FSU). BREVARD CO.: Indian 
River near Rockledge, 10-20 Mar 1916, Bartram (PH). BROWARD CO.: 
11 mi W of Davie, 31 Jan 1940, Seibert 1191 (MO, PH). COLLIER CO.: 
Pinecrest, S of Tamiami Trail between mi 40 and Ochopee, 15 Apr 
1952, Field & Lazar (US). COLUMBIA C0.: 6 mi N of Lake City, 20 
May 1964, Godfrey 63755 (FSU). DADE Co.: Everglades, W. of Miami, 
1-9 Nov 1901, Small & Nash (NY). DE SOTA CO.: Arcadia, 14 Mar 
1926, Williams (PH). DUVAL C0.: NE Forida, May, Curtis 2988 (BKL, 
F, GH, MO, NY, US). FRANKLIN CO.: 11 mi S of Sumatra, 24 Jul 1957, 
Godfrey 55715 (FSU, GH). GADSDEN CO.: 1 mi W of Ock[h]lockonee 
River, U.S. 20, 6 Jun 1956, Redfearn 2194 (FSU, GH). HAMILTON CO.: 
2 mi E of Jasper, 1 May 1959, Godfrey 58508 (FSU). HIGHLANDS CO.: 
Bear Point, Lake Childs, 10 Mar 1945, Brass 14779 (GH). HOLMES 
co.: U.S. 90, 0.2 mi E of Florida 81 in Ponce de Leon, 1 May 1968, 
Stone 2589-2591 (DUKE, NY). INDIAN RIVER C0.: St. Johns River, 10 
mi W of Vero Beach, 22 Feb 1957; Lemaire 149 (FSU). JACKSON CO.: 
2 mi E of Grand Ridge, 20 May 1960, Godfrey 59539 (FSU). LAKE 
co.: 13 mi S of Leesburg, 10 Jun 1961, Godfrey & Reinert 61036 
(FSU). LAKE or ORANGE CO.: Lake Apopka, 25 Apr 1930, O'Neill- 
Blanton 6442 (GH). LEE CO.: Fla. 80, 3 mi E of Tice, 15 Aug 1963, 
Henderson 63-1590 (FSU). LOWNDES CO.: b mi E of Valdosta, 5 May 
1963, Godfrey & Houk 62760 (FSU). MARTIN CO.: Lake Okeechobee, 
Pelican Lake to Cypress Creek, 11-25 Nov 1913, J. & G. Small 4320 
(NY). NASSAU CO.: 10 mi E of Hilliard, 23 May 1964, Godfrey 64093 
(FSU). OKEECHOBEE CO.: Kissimmee River, N end of Lake Okeecho- 
bee, 25 Nov 1913, J. & G. Small 4383 (NY). PALM BEACH CO.: Loxa- 
hatchee Wild Life Refuge, W of Delray Beach, 29-30 Mar 1952, 
Field & Lazar (F, US). POLK CO.: Mountain Lake, Lake Wales, 21 
Mar 1952, Field & Lazar (US). PUTNAM CO.: Palatka and Lake 
Ganoga, 18 Apr 1897 & Aug 1903, Williamson (PH). Georgia. 
BRANTLEY CO.: Ga. Rts. 15 & 121, 1 mi S of Hoboken, 18 Aug 1967, 
Clewell 2745 (FSU). BROOKS CO.: 1 mi NW of Morven, 6 Jun 1959, 
Adams 182 (FSU). CARLTON C0.: Near Camp Cornelia in Okefenokee 
Swamp, Jarrard 2226 (DUKE). CHATHAM CO.: Savannah, (Pontederia 


464 Rhodora [Vol. 75 


15*$ 


Figure 15. Distribution of Pontederia cordata var. cordata 
(dots), var. lancifolia (triangles) and var. ovalis (stars) in South 
America. 


1973] Pontederia — Lowden 465 


lanceolata Nutt.), (PH). CLINCH CO.: Dupont, 4 10 1935, Louett 
(DUKE). DEKALB CO.: Lakes, 24 May 1897, Eggert (MO). DOOLY CO.: 
Vienna, Apr 1845, Rugel 24 (MICH). ECHOLS CO.: Ga. Rt. 94, 5 mi 
NW of Statenville, 17 Aug 1967, Clewell 2528 (FSU). GLYNN CO.: 
7 mi NW of Brunswich, hwy. 341, 16 Apr 1961, Wright 41 (GH). 
LOWNDES CO.: Valdosta, 4 10 1938, Baker 2978 (DUKE). RICHMOND 
co.: Tubman Home, Augusta, 13 Jun 1924, Hildebrand (DUKE). WARE 
co.: Between Waycross and Ruskin, 2 Aug 1902, Harper 1469 (F, 
GH, MO, NY, US). WAYNE CO.: Jesup, 4 Jun 1893, Kearney (os). 
Indiana. LAKE CO.: Miller, 24 Jun 1896, Umbach (PH). Maine. ox- 
FORD CO.: Roxbury Pond, Byron, 20 Jul 1934, Reed 440 (DUKE, PH). 
Massachusetts. MIDDLESEX C0.: Fresh Pond, Cambridge, Ball [in 
part] (MO). WORCESTER CO.: East Templeton, 28 Jul 1886, Partridge 
(BKL). Michigan. SCHOOLCRAFT CO.: Indian Lake, 25 Jun 1937, Beck- 
man 82 (MICH). New Hampshire. CARROLL CO.: Chocorua Lake, 
Tamworth, 3 Aug 1947, Steele (NEBC). New Jersey. MORRIS CO.: 
Green Pond, 1 Aug 1894, Van Sickle (BKL). Rhode Island. WASH- 
INGTON CO.: Exeter, Ieppican Pond, 22 Sep 1920, Graves & Woodward 
(GH, NEBC). South Carolina. COLLECTON CO.: 8 mi SE of Walterboro, 
18 Jul 1927, Wiegand & Manning (GH). SUMTER CO.: Sumter vic., 
25 May 1914, Stone 438 (PH). Tennessee. COCKE CO.: Newport to 
Greenville, 5 mi E of Newport, 17 Jun 1939, Sharp & Jennison 342 
(BKL,MO). COFFEE CO.: S of Manchester, 21 Aug 1938, Svenson 8775 
[in part] (BKL, GH, PH). Texas. CHAMBERS CO.: Anahuac, 16 May 
1937, Cory 22410 [in part] (GH). HARRIS CO.: Cob Pond, Humble, 
9 10 1926, Tharp 4314 (vus). WEST INDIES. Cuba. PROV. PINAR 
DEL RIO, Coloma vic., 28 Feb-2 Mar 1911, Britton & Cowell 9693 (GH, 
NY, US) ; Cienaga de Zapata, N de la Bahia de Cochinos (Sta Clara), 
14 Aug 1920, Leon & Loustalot 9530 (NY); PROV. PINAR DEL RIO, near 
La Coloma, 1 May 1940, Leon, Victorin & Alain (GH); Plantae 
Cubenses Wrightianae, 1860-1864, Wright 3260 (GH, MO, NY, US). 
SOUTH AMERICA. Argentina. PROV. BUENOS AIRES, Buenos Aires, 
Jan 1852, Andersson (US); PROV. CORRIENTES, Dept. Ituzaingo, Salto 
Apipe, 6 X 1949, Schwarz 8130 (Mo, us). Brazil. PARANA, Desiro 
Ribas, 29 Nov 1910, Dusen 10849 (GH, NY, US); PROV. MINAS GERAIS, 
Brasilia, cidade de Caldas, 1868, Henschen (US); Rio Grande Do 
Sul, Belem Novo, Costa Rio Guahyba, 31 Jan 1948, Palacios & Cuezzo 
416 (mo); Rio De Janeiro estado, municipio Cabo Frio, Cabo Frio, 
Praia do Pontal, 17 Apr 1952, Smith 6595 (Uus). Colombia. COM- 
MISSARIA MAGDALENA, Poponte, Magdalena Valley, 2 Nov 1924, Allen 
786 (F, MO); COMMISSARIA PUTUMAYO, Umbria, Dec 1930, Klug 1877 
(F, GH, MICH, MO, NY, US). Paraguay. Iter ad Paraguariam septen- 
trionalem, superioris fluminis Apa, 1901-1902, Hassler 7849 (GH, 
MICH, MO, NY). Uruguay. Nueva Palmina [?] (Dto. Colonia), XII 
1943-I 1944. Scolnik 32 (NY). 


466 Rhodora [Vol. 75 


This lanceolate to narrowly linear-lanceolate leaved 
variety (Fig. 13) has a geographically distinct distribution 
in the southeastern portion of the United States (Fig. 11). 
The leaf width is quite variable. The extremely ovate- 
lanceolate forms, suggestive of hybridization, are predomi- 
nant throughout the overlapping ranges of this variety 
and P. cordata var. cordata. 


le. Pontederia cordata L. var. ovalis (Mart. in Roemer & 

Schultes) Solms in DC., Monog. Phan. 4: 533. 1883. 

Pontederia ovalis Mart. in Roemer & Schultes, Syst. Veg. 
7: 1140. 1830. [Martius, Fl. Bras. 3(1): 95. 1847.] 
Type: Brazil (Holotype M; photograph of holotype US!). 

Pontederia lanceolata Nutt. var. vichadens?s Hermann, 
Caldasia 5: 39. 1948. Type: Colombia: Vichada, mucky 
edge of stream in open forest on llanos bordering the 
Rio Vichada, alt. 100 m, ca. 18 km NE [E.N.E.] of San 
José de Ocuné, 21 Jan. 1944, F. J. Hermann 11045 (Holo- 
type Us!). 

Pontederia lanceolata Nutt. forma ovalis (Mart. in Roemer 
& Schultes) Castell, Rio de Janeiro Jardin Botanico 15: 
62. 1958. 


Plants up to 9 dm tall. Leaves ovate (Fig. 14), blades 
2.2-21 em wide; spathe (terminal phyllode of floral shoot 
2.2-7.4 em long; petiole of floral shoot leaf (Fig. 14, in- 
cludes sheath base) 5.3-31 cm long; petiole of other leaves 
28.7-30.6 cm long, ligule of petiole sheath up to 4 cm long; 
sheath in axil of floral shoot up to 22.3 cm long. Inflores- 
cence 2.7-16 em long; peduncle (inflorescence base to floral 
shoot leaf base) 12.3-28 cm long, densely pubescent next to 
inflorescence base. Perigone white or pale blue. 

In South America (Fig. 15) known only from Bolivia, 
Brazil, Colombia, Paraguay and Uruguay. 


Representative Specimens. SOUTH AMERICA. Bolivia. DEPART- 
MENT BENI: Reyes, 27 Oct 1921, Rusby 1389 (BKL, GH, MICH, NY, PH, 
US). Brazil. Prov. Minas Gerais, Dattos de Cima, Mun. Diamantina, 
19 Nov 1937, Barreto 9817 (F); Rio Grande Do Sul, Neu. Wurhem- 
burg, 10 11 [19]04, Bornmuller 356 (GH); Burchell A157 (GH); 
Burchell A163 [in part] (GH); 12 km S of Cristaes, Ceara, 28 Aug 


1973] Pontederia — Lowden 469 


Figure 17. Pontederia sagittata with sagittate auricled flowering 
stalk leaves, Mexico, State of Vera Cruz, R. M. King 933 (Us); at left. 

Figure 18. White inflorescences of Pontederia parviflora, with 
subcordate leaves, Panama, Province of Herrera (Lowden photo, 
Aug 1970); at right. 


Miramar, 19 Mar 1955, Sohns 1658 (MICH, NY, US). STATE OAXACA: 
Jamiltepec, Distrito Jamiltepec, Oaxaca, 10 Dec 1921, Conzatti 4434 
(NY); Chiltepee and vic., District Tuxtepec, Jul 1940-Feb 1941, 
Martinez-Calderon 76494 (GH, US); Near San Gabriel Mixtepec, 
Mpio. de Juquila, 2 km N San Gabriel, 13 Feb 1965, McVaugh 22416 
(MICH) ;Laguna Tonameca, Oaxaca, 8 Nov 1917, Reko 3451 (US); 
Foothills Sierra Madre del Sur, 53 km N of Puerto Escondido, road 
to Zimatlan km 221, 25 Jul 1965, K. & E. Roe & Mori 559 (F, Wis). 
STATE TABASCO: Popal Grande between Frontera and Villahermosa, 
Aug 1962, Barlow (MICH, Wis); W-SW of Huimanguillo, 28 May 
1963, Barlow 30/52 (wis); La Palma, Balancan, 1-6 Jun 1939, 
Matuda 3302 (GH, MICH); Curahueso, Tabasco, 6 Jan 1889, Revirosa 
334 (NY, PH). STATE VERA CRUZ: Wartenberg, near Tantoyuca, prov. 
Huasteca 1858, Ervendberg 277 (GH, PH); Vera Cruz, 12 Aug 1926, 
Fisher 108 (US); prov. Vera Cruz, Galeotti 5562 (F, GH, NY, US); 
Mun. Puente Nacional, km 394 Jalapa-Vera Cruz highway, between 
Rinconada and Puente Nacional, 18 Feb 1943, Gilly, Simpson & Dodds 
90 (MiCH); Near city Vera Cruz, 23 Jan 1906, Greenman 19 (F, 


470 Rhodora [Vol. 75 


GH); Vera Cruz, in and around, 8 Dee 1958, Jones 22660 (WIs); 
Trans-Isthmian highway Route 185, 17 km NE of Minatitlan, 1 Aug 
1958, King 933 (MICH, US); *Vera Cruz, El Coyol, Prolongacion 
Carretera Aleman, 25 Jul 1970, Lowden 6 (os); Carretera Nacional 
180, El Puente Teculapilla, 5 km NW of Lerdo Route 180, 26 Jul 
1970, Lowden 7 (os); Laguna Catemaco, S edge off Route 180, 27 
Jul 1970, Lowden 8a-b (0S); Laguna Catemaco, Arroyo Agrio, 6 km 
from Catemaco Town between Coyame and San Andres Tuxtla, 27 
Jul 1970, Lowden 9 (os); Vera Cruz, A? 1853, Müller 2145 (NY); 
Jalapa, 16 Feb 1910, Orcutt 2874 (F, GH, MO, US); Barranca de 
Panoaya, Mar 1923, Purpus 9041 (F, GH, MO, NY, US) ; Rio Maquina, 
municipio San Andres Tuxtla, 26 Mar 1964, Quintero 699 (MICH); 
Rio Los Tuxtla, municipio Santiago Tuxtla, 29 Aug 1964, Quintero 
1409 (MICH); Sontecomapan, municipio Catemaco, 5 Aug 1965, 
Rzedowshi 20368 (MicH); Rio Cosolapa, Le Gartera, Ejido San 
Agustin, 5 km NE of Campo Experimental de Hule, El Palmar, 
Zongolica, 27 Feb 1944, Santos 2821 (MICH); Vera Cruz, inter urbem 
et Santa Fe et prope los Cocos, Feb, Schiede & Deppe 981 (Mo, US) ; 
Coatzacoalcos, isthmus Tehuantepec, 8 Jan 1895, Smith 1034 (F, GH, 
MICH, MO, NY, US, WIS); Tuxpam [Tuxpan], Rio Vinasco [Rio 
Vinazco], 9 I 1903, Ex Museo botanico Berolinensi 3731 (GH, US); 
Distr. San Andres Tuxtla, Laguna Catemaco, 7 III 1907, Ex Museo 
botanico Berolinensi 5005 (GH). sTATE ?: Winter 1877, Hogg (NY); 
San Luis, 27 Feb 1899, Langlasse 928 (GH, US); Inter el Morro & 
Rancho Nuevo, 1841-43, Liebmann 1618 (US). CENTRAL AMERICA. 
Costa Rica. PROVINCE LIMON: Matina, IX 1896, Pittier 10301 (Us). 
Guatemala. DEPARTMENT ALTA VERAPAZ: Coban, M Dec 1886, Türek- 
heim 547 (GH, NY, PH, US); Cubilquitz [Gubilquitz], M Apr 1901, 
Tüerckheim 8025 (US). DEPARTMENT IZABAL: Izabal, Rio Mosinga, 
2 Jun 1919, Blake 7864 (GH, US); Jocolo, Izabal, Lago Izabal, 25 Dec 
1920, Johnson 1053 (vs); Lago Izabal, Izabal vic., 1 May 1966, Jones 
& Facey 3213 (F, NY); Livingston, 18 Jan 1905, Kellerman 5131 (0s, 
US); Quirigua, 31 Jul 1970, Lowden 10 (os). Puerto Barrios, 1 Aug 
1970, Lowden 11 (os); Puerto Barrios vic., 29 Dee 1904, Maxon & 
Hay 3061 (us); Rio Dulce, 15 May 1937, Muenscher 12623 (F); 
Livingston, 27 May 1905, Pittier 857 (Ny, US); Rio Dulce, M Mart 
1889, Smith 1652 (GH, US); Quirigua vic., 15-31 May 1922, Standley 
24029 (GH, NY, US); Puerto Barrios vic., 2-6 Jun 1922, Standley 
25005 (us); near Quirigua, 26-27 Apr 1939, Standley 72425 (F); 
Near Puerto Barrios, 25 Apr-6 May 1939, Standley 73170 (F); Be- 
tween Bananera and La Presa in Montana del Mico, 28 Mar 1940, 
Steyermark 38050 (F, vs); Lago Izabal, opp. San Felipe, between 
San Felipe and mouth of Rio Juan Vicente, 19 Apr 1940, Steyermark 
39688 (F). DEPARTMENT SAN MARCOS: Rio Suchiate, W of Ayutla, 
18 Mar 1940, Steyermark 38034 (F). DEPARTMENT PETEN: E] Paso, 
San Pedro River, 26 Apr 1932, Lundell 1564 (MICH, NY). DEPART- 


1973] Pontederia — Lowden 471 


MENT ZACAPA: Gualan, 15 Jan 1905, Deam 208 (F, GH, MICH, MO, 
NY, US). DEPARTMENT ?: Feb 1912, Cockerell 10 (vs); Eastern 
portions of Vera Paz and Chiquimula, 1885, Watson 28, 33, 386 (GH). 
Honduras. DEPARTMENT ATLANTIDA: Ceiba, 26 Sep 1916, Dyer A92 
(us); Tela, La Curva, 10 Aug 1970, Lowden 15 (os); Santiago, 11 
mi W of Tela, 10 Aug 1970, Lowden 16 (os); La Ceiba, Platanillo, 
6 Nov 1948, Molina & Becker 1 (F); Tela, 1923, Severen 5 (US); 
Tela vic., 14 Dee 1927-15 Mar 1928, Standley 53607 (F, US); Tela 
River, (Tela) Puerto Sierra, 18 Jan 1903, Wilson 70 (F, NY, US); 
Near Micos Lagoon, 12 km W of Tela, 21 Jul 1934, Ywncker 4684 
(F, MICH); Ceiba vic., 6 Jul 1938, Yuncker, Koepper & Wagner 8249 
(F, GH, MICH, MO, NY, US). DEPARTMENT CORTES: Puerto Cortes, 3 
Ave-5 Calle 0, 9 Aug 1970, Lowden 13 (os); 13.5 mi inland from 
Puerto Cortes along main road, 9 Aug 1970, Lowden 14 (os); San 
Pedro Sula, Depart. Santa Barbara, M Mart 1889, Thieme 5510 
(GH, US). 

Specimens for which identification is not positive. Guatemala. 
DEPARTMENT ALTA VERAPAZ: Route 5, between Semococh and La 
Laguna, road to Chajmayic, 10 May 1942, Steyermark 46362 (F). 
DEPARTMENT PETEN: Isabilito, 28 Mar 1932, Lundell 1445 (MICH, US); 
El Paso, San Pedro River, 26 Apr 1932, Lundell 1564 (Us); Santa 
Teresa, Subin River, 10 Apr 1933, Lundell 2709 (F, MICH); Laguna 
Peten Itza, frente Playa Blanca, Santa Elena, 3 Jan 1970, Ortiz 
512 (F). 


The smooth ridged hardened perigone bases (Fig. 1B) 
easily distinguish this species. The collection in Costa Rica 
(Fig. 16) reflects a more continuous distribution along the 
Caribbean Coastal Plain. Specimens without positive 
identification are immature vegetative specimens lacking 
mature perigones. 


3. Pontederia parviflora Alex., N. Am. Flora 19: 59. 1937. 
Type: Panama: Province of Panama, camino del Boti- 
cario, near Chepo, altitude 30 to 50 meters, October 1911, 
H. Pittier 4556 (Holotype NY!; isotype US!). Fruiting 
specimen cited with holotype, Panama, Province of Cocle, 
Aguadulce, in savannas, near sea level, 3-6 December 1911, 
H. Pittier 4915 (GH!, NY!, US!). 
Pontederia cordata L. var. parviflora (Alex.) Schery, Ann. 
Missouri Bot. Gard. 31: 156. 1944. 
Plants (Fig. 18) up to 7 dm tall, Leaves (Fig. 18) sub- 
cordate, blades 1.8-10.5 em wide; spathe (terminal phyllode 


472, Rhodora [Vol. 75 


MILES d 


Q 7 
v ar - 


Figure 19. Distribution of Pontederia parviflora in Panama and 
Colombia (stars represent sighted records). 


of floral shoot) 5-10.5 em long; petiole of floral shoot leaf 
(ineludes sheath base) 5.5-19 cm long; petioles of other 
leaves 19-45 cm long; sheath in axil of floral shoot up to 
34 cm long. Inflorescence 3-9.5 cm long, peduncle (in- 
florescence base to floral shoot leaf base) 21-35 em long. 
Perigone white to white green; hardened perigone bases 
(Fig. 1C) toothed ridged; anthers brown to black. Homo- 
stylous (Fig. 3D). Fruits and seeds (Fig. 2A-B) reniform. 

Known only from the Pacific coast of Panama, Golfo de 
Panama and the Caribbean Coast of Colombia. Figure 19. 

Specimens Examined. CENTRAL AMERICA. Panama. PROVINCE 
COCLE: Aguadulce, 3-6 Dec 1911, Pittier 4915 (GH, Ny, US); Between 
Aguadulce and Anton, 12 Jul 1938, Woodson, Allen & Seibert 1208 
(GH, MICH, MO, NY). PROVINCE HERRERA: Santa Maria, 13 Sep 1938, 
Allen 790 (MICH); Near Divisa, 10 Aug 1962, Dwyer 2468 (us); 
Santa Maria and El Escota, 24 Aug 1970, Lowden 20 (os). PROVINCE 
PANAMA: Between Panama and Chepo, 29 Nov 1934, Dodge, Hunter, 
Steyermark & Allen 16702 (MicH, MO); Between Pacora and Chepo, 


1973] Pontederia — Lowden 473 


7 mi from Chepo, Tapagara, 26 Aug 1970, Lowden 21 (os); Camino 
Boticario, near Chepo, Oct 1911, Pittier 4556 (NY, US); Swamp E of 
Rio Tecumen [Rio Tocumen], 11 Dec 1923, Standley 26495, 26656 
(US); Between Pacora and Chepo, 1 Aug 1938, Woodson, Allen & 
Seibert 1661 (GH, MICH, NY). SOUTH AMERICA. Colombia. DEPART- 
MENT MAGDALENA: Rio de Hacha, near Molino, Sa. Martha, Pur[chi], 
1845, Hooker; Costa Del Caribe, oeste de Los Venados, 31 Sep 1961, 
Dugand 5834 (us). 

This is the only homostylous member of the genus. Its 
unique style form (Fig. 3D), white inflorescence (Fig. 18) 
and brown-black anthers distinguish Pontederia parviflora 
from other members of subgenus Pontederia. A wider 
distribution is expected in South America. 

II. Pontederia subg. Reussia (Endl.) Lowden, comb. nov. 
Reussia Endl. (nom. cons.), Gen. Pl. 139. 1836. Type 
Reussia triflora Seub. in Mart. Fl. Bras. 3 (1): 96. 1847. 


Hardened perigone bases (Fig. 1D) spinulose ridged; 
floral bearing shoots prostrate; aerial stems with long 
internodes; underground rhizomes short (mature plants). 


Subgenus Reussia reaches its northern limits in British 
Honduras and extends into South America as far south as 
east central Argentina. 


4. Pontederia rotundifolia L. f., Suppl. 192. 1781. Type: 
Surinam, C. G. Da[h]lberg (Lectotype chosen, [Savage, 
407.2], LINN). 

Pontederia cordifolia Mart. in Roemer & Schultes, Syst. 
Veg. 7: 1142. 1830. Type: Brazil: “Crescit in Brasiliae 
mediterraneae stagnis" (Holotype M; photographs of 
holotype NY!, Us!). 

Pontederia brasiliensis Willd. Roemer & Schultes, Syst. 
Veg. 7: 1145. 1830. Type: Brazil: ‘In Brasilia prope 
Para. Com, de Hoffmannsegg". Type not seen. 

Unisema orbiculata Raf., Med. Fl. 2: 108. 1830. Type based 
on the type of P. rotundifolia L. f. 

Pontederia eriantha Miquel, Linnaea 17: 60. 1843. Type: 
Surinam: “Crescit Surinami, ad ripas fluminis Comme-. 
wyne, Focke" (Holotype GH!). 

?Pontederia renniformis Larranaga, Pub, Inst. Hist. Geog. 
Uruguay 2: 134. 1930. Type not indicated. 


474 Rhodora [Vol. 75 


Figure 20. Inflorescence of Pontederia rotundifolia, showing a 
bilobed spot on the largest perianth lobe, El Salvador, Laguna de 
Zapotitan (Lowden photo, Mar 1970); at left. ' 

Figure 21. Reniform leaves and short inflorescence of Pontederia 
rotundifolia, Brazil, Territory of Rondonia, Prance, Forero, Wrigley, 
Ramos & Farias 5914 (NY); at right. 


Reussia grazielae Machado, Revista Brasil. Biol. 7: 177. 
1947. Type: “Amazonas Bahia, in acquis, 1937, Ducke 
55083 (Holotype RB; photograph of holotype, Machado 
1947, Fig. 3). 

Reussia rotundifolia (L. f.) Castell., Lilloa 25: 593, 1951. 
Plants up to 12 dm tall. Leaves reniform (Fig. 21), 

sagittate (sometimes deeply auricled) or cordate; blades 

3.2-22 cm wide; spathe (terminal phyllode of floral shoot) 

2.5-4.5 em long; petiole of floral shoot leaf (includes sheath 

base) 4.7-45 cm long; petiole of other leaves 17-54 cm long, 
ligule of petiole sheath up to 7.5 em long; sheath in axil of 

floral shoots up to 24 cm long. Inflorescence (Fig. 20) 

2-7.5 em long, peduncle (inflorescence base to floral shoot 

leaf base) 6-25 cm long. Perigone pale blue or blue; har- 

dened perigone bases (Fig. 1D) spinulose ridged; anthers 


1973] Pontederia — Lowden 475 


blue, Tristylous (Fig. 3A-C). Fruits and seeds (Fig. 
2C-D) ovoid. 

In Central America (Fig. 22) a more frequent inhabitant 
of highland lagoons and lakes than the coastal plains. In 
South America (Fig. 25) primarily a floristic component 
of the tributaries of the Amazon Basin. 


Specimens Examined. CENTRAL AMERICA. British Honduras. TOLEDO 
DISTRICT: Monkey River, 9 Oct 1941, Gentle 3700 (F, GH, MICH, MO, 
NY, US). Costa Rica. PROVINCE ALAJUELA: Lago near Los Chiles, Rio 
Frio, 1 Aug 1949, Holm & Iltis 823 (GH). PROVINCE GUANACASTE: 
1 km from Arenal, 2 km from Tronadora, 18 Aug 1970, Lowden 19 
(os); Rio Arenal, 5 V 1923, Valerio 4 (US); Arenal, 5 IX 1923, 
Valerio 360 (US). PROVINCE LIMON: Barro Colorado de Norte, 8 Mar 
1965, Blaisdell 267 (FSU); Finca Montecristo, Rio Reventazon below 
Cairo, 18-19 Feb 1926, Standley & Valerio 49024 (Us). El Salvador. 
DEPARTMENT LA LIBERTAD: Laguna Zapotitan, 9 Nov 1953, Fassett 
29320 (Mo, WIS); *Laguna Zapotitan, 26 Mar 1970, Lowden 23 (os). 
DEPARTMENT SAN SALVADOR: San Salvador, 1922, Calderom 411 (NY, 
US); Asino, W end of Lago Ilopango, 30 Oct 1950, Fassett 28364 
(F, MO, US, WIS); Asino, W end of Lago Ilopango, 8 Jan 1951, Fassett 
28582 (F, GH, US, wis); San Salvador vic., 1905, Renson 273 (NY, 
US); Lago Ilopango, W Seite, bei Asino, 7 1 1951, Rohweder 566 (F); 
San Salvador vic., 20 Dee 1921-4 Jan 1922, Standley 19143 (GH, NY, 
US); San Salvador vic., 30 Mar-24 Apr 1922, Standley 23284 (GH, 
US). Guatemala. DEPARTMENT ALTA VERAPAZ: Panzos, 19 Nov 1920, 
Johnson 1028 (F, US); Panzos, 1 May 1906, Lewton 400 (vs). 
DEPARTMENT IZABAL: Izabal, Rio Izabal, 1 Jun 1919, Blake 7844 (GH, 
US). Honduras. DEPARTMENT COMAYAGUA: Pito Solo, Lake Yojoa, 
3 Sep 1932, Edwards AQ1 (F, GH); *Pito Solo, Lake Yojoa, 8 Aug 
1970, Lowden 12 (os); Pito Solo, Lago Yojoa, 18 Apr 1945, Rodri- 
guez 2917 (F); Cortez, Lake Yojoa, 7 Jul 1943, Yuncker 4840 (F, 
MICH, MO). DEPARTMENT CORTES: Agua Azul tract No. “B”, Lake 
Yojoa, 16 Aug 1951, Kamb 2094 (GH). Nicaragua. DEPARTMENT 
GRANADA: Granada, 11 Feb 1903, Baker (GH, MO, NY); Granada, 
Lake Nicaragua, J. M. & M. T. Greenman 5746 (MO); *Granada vic., 
la Terraza, Lago Nicaragua, 14, 16 Aug 1970, Lowden 17 & 18 (os). 
DEPARTMENT ?: U.S. North Pacific Exploring Expedition, 1853-56, 
Wright (GH, US). Panama. CANAL ZONE: Rio Chagres, near Gamboa, 
25 Aug 1939, Allen 1963 (GH, MICH, MO); Barro Colorado Island, 
Jan 1939, Brown 30 (F); Ahorca Lagarto to Culebra, 9 Mar 1905, 
Cowell 379 (NY); Frijoles, 17 Jun 1960, Ebinger 69 (MO); Barro 
Colorado Island, near dock, 28 Jun 1960, Ebinger 209 (Mo, US); 
Barro Colorado Island, 23 Dec 1963, Graham 212 (GH, MICH) ; Gatun 
Lake, Jul 1965, Hayden 123 (M0); Gatun Station, Panama Railroad, 


476 Rhodora [Vol. 75 


Figure 22. Distribution of Pontederia rotundifolia in Central 
America (arrow points to insert of Canal Zone). 


28 Oct 1859, Hayes 89, 93 (GH) and 99 (NY); Miraflores Lake, 
Empire, Culebra Cut and vic., 27 Feb 1935, Hunter & Allen 776 
(MO); Barro Colorado Island, 24 Jul 1927, Kenoyer 602 (Us); 
Chagres River, N of Gamboa, 2 Jun 1969, Lazor, Correa & Boreham 
3739 (FSU); Chagres River below Gatun, 17 Feb 1911, Maxon 4883 
(US); Between Gatun and Lion Hill, 26 Jan 1911, Pittier 2572 (GH, 
NY, US); Barro Colorado Island, Barbour Trail, 15 Nov 1931, Shat- 
tuck 408 (F); Barro Colorado Island, Pena Blanea Bay, 31 Jul 1934, 
Shattuck 1099 (F, Mo); Barro Colorado Island, Starry 120 (F); 
Barro Colorado Island boathouse, 25 Jun 1966, Tyson 4198 (FSU, 
MO); Gamboa, 14 Jul 1966, Tyson, Dwyer & Blum 4604 (FSU, NY, 
us); Barro Colorado Island, Chapman Trail, 28 Feb 1931, Wilson 74 
(F); Madden Lake, 3-20 Aug 1940, Woodson & Schery 949 (MO, NY, 
US); Barro Colorado Island, W of end of T. Barbour Trail, 11 Feb 
1932, Woodworth & Vestal 479 (F, GH). PROVINCE DARIEN: Rio Pirre, 
Apr 1966, Duke & Bristan 8307 (Mo). SOUTH AMERICA. Argen- 
tina. Entre Ensenada and Punta Lara, Prov. Buenos Aires, La 


1973] Pontederia — Lowden ATT 


Plata, 2 Apr 1931, Cabrera 1708 (NY); Buenos Aires, Rio Santiago, 
Prov. Corrientes, 18 Apr 1964, Cabrera 15915 (Ny); Camino de 
Santa Fe, Laguna Stubal’s Guadalupe, Prov. Santa Fe, 1 II 1936, 
Job 679 (NY); Estancia “Santa Teresa", Dep. Mburucuya, Prov. 
Corrientes, 2 4 1954, Petersen 2657 (Mo, US); Prope Barra, Prov. 
Rio Negro, May 1851, Spruce 1486 (NY). Bolivia. DEPARTMENT BENI: 
Lake Rogagua, 3 Nov 1921, Cardenas 1417 (BKL, MICH, NY, US); 
Rio Chapare-Mamore, VIII 1926, Werdermann 2198 (MO); Reyes, 
25 Oct 1921, White 1513 (BKL, NY). DEPARTMENT LA PAZ: Ixiamas, 
21 Dec 1921, Cardenas 2031 (BKL, MICH, NY). DEPARTMENT SANTA 
CRUZ: Yapacani, VI 1892, Kuntze (NY); Prov. Para, 6 Feb 1925, 
Steinbach 6977 (GH). Brazil. STATE AMAZONAS: Basin Rio Madeira, 
municipality Humayta, between Monte Christo and Santa Victoria, 
Rio Ipixuna, 15-17 Nov 1934, Krukoff 7129 (F, GH, MICH, MO, NY, 
US). STATE PARA: Belem, Mar-May 1929, Dahlgren & Sella 525 (F, 
US), 557 (F, GH); Villa Carmo, municipio Cameta, 4 Jul 1935, Drouet 
1999 (GH, MICH, US); Taperinha bei Santarem, Amazonas armes 
Ayaya, II VI 1927, Ginzberger & Zerny 757 (F); Rio Parana, Barra 
do Rio Piquiry, State of Parana, Mun. Guaira, 9 IV 1961, Hatsch- 
bach 8078 (Us); Bocca do Paru, Region Lower Amazon, 28-29 Aug 
1934, Krukoff 5935 (GH, NY); Amazonia, 28 Sep 1945, Pires & Black 
770 (GH); Amazonia, Territory Rondonia, Basin Rio Madeira east 
bank at Abuna, 12 Jul 1968, Prance, Forero, Wrigley, Ramos & 
Farias 5914 (Ny, us); Amazonia, Territory Rondonia, Basin Rio 
Madeira, Rio Pacaas Novos, 3 Aug 1968, Prance, Forero, Wrigley, 
Ramos & Farais 6764 (NY, US). Colombia. Frontera Colombo- 
Ecuatoriana, commissaria Putumayo, rio San Miguel entre los aflu- 
entes Bermeja y Conejo, 13 Dec. 1940, Cuatrecasas 11054 (F, US); 
Mocoa, camino de herradura antiquo y rio Rumiyaco, 10 Oct 1965, 
Garcia-Barriga, Hashimoto & Ishikawa 18685 (NY); Amazonas, 
Trapecio Amazonico, Loretoyacu River, Nov. 1945, Schultes 6935 
(us); Amazonas, Trapecio Amazonico, Amazon River, Leticia, Sep 
1946, Schultes 8188 (Us). Guiana. Martin, Ex Herbario Musei 
Britannici (F). Paraguay. Lag. Ypacaray, 1885-1895, Hassler 1196 
(NY); Lacus Ypacarai, Dec 1900, Hassler 3693 (NY); Lacus Ypa- 
carai regione, 1913, Hassler 11861 (F, GH, MO, NY, US) ; San Bernar- 
dino, Dept. Cordillera, 4 II 1951, Sparro & Vervoorst 2316 (GH); 
Apr-May 1845, Weddell 3156 (NY). Peru. DEPARTMENT LORETO: 
Requena, Prov. Requena, Ucayalio Fangoso, 22 Aug 1965, Sagastequi 
& Aldave 5757 (us); Pebas, Amazon River, 29 Jul 1929, Williams 
1854 (F); La Victoria, Amazon River, 5 Sep 1929, Williams 3097 
(F, US). Surinam. Focke (GH); Via secta ab Wia-bank and Grote 
Zwiebelzwamp, 23 Nov 1948, Lanjouw & Lindeman 1236 (NY). 
Venezuela. Pto. de La Ceiba, sur lago de Maracaibo, Edo. Trujillo, 
Jan 1961, Aristeguiseta 4443 (Us); C. Araguo [Boca Araguao], 
State Delta Amacuro, Dec 1952, Gines 4881 (US); 12 km SW of 


478 Rhodora [Vol. 75 


Punta de Piedra towards Sacramento, Estado Tachira, 27-30 Aug 
1966, Steyermark & Rabe 96610 (vus). 

This vigorous species is quite variable in size and leaf 
forms. In particular, specimens with sagittate leaves have 
been confused with Pontederia sagittata. The spinulose 
hardened perigone bases (Fig. 1D) and fewer flowered 
globose inflorescences (Fig. 20) readily distinguish P. ro- 
tundifolia from members of subgenus Pontederia. 


5. Pontederia subovata (Seub. in Mart.) Lowden, comb. 

nov. 

Eichhornia subovata Seub. in Mart. Fl. Bras. 3 (1): 91. 
1847. Type: Brazil: Prov. Goyazana, 1836-1841, Gard- 
ner 4022 (Lectotype chosen, NY!); prov. Bahiensi, 
Blanchet 2720. 

Reussia triflora Endl. ex Seubert in Martius, Fl. Bras. 3 
(1): 96. 1847. Type: Brazil: Pohl, Sellow, Type not 
seen. 

Pontederia schomburgkiana Klotzsch in Schomburgk., 
Vers. Faun. & Fl. v. Brit. Guiana 1118. 1848. Type: 
British Guiana: *Im See Venturu und auf stromlosen 
Stellen des Pirara in der Nähe seiner Quelle". Type not 
seen, 

Pontederia lagoénsis Warming in Videnskab. Meddel, p. 
323, T. VI. 1871. Type: Brazil: Min, Geraes, Lagoa 
Santa. Type not seen. 

Reussia subovata (Seub. in Mart.) Solms in DC., Monog. 
Phan. 4: 534. 1883. 

Reussia lagoénsis (Warm.) Castell., Arq. Jard. Bot. Rio de 
Janeiro 16: 209. 1958. 

Plants up to 30 cm tall. Leaves subovate (Fig. 23), ovate 
lanceolate or elliptic (Fig. 24); blades 0.2-4.4 em wide; 
spathe (terminal phyllode of the floral shoot) 1.9-5 cm 
long; petiole of floral shoot leaf (includes sheath base) 
4-10 cm long; petiole of other leaves 6.5-25 cm long, ligule 
(Fig. 23) of petiole sheath up to 3.5 cm long; sheath in 
axil of floral shoot up to 6.5 cm long. Inflorescence (Fig. 
23) 3-8 cm long; peduncle (inflorescence base to floral shoot 
leaf base) 5-14 cm long. Perigone ephemeral, blue purple, 


1973] Pontederia — Lowden 479 


Figure 23. Inflorescence and subovate leaves of Pontederia subo- 
vata (arrow points to a ligule of a petiole sheath), Paraguay, Ypa- 
caray, E. Hassler 12503 (Mo); at left. 

Figure 24. Narrow subovate-lanceolate leaf variant of Pontederia 
subovata, Brazil, Minas Gerais, Lagoa Santa, L. B. Smith 6702 (us); 
at right. 


blue green or pale blue; hardened perigone bases spinulose 
ridged; anthers blue. Tristylous. 

Inhabits swamps and lakes of Argentina, Bolivia, Brazil, 
British Guiana, Paraguay and Venezuela. Figure 25. 


Specimens Examined. SOUTH AMERICA. Argentina. Territorio 
de Formosa, 4 1919, Jorgensen 3347 (GH, US); Prov. Chaco, Loc. 
Colonia Benitez, 10 II 1941, Meyer 3864 (F, GH, NY); Estancia, 
"Santa Teresa", Dep. Mburucuya, Prov. Corrientes, 4 1 1952, Peter- 
sen 1427 (NY, US). Bolivia. Trinidad [Dept. of Beni], 7 Mar 1922, 
Cardenas 27 (GH, NY); Santa Cruz, Chiquitos, between El Carmen 
& Palmito-Chiquitos, II 1950, Cardenas 4488 (vus); Reis, Jun 1886, 
Rusby 553 (GH, NY). Brazil. 1836-1841, Gardner 4022 (NY); Loc. S. 
Tereziuha, Mun. Ituiutaba, Estado Mg [Minas Gerais], 23 Apr 1950, 
Macedo 2326 (Ny); Loc. S. Terezuiha, Estado Minas, Mun. Ituiutaba, 
18 Feb 1951, Marcedo 3168 (Us); Canoas, Estado do Rio Grande do 
Sul, 1 2 1949, Miguel 23 (F, GH, US); Minas Gerais, Mun. Lagoa 
Santa, Lagoa Santa, 3 May 1952, Smith 6702 (F, GH, NY, US). 


480 Rhodora [Vol. 75 


Figure 25. Distribution of Pontederia rotundifolia (dots) and 
P. subovata (triangles) in South America. 


1973] Pontederia — Lowden 481 


British Guiana. Rupununi, 20 mls. S. of Lethem, road to Wichabi, 
22 Nov 1957, Cooke 197 (NY); Twinpools South Sand Creek, Cooke 
205 (NY); Rupununi Northern Savanna, Mauritia Swamp, !4 mi N 
Waruma, 30 Sep 1963, Goodland 897 (Us). Paraguay. Iter ad Para- 
guariam Septentrionalem, prope Concepcion, Sep 1901/2, Hassler 
1352 (GH, NY); Paraguaria Centralis, in regione lacus Ypacaray, 
Feb 1918, Hassler 12503 (GH, MO, NY, US); Pilcomayo River, 1888- 
1890, Morong 859 (Ny). Venezuela. State Cojedes, near San Carlos, 
15 Jan 1939, Alston 6311 (wis); Apure, near Cunaviche, 13 Feb 1941, 
Chardon 254 (vs). 


The subovate and lanceolate leaf forms are quite variable, 
however, distinct. The narrow almost linear-lanceolate 
variants (Fig. 24) are not formally treated since material 
studied was limited. The hardened perigone bases are 
spinulose ridged. 


ACKNOWLEDGEMENTS 


This study (Paper no. 799, Botany Department, The 
Ohio State University) is a revised version of the original 
presented as partial fulfillment of the requirements for the 
Doctor of Philosophy Degree at The Ohio State Univer- 
sity. Dr. Ronald L. Stuckey was the dissertation advisor. 
The Organization for Tropical Studies, Inc. (OTS Pilot 
Research Grant F 70-23) funded field research in Mexico 
and Central America during the summer of 1970. The 
Graduate Committee of the Department of Botany and 
the Graduate School Fellowship Committee (The Ohio 
State University) awarded a University Dissertation Year 
Fellowship for the completion of this investigation. Direc- 
tors of the following herbaria are gratefully acknowledged 
for the loan of specimens studied: Arnold Arboretum 
(A), Brooklyn Botanic Garden (BKL), National Museum 
of Canada (CAN), Duke University (DUKE), Field Museum 
of Natural History (F), Florida State University (FSU), 
Gray Herbarium (GH), University of Michigan (MICH), 
Missouri Botanical Garden (Mo), New England Botanical 
Club (NEBC), New York Botanical Garden (NY), Ohio 
State University (os) and Franz Theodore Stone Labora- 
tory (FTSL), Academy of Natural Sciences of Philadelphia 


132 Rhodora [Vol. 75 


(PH), Smithsonian Institution (US), and University of 
Wisconsin (WIS). 


LITERATURE CITED 


ADANSON, M. 1763. Familles Des Plantes, pp. 54 & 581. Vincent. 

ALSTON, R. E. 1967. Biochemical Systematics. In Dobzhansky, 
Hecht and Steere (eds.), Evolutionary Biology, 1, pp. 197-9505. 
New York. 

BAKER, H. G. 1970. Evolution in the Tropics. Biotropica 2: 101- 
111. 

BowbEN, W. M. 1945. A list of chromosome numbers in higher 
plants. II. Menispermaceae to Verbenaceae. Am. J. Bot. 32: 
191-201. 

BRITTON, N. L. and H. A. Brown. 1913. An Illustrated Flora of 
the Northern United States, Canada and the British Possessions, 
2nd ed., 1, pp. 462-463. New York. 

CASTELLANOS, A. 1951. Revision de las Pontederiaceae Argentinas. 
Lilloa, Revista de Botanica 25: 585-594. 

1958. Las Pontederiaceae de Brazil. Rio de 
Janeiro Jardin Botanico 16: 149-216, tab. 1-18. 

CORE, E. L. 1966. Vegetation of West Virginia, p. 140. Parsons. 

ENDLICHER, S. L. 1836. Genera plantarum, 1, pp. 137-139. Vienna. 

FARWELL, O. A. 1924. Notes on the Michigan Flora. Part VI. Pap. 
Michigan Acad. Sci. 3: 87-109. 

1928. Contributions to the Botany of Michigan 
No. 15. Miscellaneous Notes. Am. Midl. Natl. 11: 72-85. 

FERNALD, M. L. 1925. Pontederia versus Umisema. Rhodora 27: 
76-81. 

HiTCHCOCK, A. S. and M. L. GREEN. 1929. International Botanical 
Congress, Cambridge (England), 1930, Nomenclature, Proposals 
by British Botanists, pp. 110-114, 144. London. 

KNOWLTON, F. H. 1922. Revision of the Flora of the Green River 
Formation, with descriptions of new species. U.S. Geol. Surv., 
Profess. Paper no. 131: 133-182, pl. 36, fig. 6. 

LINNAEUS, C. 1737a. Genera plantarum, 1st ed., p. 102. Leiden. 

1737b. Hortus cliffortianus, p. 133. Amsterdam. 

1753. Species plantarum, 1, p. 288. Stockholm. 

1754. Genera plantarum, 5th ed., p. 140. Stockholm. 

Masry, T. J. K. R. MARKHAM and M. B. THoMas. 1970. The 
Systematic Identification of Flavonoids, 354 pp. New York. 

NuTTALL, T. 1818. The Genera of North American Plants and a 
Catalogue of the Species, to the Year 1817, 1, pp. 215-217. 
Philadelphia. 

ORNDUFF, R. 1966. The Breeding System of Pontederia cordata 
L. Bull. Torr. Bot. Club 93: 407-416. 


1973] Pontederia — Lowden 483 


PunsH, F. 1814 [1813]. Flora Americae Septentrionalis; or, a 
Systematic Arrangement and Description of the Plants of North 
America, 1, pp. 223-224. London. 

RAFINESQUE, C. S. 1808. Prospectus of Mr. Rafinesque Schmaltz’s 
two intended works on North-American Botany. Medical Re- 
pository, second hexade, 5, no. 4, pp. 350-356. New York. 

1830. Medical Flora; or Manual of the Medical 

Botany of the United States of North America, 2, pp. 105-108. 

Philadelphia. 


. 1837. New Flora of North America. Part II 
Neophyton, 96 pp. Philadelphia. 

ScHUCHERT, C. 1935. Historical Geology of the Antillean-Caribbean 
Region, 811 pp. New York. 

SCHULZ, A. G. 1942. Las Pontederiaceas de la Argentina. Dar- 
winiana 6: 45-82, 5 pl. 

SCHWARTZ, O. 1927. Zur Systematik und Geographie der Ponte- 
deriaceen. In Engler, Bot. Jahrb. 61, Beibl. no. 139, pp. 28-50. 

1930. Pontederiaceae. Im Engler and Prantl, Die 
Naturlichen Pflanzenfamilien, ed. 2, 15a, pp. 181-188. 

ScULTHORPE, C. D. 1967. The Biology of Aquatic Vascular Plants, 
610 pp. London. 

SMITH, W. R. 1898. A Contribution to the Life History of the 
Pontederiaceae. Bot. Gaz. 25: 324-337, pl. 19-20. 

SOLMS-LAUBACH, H. 1883.  Pontederiaceae. In A. DeCandolle, 
Monographiae Phanerogamarum, 4, pp. 501-535. 

SPRAGUE, T. A. 1924. Unwarranted Changes in Generic Names. 
J. Bot. 62: 327-328. 

ToRREY, J. 1824. A Flora of the Northern and Middle Sections of 
the United States, 1, pp. 342-343. New York. 


FACULTAD DE CIENCIAS Y HUMANIDADES 
DEPARTAMENTO DE CIENCIAS NATURALES 
UNIVERSIDAD CATOLICA MADRE Y MAESTRA 
SANTIAGO DE LOS CABALLEROS 

REPUBLICA DOMINICANA, ANTILLAS (W.I.) 


184 


Rhodora [Vol. 75 


Table 2. Chromosome Counts of Pontederia Taxa Investigated 


Taxon 


Haploid 
Counts 


Vouchers (os) 


subg. Pontederia 
P. cordata var. 
cordata, 


P. sagittata 


P. parviflora 


subg. Reussia 
P. rotundifolia 


16 


U.S.: Ohio, Ottawa Co., Winous Pt., 
SW of Port Clinton, Lowden 36, 37; 
New Hampshire, Strafford Co., Bellamy 
River, Lowden 2. British Honduras: 
Belize District, 9 miles N of Belize 
City, Northern Highway, Lowden 24. 


Mexico: State Vera Cruz, Vera Cruz, 
Lowden 6, Lerdo, Lowden 7 and Laguna 
Catemaco, Lowden 9. Guatemala: Dept. 
Izabal, Quirigua, Lowden 10 and Puerto 
Barrios, Lowden 11. Honduras: Dept. 
Cortes, Puerto Cortes, Lowden 13 and 
13.5 miles inland from Puerto Cortes 
Lowden 14; Dept. Atlantida, Tela, 
Lowden 15 and Santiago, 11 miles W 
of Tela, Lowden 16. 


Panama: Province Herrera, Los Llanos 
de Santa Maria, Lowden 20; Province 
Panama, between Chepo and Pacora, 7 
miles from Chepo, Lowden 21. 


Honduras: Dept. Comayagua, Lago 
Yojoa, Pito Solo, Lowden 12. Nicaragua: 
Dept. Granada, near Granada, Lowden 
17, 18. Costa Rica: Province Guanacaste, 
1 km. from Arenal, 2 km. from Trena- 
dora, Lowden 19. El Salvador: Dept. 
La Liberatad, Laguna Zapotitan, Low- 
den 28. 


485 


1973] Pontederia — Lowden 
Table 3. Spot Colors of Phenolic Compounds in Pontederia and 
Related Genera 
Spot No Reagent Reagent NH40H 
# Visible Light UV Light UV Light 
1 aqua green (in part) 
2 aqua green (in part) 
3 purple yellow green brown 
(in part) 
4 cream-yellow 
5 blue purple 
6 green blue-cream purple 
7 white blue 
8 aqua 
9 dark blue 
10 aqua green (sometimes) 
lla rose cream 
11b blue green-pale blue 
11c blue purple 
12 green brown green yellow, red brown- 
pink purple 
13 greenbrown pink purple-bright pink 
14 purple, blue-white blue 
15a yellow green brown 
15b yellow 
16 pink (some- — bright-dull pink 
times) 
17 purple 
18 dull green blue-purple 
19 dull pink 
20 dull yellow green brown- 
green blue 
21 salmon 
22 yellow green brown 
23 yellow-gray cream 
24 pale blue 
25 dark yellow green brown 
26 dull pink purple 


[Vol. 75 


Rhodora 


186 


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AN UNUSUAL SUBSTRATE FOR POLYSIPHONIA 
PANICULATA MONTAGNE. Once in a while the curious 
juxtaposition of circumstances leads to an unusual find. 
Such a situation occurred during the summer of 1972 when 
we were exploring the algal communities along the edge 
of the San Andreas fault, at one of its drowned points. 
The fault emerges from a land position to follow Tomales 
Bay some 60 miles northwest of San Francisco, From 
Tomales Bay the fault lies under Bodega Bay and across 
Bodega Head, still on a northwesterly line. 


We were engaged in field work for the Pacific Marine 
Station at Dillon Beach, California, and while working 
the mud flats along the east side of Tomales Bay at a 
place known as Lawson's Landing came across a rather 
curious partnership. Here we found a presumably symbiotic 
partnership between the rhodophycean alga Polysiphonia 
paniculata Montagne and the Horseneck Clam, Tresus 
nuttalli Conrad. 

At the time we were collecting macroalgae, various species 
of Ulva being very common on the flats, and as we ex- 
amined a stand of Zostera marina L. in a shallow subtidal 
area, we found individual plants of Polysiphonia panicu- 
lata, scattered throughout the Zostera. Upon grasping the 
alga it became readily apparent that something unusual 
was occurring, because the alga would quite noticeably 
retract into the mud. Small plants would actually vanish 
from sight, going to depths that made any attempt to col- 
lect them futile. The first samples we collected of the alga 
lacked the lower portions. 

Further search among the Zostera revealed another 
movement; whenever several hunters gathered around one 
of the Polysiphonia plants, it would “settle” downward 
several centimeters, almost as if it were anticipating our 
next action. Never having had algae attempt to “get away" 
before, we concentrated on solving the mystery. After 
several more misses, we succeeded in getting a whole plant, 
the trick being for one person to stalk the plant, and 
moving carefully, to use a long bladed knife to cut through 


488 


1973] Polysiphonia — Colt and Swartz 489 


the mud as deeply as possible below the protruding portion 
of the plant as the grab for the plant was made. 

The cut brought to light a portion of the substrate to 
which the Polysiphonia was attached, and it proved to be 
the distal portion of the siphon of Tresus nuttalli. That 
the alga was firmly attached to the siphon was quite ap- 
parent, and subsequent microscopic examination revealed 
that the prostrate branches of the Polysiphonia had pene- 
trated into the epidermal tissue of the siphon to a depth 
of several millimeters. Most of the samples we obtained 
in this manner were evidently from older clams, being 
2 plus centimeters in the longest dimension. 

Our examination of the general area suggested that the 
Polysiphonia serves to hide the siphon from overhead 
observation, thus affording some degree of protection to 
the clam. At no time were we able to see a protruding 
siphon or any evidence that a siphon was present, although 
the alga was very evident. We collected during low tide 
periods, and this did not coincide with the clam's feeding 
times. As far as we were able to determine over the 
several acres of flats, the association was confined to the 
stands of Zostera. On the open areas of the flats, heavily 
dug over by clam diggers, only siphons or siphon holes 
were to be seen. 

We were unable to determine the significance of the 
association other than on an inferential basis. Generally in 
this region Polysiphonia can be regularly found wherever 
a firm substrate and tidal flow can be paired. It seemed 
logical to us that the clam benefited from the position of 
the alga, and that the alga would in turn benefit from the 
flow of water into and out of the siphon of the clam. This 
flow certainly would provide a source of nutrients better 
than the usual rock or wharf substrate. 

A search among specimens in the herbarium at the 
Station revealed that previous collections of Polysiphonia 
had been made in the same mud flat area, but there was 
no reference to the type of substrate from which these 
previous collections were made. Due to the lack of any 


490 Rhodora [Vol. 75 


other suitable substrate on the flats, we infer that the 
association we found is not new to this area. Polysiphonia 
paniculata was also collected at the north end of Dillon 
Beach, but in each instance the substrate was a rock sur- 
face slightly buried in sand. 

Although Tresus is reported to serve as a host for a 
variety of smaller fauna (Ricketts et al., 1968), we are 
unaware of any previous reports of an alga living in such 
a partnership with this clam. 


REFERENCES 


RICKETTS, E. F., J. CALVIN, and J. W. HEDGPETH. 1968. Between 
Pacific Tides. 4th Ed. Stanford Univ. Press. 

SMITH, G. M. 1969. Marine Algae of the Monterey Peninsula Cali- 
fornia. Stanford Univ. Press. 


L. C. COLT, JR. 
BOSTON STATE COLLEGE 
BOSTON, MASS. 02115 


STEVEN SWARTZ 
PACIFIC MARINE STATION 
DILLON BEACH, CALIF. 94929 


NEW RECORD FOR THE YELLOW LADY'S SLIP- 
PER ORCHID, CYPRIPEDIUM CALCEOLUS L. SUBSP. 
PARVIFLORUM (SALISB.) HULT., FROM ALASKA. 
In 1964, while at the University of Alaska, I was asked by 
Mr. Ross Brockman to identify a collection of plants from 
the Bettles River area about 20 miles northeast of Wise- 
man (ca. 67?36'N, 149°40’W), in the Brooks Range region 
of Alaska. Among these plants was a yellow orchid which 
at that time I mistakenly determined as Cypripedium gutta- 
tum Sw., using Hultén's (1943) Flora of Alaska and Yukon, 
part 3. A recent reexamination of this specimen clearly 
reveals it to be Cypripedium calceolus L. subsp. parviflorum 
(Salisb.) Hult. This species is reported by Hultén (Flora 
of Alaska and Neighboring Territories, 1968, p. 316) from 
as far northwest as the upper Salmon Fork River area 
(about lat. 67°), Yukon Territory near the Alaskan border. 
The present collection (Ross Brockman, 26 June 1962, 
ALA) thus constitutes the first report of this species from 
Alaska and represents about a 250 mile westward and 
nearly a 100 mile northward range extension in North 
America. The specimens were from a colony of about 100 
plants found at the well-drained, rather rocky, base of a 
limestone knoll near Bettles River. At this locality, Cypri- 
pedium calceolus occurred in a drier situation and bloomed 
earlier than the more common white lady’s slipper, C. pas- 
serinum Rich. 


VERNON L. HARMS 

FRASER HERBARIUM 
UNIVERSITY OF SASKATCHEWAN 
SASKATOON, SASKATCHEWAN 


491 


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should be selected critically especially for common species 
of broad distribution. Systematic revisions and similar 
papers should be prepared in the format of “The System- 
aties and Ecology of Poison-Ivy and the Poison-Oaks," 
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu- 
larly with reference to the indentation of keys and syno- 
nyms. Papers of a floristic nature should follow, as far as 
possible, the format of “Contribution to the Fungus Flora 
of Northeastern North America. V.," H. E. Bigelow & M. E. 
Barr, Rhodora 71: 177-203. 1969. For bibliographie cita- 
tions, a recommended list of standard journal abbreviations 
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. 
Club 85: 277-300. 1958. 


Volume 75, No. 803, including pages 325-492, was issued 
October 15, 1973. 


492 


CONTENTS: — continued 


An Unusual Substrate for Polysiphonia paniculata Mon- 
tagne 
L. C. Colt, Jr. and Steven Swartz ........... eee eerie eene entente 488 


New Record for the Yellow Lady's Slipper Orchid, Cypripe- 
dium calceolus L. subsp. parviflorum (Salisb.) Hult, from 
Alaska 
BEE PTI 2 Tl... u. ............u u erben siquid 491 


Instructions for Contributors ............. 1 :« eene nennen 492 


odora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
ALBION REED HODGDON, Editor-in-Chief 


ROLLA MILTON TRYON 7 


RADCLIFFE BARNES PIKE 
STEPHEN ALAN SPONGBERG 
GERALD JOSEPH GASTONY 
ALFRED LINN BOGLE 


> Associate Editors 


RICHARD EDWIN WEAVER y 
Vol. 75 December, 1973 No. 804 
CONTENTS: 
A Partial History of the New England Botanical Club. 
Richard A. Howard. a a S u a 498 


The Systematic Value of Trichome Complements in a 
North American Group of Vernonia (Compositae). 
W. Z. Faust and S. B. Jones, Jr. ............................................ 517 


The Morphology and Systematics of the Audouinella 
Complex (Acrochaetiaceae, Rhodophyta) in Northeastern 


United States. Wm. J. Woelkerling nousee 529 
Book Review. A. R. Hodgdom essees 622 
Instructions for Contributors. sesser 624 
MINE u s tapete eu aute s nk UELLE 625 


The Nem England Botanical Club, Jue. 
Botanical Museum, Oxford St.. Cambridge, Mass. 02188 


RHODORA. — A quarterly journal of botany, devoted primarily to the 
flora of North America and floristically related areas. Price $10.00 
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 
plants of North America, will be considered by the editorial com- 
mittee for publication. Articles concerned with systematic botany 
and cytotaxonomy in their broader implications are equally accept- 
able. All manuscripts 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. Extracted reprints, if ordered in advance, will be 
furnished at cost. 

Address manuscripts and proofs to Albion R. Hodgdon, 

Dept. of Botany, Nesmith Hall, 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 


Rhodora 


JOURNAL OF THE 
NEW ENGLAND BOTANICAL CLUB 


Vol. 75 December, 1973 No. 804 


A PARTIAL HISTORY OF 
THE NEW ENGLAND BOTANICAL CLUB 


RICHARD A. HOWARD 


The four hundred and seventy-third meeting of the New 
England Botanical Club, being the seven hundredth since 
the original establishment, was held at the Harvard Faculty 
Club, Cambridge, Massachusetts on April 6, 1973, as a 
dinner meeting. One hundred six members and guests were 
present. Dr. Wesley Tiffany presided. Dr. Gordon DeWolf, 
program chairman, arranged the evening celebration. At 
the head table were the senior members of the Club who 
were able to be present: Ralph Bean (elected 1909), 
G. Safford Torrey (elected 1912), Donald White (elected 
1913), Henry Svenson (elected 1919), W. H. “Cap” Weston 
(elected 1921), Ralph Wetmore (elected 1927), and Hugh 
Raup (elected 1929), the officers, the speaker and their 
wives. A printed menu was available, as was a souvenir 
program, prepared by Dr. DeWolf, which contained a 
reprint of an address by Emile F. Williams on the occasion 
of the tenth anniversary of the Club, a summary of meet- 
ing places, and a roster of the officers and committee 
chairmen who have served the New England Botanical 
Club during the years, I was asked to “reminisce” as the 
presentation of the evening. Although I am not the oldest 
member of the Club by a good many years, I did have a 
few kodachrome slides of earlier years, and I supple- 


495 


494 Rhodora [Vol. 75 


mented these with some investigations into the history of 
the Club. Regrettably, the first four volumes of the minutes 
of meetings of the Club have been “misplaced” for many 
years. They have since been found, so the following ma- 
terial is not exactly what was said in the course of the 
evening but amplified in part. 

In 1894 Mr. Warren H. Manning, a landscape architect 
in Boston, called a meeting at the old Horticultural Hall in 
Boston of the local botanists in order to ask their coopera- 
tion in assembling an herbarium of the plants of the 
Boston Metropolitan Parks. This may have planted the 
idea of a club of New England botanists. Manning noted 
that almost all assembled had to be introduced to each 
other. His task was to prepare a list of the plants growing 
in the local parks. Such a flora was published eventually 
by Walter Deane. 

On December 4, 1895, Professor W. G. Farlow and Dr. 
B. L. Robinson invited seventeen gentlemen to meet on 
December 10 to consider the advisabilitv of forming a 
botanical club. One of the invitations preserved in the 
Club archives specifies: “My house is 24 Quincy Street, 
next door to the Colonial Club. The cars stop at Quincy 
Square (Beck Hall)" On the site of Farlow's house now 
stands the Carpenter Center. The Colonial Club was re- 
placed by the Harvard Faculty Club. Beck Hall was on 
the location of the Gulf Station on Massachusetts Avenue, 
and the name Quincy Square is no longer used. Thus, the 
seven hundredth meeting was held next door to the original 
gathering place where these men formed the New England 
Botanical Club. A second meeting within a month reaffirmed 
the decision to form a botanical club, and the NEBC was 
officially christened on February 5, 1896, at a third meeting. 
The charter members were seventeen in number. At the 
fourth meeting in March, 1896, Messrs. Fernald, Fuller, 
Greenman, J. R. Jack, Manning, John Robinson, and Rich 
from Boston were elected members, and the first non- 
resident members, Sears and Bailey of Providence and 
Jackson of Worcester, were elected. 


1973] New England Botanical Club — Howard 495 


The early meetings were held in members’ homes, Farlow, 
Sargent, and Goodale being cited as excellent hosts. Later 
in 1896 the Club met at Young’s Hotel on Court Street; 
from November 6, 1896 to December 4, 1903 at the St. 
Botolph Club on Newbury Street; from 1904-1906 at the 
Hotel Brunswick on Boylston Street; from 1907-1923 at 
the Twentieth Century Club on Joy Street, and from 1924- 
1954 at the American Academy on Newbury Street, all in 
Boston. Since 1954 the NEBC has met at the Harvard 
University Herbaria in Cambridge. 


The Founders 


The charter members of the NEBC were seven profes- 
sional botanists and ten amateurs. Although biographical 
sketches of some are published in Rhodora, not all charter 
members are so recorded. Their contributions to the NEBC, 
to botany of New England, and to botany at Harvard are 
so great that a brief review of these founders seems to be 
in order. They lived to an average age of seventy-seven 
years, supporting the oft-quoted statistic on the longevity 
of botanists. 


Joseph Richmond Churchill — 1845-1933 
(Rhodora 36: 1-7. 1934.) 

A member of the class of 1867 at Harvard, Churchill 
studied botany under Asa Gray, but he specialized in law 
(Class of 1869) and was appointed a judge after only 
two years of private practice in 1871. He started collecting 
plants with his fiancée, Mary Cushing, and they jointly 
prepared five hundred numbers before their wedding. 
Churchill’s personal herbarium of 13,313 specimens was 
left to the Missouri Botanical Garden through the in- 
fluence of Jesse M. Greenman. However, Churchill had 
exchanged duplicates with Walter Deane, whose herbarium 
is in the NEBC. Churchill wrote a Flora of Milton (1887). 
His will contained a bequest of $1,000 for Rhodora. 


496 Rhodora [Vol. 75 


Frank Shipley Collins — 1849-1920 
(Rhodora 22: 96. 1920., Amer. Jour. Bot. 19: 54-62. 
1926.) 


An algologist with many articles on the subject to his 
credit, Collins was on the editorial staff of Rhodora for 
twenty-one years. Although an obituary was promised for 
Rhodora, only the death notice cited above ever appeared. 


George Edward Davenport — 1833-1907 
(Rhodora 10: 1-9. 1908.) 


Best known for his contributions to the study of ferns 
in nearly one hundred articles, Davenport also specialized 
on the general flora of Middlesex Fells and wrote a Flora 
of Medford (1875-1876). His collection of fern specimens 
was given to the Massachusetts Horticultural Society in 
1875, and shortly after that he began a new personal her- 
barium. This his daughter gave to the Gray Herbarium 
after his death. Subseqently the earlier collections were 
given by the Massachusetts Horticultural Society to the 
Gray Herbarium, 


Lorin Low Dame — 1838-1903 
(Rhodora 5: 121:123. 1903.) 


A principal of Lexington High School and subsequently 
of Medford High School for a total of twenty-seven years, 
Dame collected in the Lexington area and on Nantucket. 
He is best remembered for his books, “Typical Elms and 
Other Trees of Massachusetts" (1890) and the “Handbook 
of the Trees of New England" (1902). 


Walter Deane — 1848-1933 
(Rhodora 35: 69-80. 1933.) 

Deane was a graduate of Harvard, class of 1870, and 
taught in the private schools of St. Mark's and Hopkinson 
for twenty-five years. He became interested in botany in 
1880 and worked hard on the flora of the Boston Park 
System for the M.D.C., for which he compiled an herbarium 
which became the first large accession of the NEBC. He 


1973] New England Botanical Club — Howard 497 


also helped curate the ornithological collection of William 
Brewster and helped issue the “Birds of the Cambridge 
Region” (1906). His first herbarium specimen was identi- 
fied by Asa Gray as Gerardia purpurea and is in the Club 
herbarium. Deane collected with Kennedy and Churchill 
and was a long time friend of L. H. Bailey. He bought 
and exchanged specimens and built up a personal her- 
barium of 40,000 sheets. The early lists of distributions 
of New England plants were often based on his meticu- 
lous records. Walter Deane also gathered New England 
specimens which were to serve as the models for the 
Blaschka glass flowers being assembled by Goodale. 

Deane served on the first Visiting Committee of the 
Gray Herbarium and helped Robinson plan the building 
constructed in the Botanic Garden in 1915, donating his 
personal library to the Gray Herbarium in 1918. He 
assembled an extensive collection of portraits of botanists. 
On his seventy-fifth birthday (1923) the members of the 
NEBC, as a surprise to him, collected $1,000 which was 
presented to the Club as a library fund in Deane’s honor. 
His will contained bequests to the Club and to the Gray 
Herbarium. 


William Gilson Farlow — 1844-1919 
(Rhodora 22: 1-8. 1920.) 

Although Asa Gray had been asked much earlier to 
organize a club of amateurs and professionals for the study 
of the New England flora, he did not feel it was worth- 
while. It was Professor William G. Farlow who called 
the first meeting at his home, the start of the NEBC. 

Farlow studied with Gray, graduated from Harvard 
College in 1866, and obtained an M.D. degree in 1870. For 
two years he worked at the Gray Harbarium, but in 1874 
he joined the staff of the Bussey Institution. He wished to 
specialize on algae and his report of 1879 on algae in 
Massachusetts water supplies is regarded as a classic. In 
1879 Farlow announced his intentions of leaving his library 
and herbarium to Harvard. In his will, probated after 


498 Rhodora [Vol. 75 


his death in 1919, Farlow had stipulated that his collections 
were to be removed from the flammable Agassiz Museum 
and properly housed within three years or the collections 
would be given to Yale. Three months before the time 
limit expired, the University chose the Divinity School 
Library building as the location. That building and the 
collections have since been known as the Farlow Herbarium 
and Library. Until her death some years later, Mrs. Farlow 
financed the operation of this collection. 


Charles Edward Faxon — 1846-1918 
(Rhodora 20: 117-122. 1918.) 


Faxon was one of a family of linguists but is best known 
as an artist. He taught himself to draw and was making 
saleable copies of Audubon’s work at the age of 15. Al- 
though he was trained as a civil engineer at the Lawrence 
Scientific School, he took a position as instructor in botany 
at the Bussey Institution in 1879, There D. C. Eaton 
asked him to prepare drawings of ferns. By 1882 his work 
had come to the attention of Charles Sargent who lured 
him to the Arnold Arboretum where he prepared seven 
hundred and forty-four plates for the Silva of North 
America. Between 1879 and 1913 Faxon published 1,925 
plates of his botanical art, an average of fifty-seven illus- 
trations a year for thirty-four years. Faxon also trans- 
lated botanical work for Sargent from Portuguese, Italian 
and Danish, several of which are published. His personal 
writings were few, a paper on birds in the Arnold Arbore- 
tum and a note on Aster. 


Edwin Faxon — 1823-1898 
(Rhodora 2: 107-111. 1900.) 

This Jamaica Plain bachelor was a leather merchant who 
retired at the age of fifty-eight and devoted much of his 
time to collecting plants. His early botanical forays were 
with C. Pringle in Vermont, and he devoted time to the 
area of Lake Champlain, Mount Washington and Mount 
Desert Island. In 1890 Faxon translated from the German 


1973] New England Botanical Club — Howard 499 


an important monograph of Sphagnum through a contact 
with D. C. Eaton. He continued this interest, assembling 
and distributing large exsiccatae of mosses. It is reported 
that Faxon was one of the most knowledgeable individuals 
of his time on the entire flora of New England. 


George Lincoln Goodale — 1839-1923 
(Rhodora 25: 117-120. 1923.) 


A graduate of Bowdoin College, Goodale received an 
M.D. degree from Harvard in 1863. He taught at Bowdoin, 
practiced medicine, farmed, wrote on animal breeding, and 
formed a company to make condensed milk. In 1868 he 
published a catalogue of the plants around Portland, Maine, 
and in 1872 joined the Harvard faculty upon Asa Gray's 
retirement to teach classes in botany. During this period 
he also wrote the physiological section of Gray's Botanical 
Textbook. In 1879 Goodale was appointed director of the 
Botanie Garden of Harvard, succeeding Sargent in that 
role, and Curator of the Botanical Museum. His text of 
"Wild Flowers of America" (1882) was supported by 
Isaac Sprague's drawings. In 1897 Goodale tired of having 
his Botanie Garden income support the Gray Herbarium 
and asked that they be separated. The Corporation ap- 
proved, and Goodale turned his attention to the develop- 
ment of the Botanical Museum closer to the University. 


George Golding Kennedy — 1841-1918 
(Rhodora 21: 25-38. 1919.) 

A graduate of Harvard in 1864, with an M.D. in 1867, 
Kennedy had been a student of Asa Gray and was influenced 
by him. Although a successful practicing physician, Ken- 
nedy found time to travel and collect plants over the years 
with Fernald, Churchill, Collins and Williams. The early 
gatherings of the NEBC members were held at his home 
between the regular meetings. His group became known 
as the "Faxon Club" for which no records seem to be 
available, Kennedy's personal herbarium contained 14,390 
sheets of flowering plants and a large collection of mosses. 


500 Rhodora (Vol. 75 


He was known for his special knowledge of the flora of 
Willoughby, Vermont which was published in Rhodora 
(1904). 

He served as a member of the Visiting Committee of the 
Gray Herbarium and in 1896 he generously supplied funds 
for the construction of the library wing. 


Nathaniel Thayer Kidder — 1860-1938. 
(Who Was Who in America 1897-1942 p. 673. 1942.) 


No obituary of Kidder appeared in Rhodora, even though 
he was the only member of the Club to serve two distinct 
terms as president, in 1896-9 and 1902-05, and the only 
president to serve for six years. A bachelor, he lived in 
Milton. His few botanical papers covered the flora of 
Milton and the Blue Hills, with another on the plants of 
the Isle au Haut. The Kidder wing of the Gray Herbarium 
was his gift in 1910 as a member of the Visiting Com- 
mittee. 


Edward Lothrop Rand — 1859-1924 
(Rhodora 27: 17-27. 1925.) 


Rand was graduated from Harvard College in 1881 and 
obtained a degree in law in 1884. He was successful in his 
profession. As a Harvard junior, he camped at Somes 
Sound on Mount Desert Island and was one of the youthful 
naturalists who formed the Champlain Society to prepare 
a natural history catalogue of the area. The first edition 
of his Flora of Mount Desert Island omitted the difficult 
grasses and sedges, but the fourth edition published in 
1894 with J. H. Redfield, a friend of Asa Gray, was com- 
plete. A bronze plaque on a large boulder on the Jordan 
Pond trail on Mount Desert commemorates Rand’s contri- 
butions to the knowledge of that flora. 

Rand served as corresponding secretary of the NEBC 
from 1895 to 1921 and was the mainstay of the publica- 
tions committee in soliciting funds and manuscripts for 
Rhodora. His personal herbarium, left to the NEBC in 
1915, contained over 15,000 sheets. 


1973 New England Botanical Club — Howard 501 


Benjamin Lincoln Robinson — 1864-1935 
(Nat. Acad. Sci. Biographical Mem. 17: 305-330. 
1937.) 


Although he was the editor of Rhodora for the first 
thirty volumes, president of the NEBC, the Curator and 
later Director of the Gray Herbarium, no obituary of 
Robinson ever appeared in Rhodora. 

Robinson began his college career at Williams College, 
but after three months he was dissatisfied and entered 
Harvard, graduating with the class of 1887. With a young 
bride, he went to Strasburg, Germany where he received 
his Ph.D. degree in plant anatomy in 1889. Between 1891 
and 1894, he taught scientific German at Harvard. On the 
death of Sereno Watson in 1892, Robinson was appointed 
Curator of the Gray Herbarium at the age of twenty-nine. 
When the Harvard Corporation approved Goodale’s request 
that the Botanic Garden be separated from the financially 
insolvent Gray Harbarium in 1897, Robinson was forced 
reluctantly to raise money. Mrs. Asa Gray donated $20,000 
in 1899 to establish a professorship honoring her late hus- 
band, and Robinson was the first occupant of that chair. 
With the help of a “friends” organization having a basic 
membership fee of $10 in 1897, Robinson was able in his 
career to increase the endowment of the Gray Herbarium 
from $18,000 to $526,000. Since the other Harvard bota- 
nists were located in the Agassiz Museum area, Robinson 
also requested permission from the University to move the 
work and the collections of the Gray Herbarium into “the 
city" to be adjacent to the cryptogamic collections and the 
Botanical Museum. Although President Eliot approved this 
proposal in 1900, the Friends of the Gray Herbarium and 
the members of its Visiting Committee were opposed and no 
funds were raised. By 1905 Robinson drew a pathetic 
description of the housing of Gray's collections. In his 
annual report he wrote that the building was partially 
fireproof, ill-lighted even in the daytime, poorly ventilated, 
with inadequate plumbing of an archaic and wholly un- 
satisfactory type, had no basement under much of the 


502 Rhodora [Vol. 75 


building so that the lower cases were so damp as to cause 
molding of the specimens, the roof was ill-constructed 
that more than forty years of repair had never made it 
tight for more than a few weeks at a time, and so a wholly 
new struture was needed. In 1909 the University approved 
the idea of expanding and renovating the existing build- 
ings if funds could be raised for the purpose. Robinson 
and Deane drew the plans. It is worth noting that over 
half of the members of the Visiting Committee at that 
time were members of the New England Botanic Club 
whose herbarium was then housed in these same quarters. 
They responded with gifts and supported additional fund 
raising. Kidder supplied the initial gift which built the 
addition known as the Kidder Wing. A message congratu- 
lating Kidder for this gift appears in the Club records. 
A member of the Harvard Corporation offered $2,000 for 
Asa Gray's house which was unoccupied, and this was 
moved across Garden Street to its present location. The 
gavel used by subsequent presidents of the NEBC was 
turned from a corner post of this building, George Ken- 
nedy next gave $25,000 for a library wing which developed 
the Gray Herbarium toward the Garden Street side. George 
White then donated $21,500 for the construction of a new 
laboratory to replace the Hunnewell Laboratory in the 
opposite direction. In addition, $10,000 was offered for 
equipment. Kennedy again donated funds, $10,000 this 
time, to add a new facade to the old building. As the last 
construction, three members of the Visiting Committee 
supplied funds to rebuild the center part of the building 
into three galleries and a basement, completing the build- 
ing as most of us knew it before 1940, Construction was 
finished and the building occupied in February of 1915. 
The minutes of the NEBC reveal that a meeting was held 
in the building, that the members were delighted with the 
new quarters for the Club herbarium, and that the entire 
structure was “brilliantly lighted throughout with elec- 
tricity”. 


1973] New England Botanical Club — Howard 503 


Robinson's research centered on the Eupatoriae of the 
Compositae, and he made a major contribution in a study 
of the flora of the Galápagos Islands. His editorship of 
Rhodora demonstrated that he was a perfectionist in his 
writing. He was also a versatile linguist with a knowledge 
of many languages. Dr. Lily M. Perry recalls his review- 
ing his use of Hungarian, anticipating visiting with Dr. 
C. Domin because he felt it would be nice to speak to 
Dr. Domin in his own language. He is remembered to this 
day as a kind and gentle man with a formal courtesy, a 
gentleman of Harvard. 


Charles Sprague Sargent — 1841-1927 
(Jour. Arnold Arb. 8: 69-86. 1927.) 

Sargent is best remembered as the first Director of the 
Arnold Arboretum. He served in that capacity for fifty- 
four years and built up a remarkable institution soundly 
based on a library, an herbarium, and a living collection. 
Although a charter member of the NEBC, he never served 
in any official capacity. Several of the summer outings of 
the Club were under his auspices, all with the same 
itinerary: to Forest Hills by railroad, thence through 
the Arnold Arboretum and around his home in Brookline 
in carriages which he supplied. 


Roland Thaxter — 1858-1932 
(Nat. Acad. Sci. Biographical Mem. 17: 55-68. 1936.) 
Thaxter was graduated from Harvard College in the 
class of 1882. His thesis had been on entomogenous fungi, 
and this proved to be his life work. He was appointed 
assistant professor of cryptogamic botany in 1891 and 
succeeded William G. Farlow as a professor in 1901. 
Thaxter served as president of the Club, was one of the 
frequent speakers for the monthly programs, and was one 
of the first to offer a botanical travel talk involving his 
work in the West Indies. 


504 Rhodora [Vol. 75 


Benjamin Marston Watson — 1848-1918 
(Records of the Harvard class of 1870, pp. 144-145.) 
After his graduation from Harvard, Watson operated 
a nursery in Plymouth, Massachusetts, started by his father. 
In 1877 he was appointed an instructor in horticulture at 
the Bussey Institution where he taught arboriculture, 
fruiticulture and later landscape design. Although a 
charter member of the NEBC, he never held office and was 
never mentioned in the club minutes as a speaker or à 
participant. 


Emile Francis Williams — 1858-1929 
(Rhodora 33: 1-18. 1931.) 

Williams spent his childhood in France and inherited a 
profitable import business from his father. His spare time 
was spent hiking in the White Mountains where, on one 
occasion, he met G. G. Kennedy collecting plants, When 
the NEBC was organized, Williams became the original 
secretary-treasurer, a role he continued to fill from 1896 
until the Club was incorporated in 1922. He served on 
the editorial committee of Rhodora and was largelv re- 
sponsible for raising money to meet its annual deficit. In 
the early years the Club published annual booklets con- 
taining the Constitution and a list of the officers and mem- 
bers of the Club, which he edited. His obituary contained 
a detailed account of his gracious hospitality and the fine 
table he set at the dinners for members before the formal 
meetings of the Club. Throughout his life he was an active 
collector, especially on Mount Katahdin, joining all the 
club excursions and traveling with Fernald in Maine and 
Canada. His herbarium came to the NEBC in 1916. 

No mention of the New England Botanical Club and its 
early history would be complete without comments on two 
individuals who made significant contributions to its de- 
velopment: Mary Anna Day and Merritt Lyndon Fernald. 


Mary Anna Day — 1852-1924 
(Rhodora 26: 41-47. 1924.) 
The New England Botanical Club was a gentlemen’s 


1973] New England Botanical Club — Howard 505 


organization, and Miss Day was not eligible for member- 
ship. She was a teacher and served as librarian in the 
Clinton, Massachusetts, public library. She sought a posi- 
tion at Harvard just as Miss Josephine Clark left the 
library of the Gray Herbarium for a position in Wash- 
ington. Miss Day was appointed in 1893, and her first 
task was checking all the five thousand references in Gray 
and Watson’s Synoptical Flora. When the New England 
Botanical Club was formed, Miss Day compiled lists of the 
existing local floras and of the known private and insti- 
tutional herbaria of New England which were published 
in the original volume of Rhodora. For years after that 
she prepared the Index for the annual volumes of Rhodora. 
Although not a botanist, Miss Day was interested in plants 
and collected many of the early exsiccatae of the Gray 
Herbarium. In 1903 she took over the task of preparing 
the Gray Herbarium index cards and was responsible for 
the issuance of over 170,000 cards. Hers was also the task 
of organizing the library of the Gray Herbarium in the 
construction period of 1909 to 1915. She is mentioned 
occasionally in the minutes of the Club as an aid or con- 
sultant on the acquisitions for the Club library. A tribute 
to her life and contributions was published in Rhodora. 


Merritt Lyndon Fernald — 1873-1950 
(Rhodora 53: 33-65. 1951.) 

Young Merritt Fernald came to the Gray Herbarium 
from his home in Maine in 1891 at the age of 18 and served 
as an assistant to Sereno Watson, curator of the Gray 
Herbarium until 1902. He was not a charter member of 
the New England Botanical Club but was among the first 
new members elected during the fourth meeting on March 4, 
1896. Fernald immediately became active in the Club, 
working hard to establish the herbarium and contributing 
his observations regularly to the Club meetings, even 
guiding the amateurs in the proper preparation of speci- 
mens and in the identification of plants. At the tenth 
annual meeting of the Club, December 1, 1905, Emile Wil- 
liams was asked to give “a little history of the Club, its 


506 Rhodora [Vol. 75 


origin and development." This was printed separately and 
is a memento in the Club records. Of the herbarium 
Williams wrote, “The herbarium of the Club is one of its 
best institutions. Dr. Robinson can tell you how useful it 
has become. We owe a large debt of gratitude to Mr. Fer- 
nald for his labors in this direction, as well as in every 
other. How could we get along without Mr, Fernald? The 
devoted Mr. Fernald, always ready to stand in the breach, 
to supply an extra number for Rhodora, to deliver an 
address at one of our meetings." 

Fernald was officially the ‘“Phanerogamic Curator" 
from 1899 to 1911 and served then in succession as Presi- 
dent (1911-1914), as Councillor (1914-1928), and as Editor 
of Rhodora from 1929 until his death in 1950. Short and 
paunchy, as I first knew him, Fernald remained bouncy 
and confident, a showman to the end. He livened the 
meetings of the New England Botanical Club through his 
contributions for over fifty-four years. 


The Meetings 


The goals of the original New England Botanical Club 
were many — to meet in a fraternity of gentlemen with 
a mutual interest in the plants of New England; to as- 
semble an herbarium representing plant specimens from 
each county in the New England states; to prepare dis- 
tribution lists of taxa to determine the ranges and the 
gaps of New England plants, and to publish a journal, 
Rhodora, for contributions on New England botany. 

Initially the Club accepted nominations from its charter 
members for additional resident and non-resident members 
to be elected by written and secret ballot with a corps of 
"serutineers" to count and report the ballots, a practice 
continued to this day. An induction fee was charged and 
dues were regularly assessed. 

The fraternity of the meetings was obvious in good food, 
and often wines, served by a caterer following the formal 
meeting. At the June and October meetings all of the 
members were called upon, in order, to report on the 


1973] New England Botanical Club — Howard 507 


specimens of interest which they had found. Informal 
meetings for the insertion of specimens into the herbarium 
were well attended. Special conferences were conducted 
in the early years by Professor Fernald and others when 
groups of plants such as grasses, Ranunculus, sedges, 
ferns, aquatic plants were discussed in considerable detail, 
alerting the members to the characteristics and informa- 
tion to be sought in living specimens. Field trips were 
regularly scheduled to remote places needing concentrated 
collecting. Schedules of transportation, hotel reservations, 
etc., were issued in advance, and field work was well 
organized in assigned areas or details. 

The NEBC continued its informal existence through two 
hundred twenty-seven meetings, until the tax laws of the 
Commonwealth became a concern and the club was officially 
incorporated in January of 1921. Since 1922 the secre- 
tary’s minutes have begun as they do today, “‘The four 
hundred and seventy-third meeting of the New England 
Botanical Club being the seven hundredth since its estab- 
lishment. .... ". Recording secretaries have been many, 
yet the quality of the reports through the years provides 
uniformly interesting reading of the activities of the Club; 
the election and comments of the members; the accounts 
of the principal speakers; the introduction of the first 
lantern slides or the first use of colored movies; the war- 
time trials and limitations such as cold meeting rooms to 
conserve coal or the substitution of cheese and crackers for 
luncheon plates in 1918; of deaths and touching tributes 
to members; of appeals for new members; or of the rising 
costs and budget deficits. 


The Herbarium 


The herbarium was begun with the original meeting 
and grew with regularity as members added specimens. 
Collections in private hands were sought out and acquired 
by the Club, and members usually bequeathed their own 
collections to the Club. A mounter was hired but club 
members inserted the specimens. Fernald once requested 


508 Rhodora [Vol. 75 


paid help for the herbarium, but the suggestion was re- 
ferred to the Council, which, according to the following 
records, never took any action, In 1926 the treasurer 
reported that the real cost of mounting and inserting a 
sheet in the herbarium was “well in excess of 5 cents" 
and therefore the investment in the herbarium was over 
$7,000. Today the Club herbarium houses 243,189 speci- 
mens. 

The Club herbarium was originally housed in the Agassiz 
Museum. There was a long debate on where the key should 
be kept, in Dr. Farlow's office or with the janitor, and 
this was finally settled by having it in Farlow's home. The 
herbarium was available to members on a regular basis, 
including weekly evening hours. In 1905 the herbarium 
was moved to the original Gray Herbarium on Garden 
Street and in 1915 to new quarters in the White wing of 
the renovated building. Finally in 1954 the herbarium 
was moved to its present quarters in the Harvard Uni- 
versity Herbaria building, 22 Divinity Avenue, Cambridge, 
where the monthly meetings are also held. 

The survey of distribution of New England plants pub- 
lished periodically in the pages of Rhodora was begun by 
naming a family of plants with the request that the indi- 
vidual members check their private herbaria and bring the 
distribution records to a stated meeting. In later years 
the NEBC, by additions to its own herbarium, depended 
upon the faithful work of members to compile such listings. 
How well remembered is the work of Richard Eaton, 
Stuart Harris, Ralph Bean, Frank Seymour and Donald 
White to complete the lists published in the last decade! 


Rhodora 


The charter members of the NEBC considered carefully 
the formation of an official publication. By circulation of 
an invitation to subscribe, an original mailing list of six 
hundred was obtained. An editorial announcement in 
Volume 1, Number 1, January 1, 1899, stated, "Its publi- 
cation has not been undertaken without mature considera- 


1973] New England Botanical Club — Howard 509 


tion, nor until through keen and helpful interest of New 
England botanists, a sufficient subscription list has been 
secured to assure its monetary support. The name Rhodora, 
although the designation of one of our most attractive New 
England plants, has been chosen, not from sentiment but 
primarily from a desire to have a distinctive and euphonious 
one-word title, experience having amply shown that similar 
titles (e.g, Linnaea, Grevillea, Helios, Erythrea, Auk, 
Ibis, etc.) soon become familiar, and possess great merit 
in their brevity and ease of citation." In fact, this decision 
and choice was reported amply in the records of the Club. 
B. L. Robinson was the spokesman for the one-word title, 
and a vote was taken on the choice of a name. Ballots were 
east for Rhodora, Oakesia, Watsonia, Bigelovia, Graia 
and Nova Anglia, in that order. 

The original subscription to Rhodora was one dollar a 
vear, with single copies priced at fifteen cents. Authors 
of more than one page were to receive twenty-five copies 
of the issues with reprints at cost, The early copies carried 
advertising. Members were invited to subscribe; the dues 
did not automatically include a subscription. 


From the beginning Rhodora was a deficit operation. 
The initial subscription list dropped rapidly at the time 
of renewal, for many expressed the opinion that the 
journal was too technical. When the treasurer reported 
that the cost of Rhodora was four times the receipts from 
subscriptions, an appeal was made, at first annually and 
later triennially, for special gifts to subsidize the publica- 
tion without the annual draft against the general funds of 
the Club. For a period manuscripts were rejected due to 
excessive maps or plates. As late as 1933 M.L. Fernald 
reported to the club, “Technical scientific journals do not 
pay their way. Intended for the permanent record of newly 
discovered truth, they must be issued at a price within 
reach of any who need access to them. Temporarily we 
are most fortunate in our support; but we must look to 
the future. The bequests already made by members of the 
Club who while with us took a keen interest in Rhodora 


510 Rhodora [Vol. 75 


are comfortable nest eggs which greatly relieve the worries 
of the Treasurer and of the Editorial Board. Provisions 
in our wills for their multiplication will insure the future 
of Rhodora; but, since botanists as a group are long-lived, 
it is suggested that earlier additions to the permanent 
endowment of Rhodora may bring prompter and greater 
returns in satisfaction to the investors." Eventually several 
endowed funds for the support of Rhodora were received 
by the Club. Rhodora was edited by B. L. Robinson for 
Volumes 1-30, M. L. Fernald for Volumes 31-52, R. C. 
Rollins for Volumes 53-63 and A. R. Hodgdon for Volumes 
64-15. The Club has been fortunate in having this small, 
dedicated group of voluntary and unpaid editors. 


The Archives 


The Club has a long history partly preserved in the 
minutes of the meetings and partly as archival material. 
Various people over the years contributed newspaper clip- 
pings, photographs, cartoons, poetry, letters or miscella- 
neous mementos. Recently the current Club librarian. 
M. Canoso, has attempted to organize the material on hand. 
A series of undated photographs of members appear to be 
associated with an appeal made by Mr. Bullard, as recorded 
in the minutes of October 4, 1934 for “all members who 
had not yet had their photographs taken for the club 
records to send to him a suitable likeness so as to complete 
this branch of the Historical Committee's work." Koda- 
chrome slides made of these early photographs accom- 
panied this presentation on the occasion of the seven hun- 
dredth meeting. 

One of the oldest of the “poems” long associated with 
the New England Botanical Club was attributed to Edward 
L. Rand in 1900 and is available in a much duplicated 
form. Although it has been read on occasions, the original 
was considered lost with the older records. Now it can be 
dated in the minutes, having been read first on March 2, 
1900. “Verses by Edward L. Rand inspired by the follow- 
ing statement of Mr. Fernald in Rhodora 11-39. ‘Mr. 


1973] New England Botanical Club — Howard 511 


Walter Deane informs me that in his youth he was familiar 
with Old Orchard Beach and that at that time this Arte- 
mesia (A. stelleriana) was not seen'." 


Me Judice 


In the youth of Walter Deane, in those glorious ancient 
days, 
Foreign plants had not crept in with insinuating ways. 
Every plant was then our own, from each rootlet tip so 
small 
And the old world floral tramps did not cut a dash at all; — 
All our plants were indigene 
In the youth of Walter Deane. 
In the youth of Walter Deane, gardens were not hard to 
weed ; 
Our plants were too polite to promiscuously seed, 
And profanity produce. Foreign weeds grew only then 
In ash barrels far remote, — rarities were they to men. 
Shepherd's-purse grew not, I ween, 
In the youth of Walter Deane. 
Little Walter on the wharves used to sit from day to day, 
Waiting for the ships to bring plants from lands so far 
away, 
Dandelions, buttercups, white weed, chickweed, — all were 
new, — 
With a thousand other things, well known plants to me 
and you. 
These, remember, were first seen 
Since the youth of Walter Deane. 
Long ago these days have fled. Walter to a man has grown 
All the floras of the world now contribute to our own. 
Yet confusion can't arise, all new comers one by one 
Have been noted by our friend since their inroad first 
begun ; 
So we know what’s indigene 
From the words of Walter Deane. 


The twentieth anniversary meeting of the Club was re- 


512 Rhodora [Vol. 75 


ported in the minutes of February 4, 1916. "Mr. Edward 
L. Rand, the recognized bard of the Club, an office not in 
the Club book but which so far no one has ventured to com- 
pete for against him, read the following verses which he 
too modestly calls a skit." 


As I was walking through a field 
The habitat was wet 

I chanced to see a winsome thing 
The modest Violet. 


She looked familiar. Once I knew 

Her given name — I think 

But now we know these plants seem named 
By influence of drink. 


Her eolor was what poets call 
"A bit of Heaven's own blue" 
But this is very, very wrong 
As Mathews shows to you. 


*Dear Flower," I said, "I love you well. 
Yourself pray introduce." 

She bowed her head and murmured low, 
“It would not be much use. 


"A man who lives up in Vermont 
‘Freakish’ calls me and ‘cross’ 
And says no one respectable 
Can guess what is my source. 


"A nameless nothing sure am I. 
Forget your love for me 

Until I find out who I am 

In plant society." 


I could not comfort her, alas, 

For specialistic ways 

Must hurt the feelings of the plants 
In these intensive days. 


1973] New England Botanical Club — Howard 513 


I had to leave her to her grief, 

For versed not am I 

In dead names that are brought to life 
And living names that die. 


No comfort lies in synonyms, — 
But how nice it would be 

If she could have the novum nom 
of V. brainerdii. 


This poem ribbing the very active members of the Club, 
Vermont specialist on Viola, Ezra Brainerd, and F. 
Schuyler Mathews, of “Wild Flower" fame was followed 
by a song “which will doutless become a classic at celebra- 
tions of the New England Botanical Club." It is entitled 
“Segregation Song", and needless to say it is the work of 
Edward L. Rand. This was sung by the whole club under 
the leadership of Harold St. John. Sung to the tune of 
"My Bonnie”, the first of six verses is: 


The deuce of a genus is Aster, intermediates come by the 
score 

"Destroy them at once", says the wise man, else there will 
be species galore. 


Chorus 
Segre —, Segre —, Segregation is a terrible bore 
Bring back, bring back, oh, bring back our species of yore. 


At this same meeting Mr. Rand also made the following 
announcement in verse: 


If anyone knows a man who would enjoy the Club 

And whom the members might enjoy, whose home is near 
the Hub, 

Throw off this girlish bashfulness, and send his name to 
me. 

Pll take it to the Council, so he may elected be 

(We really need a lot of names, for it would be a winner 

If we could make new members pay the bill for this great 
dinner.) 


514 Rhodora [Vol. 75 


The twentieth meeting concluded with another song with 
words by Rand and sung to the tune of John Brown's Body. 
The verses were a parody on the many talks given by M. L. 
Fernald, and two of them are worthy of quotation. 


In old times the Bryan Vikings voyaged far across the 
foam 

Just to find the genial grapejuice, as is stated in the poem. 

How that juice must have fermented — do you think they 
brought it home? 

Rhodora tells it all. 


Chorus 


Glory, glory to New England 

Glory, glory to New England 

Glory, glory to New England 
and our N.E.B.C, 


Now for all these great achievements which our Club may 
justly claim 

We must give great praise and credit, else we should be 
much to blame, 

To that man who led us onward, M. L. Fernald is his name. 

Rhodora tells it all. 


Although a “bard” has not appeared in the Club in recent 
years, there is an anonymous short poem often quoted at 
the Club even in his time, to the delight of its victim the 
Latin scholar and ex-college president, Arthur Stanley 
Pease. George R. Cooley recorded this in a letter to Richard 
Eaton in 1959. 


Here lies the body of Stanley Pease 
Buried beneath the venerable trees 
What lies here is only the pod 

Pease shelled out and went to God. 


Although the New England Botanical Club was originally 
a gentlemen's organization, there is a reference in the 
minutes of the meeting of May 7, 1915, to “our one lady 
member." Who this female member might be cannot be 


1973] New England Botanical Club — Howard 515 


determined by a check of the club books of the year. On 
the occasion of the fifieth anniversary of the Club in 1946 
greetings were received from the California Botanical 
Society which prompted the following poetic response from 
E. J. Palmer. 


We thank you for the good intent 
Of the kind sentiments you sent 
When this distinguished club of Men 
Had reached the age of two-score-ten. 


Back in the days when men were men 
And botanists were gentlemen, 

They loved their wives and sisters too — 
As we do still, as well as you. 


But there were times — the saints be thanked 
Our fathers felt were sacrosanct, 

And spots to which they might retreat 

That were forbid to female feet; 


Where they might gather and regale 
On wisdom spiced with salty tale, 

Secure and sure they wore the pants 
And not the wives and spinster aunts. 


We, sons of sires of those brave days, 
Still follow in their prudent ways; 
And so as was the founders’ plan, 
There are no sisters in our clan. 


So pardon us, if we should quote 

One slight faux pas in your kind note, 
When you address — by slip of pen — 
As a “sister club", this club of Men. 


1973 


However, times have changed, and in 1968, after much 
agonizing debate, the New England Botanical Club amended 
its by-laws to admit women. With good taste, recent presi- 
dents have not asked the lady members to serve on the 


516 Rhodora [Vol. 75 


refreshment committee or to pour tea. In fact, one of them 
is now the recording secretary and another the phanero- 
gamic curator. Meetings in 1973 close not with dinners, 
or the tub of ice cream so popular with the student members 
when I joined the club, but with beer of many varieties 
and ginger ale for those who so choose, accompanied by 
doughnuts and cheese and crackers. Rhodora is now a 
quarterly containing a larger number of extra-territorial 
papers. Council meetings precede the regular meetings 
since many of the officers travel distances to attend which 
would be beyond the comprehension of the founders of 
1895. The herbarium still needs volunteers to insert speci- 
mens or to check distribution records. The June and 
October meetings have been termed “show and tell" ses- 
sions but are still enjoyed by those attending. Above all, 
good fellowship prevails and a mutual interest in the flora 
of New England forecasts a long life for the New England 
Botanieal Club. 


ARNOLD ARBORETUM OF HARVARD UNIVERSITY 
JAMAICA PLAIN, MASS. 02130 


THE SYSTEMATIC VALUE OF TRICHOME 
COMPLEMENTS IN A NORTH AMERICAN GROUP 
OF VERNONIA (COMPOSITAE) 


W. Z. FAUST AND S. B. JONES, JR. 


Trichomes are among the most useful of all anatomical 
features for systematic comparisons of angiosperms. This 
is because of their variety, their wide occurrence, their 
ease of preparation for study, and sometimes their close 
correlations to variation patterns among the taxa (Carl- 
quist, 1961). The characteristic forms of trichomes are 
often of considerable value in recognition of genera 
(Heintzelman & Howard, 1948), and certain distinct types 
of trichomes may be used as criteria for distinguishing 
species (Cowan, 1950; Goodspeed, 1954). In Nicotiana, 
Goodspeed (1954) found that the trichome complement is 
of phyletic significance because the types of trichomes 
point to species origins and relationships. The trichome 
complements in Nicotiana are correlated with patterns of 
morphology, geographical distribution, and cytogenetics. 
Cowan (1950) demonstrated that trichomes are a reliable 
guide to relationships and classification in Rhododendron. 

Uniseriate trichomes as well as a variety of multicellular 
glandular and non-glandular trichomes are widely dis- 
tributed in Vernonia (Solereder, 1908). Hunter and Aus- 
tin (1967) demonstrated the hybrid origin of a Vernonia 
taxon by a comparison of the trichome complement of the 
two parental species and their hybrid progeny. They sug- 
gested that a study of the trichomes of other species of 
Vernonia might be of taxonomic significance. Urbatsch 
(1972), Faust (1972), and King (1971), working on sys- 
tematic problems among the species of Vernonia native 
to the United States, found evidence that trichomes are of 
systematic value in the genus. In 1923, Gleason erected a 
series of hypotheses on migration and evolution of Ver- 
nonia in North America. These hypotheses provide a basis 
of comparison for studies of trichome patterns and schemes 


517 


518 Rhodora [Vol. 75 


of phyletie groupings. More recently, Vernonia has been 
the subject of a series of biosystematic and biochemical in- 
vestigations by Jones and his students and by Abdel-Baset 
and Padolina in Dr. Tom Mabry's laboratory at the Uni- 
versity of Texas. These later studies have provided 
additional data and an opportunity to refine Gleason's 
hypotheses. 

Living material of 21 taxa of a closely related group of 
Vernonia, the Paniculatae verae, from the Sierra Madre 
Oriental of Mexico and from the eastern one-half of the 
United States were available in the greenhouse. This pro- 
vided an opportunity to determine whether or not trichome 
complements might provide information useful in the classi- 
fication of these Vernonia species. This evidence along 
with other data from biochemical systematies, transplant 
studies, hybridization experiments, and population analyses 
will eventually be combined into a taxonomic revision of 
the subsection Paniculatae verae. 


MATERIALS AND METHODS 


Mid-cauline leaves of each taxon collected from green- 
house grown transplants were fixed in FPA (formaldehyde, 
propionie acid, and alcohol) for anatomical comparisons. 
Whole mounts of leaves were cleared in 1% aqueous NaOH 
at 60°C and transferred to 75% lactic acid at 60°C until 
completely cleared. The leaves were then stained in safra- 
nin and fast green and mounted in Hoyer’s solution. Leaves 
from herbarium specimens were cleared in the same man- 
ner for additional observations. Leaf cross-sections were 
prepared by dehydrating in an ethyl alcohol-butanol series, 
embedding in paraffin, and sectioning at 12 ». The sections 
were stained with safranin and fast green. Fresh whole 
mounts of the leaves, 3 mm’, were examined by scanning 
electron microscopy. The whole mounts were glued onto 
the specimen stubs with a mixture of Elmer's Glue-All and 
silver conductive paint. Micrographs were made of the 
trichome types. Paired affinity indexes were constructed 
for the trichome complement of the lower leaf surfaces as 


519 


Vernonia — Faust & Jones 


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Figure 1. Trichomes on lower leaf surfaces of Vernonia: (A) 
SEM of V. missurica 100X, (B) SEM of V. marginata bilobed and 
awl-shaped types 100X, (C) SEM of V. marginata bilobed type 
500X, (D) cross-section of V. marginata bilobed type illustrating 
basal cells 500%, (E) whole mount of V. baldwinii ssp. interior 
T-shaped type 400X, (F) whole mount of V. baldwinii ssp. bald- 
winii L-shaped glandular type 400. 


1973] Vernonia — Faust & Jones 521 


TN 


Figure 2. Line drawings of populations of the trichome types of 
Vernonia: (A) bilobed, (B) awl-shaped, (C) awl-shaped glandular, 
(D) L-shaped, (E) L-shaped glandular, (F) T-shaped, (G) uni- 
seriate, (H) longhorn. Trichomes selected include typical and ex- 
treme forms. 


522 Rhodora [Vol. 75 


they usually are the most reliable guide to relationships 
(Cowan, 1950) : 
Paired affinity index — 
trichome types common to both species x 100 
total trichome types of both species I 


RESULTS AND DISCUSSION 


The trichome complements of 21 taxa of Vernonia are 
presented in Table 1 along with their distributions on the 
leaves, Photomicrographs and scanning electron micro- 
graphs of selected trichomes are shown in Figure 1. Popu- 
lations of the trichome types are illustrated by camera 
lucida drawings in Figure 2. Comparison of possible ar- 
rangements of the taxa, based upon trichome complements 
with those groupings suggested by Gleason (1906, 1922, 
1923) indicated that trichomes in Vernonia may indeed 
have some phyletic significance. Analysis of the trichome 
complement on both the upper and lower leaf surfaces sug- 
gested that certain species or groups of species stand apart 
from the other species (see Table 1). For example, V. 
larsenii and V. lindheimeri are closely related based upon 
gross morphology as well as upon trichome complement 
(King, 1971). They are the only taxa with the longhorn 
trichome type. Vernonia larsenii and V. lindheimeri also 
possess the same 15 flavonoid compounds and the same 
major sesquiterpene lactone, Glaucolide-F (Abdel-Baset and 
Padolina, per. com.). Vernonia flaccidifolia is the only 
species which does not have the bilobed trichome type. The 
taxon contains only one flavonoid compound and no sesqui- 
terpene lactones (Abdel-Baset and Padolina, per. com.). 
Therefore, as Gleason (1923) suggested, V. flaccidifolia 
may be more advanced and evolutionarily specialized than 
some of the other species of Vernonia. Vernonia fasciculata 
and V. marginata, placed in the Fasciculatae group by 
Gleason (1923), possess the same types of trichomes on 
both the upper and lower surfaces of leaves (Jones, 1972). 
Chemical evidence indicates a close relationship; V. mar- 
ginata and V. fasciculata have identical flavonoid com- 


523 


Vernonia — Faust & Jones 


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524 Rhodora [Vol. 75 


pounds (Abdel-Baset and Padolina, per. com.). Also, these 
two species along with V. arkansana contain Glaucolide-M 
as the major sesquiterpene lactone compound. 


Examination of the paired affinity indexes of the tri- 
chome complement of the lower leaf surfaces (Table 2) 
indicate several natural groupings. Vernonia schaffneri, 
V. greggii, V. arkansana, V. angustifolia, V. fasciculata, 
and V. marginata each have a paired affinity index of 100 
to each other. Two of these species, V. schaffneri and V. 
greggii of Mexico, were considered by Gleason (1923) to 
be from the most primitive stock based upon morphology 
and phytogeography. Vernonia schaffneri, V. greggii, 
V. ervendbergii, and V. texana contain the same major 
sesquiterpene lactone, Glaucolide-A, and many identical 
flavonoid compounds (Abdel-Baset and Padolina, per. 
com.). Vernonia gigantea and V. noveboracensis have 
a paired affinity index of 100. They also possess the same 
flavonoid compounds and the same major sesquiterpene 
lactone compound, Glaucolide-A (Abdel-Baset and Pado- 
lina, per. com.). Gleason suggested that these two taxa 
evolved from the same ancestral line. The two species are 
similar in vegetative features such as leaf size and shape 
and stem height. Vernonia noveboracensis is a plant of 
wet soil, whereas V. gigantea grows in well drained soil. 
There are also differences in the number of flowers per 
head and in the length of the bract tips. On the other 
hand, it is difficult to distinguish vegetatively V. gigantea 
and V. noveboracensis grown in the greenhouse or the 
transplant garden unless the flowers are present. 

Gleason (1923) proposed that the ancestral home of the 
genus Vernonia, as far as North American species are con- 
cerned, is tropical South America. He suggested that 
migration occurred from tropical South America through 
Central America and Mexico into Texas. From Texas 
migration accompanied by evolution proceeded in two di- 
rections, northward through the prairie region and east- 
ward along the coastal plain. Speculation concerning the 
evolution of the taxa utilizing morphological and ecological 


1973] Vernonia — Faust & Jones 525 


sisuə5SgioqəAou 


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Lines show distribution of the taxa by their 


Diagrammatic scheme of migration and evolution of 


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526 Rhodora [Vol. 75 


information and chemical data (Abdel-Baset and Padolina, 
per. com.) is presented in Figure 3. The western species 
of Vernonia have more flavonoids than do the eastern 
species of Vernonia. Generally, the western taxa contain 
from three to five compounds that are lacking in the eastern 
taxa. The eastern species have the same major sesqui- 
terpene lactone, Glaucolide-A, that occurs in the species 
of Vernonia from the Sierra Madre Oriental of Mexico. 


The use of the trichome complement as an aid to estab- 
lishing relationships in Vernonia does have its limitations. 
For example, V. blodgettii and V. missurica have a paired 
affinity index of 100. Yet based upon morphological char- 
acters and geographic criteria they are very different. 
Vernonia blodgettii is a low, sparsely pubescent plant 
endemic to southern Florida, whereas V. missurica is a 
tall, extremely pubescent, wide-ranging species of the 
Mississippi Valley region. Another problem is presented 
by V. baldwinii ssp. baldwinii and ssp. interior. Evidence 
from morphological studies suggests close relationships 
(Faust, 1972). Populations of V. baldwinii ssp. baldwinii 
are found throughout the Ozark Plateaus province and the 
Ouachita province. Vernonia baldwinii ssp. interior popu- 
lations oecur primarily on the prairies and pastures of the 
central plains. The ranges of these two taxa overlap along 
the edges of the Ozarks and Ouachitas and a broad transi- 
tion zone, or zone of intergradation is present (Faust, 
1972). The characteristies of the intergradation zone sur- 
rounding the Interior Highlands suggest that the genomes 
of V. baldwinii ssp. baldwinii and ssp. interior are highly 
compatible and that the volume of gene flow between the 
two taxa may be relatively high. Additional evidence from 
biochemical systematics (Abdel-Baset and Padolina, per. 
com.) indicates a close relationship ; the two subspecies 
contain the same major sesquiterpene lactone, Glaucolide-B. 
However, the trichome complement shows differences in 
the two subspecies. Vernonia baldwinii ssp. baldwinit has 
the L-shaped glandular trichome, whereas ssp. interior 
does not have this type. Also, uniseriate trichomes were 


1973] Vernonia — Faust & Jones 527 


not found on the lower leaf surfaces of V. baldwinii ssp. 
baldavinii. 

The evidence from this study suggests that trichomes 
can be diagnostic characteristics in Vernonia. However, 
their value as taxonomic criteria will be greatly increased 
if the information can be interpreted with reference to 
other lines of evidence such as that obtained from bio- 
chemical systematies, distributional studies, gross morphol- 
ogy, and hybridization experiments. All of the 21 taxa 
in this study form fully fertile F, hybrids but sometimes 
have hybrid breakdown in the F, and F, generations 
(Urbatsch, 1972; Faust, 1972; King, 1971; Jones, un- 
published). Thus, trichome complement differentiation in 
this group of closely related plants must in some way be 
related to natural selection and the habitat that they occupy. 


ACKNOWLEDGMENTS 


This study was supported by the University of Georgia 
and by N.S.F. grant GB-20687. We thank Dr. W. J. 
Humphreys for the use of the facilities of the University 
of Georgia Electron Microscope Laboratory. We are grate- 
ful to Dr. Tom Mabry of the University of Texas and two 
of his students, Mrs. Zeinab Abdel-Baset and Mr. Bill 
Padolina, for the use of some unpublished biochemical 
data. Dr. Lowell Urbatsch, Mr. Bruce King, and Mr. Cay- 
wood Chapman have also worked with us on our studies 
in Vernonia. 


LITERATURE CITED 


CARLQUIST, S. 1961. Comparative plant anatomy. Holt, Rinehart, 
and Winston, New York. 29-36. 

Cowan, J. M. 1950. The Rhododendron leaf: a study of epidermal 
appendages. Oxford and Boyd, London. 

Faust, W. Z. 1972. A biosystematie study of The Interiores species 
group of Vernonia (Compositae). Brittonia 24: 363-378. 
GLEASON, H. A. 1906. A revision of the North American Vernonieae. 

Bull N. Y. Botan. Gard. 4: 144-243. 
1922. Vernonieae. North American Flora 33: 52- 


95. 


528 Rhodora [Vol. 75 


1923. Evolution and geographical distribution of 
the genus Vernonia in North America. Amer. J. Bot. 10: 187- 
202. 

GOODSPEED, T. H. 1954. The genus Nicotiana. Chronica Botanica, 
Waltham, Mass. 109-131. 

HEINTZELMAN, C. E., JR., and R. A. HOWARD. 1948. The compara- 
tive morphology of the Icacinaceae. V. The pubescence and 
crystals. Amer. J. Bot. 35: 42-52. 

HUNTER, G. E., and D. F. AusTIN. 1967. Evidence from trichome 
morphology of interspecific hybridization in Vernonia ( Compos- 
itae. Brittonia 19: 38-41. 

Jones, S. B. 1972. A systematic study of the Fasciculatae group 
of Vernonia (Compositae). Brittonia 24: 28-45. 

KING, B. L. 1971. The systematics of the Vernonia lindheimeri 
complex. M. S. Thesis. Univ. Georgia, Athens. 

SoLEREDER, H. 1908. Systematic anatomy of the Dicotyledons. Vols. 
I & II (with addenda). Transl. by L. A. Boodle and F. E. Fritsch. 
Clarendon Press, Oxford. 

UnBATSCH, L. E. 1972. Systematic study of the Altissimae and 
Giganteae species groups of the genus Vernonia (Compositae). 
Brittonia 24: 229-238. 


DEPT. OF BIOLOGY 

COLUMBUS COLLEGE 

COLUMBUS, GEORGIA 31907 
and 

DEPT. OF BOTANY 

UNIVERSITY OF GEORGIA 

ATHENS, GEORGIA 30601 


THE MORPHOLOGY AND SYSTEMATICS 
OF THE AUDOUINELLA COMPLEX 
(ACROCHAETIACEAE, RHODOPHYTA) 
IN NORTHEASTERN UNITED STATES 


WM. J. WOELKERLING 


Our knowledge of the marine species of the Audovwinella 
complex along the New England and adjacent coasts of 
North America (including New York and New Jersey) 
comes mainly from the papers of Collins (1906, 1908, 
1911), Davis (1913), Farlow (1881), Harvey (1853), 
Hehre and Mathieson (1970), Jao (1936), and Woelkerling 
(1972). Although many of the earlier records (which 
leave a number of points to be clarified) are incorporated 
in the descriptive catalogues of Taylor (1937, 1957), a 
comprehensive morphotaxonomic study of New England 
marine audouinelloid algae has not appeared to date, and 
the only monographic attempt involving the New England 
species has been that of Collins (1906) which included all 
of North America and is now out of date. 

The present account reviews the systematics, morphology, 
ecology, and distribution of New England marine audoui- 
nelloid algae and provides a taxonomie treatment of all 
representatives based on extensive field and herbarium 
studies including the examination of many of the type 
collections. Canadian representatives of the complex (see 
South and Cardinal, 1970) have not been considered in 
detail because of inadequate material and the questionable 
nature of a number of records, A brief discussion on these 
taxa appears at the end of this report. 

The techniques employed in these studies have been de- 
tailed elsewhere (Woelkerling 1970, 1973). Abbreviations 
for herbaria follow Lanjouw and Stafleu (1964) ; collec- 
tions designated by WJW refer to specimens in the author's 
personal herbarium, currently housed at Wis. 

A comprehensive review of the taxonomic history of 
audouinelloid algae has appeared recently (Woelkerling 


529 


530 Rhodora [Vol. 75 


1971), and the systematic conclusions reached in that 
paper are supported and adopted here. D'Lacoste and 
Ganesan (1972), however, maintain a classification of 
genera based on chromoplast morphology and criticize the 
placement of Rhodochorton in synonomy with Audouinella 
on grounds that Woelkerling (op. cit.) did not examine 
chromoplasts in living material of Rhodochorton pur- 
pureum, the type species of Rhodochorton. Cells of living 
R. purpureum (referred here to Audouinella) have been 
examined during the present New England study and found 
to show the same variation in chromoplast morphology 
described by Drew (1928, pp. 156, 177, fig. 33). 

D’LaCoste and Ganesan (1972, p. 235) also point out 
(citing the findings of West, 1968, p. 98) that chromoplast 
shape can change as a result of fixation of material and that 
studies on the latter (including, presumably, those of Drew, 
1928) can yield misleadng results. They apparently over- 
looked the other comments of West (op. cit.) in the same 
paragraph where he indicates the “. .. . inadequacy of 
plastid form as a taxonomic criterion in certain in- 
stances .. .”. Thus the studies of West (1968), Woelkerling 
(1971), and others (see Woelkerling op. cit., p. 14) clearly 
show that chromoplast morphology is so variable in certain 
taxa that any generic segregation based on differences in 
plastid shape would result in a situation where certain 
species could be placed in several genera simultaneously. 
As a result, the present author has abandoned the use of 
chromoplast morphology as a generic criterion and has 
referred the genus Rhodochorton to the synonomy of Au- 
douinella. For further comments, see the discussion of 
A. purpurea below and Woelkerling 1971, p. 4. 

Taylor (1937, 1957) records 27 species in 5 genera from 
the northeastern coast of North America, all belonging to 
the family Acrochaetiaceae, (G. F. Papenfuss [personal 
communication] and P. C. Silva [ personal communication] 
have indicated that the family name Acrochaetiaceae can 
be used correctly for the Audouinella complex even though 
the genus Acrochaetium is considered congeneric with 


1973] Audouinella — Woelkerling 531 


Audouinella; therefore the family name Audouinellaceae 
[Woelkerling 1971, p. 7, 22] is superfluous.) 


Taylor's list may be enlarged to include the New England 
records of Giard (1890) and Hehre and Mathieson (1970) 
and the Canadian reports summarized by South and Cardi- 
nal (1970) to yield a total of 36 taxa for the region, 
apparently a greater number than in any other family of 
Rhodophyta. Of these, 29 are reported to occur in New 
England waters. 


This account, however, recognizes only 11 species of 
audouinelloid algae as definitely occurring in New England; 
7 of these are referred to the genus Awdouinella and 4 
to the genus Colaconema. Moreover, the names and(or) 
taxonomie status of all 29 taxa previously reported from 
New England have been changed; a summary appears in 
Table 1. 


MORPHOTAXONOMIC FEATURES 


The Acrochaetiaceae is generally circumscribed to in- 
clude those Florideophycidae with simple or branched mono- 
siphonous filaments usually less than 5 (rarely to 25) mm 
tall, with asexual reproduction by monospores, bispores, 
tetraspores, and/or multipartite spores, with carpogonia 
sessile on cells of vegetative filaments, intercalary in vege- 
tative filaments or terminating one to several celled stalks, 
with gonimoblasts, when present, forming directly from 
the fertilized carpogonium, and without a pericarp. 
Audouinella Bory is the type genus. 

Following Woelkerling (1971), the presence (Awudowi- 
nella) or absence (Colaconema) of sexual reproduction is 
considered to provide a useful separation of the New Eng- 
land taxa into genera, with Colaconema regarded as a 
form genus similar to those of the Fungi Imperfecti. Taxa 
referrable to Kylinia Rosenvinge (in its original sense) 
and Liagorophila apparently do not occur in New England, 
and the systematic status of these genera remains in doubt 
(see Woelkerling 1971, p. 6). 


532 Rhodora [Vol. 75 


Within the genera Audouinella and Colaconema, species 
have been grouped into three sections based on the absence 
of, the presence of one, or the presence of more than one 
pyrenoid per chromoplast. Species possessing more than 
one pyrenoid per chromoplast have not been collected in 
New England waters to date. 

Additional features which appear to be of value in de- 
fining species limits include eell and spore dimensions, and 
in some eases chromoplast morphology, prostrate system 
morphology, and spore arrangement. Certain other char- 
acteristics (e.g. height of erect filaments, degree of branch- 
ing, habitat preferences) sometimes are useful features 
for purposes of taxon recognition and in keys, but are too 
variable to be of value in defining species limits. 

Considerable confusion has hitherto surrounded the 
identification of species in New England waters, and a 
number of records based on incorrect determinations have 
crept into and subsequently have been perpetuated in the 
literature. Thus, for example, collections originally identi- 
fied by Collins (1906, 1911) as Acrochaetium (= Chan- 
transia) flexuosum Vickers and subsequently recorded by 
Taylor (1937, 1957) have been re-examined during this 
study and found to belong unmistakably to Colaconema 
secundata. Likewise collections hitherto referred to Acro- 
chaetium sagraeanum involve at least three different 
audouinelloid taxa, and the type specimen itself is here ex- 
cluded from the Rhodophyta (see “Species Excludendae"). 

To help minimize confusion of a similar nature in the 
future, the distinguishing features of each taxon known 
to oecur in New England waters have been outlined at the 
onset of the discussion section of each species in the taxo- 
nomic accounts. Available distributional and ecological 
data are also included. While each species recognized here 
possesses a distinctive set of morphological features, most 
taxa exhibit considerable variation, and numerous plants 
of a collection should, therefore, be examined to build a 
composite picture of the population before identification 
is attempted, Collections of single specimens whose status 
is in doubt are best left unidentified. 


1973] Audouinella — Woelkerling 533 


KEY TO SPECIES OF NEW ENGLAND AUDOUINELLOID ALGAE 


This key includes only species definitely known to occur 


in New England waters. Records of questionable occur- 
rence, of taxa whose systematic status is uncertain, etc. are 
discussed in a separate section at the end of this report. 
Characters relating to chromoplasts and pyrenoids often 
are not preserved in dried or spirit material, and in these 
cases supplementary criteria are given in the key. 


Le 


Multicellular prostrate system present; filamentous, 
parenchymatous, or pseudoparenchymatous. ...... 2 
Multicellular prostrate system absent; plants attached 
to substrate by a single cell which may rarely give rise 
to one or several accessory cells, ................ 7. 
2. Cells with a single chromoplast containing a sin- 
gle pyrenoid; plants generally (but not invariably) 
epiphytic or endophytic. ................565 € 2. 

2. Cells with one to a number of chromoplasts with- 
out pyrenoids; plants generally (but not invari- 
ably) saxicolous or endozoic. ..  .......... 10. 
Erect filaments generally over 500 „m long and usually 
exceeding the prostrate filaments in length; mono- 
sporangia commonly over 15 um long. ............ 4. 
Erect filaments generally under 250 um long or absent, 
usually shorter than or equal to the prostrate filaments ; 
monosporangia under 15 am long. .............. 11. 
4. Chromoplasts distinctly stellate; prostrate system 
developing from an orbicular, parenchymatous 
group of cells which forms a small dise that may 
later proliferate and become obscured; lateral 
branches commonly but not always virgate or se- 
cundate sic. 11. Colaconema secundata. 

4. Chromoplasts parietal lobate curved plates; pros- 
trate system filamentous or pseudoparenchyma- 
tous, not developing from a distinctive paren- 
chymatous disc; lateral branches not virgate. . 5. 


5. Monosporangia borne, at least in part, in clusters of 


3 or more on branched stalks. 4. Awdowinella daviesii. 


534 


11. 


11. 


Rhodora [Vol. 7 


Monosporangia borne singly or in pairs but not in 
Clusters. ..................................... 6. 
6. Prostrate system composed of an enlarged, central, 
more or less panduriform to pyriform cell and 
aecessory cells or filaments which arise from it. 
Ce eee eee ee eee eee 3. Audouinella dasyae. 
6. Prostrate system filamentous or pseudoparenchy- 
matous, without an enlarged, central panduriform 


or pyriform cell. ........ 6. Audouinella saviana. 
Cells commonly over 20 jm long; plants commonly over 
500 um tall .................................. 8. 
Cells under 20 ¿im long; plants rarely over 100 „m 
tall... eee Re 9. 
8. Basal cell elongate, panduriform to pyriform; 

usually bearing accessory cells or filaments. .... 


........................ 3. Audouinella dasyae. 

8. Basal cell globose or subglobose; lacking accessory 
cells or filaments. ........ 2. Audouinella alariae. 
Cells commonly over 10 „m long; filament(s) of erect 
system procumbent; protoplast of basal cell hemi- 
spherical and distinctly flattened on side in contact 
with substrate................. 7. Audouinella unifila. 
Cells generally 10 „m or less long; filament (s) of erect 
system upright to arcuate; protoplast of basal cell 
more or less globose. .... 5. Audouinella microscopica. 
10. Sporangia (i.e. carpotetrasporangia) borne most- 
ly in clusters on branched stalk-like gonimoblast 
filaments; plants commonly but not invariably 
saxicolous. ........... 1. Audowinella purpurea. 

10. Sporangia (i.e. tetrasporangia) solitary, sessile 
or on unbranched stalks; plants commonly but 
not invariably endozoic. ...................... 


Cells commonly over 2 diameters long; prostrate fila- 
ments widely creeping, commonly over 1 mm long. 
DENDUM 10. Colaconema minima. 
Cells rarely over 2 diameters long; prostrate filaments 
rarely over 500um long. ...... 9. Colaconema humilis. 


1973] Audouinella — Woelkerling 535 


AUDOUINELLA Bory 


Audcuinella Bory 1823: 340 (as Auduinella). Woelkerling 
1971: 22. Acrochaetiwm Naegeli 1861: 402. Balbiania 
Sirodot 1876: 149. Chromastrum Papenfuss 1945: 320. 
Grania Kylin 1944: 26. Rhodochorton Naegeli 1861: 
355. Thamnidium Thuret In Le Jolis 1863: 110. Trente- 
pohlia Pringsheim 1862: 29. 


Note: Species now referrable to Audowinella have also 
been placed in the past in Byssus, Callithammion, Cera- 
mium, Chantransia, Conferva, Kylinia, and Trentepohlia 
Martius (1817). In addition, some species hitherto placed 
in Acrochaetium, Audouinella, Callithamnion, Cermium, 
Chantransia, Chromastrum, Kylinia, Rhodochorton, and 
Trentepohlia are referable here to the form genus Cola- 
conema. 

Plants epibiotic, endobiotic, or saxicolous; attached to 
or suspended in the substrate by a single basal cell, by a 
prostrate system of simple or branched filaments which 
may or may not become pseudoparenchymatous, or by a 
parenchymatous disc. Erect filaments simple or branched, 
up to 25 mm long; cells containing one to many variously 
shaped chromoplasts with or without pyrenoids. 

Asexual reproduction by monosporangia, bisporangia, 
tetrasporangia, and/or multipartite sporangia; sporangia 
sessile or stalked and borne on erect or prostrate filaments. 

Plants monoecious or dioecious. Spermatangia in clusters 
or oceasionlly single or in pairs, terminal or lateral on 
simple or branched stalks or sessile on ordinary vegetative 
cells. Carpogonia intercalary or more commonly sessile or 
terminating 1-2 celled stalks on vegetative cells, solitary 
cr rarely in groups of 2-3; remaining undivided or dividing 
transversely after fertilization and giving rise directly to 
gonimoblast filaments bearing carposporangia or carpo- 
tetrasporangia. 

Type Species: Audouinella hermanni (Roth) Duby. 


Section I 


Species in this section are not known to contain pyre- 


536 Rhodora [Vol. 75 


noids in their chromoplasts. Within each section species 
are discussed in alphabetical order by specific epithet. Cell 
and spore dimensions are based only on the New England 
populations studied and may not reflect the total range 
of variation found in populations elsewhere. The ratio of 
greatest cell length to greatest cell diameter (width) is 
denoted by L/D in all species accounts. 


1. Audouinella purpurea (Lightfoot) comb. nov. Figs. 
52-55. 

Byssus purpurea Lightfoot 1777: 1000. 

Callithamnion purpureum (Lightfoot) Harvey 1841: 
116, 1849: 183. 

Conferva purpurea (Lightfoot) Dillwyn 1806: 56, pl. 
43. C. Agardh 1817: XXIX. 

Rhodochorton purpureum (Lightfoot) Rosenvinge 1900: 
75. Collins, Holden, and Setchell 1895: 49. Conway and 
Knaggs 1966: 195 et. seq., Figs. 1-3. DeToni 1903: 1510. 
Edelstein and McLachlan 1966a: 1041, 1052. Edelstein 
et al. 1970: 626. Hamel 1927: 57, 108; 1928: 201; 1928a: 
151. Hehre and Mathieson 1970: 207. Knaggs 1965: 
499 et seq., 1966b: 521 et seq., pl. 102-107; 1967: 139 et 
seq.; 1967a: 549 et seq., pl. 134-139; 1968: 449 et seq.. 
pl. 170-174. Mathieson et al. 1969: 132. Papenfuss 1945: 
327. South 1970: 1. South and Cardinal 1970: 2079. Stone 
et al. 1970: 325. Taylor 1957: 226, pl. 45, Figs. 1-2. West 
1967: 11; 1969: 12 et seq., Figs. 1-22; 1970: 368 et seq., 
Figs. 1-8. 

Trentepohlia purpurea. (Lightfoot) C. Agardh 1824: 36. 
Harvey 1833: 382. Harvey and MacKay 18536: 218. 

Rhodochorton islandicum Rosenvinge 1900: 75, Figs. 
1-4. Papenfuss 1945: 327. 

Rhodochorton intermedium (Kjellman) Kjellman 1883: 
184, pl. 15, Fig 8. DeToni 1903: 1509. Hamel 1927: 107; 
1928: 200. 

Thamnidium intermedium Kjellman 1875: 28, Fig. 10. 

Rhodochorton parasiticum Batters 1896: 389. Collins 
1900: 51; 1900a: 12; 1911: 281. Collins, Holden and 
Setchell 1901: 848. 


1973] Audouinella — Woelkerling 537 


Callithamnion rothii (Turton) Lyngbye 1819: 129, pl. 
4la. J. Agardh 1851: 17. Bailey 1847: 85. Collins 1880: 
162. Eaton 1873: 348. Farlow 1875: 376; 1876: 704; 
1879: 169; 1881: 121. Harvey 1846: pl. 120B; 1853: 243. 
Hay 1887: 66. Hay and MacKay 1888: 173. Jelliffe 1899: 
15. Jordan 1874a: 488. Klugh 1917: 83. Kuetzing 1861: 
XI, pl. 621. Olney 1871: 8; 1872: 132. Pike 1886: 110. 


Rhodochorton rothii (Turton) Naegeli 1861: 356, pl. 1, 
Figs. 1, 3. Bell and McFarlane 1933: 270. Boergesen 1902: 
390, Figs. 61-64. Collins 1894: 230; 1900: 51; 1905: 234; 
1911: 280; 1914:4. Davis 1913: 818. DeToni 1903: 1507. 
Drew 1928: 177. Gibson 1891: 201, Fig. 38. Hamel 1927: 
54, 106, Fig. 38; 1928: 199; 1928a: 147, Fig. 38. Hylander 
1928: 169. Kjellman 1883: 185. Kuckuck 1897: 345, Fig. 5. 
Kylin 1944: 28, Fig. 25. Nakamura 1941: 282, Figs. 8-9. 
Rosenvinge 1923-24: 390, Figs. 328-330. Taylor 1937: 239, 
pl. 45, Figs. 1-2. Whelden 1947: 119. 

Thamnidium rothii (Turton) Thuret In LeJolis 1863: 
111, pl. 5. 


Rhodochorton tenue Kylin 1925: 44, Figs, 25 b-e. Drew 
1928: 177, pl. 40, Fig. 33. Papenfuss 1945: 328. West 
1969: 12 et seq., Figs. 1-22. 

Conferva violacea Roth 1797: 190, pl. 4, Fig. 1. Non 
Conferva violacea Hudson 1778: 592, nec Ceramium viola- 
ceum Roth 1797: 150, nee Conferva rothii Turton 1806: 
1809. 


Plants saxicolous or occasionally epibiotic, caespitose or 
forming conspicuous matted expanses with erect filaments 
up to 25 mm tall; original spore non-persistent, Prostrate 
system composed of branched, creeping filaments which 
may be loosely entangled or more compactly pseudoparen- 
chymatous, but not forming distinct discoid holdfasts. 
Erect filaments moderately to freely and_ irregularly 
branched and usually tapering towards the tips. Cells 
cylindrical, 10-20 um wide and 15-75 m long, L/D (1-) 2.5; 
tapering to 7-12 um wide and 40-175 m long (L/D up to 
15) near and at the tips of branches; each cell containing 


538 Rhodora [Vol. 75 


one to a number of irregularly lobate to reticulate to dis- 
coid chromoplasts without pyrenoids. 

Monosporangia, bisporangia, parasporangia, and sex 
organs not observed. Carposporphytes consisting of short 
(up to 10 cells long), simple or usually branched gonimo- 
blast filaments bearing terminal or occasionally lateral, 
ovoid, solitary or paired carpotetrasporangia 15-25 „m wide 
and 25-40 um long. Other reproductive structures not 
observed, 


Type Locality: Ruined Abbey, Island of Iona, Scotland. 

Type: Possibly destroyed (see Conway and Knaggs 
1966). 

Distribution: Nearly cosmopolitan in temperate and 
polar marine waters; also known from terrestrial and 
freshwater sites. 

Hosts: Most common on rocks but also on algae and 
invertebrates. 


Specimens examined: 
CONNECTICUT: Bridgeport (Harbor) 4. IX. 1893, Johnson (NY); 8. HI. 
1891, Holden (NY). 
MAINE: Appledore Island (Isles of Shoals), 28. VII. 1938, Croasdal« 
(NHA). Haley's Cove (Smuttynose Is., Isles of Shoals), 16. VI. 1966, 
Shipman (NHA). Harpswell, 6. VII. 1905, Collins (NY). Inner Mark 
Island, 29. VIII. 1903, Collins (NY). Lubec (Bailey's Mistake) 22 V. 
1966, Femino (NHA). Lubec (West Quoddy Head), 21. V. 1966, Ball 
and Lavise (NHA). Peaks Island, VII. 1881, Collins (NY); 22. V. 
1904, Collins (NY). Penobscot Bay, prior to 1853, Hooper (TCD). 
Popham, 7. VIII. 1900, Chernington (NY). South Harpswell, 17. VII. 
1902, Collins (NY); 8. VII. 1903, Collins (NY). York (Nubble Light), 
19. V. 1966, Pollock (NHA). 
MASSACHUSETTS: Magnolia, 24. VI. 1896, Collins (NY). Nahant, 18. 
IV. 1879, Collins (?) (NY); 9. VI. 1889, Collins (NY) ; 30. III. 1890. 
Collins, (NY) ; 22. II. 1891, Collins (NY, — Collins, Holden, and Setch- 
ell 1895, No. 49); 30. V. 1900, Collins (NY, — Collins, Holden, and 
Setehell 1901, No. 848) ; 13. IV. 1907, Collins (NY). Penikese Island, 
9. VII. 1926, Taylor (NY). Sandwich (jetty), 7. IV. 1968, Kastelowitz 
(NHA); 22. I. 1971, Woelkerling (wJw 3297). Scusset Beach, 7. IV. 
1968, Bosworth (NHA); 29. III. 1969, Mills (NHA). 
NEW HAMPSHIRE: Hampton, VIII. 1884, Collins (NY). North Wallis 
Sands, 16. VII. 1967, Hehre (NHA). Rye Ledge, 18. VIII. 1966, Con- 
way and Shipman (NHA); 15. X. 1966, Hehre (NHA); 23. XII. 1966, 


1973] Audouinella — Woelkerling 539 


Conway and Hehre (NHA); 10. I. 1967, Conway and Hehre (NHA); 
26. II. 1967, Conway and Hehre (NHA); 25. IV. 1967, Hehre (NHA); 
25. V. 1967, Hehre (NHA); 22. VI. 1967, Hehre (NHA); 24. VII. 
1967, Hehre (NHA); 12. VIII. 1967, Mathieson (NHA), 15. III. 1969, 
Hutchinson (NHA). 

NEW JERSEY: Bay Ridge, 17. III. 1893, Collins (NY). 

NEW YORK: Fort Hamilton (Long Island), 16. V. 1866, Pike (NY). 
RHODE ISLAND: Newport, prior to 1853, Bailey (TCD). 


Audowinella purpurea is distinguished from other New 
England audouinelloid algae in having a diphasic life 
cycle involving the production of carpotetraspores (Fig. 52) 
and in having erect filaments commonly over 10 mm long 
with cells containing 1 to a number of irregularly shaped 
to discoid chromoplasts without pyrenoids (Figs. 53-55). 
It is the only known New England species of audouinelloid 
algae which commonly develops on rocks and forms mat- 
like expanses. 

A. purpurea grows throughout the year in New England 
and has been collected as far south as New Jersey. Most 
commonly it occurs in shaded habitats in the midlittoral, 
especially underneath Ascophyllum or Fucus, but it has 
been found throughout the intertidal and in the sublittoral 
zones. In sublittoral communities it apparently grows 
primarily as an epiphyte on kelp stipes. This species is 
widely distributed throughout the cooler regions of the 
northern hemisphere, and because it sometimes becomes an 
ecological dominant in the intertidal zone, it has been re- 
ported more frequently from northern regions than any 
other audouinelloid species. 

Its vegetative morphology can vary considerably, de- 
pending upon the particular ecological niche it occupies, 
and this morphological variation has been the subject of 
intensive studies (Conway and Knaggs 1966; Knaggs 
1965, 1966, 1966a, 1967, 1967a, 1968). In general, 
plants growing under stressed conditions tend to have 
shorter and less branched erect filaments. Knaggs (1965, 
1966, 1966a) and Conway and Knaggs (1966) have re- 
corded this species from freshwater and terrestrial habi- 
tats as well as intertidal and sublittoral habitats and have 


540 Rhodora [Vol. 75 


recognized a number of distinct taxonomic forms. These 
forms, however, appear to have little taxonomic value in 
so far as New England populations are concerned and 
consequently are not recognized here as distinct systematic 
entities. 

Reproductive structures have been found only in the 
winter and early spring; sterile plants, however, occur 
throughout the year. The latter bear some superficial re- 
semblance to Spermothamnion but can easily be distin- 
guished from that genus by the possession of a true hetero- 
trichous habit. Sex organs have not been found in New 
England populations and to the author's knowledge have 
never been found in field collected samples. West (1969, 
1970) has given an excellent account of these and other 
stages in the life history based on culture studies; Knaggs 
(1968) also reported sexual stages in cultural material. 
Mature carposporphytes are somewhat more loosely organ- 
ized in comparison with other New England species and 
often greatly resemble ordinary vegetative branches bear- 
ing sporangia. Only carpotetrasporangia have been seen 
during this study, but bisporangia and parasporangia are 
reported from elsewhere (Conway and Knaggs, 1966; 
Knaggs, 1967). 

Audouinella purpurea was the first described member of 
the Audouinella complex (Lightfoot, 1777), and because 
of its distinctiveness as a species (primarily in terms of 
habit and chromoplast morphology), it traditionally has 
been referred (as the type species) to a separate genus 
Rhodochorton Naegeli (1861, p. 355). While recognizing 
its distinctness as a species, no reliable characteristics of 
generic significance (see Woelkerling 1971, p. 4) are ap- 
parent at present; consequently Rhodochorton is consid- 
ered congeneric with Awdowinella, and the type species, 
R. purpureum, is also referred to Audouinella. 

Following Papenfuss (1945) and West (1969), the taxa 
originally described as Rhodochorton parasticum Batters 
(1869), R. rothii (Turton) Naegeli (1861), R. tenue Kylin 
(1925), and Thamnidium intermedium Kjellman (1875) 


1973] Audouinella — Woelkerling 541 


are here considered conspecific with Audouinella purpurea. 
In agreement with Conway and Knaggs (1966), Rhodo- 
chorton islandicum Rosenvinge is also regarded as con- 
specific with Audouinella purpurea. 


Section II 


Species in this section have chromoplasts with only one 
pyrenoid each. 


2. Audouinella alariae (Jonsson) comb. nov. Figs. 1-9. 

Acrochaetium alariae (Jonsson) Bornet 1904: XIX. 
Collins 1906: 192. Croasdale 1941: 214. Erskine 1955: 
151. Hamel 1927: 84; 1928: 177. South 1970: 1. South 
and Cardinal 1970: 2079. Taylor 1937: 229. 

Chantransia alariae Jonsson 1901: 132, Fig. 1. Adams 
1904: 351. Boergesen 1902: 356. Boergesen and Jonsson 
1905: Appendix, p. III. Collins 1911: 276, Cotton 1912: 
98, 133. DeToni 1924: 45. Levring 1937: 86, Fig. 12. 

Chromastrum alariae (Jonsson) Papenfuss 1945: 320. 

Kylinia alariae (Johnsson) Kylin 1944: 13. Hehre and 
Mathieson 1970: 206. Mathieson et al. 1969: 132. Papen- 
fuss 1947: 436. Taylor 1957: 213. 

Acrochaetium rhipidandra (Rosenvinge) Hamel 1927: 
25, 82, Fig. 23; 1928: 176; 1928a: 119, Fig. 23. 

Chantransia rhipidandra Rosenvinge 1909: 91, Figs. 19- 
20. DeToni 1924: 37. Kylin 1928: 5, Figs. 1-2. Levring 
1937: 83. 

Chromastrum | rhipidandra (Rosenvinge)  Papenfuss 
1945: 322. 

Kylinia  rhipidandra (Rosenvinge) Kylin 1944: 13. 
Papenfuss 1947: 437. 

Rhodochorton rhipidandra (Rosenvinge) Drew 1928: 
151. 

Chantransia secundata auct. non. (Lyngbye) Thuret: 
Collins, Holden, and Setchell 1896a: 236. Collins 1900: 
49 (pro parte). 

Chantransia virgatula auct. non. (Harvey) Thuret: 
Collins 1894: 233 (Holden collection). 


542 Rhodora [Vol. 75 


Plants epiphytic, up to 1.0 mm tall; original spore per- 
sisting as a globose to subglobose unicellular base up to 
25 um in diameter and usually larger than other cells. Basal 
cell bearing 1-2 (-3) erect filaments which are almost 
always branched; laterals irregularly arranged, few in 
number and nearly simple to numerous and frequently 
further subdivided; laterals generally tapering towards 
the tips. Cells cylindrical or occasionally somewhat barrel- 
shaped near the base, 12-18 „m wide and 15-60 „m long 
near the base, L/D (1-) 1.5-4; (4-) 7-10 (-15) „m wide and 
15-65 »m long in the laterals, L/D 2-6 (-9) ; tapering to 
4-9 um wide near the tips; each cell containing a single 
chromoplast and one pyrenoid. Unicellular hairs not ob- 
served. 

Monosporangia ovoid, 8-12 „m wide and 12-20 „m long, 
solitary or in pairs (occasionally in groups of 3), sessile 
or stalked, scattered to secundate along lateral branches 
or occasionally opposite or terminal on the branchlets. 

Other reproductive structures not observed in New Eng- 
land collections. 


Type Locality: The Maelstrom, Hvammsfjordur, Ice- 
land. 

Type: c (Jonsson 597). 

Distribution: Northern Massachusetts northward; Den- 
mark, Faeroes Islands, France, Great Britain, Iceland, 


Norway. 
Hosts: Alaria in New England; Alaria and Porphyra 
in Europe. 


Specimens examined : 

MAINE: Bald Head Cliffs (Ogunquit), 17. VIII. 1966, Shipman and 
Conway (NHA); 7. XI. 1967, Stone and Hehre (NHA). Boon Island, 
26. VII. 1938, Croasdale (NHA). Burnt Island (Cushing), 17. VII. 
1888, Collins (Ny). Cushing, no date, Collins (Ny, Collins 446A). 
Casco Bay, VII. 1939, Weatherhill (FH). Hardhead Island (Penob- 
scot Bay), VII. 1894, Collins (Ny, loose duplicate of P.B.A. No. 236; 
FH, P.B.A. No. 236; WIS, P.B.A. No. 236). Mt. Desert Island, 12. VIII. 
1890, Holden (NY, Holden 134). Nubble Light, 19. VIII. 1969, Here 
(wJw 2275). Pemaquid Point, 18. VII. 1901, Collins (NY). South 
Harpswell, 8. VII. 1903, Collins (NY). 


1973] Audouinella — Woelkerling 543 


MASSACHUSETTS: Pigeon Cove (Cape Ann), 25. IX. 1966, Hehre 
(NHA). 

NEW HAMPSHIRE: Hampton, VII. 1894, Collins (NY); VIII. 1894, 
Collins (NY). Star Island (Isles of Shoals), 22. VII. 1966, Conway 
and Shipman (NHA). 

CANADA: Ferryland, Avalon Penn. Newfoundland, 4. XII. 1968, 
South (CANA 7583). Ketch Harbor, Nova Scotia, 7. II. 1966, Edel- 
stein (2) (Edelstein 2239, Herb. Nat. Res. Council, Halifax). Mul- 
holland’s Bend, Campebello Island, New Brunswick, 15. VI. 1967, 
Hehre and Stone (NHA). 

DENMARK: Frederikshavn, Kattegat, 25. VIII. 1891, Rosenvinge (C, 
type of Chantransia rhipidandra) . 

ICELAND: The Maelstrom, Hvammsfjórdur, 28. VI. 1897, Jonsson (C, 
Jonsson 597, type of Audouinella alariae). 


Audouinella alariae is distinguished from other New 
England audouinelloid algae by the presence of a unicellu- 
lar, more or less globose base (Figs. 1-3) bearing erect 
filaments whose cells are commonly over 2 diameters and 
20 um long. It has been collected in New England from 
July through November on Alaria esculenta (L.) Greville, 
but also has been collected along the Canadian coast in 
February, June, and December, Further investigation may 
show that Audouinella alariae not only develops at other 
times of the year, but also grows on other substrates. 

New England populations of A. alariae exhibit consider- 
able morphological variation. Some mature plants reach 
heights of 1 mm while others never exceed 500 „m. A few 
plants consisting of a single, nearly simple erect filament 
have been observed (Fig. 7), but in most cases lateral 
branches develop. The laterals themselves may be little 
further branched, long, and bear a few, scattered sporangia 
(Figs. 8-9), or they may be more densely subdivided, rela- 
tively short, and bear numerous, rather crowded sporangia 
(Figs. 4-5). All intermediate situations can occur within 
a single population. 

Although pyrenoids have been observed during this in- 
vestigation, the chromoplast shape could not be definitely 
determined. Jonsson (1901, p. 133) and Levring (1937, 
p. 87) both report a stellate chromoplast, but further study 
is needed to determine to what extent, if any, the plastid 


544 Rhodora [Vol. 75 


shape might vary in this species. Collins (1906, p. 192) 
and Taylor (1937, p. 229; 1957, p. 213) report unicellular 
hairs, but such hairs have not been observed during this 
study and have not been recorded by other authors. 

Collins (1906), Jonsson (1901), and Taylor (1937, 1957) 
report the monosporangia to be sessile and opposite; Adams 
(1904) records mostly alternate sporangia; and Levring 
(1937) states that the sporangia are often scattered. Plants 
in New England populations show a combination of all 
such arrangements on the same individual (Figs. 4, 5, 7-9), 
with oppositely arranged sporangia occurring only occa- 
sionally. They may be either sessile or on 1-2 celled stalks 
and sometimes terminate short lateral branches. 

Critical comparisons (Table 2) of type collection ma- 
terial of Audouinella alariae and Chantransia rhipidandra 
Rosenvinge strongly indicate that the two taxa are con- 
specific. Rosenvinge (1909, p. 91) stated that C. rhipi- 
dandra differed from Audouinella alariae in *. . . having 
much thicker and more branched filaments, and further 
by the branches being often opposite . . ."; he also noted 
the presence of sex organs in the former and their absence 
in the latter. 

However, plants examined during this study, including 
the types of both taxa, indicate considerable overlap in 
filament diameters (Table 2), and the degree of branching 
and branch arrangement (Figs. 4, 5, 7-9) vary consider- 
ably. Thus these criteria do not appear reliable for sepa- 
rating the two taxa. The presence or absence of sex organs 
alone also appears unreliable as a criterion of species 
separation (Woelkerling 1971), and since there appear to 
be no other criteria by which the two taxa can be reliably 
separated, they are regarded here as conspecific, 

Hamel (1927, 1928), Kylin (1928), and Rosenvinge 
(1909) have described and illustrated sexual and carpo- 
sporophyte stages of this species (all using the specific 
epithet “rhipidandra’’). 

Chantransia unilateralis Kjellman (1906, p. 11, Taf. II, 
Figs. 1-4) is almost certainly conspecific with Audowiunella 


1973] Audouinella — Woelkerling 545 


alariae, but the type collection of Kjellman’s species ap- 
parently is not at UPS (personal communication), and final 
judgement must, therefore, be deferred until the type can 
be located and examined. 

The relationships of A. alariae to at least 15 other taxa 
of similiar morphology await clarification. Once critical 
examinations of all the types and other populations of these 
taxa are completed, it appears likely that a number will 
prove to be conspecific with A. alariae. 

Specimens referred to Chantransia secundata by Collins 
(1900) and Collins et al. (1896a) and to Chantransia 
virgatula by Collins (1894) have been examined and found 
to contain only plants of Audowinella alariae. In the case 
of the former specimens, this confirms the opinions of 
Jonsson (1901) and Collins (1906). 

Young plants of Audouinella dasyae bear many resem- 
blances to A. alariae but usually can be recognized by the 
presence of a panduriform or pyriform basal cell rather 
than a globose one. 


3. Audouinella dasyae (Collins) comb. nov. Figs. 10-31. 

Acrochaetiwm dasyae Collins 1906: 191. Aziz 1967: 408. 
Davis 1913: 813. Edelstein et al. 1967: 195, Fig. 9. Hamel 
1927: 77, 95, Figs. 47b-g; 1928: 171, 189, Figs. 47b-g. 
Papenfuss 1945: 308. Taylor 1937: 231; 1957: 217. 

Chantransia dasyae (Collins) Collins 1911a: 186. DeToni 
1924: 40. 

Acrochaetium intermedium Jao 1936: 242, pl. 11, Figs. 
1-4, Papenfuss 1945: 314. Taylor 1937: 231, pl. 33, Figs. 
1-4; 1957: 218, pl. 33, Figs. 1-4. 

Acrochaetium subseriatum Jao 1936: 243, pl. 11, Figs. 
5-7 (Non Acrochaetium subseriatum Boergesen 1932: 118, 
Figs. 8-1). Taylor 1937: 232, pl. 8383, Figs 5 T 

Acrochaetium zosterae Papenfuss 1945: 307. Taylor 
1957: 218, pl. 33, Figs. 5-7. 

Callithamnion virgatulum auct non Harvey: Harvey 
1853:5243; 


Plants epiphytic, caespitose, up to 3 mm tall. Prostrate 


546 Rhodora [Vol. 75 


system usually consisting of an enlarged, central, more 
or less panduriform to pyriform cell — the original spore 
— bearing several smaller accessory cells or short fila- 
ments; central cells becoming obscured by accessory fila- 
ments in robust plants or rarely remaining unicellular in 
young or small plants. Erect filament(s) arising from the 
central cell, moderately to freely and irregularly branched; 
sometimes tapering towards the tips. Cells cylindrical, 
25-70 (-90) ¿m long and 7-12 (-16) „m wide [L/D 2-6 
(-10)] in main axes, 15-60 „m long and 6-10 ¿m wide 
(L/D 2-7) in the laterals; each containing a single parietal 
chromoplast with one pyrenoid. Unicellular hairs not 
observed. 

Monosporangia ovoid, 16-24 um long and 9-12 (-16) „um 
wide, solitary or occasionally in pairs, sessile or occasion- 
ally stalked, in secundate series along the laterals or more 
scattered. Bisporangia and tetrasporangia not observed. 

Spermatangia globose to ovoid, up to 3um wide and 5 „m 
long, borne terminally or laterally on vegetative cells or 
unicellular stalks, or usually in clusters of varying size on 
multicellular simple or branched stalks. Carpogonia ses- 
sile or stalked, scattered over the lateral filaments; fer- 
tilized carpogonium dividing transversely and eventually 
giving rise to a branched gonimoblast bearing terminal 
carposporangia 16-24 „m wide and 9-12 „m long. 


Type locality: Woods Hole, Massachusetts. 

Type: FH (accompanied by a card in Collins script and 
numbered 5370). Isotypes have been distributed as No. 
1342 in Phycotheca Boreali Americana (Collins, et al. 
1906). 

Distribution: Atlantic Coast of North America. 

Hosts: Dasya (Rhodophyta) and Zostera (Angio- 
spermae). 

Specimens examined : 

MASSACHUSETTS: Edgartown (Martha's Vineyard), 19. XII. 1969, 
Woelkerling (waw 2245). Great Rip (1.25 km E. of Great Point, 
Nantucket Is.), 14. IV. 1970, Woelkerling (wyw 2500). North East- 
ham, 9. VIII. 1959, Lamb (FH, filed under the host, Dasya pedicel- 


1973] Audouinella — Woelkerling 547 


lata). Old Silver Beach, 31. X. 1970, Woelkerling (wJw 2853); 15. 
XI. 1970, Woelkerling (waw 2856); 30. XII. 1970, Woelkerling (WJW 
2977). Waquoit Bay (Falmouth), 16. X. 1969, Conway (wJw 2293); 
27. IV. 1970, Woelkerling (wyw 2534). West Falmouth Harbor, 6. 
I. 1970, Woelkerling (wyw 2284); 18. VII. 1970, Woelkerling (wJw 
2751). West Yarmouth, 17. IX. 1969, Woelkerling (wyw 2252); 3. X. 
1969, Woelkerling (wyw 1852), (waw 2255); 16. XI. 1969, Woelker- 
ling (ww 2249), (wyw 2250); 12. XII. 1969, Woelkerling (WJW 
2260). Woods Hole, 12. VIII. 1894, Holden (FH, filed under the host, 
Dasya pedicellata) ; 2. IX. 1905, Collins (FH, type) ; 29. VIII. 1944, 
Taylor (rH, filed under the host, Dasya pedicellata) ; 4. IT. 1970, 
Woelkerling (wyw 2330); 5. VII. 1970, Wilce (wJw 2690); 17. VII. 
1970, Woelkerling (wyw 2716); 13. X. 1970, Woelkerling (wJw 
2820); 29. I. 1971, Woelkerling (wyw 3298); 16. II. 1971, Woelker- 
ling (wyw 3311); 25. VIII. 1933, Jao (MicH, Woods Hole No. 278, 
type of Acrochaetium zosterae Papenfuss). Pine Island (Woods 
Hole), 7. VIII. 1934, Jao [MiCH, Woods Hole No. 277, type of Acro- 
chaetium intermedium Jao, non Thamnidium intermedium. Kjellman 
= Audouinella purpurea (Lightf.) comb. nov.]. 

NEW JERSEY: Great Bay (.4 km SSW of Wells Is.), 17. VII. 1963, 
Moeller (RUT, filed under the host, Dasya pedicellata). 

NEW YORK: Hell Gate (Long Island), 29. IX. 1850, Walters (TCD). 
CANADA: Malpeque Bay (Prince Edward Island), 5. VIII. 1966, 
Edelstein (Nat. Res. Coun. Herb., Halifax). 


Audouinella dasyae is distinguished from other New 
England audouinelloid algae by the presence of an enlarged, 
central, more or less panduriform to pyriform cell in the 
prostrate system (Figs. 12-17) which bears accessory pros- 
trate cells or filaments and which gives rise to the erect 
filaments. A. dasyae appears to be confined to the sub- 
littoral and has been collected in New England all months 
except March, May, and June. It undoubtedly occurs 
throughout the year. Sexual plants have been found from 
July through November; tetrasporangial plants have not 
been observed but probably occur and are to be expected 
in deeper waters in late spring and early summer. 

Like many audouinelloid algae, A. dasyae exhibits con- 
siderable variation [more than indicated by Collins (1906) 
or Taylor (1957)] in cell width, height, spore size, and 
other features. On Dasya for example, it is not uncommon 
to find mature plants under 500 ¿m tall anchored to the 
monosiphonous filaments of the host and plants up to 


548 Rhodora [Vol. 75 


3 mm attached to the more robust multilayered primary 
axes of the the host. Likewise branching of A. dasyae 
may be dense or relatively sparse (Figs. 20, 21), depending 
upon the “growing room" available. In all cases the en- 
larged central cell representing the original spore was 
panduriform or pyriform; globose cells described by Taylor 
(1957) and illustrated by Hamel (1927, Fig. 47b; 1928, 
Fig. 47b) have not been observed except in sporelings 
(Figs. 10-11). In addition, the range of cell and spore 
dimensions recorded during this study are considerably 
greater than those reported by Collins (1906) or Taylor 
(1937, 1957). 

Sexual plants have been reported before only by Aziz 
(1967). Spermatangial clusters vary considerably in size 
both within and between populations. In some cases the 
cluster is densely branched and bears numerous sperma- 
tangia, while in others it remains small, and in some cases 
spermatangia are even borne directly on vegetative cells 
or on unicellular stalks (Figs. 29-31). 

It appears that the trichogynes of unfertilized carpo- 
gonia continue to elongate until a spermatium lands on 
the tips (Fig. 22). Only several fertilized carpogonia with 
clear transverse divisions have been observed (Fig. 25), 
and further study is needed to determine whether gonimo- 
blast cells are cut off without such a division first occurring. 

Critical comparisons of the type and other collections 
of Audouinella dasyae, Acrochaetium | intermedium Jao 
(1936, p. 242, pl. 11, Figs. 1-4), and Acrochaetium zosterae 
Papenfuss (1945, p. 307 — A. subseriatum Jao 1936, p. 243, 
pl. 11, Figs. 5-7, non A. subseriatum Boergesen 1932, p. 118) 
indicates that all three taxa are conspecific (Table 3). 

Jao (1936, p. 243) distinguished A. intermedium from 
Audouinella dasyae on the bases of an elongate rather than 
globose central cell, the presence of endophytic prostrate 
filaments, longer cells, more extensive branching, and the 
presence of bisporangia. However, all collections of A. 
dasyae examined, including the type, have panduriform 
or pyriform central cells at maturity, may or may not 


1973] Audouinella — Woelkerling 549 


produce accessory prostrate filaments, have cells of similar 
dimensions (Table 3), and contain plants with varying 
degrees of branching. Bisporangia have not been observed 
in the type collection material of A. intermedium examined 
during this study; however, the presence or absence of 
bisporangia (possibly incompletely divided tetrasporangia) 
alone does not warrant specific distinction (Woelkerling 
1971). Since no other reliable distinguishing criteria 
could be found, the two taxa are considered conspecific. 
Aziz (1967) previously proposed that the two entities be- 
longed to the same species. 

Acrochaetium zosterae reportedly (data from Jao 1936, 
p. 244) differs from Awdowinella dasyae in the shape of 
the central cell and in the larger size of monosporangia. 
Central cell shape of the two taxa is essentially the same 
(see above), and an examination of type material of 
Acrochaetium zosterae indicates that most monospores are 
18-24 um long and 8-11 um wide rather than 22-31 um long 
and 6.5-9.5 „m wide as reported by Jao (1936). The first 
set of dimensions falls within the range found for other New 
England populations of Audouinella dasyae. Since differ- 
ences in host (Dasya vs. Zostera) are not taxonomically 
reliable (Woelkerling 1971) and since no other reliable 
criteria could be found to distinguish the two taxa, they 
are also considered conspecific. 

Acrochaetium opetigenum Boergesen (1915, p. 38, Figs. 
35-37), A. robustum Boergesen (1915, p. 40, Figs. 38-40), 
and A. unipes Boergesen (1915, p. 35, Figs. 31-35) are 
very similar to Awdowinella dasyae, and a comparative 
examination of the type collections will probably show that 
all are conspecific. Boergesen (1915) separated all of 
these taxa on slight differences in prostrate system struc- 
ture, but none appears to be taxonomically reliable (see 
Woelkerling 1971). 

The relationships between A. dasyae and Chantransia 
macountt Collins (1913, p. 113) require further clarifica- 
tion. Collins (1913) states in the original description that 
the cell representing the original spore does not remain 


550 Rhodora [Vol. 75 


distinct; Drew (1928, p. 184, pl. 43, Figs. 47-52, pl. 44, 
Fig. 53), however, describes and illustrates a distinct cen- 
tral cell which she says becomes obscured by other prostrate 
filaments. Both Drew’s material and the type collection 
of Collins need to be re-examined and compared with 
Audouinella dasyae to determine whether any or all are 
conspecific. 

Collins (1906, p. 192) suspected that specimens of Harvey 
(1853, p. 243) collected at Hell Gate, Long Island, New 
York in 1850 and referred to Callithamnion virgatulum 
Harvey were in reality plants of Audouinella | dasyae. 
Harvey's original material from TCD has been examined 
and indeed contains only plants of A. dasyae, as suspected 
by Collins. 

As noted previously, young plants of A. dasyae can under 
some circumstances become confused with specimens of 
A. alariae. The two can usually be distinguished on dif- 
ferences in basal cell shape. 


4. Audouinella daviesii (Dillwyn) Woelkerling 1971: 28, 
Figs. 7, 22; 1973: 81. Figs. 32-43. 

Acrochaetium daviesii (Dillwyn) Naegeli 1861: 405, 
Figs. 26-7. Baardseth 1941: 42, Fig. 18. Collins 1906: 
194 (2); 1908: 134. Davis 1913: 813. Doty 1948: 263. 
Edelstein et al. 1970: 634. Hamel 1927: 39, 98, Fig. 31; 
1928: 192; 1928a: 133, Fig. 31. South and Cardinal 1970: 
2079. Taylor 1937: 234, pl. 31., Figs. 8-10; 1957: 221, 
pl. 31, Figs. 8-10. 

Callithamnion daviesii (Dillwyn) Lyngbye 1819: 129 (only 
as to binomial). J. Agardh 1851: 11; 1876: 8. Jordan 
1874: 197; 1874a: 488. 

Ceramium daviesii (Dillwyn) C. Agardh 1817: XXVII. 

Chantransia daviesii (Dillwyn) Thuret In LeJolis 1863: 
106. Collins 1880: 162 (?) ; 1894: 233 (?) ; 1900: 49 (7) ; 
1911: 276 (2). DeToni 1897: 67; 1924: 55. Farlow 1875: 
376: 1876: 705. Kylin 1907: 117, Fig. 27. Rosenvinge 
1909: 104, Fig. 34. 

Conferva daviesii Dillwyn 1809: 73, Suppl. pl. F. 


1973] Audouinella — Woelkerling 551 


Rhodochorton daviesii (Dillwyn) Drew 1928: 172. Naka- 
mura 1944: 106, Fig. 5. 

Trentepohlia daviesii (Dillwyn) Areschoug 1847: 338. 
Farlow 1881: 109. Martindale 1889: 100. 

Acrochaetium amphiroae (Drew) Papenfuss 1945: 312. 
Doty 1948: 263. Edelstein and McLachlan 1968: 993, Figs. 
49-50 (?). Mathieson et al. 1969: 131. South 1970: 1. 
South and Cardinal 1970: 2079. Taylor 1957: 223. 

Rhodochorton amphiroae Drew 1928: 179, pl. 40, Figs. 
34-37. 

Acrochaetium alcyonidti Jao 1936: 245, pl. 12, Figs. 2-4. 
Papenfuss 1945: 312. Taylor 1937: 234, pl. 34, Figs. 2-4; 
1957: 200, pl. 34, Figs. 2-4. 

Acrochaetium alcyonidii Jao var. cylindricum Jao 1936: 
245, pl. 12, Fig. 5. Taylor 1937: 234, pl. 34, Fig. 5; 1957: 
221, pl. 34, Fig. 5. 

Acrochaetium sagraeanum auct. non. (Montagne) Bor- 
net: Boergesen 1915: 35 (pro parte). Bornet 1904: XXI 
(pro parte). Collins 1906: 192 (pro parte), Hamel 192": 
77, 99 (pro parte) ; 1928: 173, 191 (pro parte). Jao 1936: 
244 (pro parte). Papenfuss 1945: 311 (pro parte). Taylor 
1960: 309 (pro parte). Vickers 1905: 60. 

Chantransia sagraeana auct. non. (Montagne) DeToni: 
DeToni 1924: 51 (pro parte). 

Chantransia corymbifera auct. non. Thuret In LeJolis 
(pro parte; see Papenfuss 1945, p. 313 under Acrochaetium 
bornetü): Collins 1896: 5. Collins, Holden, and Setchell 
1896: 192. 

Chantransia efflorescens var thuretii auct. non. Bornet: 
Collins 1906: 196. Davis 1913: 813. 

Acrochaetium  thuretii auct. non. (Bornet) Collins et 
Hervey: Taylor 1937 : 236; 1957: 222. 

Note: Additional synonomy is presented by Woelkerling 
(1971). 


Plants partly to entirely epiphytic or epizoic, caespitose, 
up to 6 mm tall; original spore non-persistent. Prostrate 
system consisting of branched epi- or endophytic or epi- 
or endozoic filaments forming a pseudoparenchymatous 


552 Rhodora [Vol. 75 


disc or an entangled funiform mass. Erect filaments 
moderately to freely and irregularly branched, sometimes 
tapering towards the tips or attenuate and ending in multi- 
cellular hair-like prolongations. Cells cylindrical, (6-) 
9-12 (-20) um wide and (8-) 15-50 (-70) um long, L/D 
(1-) 2-4 (-6) in main axes and laterals; about 4 um wide 
and up to 75 um long in hair-like prolongations ; each cell 
containing a single parietal lobate chromoplast with one 
pyrenoid. 

Monosporangia ovoid, 7-13 »m wide and (8-) 12-20 um 
long, in clusters of 3 or more on branched stalks or singly 
or in pairs on 1-2 celled stalks, situated on the lowermost 
cells of laterals or sometimes more scattered. Tetraspor- 
angia ovoid, 16-22 „m wide and 24-36 „m long, borne in 
pairs on unicellular stalks or occasionally solitary or in 
groups of 3, situated on the lowermost cells of laterals 
or more scattered. 

Spermatangia ovoid to spherical, up to 4 „m wide and 
5 um long, borne terminally or laterally in small clusters 
on branched stalks. Carpogonia terminal on unicellular 
stalks; immediate post-fertilization stages not observed. 
Mature carposporophyte consisting of branched gonimo- 
blast filaments bearing terminal, ovoid carposporangia 9-18 
um wide and 18-26 „m long. 


Type Locality: Bantry Bay, Ireland (Hutchens) ; lo- 
cality for H. Davies collection not given by Dillwyn 
(1809). 

Type: NMW. 

Distribution: Nearly cosmopolitan, 

Hosts: A wide variety of algae, and invertebrates. 


Specimens examined : 

MAINE: Eastport, 1873, Averill (?) (NY). 

MASSACHUSETTS: Gay Head (Martha’s Vineyard), VIII. 1875, Far- 
low (FH); (NY). Gloucester, IX. 1878, Farlow (FH). Marblehead, 
16. IX. 1888, Collins (NY); 27. VIII. 1895, Collins (NY). Nantucket 
Center (Nantucket Is.), 14. IV. 1970, Woelkerling (wJw 2577). Old 
Silver Beach, 31. X. 1970, Woelkerling (wyw 2855); 30. XII. 1970, 
Woelkerling (wsw 2978), (wJw 2976). Salisbury Beach, 31. V. 1909, 
Collins (?) (NY). Waquoit (Harbor Entrance), 27. IV. 1970, Woelk- 


1978] Audouinella — Woelkerling 553 


erling (WJW 2536). West Falmouth Harbor, 3. X. 1970, Woelkerling 
(wJw 2769). West Yarmouth, 16. XI. 1969, Woelkerling (waw 2248). 
Woods Hole (Butlers Point), 15-20. VII. 1895, Nott (rH [P.B.A. 192], 
NY, WIS [P.B.A. 192]). Woods Hole (Nobska Point), 4. II. 1970. 
Woelkerling (wyw 2332), (wyw 2322); 2. VII. 1970, Woelkerling 
(wJjw 2678); 17. VII. 1970, Woelkerling (wJw 2719); 29. I. 1971, 
Woelkerling (wyw 3302); 12. II. 1971, Fiore (wyw 3320); 16. II. 
1971, Woelkerling (wyw 3314). Woods Hole (Pine Island), 9. VII. 
1938, Taylor (NY). Woods Hole (Sheep Pen Harbor), 1. IX. 1934, 
Jao (MICH, apparently a portion of type collection of Acrochaetium, 
alcyonidii Jao). 

NEW YORK: Montauk Point (Long Island), 20. I. 1970, Woelkerling 
(wJw 2198). 

CANADA: Paddy’s Head (Nova Scotia), 13. X. 1965, Edelstein (Herb 
Nat. Res. Council, Halifax, 2110). 

IRELAND: Bantry Bay, prior to 1809, Hutchins (NMW, Dillwyn col- 
lection, type). 


Audouinella daviesii is distinguished from other New 
England audouinelloid algae by the arrangement (at least 
in part) of monosporangia in clusters of 3 or more on 
branched stalks which are usually situated on the lower- 
most cells of lateral branches (Figs. 32-35). On some plants 
all monosporangia are borne in such clusters while on 
others, notably sexual and tetrasporangial plants, clustered 
monosporangia may not be very evident or numerous. 

A. daviesii appears to be mainly a sublittoral plant al- 
though several populations have been found on Fucus in 
the lowermost littoral. It has been collected in New Eng- 
land in all months except March and June, but almost 
certainly occurs throughout the year. Sexual plants have 
been collected in July, August, and September (wJw 2678, 
FH, NY) ; tetrasporangial plants have appeared in January 
(wJW 2198). 

The vegetative appearance of Audouinella daviesii varies 
considerably. When growing on Codium, for example, the 
prostrate system becomes an endophytic funiform mass of 
filaments lodged between the utricles of the host. When 
growing on Chondria, in contrast, the prostrate system 
forms an epiphytie pseudoparenchymatous dise, Similar 
variation has been found in Australian populations of 
A. daviesii (Woelkerling 1971). Intergrades between these 


554 Rhodora [Vol. 75 


forms also occur. The ratio of cell length to cell diameter 
also shows considerable variation. In some populations, most 
plants have cells with an L/D over 3 (Fig. 32) while in 
other populations, the ratio of cell length to cell diameter 
is almost always less than 3 (Fig. 33). Intergrades always 
occur, and sometimes transitions from short cells to longer 
cella can be found along single filaments (Fig. 34). 

Other vegetative features which vary include the degree 
of branching and the occurrence of multicellular hair-like 
prolongations. In some plants the degree of branching is 
only moderate; laterals arise at infrequent intervals and 
are little further divided. In other plants, however, branch- 
ing is abundant and laterals often arise in groups in close 
proximity to one another in a more or less fasciculate 
manner. Hair-like prolongations occur in some popula- 
tions but not in others. Often these prolongations were 
broken off giving the lateral branches a stubby appearance. 

Several sexual and one tetrasporangial collection have 
been made in New England waters to date. Unfortunately, 
immediate post-fertlization stages have still not been ob- 
served (see Woelkerling 1971) and it is not known whether 
the fertilized carpogonium divides transversely, longitudi- 
nally, or gives rise directly to gonimoblast filaments. 
Carposporangia in the New England populations tend to 
be larger than those found in Australia (Woelkerling 
1971). 

Acrochaetium alcyonidii Jao (1936, p. 245, pl. 12, Figs. 
2-4) and A. aleyonidii var. cylindricum Jao (1936, p. 245, 
pl 12, Fig. 5) are here considered conspecific with 
Audouinella daviesii. Two slides labelled *aleyonidii" by 
Jao in MICH, which are presumably from the original 
collection but do not bear the number Woods Hole 279 
(see Jao 1936, p. 245), have been examined and found to 
contain only the basal portions of several plants without 
any monosporangia or other reproductive structures. Thus 
the material is unidentifiable. 

The published accounts and illustrations of Jao (1936) 
and Taylor (1937, 1957) leave little doubt about the con- 


1973] Audouinella — Woelkerling 555 


specificity of Acrochaetium aleyonidii and Audouinella 
daviesii. Jao (1936) separated the former from the latter 
*. .. in having a strongly marked endozoic habit and very 
short lateral branches." However, host differences, habit, 
and length of lateral branches appear to be of little taxo- 
nomie significance (Woelkerling 1971). Moreover several 
New England populations (Taylor, Pine Island Coll. in 
NY; WJW 3320) growing on Alcyonidium (the host of 
Acrochaetium alcyonidii) have been examined and found 
to agree in all essential respects with Audouinella daviesii. 
In light of these considerations, and because no other cri- 
teria of taxonomic significance are apparent (the presence 
of bisporangia, reported by Jao (1936) but not seen during 
this study, does not alone appear to be sufficient grounds 
for species separation), Awdowinella daviesti and Acro- 
chaetium alcyonidii are considered conspecific. 


Acrochaetium amphiroae (Drew) Papenfuss has already 
been referred to the conspecificity of Audowinella daviesii 
(Woelkerling 1971); consequently the records of Doty 
(1948), Mathieson et al. (1969), South (1970), and Taylor 
(1957) are referred (at least as to name) to A. daviesi. 
None of the collections upon which these records are based, 
however, has been available for examination. 

A number of collections of other investigators have been 
examined and apparently found not to contain plants of 
A. daviesiti, Specimens which Harvey (1853, p. 243) re- 
ferred to Callithamnion daviesii belong to Colaconema 
secundata. With one exception (see specimens examined 
listing), all the collections of Collins examined, both in 
FH and NY, including Phycotheca Boreali Americana 
Specimen No. 880, have been found not to contain plants 
of A. daviesii but rather plants referable to Colaconema 
secundata. Since there is a slight possibility that both 
taxa are present in Collins material, Collins references 
are listed with question marks in this account. The un- 
certainty over Collins material leads to uncertainty over 
the reported distribution of A. daviesii in the New England 
region; in addition to a number of stations in Massachu- 


556 Rhodora [Vol. 75 


setts, A. daviesii is also definitely known from only one 
locality in Maine and one in New York as a result of the 
present study. 

Plants from Nova Scotia referred to Acrochaetium 
aleyonidii (Edelstein et al. 1967) and A. amphiroae (Edel- 
stein and McLachlan 1968) should be checked; one collec- 
tion sent from Dr. Edelstein and identified as A. aleyonidii 
has been examined and found to contain only plants of 
Colaconema membranacea (Magnus) comb. nov. Likewise, 
the illustrations (Edelstein and McLachlan 1968, Figs. 
49-50) of plants referred to Acrochaetium amphiroae leave 
considerable doubt as to their true relationship to Audowi- 
nella davies. 

A number of specimens hitherto referred to Acro- 
chaetium sagraeanum (Montagne) Bornet have been ex- 
amined and found to be Audowinella daviesii; citations are 
appended. The type of Acrochaetium sagraeanum has been 
excluded from the Rhodophyta (see “Species Excluden- 
dae’’). 

Massachusetts specimens collected by Collins in 1888 
and 1895 from Marblehead, by Farlow in 1875 from Gay 
Head (Martha’s Vineyard; host is Cystoclonium), and 
by Nott in 1895 from Butler’s Point (Woods Hole) in FH 
and NY have been examined during this study and found 
to contain sexual plants of Audouinella daviesi and, in 
several instances, a plant or two of Colaconema secundata. 
In the literature, these collections have been referred 
erronously to Chantransia corymbifera (Collins 1896 ; 
Collins, Holden, and Setchell 1896, No. 192), C. efflorescens 
var. thuretii (Collins 1906, Davis 1913), and/or Acro- 
chaetium thuretii (Taylor 1987, 1957). The vast majority 
of plants examined in the above populations contained 
numerous carposporophytes and very few monosporangia. 
In each case, however, the few monosporangia found tended 
to be arranged in axial clusters of 3-4 on branched stalks, 
and this is the characteristic arrangement found in Audoui- 
nella daviesii. In other respects these populations agree 
well with sexual plants collected during this study. For 


1973] Audouinella — Woelkerling 557 


additional information, see comments below under Audoui- 
nella efflorescens. 


5. Audouinella microscopica (Naegeli In Kuetzing) Woel- 
kerling 1971: 33, Figs. 10, 23A. 1972: 85 et seq., Figs. 
1-14; 1973: 86. Figs. 46-51. 

Acrochaetium microscopicum (Naegeli In Kuetzing) 
Naegeli 1861: 407, Figs. 24-25. 

Callithamnion microscopicum Naegeli In Kuetzing 1849: 
640. 

Chantransia microscopica (Naegeli In Kuetzing) Batters 
In Schiffner 1916: 136, Figs. 13-18. 

Chromastrum microscopicum (Naegeli In Kuetzing) 
Papenfuss 1945: 322. 

Kylinia microscopica (Naegeli In Kuetzing) Kylin 1944: 
13. Papenfuss 1947: 437. 

Rhodochorton microscopicum | (Naegeli In Kuetzing) 
Drew 1928: 151, 163. 

Acrochaetium catenulatum Howe 1914: 84, pl. 31, Figs. 
12-18. 

Chantransia catenulata (Howe) DeToni 1924: 44. 

Kylinia catenulata (Howe) Kylin 1944: 13. 

Rhodochorton catenulatum (Howe) Nakamura 1941: 
273, 280, Fig. 1. 

Acrochaetium | collopodum (Rosenvinge) Hamel 1927: 
81; 1928: 175. 

Chantransia collopoda (Rosenvinge) Rosenvinge 1909: 
81. 

Chromastrum collopodum (Rosenvinge) Papenfuss 1945: 
320. 

Kylinia collopoda (Rosenvinge) Kylin 1944: 13, 15, 
Fig. 6. 

Acrochaetium compactum Jao 1936: 241, pl. 10, Figs. 
6-14. Taylor 1937: 228, pl. 32, figs. 6-14. 

Chromastrum compactum (Jao) Papenfuss 1945: 321. 

Kylinia compacta Papenfuss 1947: 436. South 1970: 1. 
Taylor 1957: 212, pl. 32, Figs. 6-14. 

Acrochaetium crassipes (Boergesen) Boergesen 1915: 
20, Figs. 11-13. Boergesen 1927: 12, Fig. 5. Collins and 


558 Rhodora [Vol. 75 


Hervey 1917: 96. Howe 1918: 511. Taylor 1941: 75. 

Chantransia crassipes Boergesen 1909: 1, Fig. 1. Taylor 
1928: 134, pl. 28, Fig. 16. 

Chromastrum crassipes (Boergesen) Papenfuss 1945: 
321. 

Kylinia crassipes (Boergesen) Kylin 1944: 18. Taylor 
1960: 300. 

Acrochaetium microfilum Jao 1936: 240, pl. 10, Figs. 1-5. 
(Non A. microfilum Levring 1945: 12, Fig. 4. = A. levringti 
Papenfuss 1947: 436). Taylor 1937: 232, pl. 32, Figs. 1-5: 
1957: 219, pl. 32, Figs. 1-5. 

Acrochaetium | moniliforme (Rosenvinge)  Boergesen 
1915: 22. Jao 1936: 241, pl. 10, Figs. 15-17. Taylor 1937: 
227, pl. 32, Figs. 15-17. 

Chantransia moniliformis Rosenvinge 1909: 99, Figs. 
28-29. 

Chromastrum | moniliforme (Rosenvinge)  Papenfuss 
1945: 322. 

Kylinia moniliformis (Rosenvinge) Kylin 1944: 13. 
South 1970: 1. Taylor 1957: 211, pl. 32, Figs. 15-17. 

Rhodochorton moniliforme (Rosenvinge) Drew 1928: 
151, 164. 

Chantransia secundata auct. non. (Lyngbye) Thuret: 
Hauck and Richter 1892: 454. 

Plants epiphytic, up to 100 „m tall exclusive of hairs; 
original spore persisting as a unicellular base slightly 
smaller to slightly larger than other cells. Filaments of 
erect system 1-4, commonly arcuate, simple or with a few 
secundly to irregularly arranged lateral branches. Cells 
barrel-shaped to cylindrical, 3-10 „m wide and 3-11 „m 
long, L/D 0.75-2; each cell containing a single parietal 
lobate to stellate chromoplast with one pyrenoid. Terminal 
hairs up to 40 »m long occur occasionally. 

Monosporangia ovoid, 4-9 „m wide and 6-15 „m long. 
terminal or lateral, single or rarely in pairs, sessile or 
stalked, adaxially seriate or occasionally more scattered. 

Other reproductive structures not observed in New Eng- 
land populations. 


1973] Audouinella — Woelkerling 559 


Type locality: Torquay, England. 

Type: L, No. 940285... 306. 

Distribution: Nearly cosmopolitan. 

Hosts: Chaetomorpha, Chordaria, Cladophora, Entero- 
morpha, Polysiphonia, and Sphaerotrichia in New Eng- 
land; a wide variety of algae and bryozoans elsewhere. 


Specimens examined: 

MASSACHUSETTS: Black Rock, Sconticut Neck, New Bedford, 25. VII. 
1934, Jao (MICH, Woods Hole, No. 275, type of Acrochaetium com- 
pactum Jao). Cape Codder Point, Falmouth 19. XI. 1969, Woelker- 
ling (WIW 2292). Norton Point, Martha's Vineyard, 3. VIII. 1934, 
Jao (MICH, Woods Hole, No. 280 [not 274 as reported by Jao 1936, 
p. 240], type of Acrochaetium microfilum Jao). West Falmouth 
Harbor, 17. X. 1970, Woelkerling (waw 2826). Woods Hole (Nobska 
Point), 4. II. 1970, Woelkerling (wyw 2320). 

ENGLAND: Torquay, 1845, Naegeli (L 940285 . . . 306, type of Au- 
douinella microscopica (Naegeli) Woelkerling). Torquay, 1845, 
Naegeli (FH = Hauck and Richter 1892, No. 454, isotype of A. micro- 
scopica, which is labeled Chantransia secundata). Torquay, 1845, 
Naegeli (NY — Hauck and Richter 1892, No. 454, isotype of A. micro- 
scopica, which is labeled Chantransia secundata). 


Audouinella microscopica is distinguished from other 
New England audouinelloid algae with a unicellular base 
1) in commonly having cells isodiametric or broader than 
long; 2) in having a more or less globose basal cell proto- 
plast that is not markedly flattened on the side in contact 
with the substrate; and 3) in having erect or ascending 
rather than procumbent filaments. 

A detailed morphotaxonomic account of this species in 
New England and adjacent regions has recently appeared 
in the literature (Woelkerling 1972). To date this species 
has been collected only from Massachusetts along the New 
England coast, but it is known from Nova Scotia (Edel- 
stein and McLachlan 1966, 1968; Edelstein et al. 1967: 
South and Cardinal 1970) and, therefore, is to be expected 
at intermediate points. Although collection data is incom- 
plete, A. microscopica probably is present throughout the 
year and is to be sought on any older, epiphytized algae. 
Its microscopic size may account for its being generally 
overlooked. 


560 Rhodora [Vol. 75 


6. Audouinella saviana (Meneghini) comb. nov. Figs. 
56-60. 

Acrochaetium savianum (Meneghini) Naegeli 1861: 
405. Feldmann 1942: 218. Hamel 1927: 41, 98, Fig. 32; 
1928: 192: 1928a: 135, Fig. 32. Papenfuss 1945: 311; 
1947: 435. 

Callithamnion savianum Meneghini 1840: 511. J. Agardh 
1851: 14; 1876: 6. Kuetzing 1849: 641. 

Chantransia saviana (Meneghini) Ardissone 1883: 276 
(pro parte). DeToni 1897: 68. 

Chantransia efflorescens var. thuretii Bornet 1904: XVI, 
pl. 1. 

Acrochaetium thuretii (Bornet) Collins et Hervey 1917: 
98. Doty 1948: 263. Hamel 1927: 37, 97, Fig. 30. 1928: 
191; 1928a: 131, Fig. 30. Kylin 1944: 21, Fig. 14. Papen- 
fuss 1945: 311. 

Audouinella thuretii (Bornet) Woelkerling 1971: 36, 
Figs. 12, 24; 1973: 88. 

Chantransia thuretii Bornet. Collins 1900: 49 (nom. 
nud.). 

Chantransia thuretii (Bornet) Kylin 1907: 119, Fig. 28. 
Rosenvinge 1909: 100, Figs. 30-33. 

Rhodochorton thuretii (Bornet) Drew 1928: 171. 

Acrochaetium, sagraeaum auct. non. (Montagne) Bornet: 
Boergesen 1915: 35 (pro parte). Bornet 1904: XXI (pro 
parte). Collins 1905: 231 (pro parte); 1906: 192 (pro 
parte). Collins and Hervey 1917: 97. Collins, Holden, et 
Setchell 1917: 2181. Hamel 1927: 77, 99 (pro parte) ; 
1928: 173, 191 (pro parte). Hylander 1928: 159 (pro 
parte) Jao 1936: 244 (pro parte). Papenfuss 1945: 311 
(pro parte). Taylor 1937: 233 (pro parte); 1957: 220 
(pro parte); 1960: 309 (pro parte). 

Chantransia sagraeana auct, non. (Montagne) DeToni: 
DeToni 1924: 51 (pro parte). 

Chantransia virgatula auct. non. (Harvey) Thuret In 
LeJolis: Collins 1900: 49 (pro parte). Collins, Holden, 
et Setchell 1895: 39. Hylander 1928: 158. 


Plants partly to entirely epipytic, caespitose, up to 4 mm 


1973] Audouinella — Woelkerling 561 


tall; original spore non-persistent. Prostrate system com- 
posed of short, simple or branched filaments free from one 
another or united into an irregularly shaped pseudoparen- 
chymateous disc. Erect filaments moderately to freely and 
irregularly branched. Cells cylindrical, (7-) 8-12 (-14) 
um wide and 20-60 »m long [L/D 2-6(-8)] in main axes 
and laterals, sometimes tapering to 4-6 „um wide near the 
tips; each cell containing a single parietal lobate chromo- 
plast and one pyrenoid. Unicellular hairs unknown. 

Monosporangia ovoid, 10-15 „m wide and 18-27 um long, 
sessile or stalked, single or in pairs, in a secund series along 
the laterals or more scattered. Tetrasporangia ovoid, 17-24 
um wide and 26-34 um long, sessile or stalked, single or in 
pairs, scattered on the erect filaments. 

Other reproductive structures not observed. 


Type locality: Genoa, Italy. 

Type: FL. 

Distribution: Nearly cosmopolitan. 

Hosts: A variety of algae and marine angiosperms. 


Specimens examined: 

MASSACHUSETTS: Sandwich Jetty (Cape Cod Canal), 13. X. 1970, 
Woelkerling (WIW 2818). West Yarmouth, 17. IX. 1969, Woelkerling 
(WJW 2253); 3. X. 1969, Woelkerling (wyw 2256). Waquoit Bay 
(Falmouth), 30. VI. 1969, Conway (wJw 1839); 15. IX. 1969, Con- 
way (wJw 1840). Woods Hole (Eel Pond), 21. VII. 1970, Wilce 
(WJW 2753). Woods Hole (Nobska) 17. VII. 1970, Woelkerling 
(ww 2722). Woods Hole, 1876, Dudley (wsw 3284). 

FRANCE: Cherbourg, 1. IX. 1856, Bornet [PC, possibly the type of 
Acrochaetium thuretti (Bornet) Collins et Hervey]. 

ITALY: Genoa, (8. VI ?). 1839, Meneghini (Ft, type of Callithamnion 
savianum Meneghini). 


Audouinella saviana is distinguished from other New 
England audouinelloid algae of similar morphology in 
having a filamentous to pseudoparenchymatous prostrate 
system without an enlarged panduriform or pyriform cell 
and having the monosporangia solitary or in pairs on sim- 
ple stalks rather than in clusters on branched stalks. 

Audouinella saviana appears to be confined to the sub- 
littoral and lowermost littoral in New England waters and 


562 Rhodora [Vol. 75 


has been collected in June, July, September and October 
as an epiphyte on various algae including Champia, Chon- 
dria, and Stilophora. Tetrasporangial plants have been 
colleeted in October; sexual plants have not been found 
in New England waters, but are described in detail by 
Kylin (1907) and Rosenvinge (1909) [both as Chantransia 
thuretii]l. Additional study may reveal that the species is 
present in New England during all or most of the year. 

The vegetative system shows considerable variation in 
New England populations. In some cases, the prostrate 
system is quite reduced while in others it is comparatively 
robust. The L/D ratio of cells also varies considerably. 
In some plants, most cells are 4-8 diameters long; in others 
they are mostly 2-4 diameters long ; and in still others they 
show a combination of both of the above (Figs. 57, 59-60). 

Acrochaetium thuretii (Bornet) Collins et Hervey is 
here considered conspecific with Audouinella saviana after 
comparing specimens from the type collections of the two 
taxa. Material from FI and PC agree in all essential re- 
spects with one another and with New England popula- 
tions (Table 4; also see Addendum). 

Plants referable to Awdouinella saviana appear to be 
widely distributed, but have usually been reported under 
the specific epithet "thuretti" while the epiphet “saviana”’ 
has persisted in relative obscurity. Hamel (1927, 1928a) 
attempted to distinguish the two taxa on the bases of 
presence or absence of sexual structures and on the rela- 
tive L/D ratio of cells, but neither of these criteria appears 
systematically reliable. 

A number of specimens hitherto referred to Acrochaetium 
sagraeanum (Montagne) Bornet have been examined and 
found to be Audouinella saviana; citations have been 
appended. The type specimen of Acrochaetium sagraeanum 
has been excluded from the Rhodophyta (see “Species 
Exludendae"). 

The literature references of Collins (1896, 1906), Col- 
lins, Holden, and Setchell (1896), Davis (1913), and Taylor 
(1937, 1957) to specimens under the names of Chan- 


1973] Audouinella — Woelkerling 563 


transia corymbifera, C. efforescenes var. thuretii, and/or 
Acrochaetium thuretii are based on misidentifications. 
These collections have been re-examined and found to con- 
tain plants of Audouinella daviesii; for further discussion, 
see the account of that species. 

The relationships of A. saviana to a number of taxa of 
similar morphology await clarification. These include 
Acrochaetium avrainvillae Boergesen (1915, p. 48, Figs. 
47-49), A. hypneae Boergesen (1909), and A. pallens 
(Zanardini) Naegeli (See Hamel 1927, p. 48). All of these 
taxa appear to be very similar to Audouinella saviana and 
a comparison of type collections may show some or all to 
be conspecific. 

The precise publication date of Meneghini's original 
diagnosis of Callithamnion savianum requires further com- 
ment. The issue of Flora (Regensburg) containing the 
diagnosis is dated 28 August 1840; however reference is 
made on p. 511 of that issue to prior publication of diag- 
nosis on 23 May 1840 in a Pisa “Giornale?” (= journal?, 
newspaper?), 

Copies of a three page type-set document dated 23 May 
1840 which resemble a journal reprint and are entitled 
"Lettera del Prof. Giuseppe Meneghini at Dott. Jacob 
Corinaldi a Pisa" are located at FI and pc (Dr. Paul Silva, 
personal communication) ; Prof. Peter S. Dixon has kindly 
sent me a photocopy of the PC document. Hamel (1927, 
1928) also has made reference to this document. The Pc 
copy unfortunately provides no clues as to the original 
place of publication other than three words — “Pisa Tipo- 
grafia Prosperi" — written in longhand at the bottom of 
the first page, and until further light can be shed on the 
matter of its original place of publication (if, indeed, 
there was one), it seems wise to continue to refer to the 
"Flora" article when citing Meneghini's original diagnosis 
of Callithamnion savianum since the wording of the diag- 
nosis in *Flora" is identical to that in the PC document. 


7. Audouinella unifila (Jao) comb. nov. Figs. 44-45. 
Acrochaetium unifilum Jao 1936: 239, pl. 10, Figs. 26-32. 


564 Rhodora [Vol. 75 


Taylor 1937: 228, pl. 32, Figs. 26-32. Non. A. unifilum 
Levring 1953: 472, Fig. 9 ( — Colaconema nakamurai 
Woelkerling 1971: 46, Fig. 16). 

Chromastrum unifilum (Jao) Papenfuss 1945: 322. 

Kylinia unifila (Jao) Papenfuss 1947: 437. Taylor 1957: 
212, pl. 32, Figs. 26-32. 

Audouinella australis (Levring) Woelkerling 1971: 25, 
Figs. 4-5. 

Kylinia australis Levring 1953: 487, Figs. 21A-C. Boney 
and White 1967: 595. 


Plants epiphytic, up to 150 „m tall; original spore per- 
sisting as a hemispherical unicellular base appressed to the 
substrate and usually giving rise to a single erect filament. 
Erect filament(s) procumbent to semi-upright, simple or 
rarely with 1-2 celled laterals. Cells cylindrical, 6-8 „m 
wide and 8-20 „m long, L/D 1-3, each cell containing a 
single parietal lobate to stellate (Jao 1936) chromoplast 
and one pyrenoid. Terminal and pseudolateral hairs up to 
50 um long occur. 

Monosporangia ovoid, 5-7 „m wide and 8-14 „m long, 
sessile or occasionally on unicellular stalks, scattered along 
the erect filament (s). 

Other reproductive structures not observed in New Eng- 
land populations. 


Type Locality: Norton Point, Martha's Vineyard, Massa- 


chusetts. 
Type: MICH, Woods Hole No. 274. 
Distribution: Massachusetts; South Australia. 
Hosts: Arthrocladia villosa Duby, Audouinella sp. 


Specimens examined : 
MASSACHUSETTS: Norton Point (Martha's Vineyard), 3. VIII. 1934, 
Jao (Michigan, holotype). 
AUSTRALIA: Pennington Bay (Kangaroo Island), 7. II. 1947, Wo- 
mersley (ADU, A31373, isotype of Kylinia australis Levring). Note: 
Other Australian collections examined are cited in an earlier paper 
( Woelkerling, 1971, p. 26). 


Audouinella unifila is distinguished from other New 
England audouinelloid algae with a unicellubar base by 


1973] Audouinella — Woelkerling 565 


the procumbent development of erect filaments and the 
characteristically flattened (compressed)  hemispherical 
basal cell. D. 

The one known New England collection of this taxon is 
the type collection, plants of which are on the permanent 
slides in MICH. Only monosporangial specimens occur, and 
the range of cell and spore dimensions of these plants is 
somewhat greater than reported originally by Jao (1936). 
Chromoplasts were not recognizable, but at least one 
pyrenoid was observed. 

Critical comparisons of the type collections of A. wnifila 
and A. australis strongly indicate that the two taxa are 
conspecific. Monosporangial plants of both taxa show 
virtually identical ranges of cell and spore dimensions and 
in general have the same habit. Apparent differences in 
chromoplast shape (Jao, 1936, reported stellate plastids 
in A. wnifila; Levring (1953) recorded parietal plastids in 
A. australis) do not appear to be taxonomically significant 
(see Woelkerling 1971, p. 14). Woelkerling (1971, p. 26) 
previously noted similarities in the two taxa. 

Although sexual plants do not occur in the type collec- 
tion, this taxon is referred to Audouinella on the basis of 
the rather common occurrence of sexual plants in South 
Australian populations. 

Some questions may be raised as to the relationships 
between Audouinella unifila and A. microcopica. The 
author’s experience to date indicates that the two have 
quite distinct basal cells and modes of erect filament de- 
velopment, and no intermediates have been observed. How- 
ever, the possibility exists that such intermediates do occur 
and they should be watched for. 


COLACONEMA Batters 


Colaconema Batters 1896: 8. Woelkerling 1971: 40. Non 
Colaconema Schmitz In Schmitz and Falkenberg 1897: 452. 


Note: Species not referable to Colaconema have been 
placed in the past in Acrochaetium, Audouinella, Calli- 


566 Rhodora [Vol. 75 


thamnion, Ceramium, Chantransia, Chromastrum, Kylinia, 
Rhodochorton, and/or Trentepohlia. 


Plants epibiotic, endobiotic, or saxicolous; attached to 
or suspended in the substrate by a single-celled holdfast 
or more commonly by a prostrate system of simple or 
branched filaments, which may or may not become pseudo- 
parenchymatous, Erect filaments, when present, simple or 
branched, up to 10 mm tall; cells containing one to a 
number of variously shaped chromoplasts with or without 
pyrenoids, 

Asexual reproduction by sessile or stalked monospor- 
angia, bisporangia, tetrasporangia, and/or multipartite 
sporangia borne on the erect and/or prostrate filaments. 

Sexual reproduction unknown. 

Type Species: Colaconema bonnemaisoniae Batters. 


Section I 


The species in this section are not known to contain 
pyrenoids in their chromoplasts. 


8. Colaconema membranacea (Magnus) comb. nov. Figs. 
64-65. 

Audouinella membranacea (Magnus) Papenfuss 1945: 
326; 1947: 438. Edelstein and McLachlan 1966a: 1052. 
Taylor 1957: 224, pl. 31, Figs. 11-12. 

Callithamnion membranaceum Magnus 1874: 67, tab. II. 
Figs. 7-15. Collins 1883: 56; 1888: 10. 

Rhodochorton membranaceum (Magnus) Hauck 1885: 
69 (only as to binomial). Collins 1894: 230; 1900: 51, 
1911: 280. Collins, Holden and Setchell 1895a: 99. Davis 
1913: 818, DeToni 1903: 1513. Drew 1928: 186. Hamel 
1927: 59, 109, Fig. 39A-E; 1928: 202; 1928a: 152, Fig. 
39A-E. Hauck and Richter 1888: 154. Hylander 1928: 169. 
Kuckuck 1897: 337 et seq., Figs. 1-7. Taylor 1937: 240, 
pl. 31, Figs. 11-12. 


Plants endozoic; original spore non-persistent, Prostrate 
system consisting of irregularly branched filaments of 


1973] Audouinella — Woelkerling 567 


indefinite length which may be free from one another or 
pseudoparenchymatously united into a sheet-like endozoic 
thallus. Cells of prostrate filaments cylindrical to irregular 
in shape, 8-60 „m long, and 5-11 „m wide, L/D 0.75-5(-8) : 
each cell containing several to many discoid or irregularly 
shaped chromoplasts (sometimes becoming more or less 
spirally twisted) without pyrenoids. Erect filaments, when 
present, simple or sparingly and irregularly branched, 
rarely more than 25 cells long. Cells 10-45 ¿m long and 
7-13 um wide, L/D 1-4. 

Tetrasporangia ovoid to globose, 10-15 „m wide and 
16-25 »m long, solitary, sessile or occasionally stalked, ter- 
minal or rarely lateral on erect filaments or on prostrate 
filaments. 


Type locality: Store Baelt Channel, between Sporogoe 
Island and Korsoer, Denmark. 

Type: ? 

Distribution: Europe and cooler waters of both coasts 
of North America. 


Hosts: A variety of marine invertebrates, especially 
hydroids. 


Specimens examined : 
CONNECTICUT: Charles Island (Milford Channel), 22. IV. 1889, 
Holden (NY). 
MAINE: Eagle Island, VII. 1894, Collins (NY). 
MASSACHUSETTS: Brandt Point (Marshfield), 7. IV. 1968, Greenwich 
(NHA). Gay Head (Martha's Vineyard), 3. VIII, 1948, Doty (U. 
Hawaii). Marblehead, 30. V. 1884, Collins (NY). Martha's Vineyard, 
16. VII. 1941, Taylor (FH). Nahant, 14. V. 1882. Collins (NY); 1. V. 
1970, Woelkerling (wyw 2545). Revere Beach (Boston), IV. 1885, 
Collins (NY); 29. V. 1886, Collins (NY, FH = Hauck and Richter 
1888, No. 154); 10. VI. 1894, Collins (NY). Sandwich Jetty (Cape 
Cod Canal), 13. X. 1970, Woelkerling (wa3w 2812). Sconset Beach, 
13. VIII. 1886, Collins (NY). Scusset Beach, 29. III. 1969, LaPlante 
(UNH). Woods Hole, 7. IV. 1968, Logan (NHA); 6. IV. 1968, Kas- 
telowitz (NHA). 
NEW HAMPSHIRE: Cedar Point (Dover), 21. XII. 1966, Hehre and 
Conway (NHA); 3. III. 1967 Hehre (NHA). Hilton Park (Dover 
Point), 19. III. 1967, 'Hehre and Conway (NHA); 27. IV. 1967, Hehre 
and Stone (NHA). Rye Ledge, 5. II. 1970, Woelkerling (wJw 2336). 


568 Rhodora [Vol. 75 


NEW YORK: Montauk Point (Long Island), 20. I. 1970, Woelkerling 
(wsw 2258). 


Colaconema membranacea is distinguished from other 
New England audouinelloid algae in having several to 
many irregular to discoid to spiral chromoplasts without 
pyrenoids and in having solitary tetrasporangia that are 
borne in a sessile condition or on simple stalks rather than 
in clusters on branched gonimoblast filaments. In addition, 
this species is known to occur only as an endozoophyte in 
hydroids and several other types of invertebrates, and 
while this in itself is no criterion of specific distinction, 
it can serve as a useful ecological guide for identification 
purposes. 

Colaconema membranacea. has been collected throughout 
the year in New England waters and commonly occurs on 
hydroids in both the intertidal zone (especially hydroids 
attached to Ascophyllum or Fucus in shaded habitats) and 
the sublittoral zone, Often the hydroids become reddish 
as a result of the algal infestation. 

Among New England populations, the nature of the vege- 
tative system varies considerably. In lightly infested host 
animals, the prostrate filaments tend to remain largely 
free from one another, but under more crowded conditions, 
the prostrate system becomes pseudoparenchymatous and 
all traces of a filamentous character are lost (Fig. 64). 
In most populations the erect system is entirely suppressed 
and sporangia, when present, are borne directly on the 
prostrate filaments. In a few instances, however, the erect 
systems were more highly developed and consisted of 
branched filaments up to 750 ¿m long. 

The nature of the life cycle of this species awaits clari- 
fieation by means of culture studies, and until such time, 
it is difficult to suggest definite relationships between 
C. membranacea. and other audouinelloid algae. Thus, for 
example, Rhodochorton concrescens (see West 1970a) and 
Audouinella purpurea (see West 1969, 1970) share a num- 
ber of morphological features with Colaconema membra- 
nacea, but apparently they have quite different types of 
life cycles. 


1973] Audouinella — Woelkerling 569 


The relationships of C. membranacea. to Callithamnion 
entozoicum Reinsch in Giard (1890) are discussed under 
the latter species in the section on Species Inquirendae. 


Section II 


Species in this section have chromoplasts each with only 
one pyrenoid, 


9. Colaconema humilis (Rosenvinge) Woelkerling 1971: 44, 
Figs. 15J-O. Figs. 66-73. 

Acrochaetium humile (Rosenvinge) Boergesen 1915: 23. 
Baardseth 1941: 41. Jorde et Klavestad 1963: 76. Kylin 
1944: 22, Fig. 17. Levring 1953: 478. Schiffner 1931: 143. 
Sundene 1953: 185. 

Chantransia humilis Rosenvinge 1909: 117, Figs. 44, 45. 
Boergesen 1927: 21. Levring 1935: 37, Figs. 7F-S; 1937: 
89; 1940: 78, Figs. 23A-B; 1942: 8. 

Chromastrum humile (Rosenvinge) Papenfuss 1945: 
323. 

Kylinia hwmile (Rosenvinge) Papenfuss 1947: 437. 

Rhodochorton humile (Rosenvinge) Drew 1928: 151, 
169. 

Acrochaetium radiatum Jao 1936: 246, pl. 10, Figs. 
18-25. Edelstein et al. 1967: 196. Papenfuss 1945: 310. 
South and Cardinal 1970: 2079. Taylor 1937: 237, pl. 32, 
Figs. 18-25; 1957: 223, pl. 32, Figs. 18-25. 


Plants epiphytic, up to 60 „m tall exclusive of hairs; 
developing from septate or occasionally aseptate spores. 
Prostrate system consisting of 2-6 simple or sparsely 
branched filaments up to 300 »m long, arising from the 
spore and creeping or forming an irregular, more or less 
pseudoparenchymatous disc. Cells of prostrate filaments 
cylindrical or occasionally somewhat subglobose, 6-14 „m 
long and 4-8 um wide, L/D 1-2; each cell containing a 
single chromoplast and one pyrenoid. Erect filaments absent 
to numerous, simple or sparsely irregularly branched, 
generally less than 10 cells long; cell dimensions similar 


510 Rhodora [Vol. 75 


to those of prostrate filaments. Unicellular hairs up to 
100 4m long occur. 

Monosporangia ovoid, 7-14 um long and 5-8 ¿m wide, 
solitary or occasionally in pairs, sessile or stalked, scattered 
on prostrate or erect filaments. 


Other reproductive structures unknown. 
Type locality: Spodobjerg, Langeland, Denmark. 
Type: C. 


Distribution: Massachusetts; Nova Scotia; Australia: 
Atlantic and Mediterranean shores of Europe. 


Hosts: Ectocarpus and Polysiphonia in North America: 
a variety of algae elsewhere. 


Specimens examined : 


MASSACHUSETTS: Norton Point (Martha's Vineyard), 3. VIII. 1954. 
Joa (Micu, Woods Hole No. 280, type of Acrochaetium radiatum 
Jao). West Falmouth (Harbor), 3. X. 1970, Woelkerling (WJW 
2767). West Yarmouth, 12. XII. 1969, Woelkerling (waw 2259). 
Woods Hole (Nobska Point), 26. VI. 1969, Conway (wJw 1838); 
4. II. 1970, Woelkerling (WIw 2329). 

NOVA SCOTIA: Ketch Harbor, 31. V. 1965, Edelstein (Herb. Nat. Res. 


Coun., Halifax). 


Colaconema humilis can be distinguished from other New 
England audouinelloid algae by the following combination 
of features (as found in the collections examined): 1) A 
reduced erect system with most filaments of 10 or fewer 
cells; 2) Cells of erect and prostrate filaments mostly less 
than two diameters long; and 3) Prostrate system more or 
less compact; filaments rarely over 500 ¿m long. The first 
and third criteria probably do not represent very soundly 
based systematic characters, but until they can be re- 
examined in light of additional New England and other 
collections, they have been offered as useful guides for 
species recognition in this instance. 


C. humilis is known in New England waters from col- 


lections made in February, June, August, October, and 
December. Edelstein et al. (1967) report it from May 


1973] Audouinella — Woelkerling 571 


through September in Nova Scotia, and judging from this, 
it seems reasonable to think that it may occur throughout 
the year in this region. 


The erect system varies considerably and may be sup- 
pressed altogether (Fig. 67), consist of a few several celled 
filaments (Figs. 69-70), or of a number of short rather 
densely crowded filaments (Fig. 73). Likewise the extent 
of the prostrate system varies according to available sub- 
strate space; on the finely filamentous Ectocapus, it con- 
sists of a very few cells (Fig. 71) but on a larger species 
of Polysiphonia it can spread out more (Fig. 73). Most 
germinating spores divide into two distinct daughter cells 
(Fig. 68) but others do not (Fig. 66). Chromoplasts have 
not been observed; Jao (1936) reports a parietal plastid, 
Rosenvinge (1909) found a stellate plastid, and Woelkerling 
(1971) found a variable plastid shape. 


Several investigators (e.g. Baardseth 1941, p. 41; Woel- 
kerling 1971, p. 45) have commented upon the possible 
conspecificity of a number of audouinelloid algae whose 
morphology is similar to Colaconema humilis. On the 
basis of the examination of the type and other populations, 
Acrochaetium radiatum Jao has been referred to the 
synomomy of Colaconema humilis, Jao (1936, p. 247) 
separated the two taxa on the basis of the former “ 
having more than two main filaments arising from the 
divided germinating spore and all the filaments arranged 
radiately and densely from the center of a fully developed 
thallus.” The type collection of Acrochaetium radiatum, 
however, contains plants that have 2 as well as more than 
two filaments emanating from the germinating spore (Fig. 
68), and has plants that are not densely radiate (Fig. 69) 
as well as ones that are (see Jao, 1936, pl. 10, Figs. 22-25). 
Thus the criteria of specific distinction offered by Jao do 
not appear to be taxonomically reliable, and the two taxa 
ean be considered conspecific. 


The relationships of Colaconema humilis to a number 
of similar taxa (see Baardseth 1941, p. 41 and Woelkerling 


572 Rhodora [Vol. 75 


1971, p. 45 for names) remains unclear, but there is in- 
creasing doubt as to whether many of these are really 
distinct species. In the Nova Scotia population examined 
during this study, plants with varying degrees of erect 
system development occur side by side and agree morpho- 
logically with one or several of the taxa cited by Baardseth 
(1941) and/or Woelkerling (1971). Similar variation 
occurs among the New England populations. Final clari- 
fication must await a comparison of the various type col- 
lections (which in many cases represent the only reported 
collections), but is seems likely that a number of these 
taxa will eventually be reduced to synonomy. 


10. Colaconema minima (Collins) comb. nov. Figs. 74-76. 


Acrochaetium minimum Collins 1908: 133. Collins, 
Holden, and Setchell 1908: 1493. Davis 1913: 813. Hamel 
1927: 89; 1928: 183. Papenfuss 1945: 316. Taylor 1937: 
237; 1957: 222. 


Chantransia minima (Collins) Collins 1911a: 186, De- 
Toni 1924: 62. 


Acrochaetium emergens (Rosenvinge) Weber van Bosse 
1921: 194. Hamel 1927: 93; 1928: 186. Jao 1936: 247, 
pl. 13, Figs. 7, 7B. Papenfuss 1945: 314. Taylor 1937: 
235; 1957: 221. 


Chantransia emergens Rosenvinge 1909: 128, Fig. 55. 
DeToni 1924: 67. 


Rhodochorton emergens (Rosenvinge) Drew 1928: 151, 
188. 


Plants epi- to endophytic; sporelings not observed. Pros- 
strate system consisting of irregularly branched filaments 
of indefinite length creeping on the surface or between the 
superficial cells of the host. Cells of prostrate filaments 
cylindrical to irregular in shape, 5-25 (-44) „um long and 
3-8 (-14) um wide, L/D 1-6 (-8) ; each cell containing a 
parietal chromoplast and one pyrenoid. Erect filaments 1-5 
(-25) cells long, simple or with a few short, irregularly 


1973] Audouinella — Woelkerling 573 


arranged laterals; cell dimensions similar to those of pros- 
trate filaments. 

Unicellular hairs not observed. 

Monosporangia ovoid to globose to hemispherical, 5-12 
um long and 4-10 „m wide, solitary or occasionally in pairs, 
sessile or on 1-2 celled stalks, scattered on the prostrate or 
erect filaments. 

Other reproductive structures unknown. 


Type locality: Robinson's Hole, Elizabeth Islands, Mas- 
sachusetts. 


Type: FH. 
Distribution: Massachusetts; Denmark; Norway. 
Hosts: Asparagopsis, Desmarestia, Polysiphonia. 


Specimens examined: 

MASSACHUSETTS: Robinson’s Hole (Elizabeth Islands), VIII. 1907, 
Collins (FH, type); (wis, isotype). [Isotypes have been distributed 
as No. 1493 in Collins, Holden, and Setchell (1908).] Sandwich 
Beach (Cape Cod Canal), 30. VI. 1970, Wilce (wyw 2693); 11. VII. 
1970, Woelkerling (wyw 2699). 

DENMARK: Mollegrund (Hirshals), 8. VIII. 1899, Rosenvinge (C, 
Algae Marinae Danicae No. 6574, type of Chantransia emergens 
Rosenvinge). 


As circumscribed here, Colaconema minima can be dis- 
tinguished from other New England audouinelloid algae 
by the following combination of features: 1) A prostrate 
system of indefinite length bearing no or short (1-5 (-25)) 
celled erect filaments, 2) Sporangia under 15 „m long and 
12 „m wide; 3) Cells with single chromoplasts with one 
pyrenoid each; commonly over two diameters long. 


Colaconema minima is known from three Cape Cod, 
Massachusetts collections. The type and the WIS isotype 
of the species consist of dried specimens that have proven 
diffieult to work with because of the matted and rather 
unresponsive (to resoaking and slide making) condition. 
Nevertheless it has been possible to study and illustrate 
(Fig. 75) a portion of type material. Cells up to 15 um 
long and 7 um wide and spores up to 11 „m long and 


574 Rhodora [Vol. 75 


6 um wide have been found and thus extend the maximum 
limits recorded for this material by Collins (1908) and 
Taylor (1937, 1957). Erect filaments of more than 8 cells 
and cells over 6 diameters long were not encountered. 


The two other collections were found growing on Aspara- 
gopsis and showed over all agreement with the type, 
although a number of cells were over 15 „m long and 
sporangia occurred on both prostrate and erect filaments. 
The relationship of these collections to Colaconema ameri- 
cana Jao is discussed under that taxon, which is regarded 
here as a “Species Inquirendae.” 


Critical comparisons of the type collections of C. minima 
and Chantransia emergens Rosenvinge (1909, p. 128, Fig. 
55) have resulted here in regarding the two taxa as con- 
specific. They agree in all essential respects (Figs. 74-76: 
Table 5). Jao (1936, p. 247, pl. 18, Figs. 7, 7B) recorded 
the latter from New England waters, and Taylor (1937, 
p. 226; 1957, p. 217) separated it from Colaconema minima 
on the basis of slight differences in spore width. A com- 
parison of the type collections indicates that this apparent 
difference is of no taxonomic significance since spores 
3-5 um wide occur in both taxa. 


The relationships of Colaconema minima to other audo- 
uinelloid algae of similar morphology require considerable 
clarification. Acrochaetium endophyticum Batters (1896, 
p. 386; see also Baardseth 1941, p. 45, Figs. 19E-G) [non 
Rhodochorton endophyticum Kylin 1907, p. 188, Fig. 40 = 
Acrochaetium kylinti Hamel 1927, p. 93; 1928, p. 187 [nec 
Liagorophila endophytica Yamada 1944, p. 16, Fig. 4 (see 
also Abbott 1966)] appears to have a virtually identical 
morphology, and when the types can be compared, the 
two taxa will almost certainly prove to be conspecific, with 
the specific epithet *endophyticum" having priority. Like- 
wise Colaconema americana Jao and a number of other 
similar taxa almost certainly will be relegated to con- 
specificity when critical comparisons of types can be made. 


1973] Audouinella — Woelkerling 575 


11. Colaconema secundata (Lyngbye) Woelkerling 1973: 
94, Figs. 7-8. Figs. 77-83. 

Callithamnion daviesii var. secundatum Lyngbye 1819: 
129, pl. 41, Figs. 84-6. 

Acrochaetium luxurians (J. Agardh) Naegeli 1861: 405. 

Callithamnion luxurians J. Agardh 1851: 14. Hall 1876: 
111. Harvey 1853: 242. Jordan 1874: 197; 1874a: 488. 

Chantransia luxurians (J. Agardh) Kylin 1907: 117, 
Fig. 26. 

Acrochaetium secundatum (Lyngbye) Naegeli 1861: 
405. Collins 1906: 194. Davis 1913: 813; 1913a: 477. 
South 1970: 1. South and Cardinal 1970: 2079. Taylor 
1937: 230, pl. 31, Figs. 1-3. 

Callithamnion secundatum (Lyngbye) C. Agardh 1828: 
187. 

Ceramium secundatum (Lyngbye) C. Agardh 1824: 132. 

Chantransia secundata (Lyngbye) Thuret In LeJolis 
1863: 106. Collins 1900: 49; 1911: 276. Collins, Holden, 
and Setchell 1903: 1088. Davis 1913a: 462, 473, 474. Far- 
low 1875: 376; 1876: 705. Hylander 1928: 158. Kylin 
1910: 28. 

Chromastrum secundatum (Lyngbye) Papenfuss 1945: 
25223. 


Kylinia secundata (Lyngbye) Papenfuss 1947: 487. 
Edelstein and McLachlan 1966a: 1052, Fig. 18. Hehre 
and Mathieson 1970: 206, 237. Mathieson et al. 1969: 132. 
Taylor 1957: 214, pl. 31, Figs. 1-3. 

Acrochaetium subsimplex Levring 1953: 473, Figs. 


‘According to the “Programme and Abstracts of the Twenty-first 
Annual Meeting of the British Phycological Society" (3-5 January 
1973), W. J. Borsje, Vrije University, Amsterdam, reported sexual 
stages of Acrochaetium virgatulum in culture. If his plants are the 
same as New England plants (which, no doubt, will prove to be the 
case), Colaconema secundata (syn. Acrochaetium virgatulum) will 
then be transferred to the genus Audouinella as A. secundata (Lyng- 
bye) comb. nov. Borsje’s abstract also appears in Br. phycol. J. 
8: 204-5, 1978. 


576 Rhodora [Vol. 75 


10A-D, 11. Non Rhodochorton subsimplex (Harvey) 
DeToni 1897: 1515. 

Acrochaetium tenuissimum (Collins) Papenfuss 1945: 
319. 

Chantransia tenuissima (Collins) Kylin 1941: 5, Figs. 
le, f. 

Colaconema tenuissima (Collins) Woelkerling 1971: 
51, Fig. 21. 

Rhodochorton tenuissimum (Collins) Drew 1928: 170, 
pl. 38, Figs. 26, 27. 

Acrochaetium virgatulum (Harvey) Bornet 1904: XXII. 
Collins 1906: 193. Davis 1913: 813; 1913a: 477. Doty 
1948: 264. Edelstein et al. 1970: 625. South and Cardinal 
1970: 2079. Taylor 1937: 230. Taylor In Lewis 1924: 
214. 

A. virgatulum f. luxurians (J. Agardh) Collins 1906: 
194. Collins, Holden, and Setchell 1906: 1393. Hylander 
1928: 159, Taylor 1937: 230, pl. 31, Figs. 4-7. 

A. virgatulum f. tenuissimum (Collins) Collins 1906: 
194. 

Callithamnion virgatulum Harvey In Hooker 1833: 349. 
Jordan 184a: 17. 

Chantransia virgatula (Harvey) Thuret In LeJolis 1863: 
106. Collins 1880: 162; 1894: 233; 1900: 49; 1911: 276. 
Davis 1913a: 462, 473, 474. Farlow 1875: 376; 1876: 705. 
Farlow, Anderson, and Eaton 1881: 157. Grier 1925: 296. 
Jelliffe 1904: 98. Hylander 1928: 158. Kylin 1910: 28. 
Rosenvinge 1909: 109, Figs. 37-41. 

C. virgatula var. luxurians (J. Agardh) Rosenvinge 
1909: 110. Collins 1911: 276. 

C. virgatula f. tenwissima Collins In Collins, Holden, 
and Setchell 1900: 741. 

Chromastrum virgatulum (Harvey) Papenfuss 1945: 
323. Doty 1948: 264. 

Kylinia virgatula (Harvey) Papenfuss 1947: 437. Hehre 
and Mathieson 1970: 206, 237. Mathieson et al. 1969: 132. 
Taylor 1957: 214. 

K. virgatula f. luxurians (J. Agardh) Collins 1906: 194. 


1973] Audouinella — Woelkerling 577 


Hehre and Mathieson 1970: 206, 237, Taylor 1957: 214, 
pl. 31, Figs. 4-7. 

Rhodochorton virgatulum (Harvey) Rosenvinge 1935: 7. 

Trentepohlia virgatula (Harvey) Farlow 1881: 109, pl. 
X, Fig. 3. Collins 1888: 312; 1888a: 10. Martindale 1889: 
100. Pike 1886: 109. 

T. virgatula var. secundata (Lyngbye) Farlow 1881: 109. 
Pike 1886: 109. 

Acrochaetium daviesii auct, non. (Dillwyn) Naegeli: 
Collins 1906: 194 (pro parte?). 

Callithamnion daviesii auct. non. (Dillwyn) Lyngbye: 
Harvey 1853: 243. 

Chantransia daviesii auct. non. (Dillwyn) Thuret In 
LeJolis: Collins 1880: 162 (pro parte?) ; 1894: 233 (pro 
parte?) ; 1900: 49 (pro parte?) ; 1911: 276 (pro parte?). 

Trentepohlia daviesii auct. non. (Dillwyn) Pringsheim: 
Pringsheim 1862: 26, Taf VIII, Figs. 1-6. 

Acrochaetium flexuosum auct. non. Vickers: Collins 1906: 
192. Collins, Holden, and Setchell 1910: 1696. Taylor 1937: 
209; 1957: 219. 

Chantransia flexuosa auct. non. (Vickers) Collins: Col- 
lins 1911: 186. 

Acrochaetium sagraeanum auct. non. (Montagne) Bor- 
net: Boergesen 1915: 35 (pro parte). Bornet 1904: XXI 
(pro parte). Collins 1905: 231 (pro parte) ; 1906: 192 (pro 
parte). Hamel 1927: 77, 99 (pro parte); 1928: 173, 191 
(pro parte). Hylander 1928: 159 (pro parte) Jao 1936: 
244 (pro parte). Papenfuss 1945: 311 (pro parte). Taylor 
1937: 233 (pro parte) ; 1957: 220 (pro parte) ; 1960: 309 
(pro parte). 

Chantransia sagraeana auct. non. (Montagne) DeToni: 
1924: 51 (pro parte). 


Plants epiphytic or epizoic, caespitose, up to 3 mm tall; 
original spore non-persistent. Prostrate system at first 
forming a small parenchymatous disc usually composed of a 
central cell and 3-4 peripheral cells; later proliferating in 
some plants to form a unistratose (rarely a partially bistra- 
tose) pseudoparenchymatous disc, Erect filaments sparsely 


578 Rhodora [Vol. 75 


or moderately to freely and irregularly branched; laterals of 
variable length and some populations only 1-5 cells long. 
Cells cylindrical 8-15 (-20) um wide and 15-100 «m long, 
L/D (1-) 2-6; each containing an axial or parietal stellate 
chromoplast and a central pyrenoid. Unicellular hairs up 
to 300 um long abundant in some populations,, pseudolateral 
or terminating 1-3 celled lateral branches ; sparse or absent 
in other populations. 

Monosporangia ovoid, 10-20 „m wide and 15-26 (-32) „m 
long, solitary or in pairs, sessile or stalked, scattered on 
the erect filaments or sometimes densely erowded on short 
lateral branches. 

Other reproductive structures not observed. 

Type locality: Kvivig, Faeroes Islands (on “Conferva 
rupestris”). 

Holotype: C. 

Distribution: Atlantic Coast of North America, Austra- 
lia, Canary Islands, Europe, Sargasso Sea. 

Hosts: A wide variety of algae and on hydroids, 


Specimens examined : 

CONNECTICUT: Black Rock Beacon (Long Island Sound), 20. VII. 
1892, Holden (NY, Holden 646) New Haven, prior to 1853, Hoope) 
(TCD). Woodmont, 27. VII. 1893, Holden (NY). 

MAINE: Eagle Island (Penobscot Bay), 16. VII. 1905, Collins (NY). 
Fox Island, VIII. 1880, Collins (NY). Inner Mark Island, 8. VII. 
1903, Collins (NY, PBA 1088). Kennebec River Mouth, VIII. 1880. 
Booth (NY). Peak's Island, IX. 1874, Farlow (FH). 

MASSACHUSETTS: Eastham, 10. VII. 1907, Collins (Ny). Falmouth 
(Cape Codder Point), 30. X. 1966, Conway et al. (NHA) ; 19. XI. 1969, 
Woelkerling (waw 2237) ; 7. I. 1970, Conway (wyw 2294). Falmouth 
(Monauhant Beach), 12. III. 1969, Conway (wsw 1850); (wyw 1851). 
Falmouth (Waquoit Bay), 21. I. 1969, Conway (wJw 1842); 11. III. 
1969, Conway (wJw 2273); 30. VI. 1969, Conway (WIW 2274); 7. 
VII. 1969, Conway (WJW 1845); 15. IX. 1969, Conway (WJW 1844) ; 
1. X. 1969, Conway (WJW 1841). Gloucester (Niles Beach), IX. 1875, 
Farlow (FH); no date, Farlow (Ny, Algae Exsic. Am. Bor. — Far- 
low, Anderson, and Eaton, No. 157). Hingham, prior to 1853, Brewer 
(tcp). Marblehead, 17. VI. 1902, Collins (NY). Mattapoisett, 30. V. 
1905, Collins (NY); 20. X. 1906, Collins (NY, PBA 1393). Nantucket 
(Brauts Point), 16. VIII. 1898, Collins (NY). Nantucket (West Jetty, 
Nant. Center), 14. IV. 1970, Woelkerling (waw 2476). Nantucket 
(?), prior to 1853, Durkee (TCD). New Bedford (Scontient Point), 


1973] Audouinella — Woelkerling 519 


13. VII. 1927, Taylor (NY). Penikese Island, 1. X. 1970. Woelkerling 
(wsw 2808). Sandwich (Cape Cod Canal Jetty), 7. IV. 1968, Kas- 
telowitz (NHA); 18. X. 1970, Woelkerling (wyw 2816). West Fal- 
mouth (Harbor entrance), 10. XII. 1969, Conway (wsw 2131); 6. I. 
1970, Woelkerling (waw 2276). Woods Hole, VII. 1875, Farlow (FH); 
VII. 1903, Collins (Ny); 10. VIII. 1947, Abbott (Abbott 1638); 3. 
VI. 1966, Conway (WJw 2168); 4. II. 1970, Woelkerling (WJW 2316); 
2. VII. 1970, Woelkerling (wJw 2680) ; 29. I. 1971, Woelkerling (WJW 
3299); 16. II. 1971, Woelkerling (wJw 3315). 

NEW HAMPSHIRE: Fort Constitution, 20. IX. 1966, Conwa; and Hehre 
(NHA). Fox Point, 21. VII. 1966, Conway and Shipman (NHA). 
Great Boar's Head, 15. X. 1966, Hehre and Conway (NHA); 11. XII. 
1966, Hehre and Conway (NHA). Little Boar's Head, 14. IX. 1966, 
Hehre (NHA); 25. XI. 1966, Conway and Hehre (NHA). Rye Ledge, 
29. III. 1967 Mathieson and Murphy (NHA). 

NEW JERSEY: Atlantic City, 16. IV. 1892, Morse (Ny, No. 1696 in the 
Phycotheca Boreali Americana); 1. IX. 1925, Rose (NY). 

NEW YORK: Cold Spring Harbor, 21. VII. 1893, Howe (?) (NY). 
Little Neck Bay (Long Island), 7. IV. 1966, Keenan (NHA). 

RHODE ISLAND: Napatree Point, 9-11. X. 1872, Eaton (NY); (FH). 
CALIFORNIA: San Pedro, XI. 1898, Monk (FH, type of Chantransia 
virgatula F. tenuissima Collins; ADU A32706, isotype; NY, isotype; 
WIS, isotype). 

EUROPE: England (Torquay), prior to 1833, Griffiths (TCD, type of 
Acrochaetium virgatulum (Harvey) Bornet). Faeroes Islands (Kvi- 
vig), 19. VI. 1817, ? (C, Herb. Lyngbye, type). Sweden (Kattegat 
Channel), no date, ? (LD 35117, type of Callithamnion luxurians J: 
Agardh). 

AUSTRALIA: Musselroe Bay (Tasmania), 7. II. 1948, Levring (ADU, 
A19846, isotype of Acrochaetium subsimplex Levring). 


Colaconema secundata is distinguishable from other 
New England audouinelloid algae in 1) having spores 
germinating to form a distinctive, orbicular parenchy- 
matous group of cells (Fig. 77) which later may proliferate 
(Figs. 80-81), and in 2) having distinctly stellate chromo- 
plasts (Fig. 83), each with one pyrenoid, in cells of the 
erect filaments. In addition, most, but not all, New Eng- 
land populations possess a somewhat twiggy (virgate) 
habit as a result of having numerous 1-3 (-5) celled lateral 
branches terminated by unicellular hairs (Fig. 82). 

Colaconema secundata occurs throughout the year in 
New England on a variety of algae or invertebrates in the 
sublittoral, and occasionally appears in the lower littoral, 


580 Rhodora [Vol. 75 


particularly on Porphyra. It probably is found in greater 
numbers than any other audouinelloid alga in New England 
waters and most frequently is encountered as a dense fringe 
on blades of the sea grass Zostera. 

Rosenvinge (1909) has provided an excellent account of 
this species (as Chantransia virgatula) and of the varia- 
tion it exhibits; his concept of this species is recognized 
here (see also Hamel 1927, 1928). As noted previously 
(Woelkerling 1973, p. 96), the specific epithet *secundata" 
has nomenclatural priority over the specific epithet *'vir- 
gatula". The form illustrated in this study (Fig. 82) is 
by far the most common in New England waters, although 
individuals of all forms discussed by Rosenvinge (1909) 
have been observed. 

Most previous aecounts (e.g. Hamel 1927, Kylin 1907, 
Rosenvinge 1909) state that the germinating spore divides 
to form a central cell and three surrounding cells, In all 
New England plants examined, four cells surround the 
central cell (Fig. 77); this difference, however, does not 
appear to be of any taxonomic consequence. 

Some variation in the number of cell layers in the basal 
disc has also been reported. Hamel (1927), Kylin (1907), 
and Rosenvinge (1909) all record only single layered basal 
discs. Boergesen (1902) separates Chantransia secundata 
from C. virgatula on the basis that the former always has 
à two layered basal disc while the latter has only a single 
layered disc, Collins (1906) states that the basal dises are 
usually two layers thick; Taylor (1937, 1957) follows 
Boergesen (1902) in recognizing two species in his keys 
(Taylor 1937, p. 227; 1957, p. 211), but states in his de- 
scriptions (Taylor 1937, p. 230; 1957, p. 214) that the 
base is “. .. a multicellular disc 1-2 layers or more in thick- 
ness .... 

During this study, several dises (Fig. 80) that are 
partially two layered have been seen, but the majority are 
single layered. None of the sort illustrated by Boergesen 
(1902, p. 350, Fig. 51) or Taylor (1957, pl. 31, Fig. 3) 
have been encountered. It also appears evident that some 


1973] Audouinella — Woelkerling 581 


variation can occur in the number of layers in the base 
within single populations, and therefore, this variation has 
little taxonomic significance. Indeed, all plants examined 
in the type collections of Colaconema secundata and the 
other taxa here considered conspecific have single layered 
basal discs. 

Only monosporangial plants have been observed during 
the present study; tetrasporangial individuals have, how- 
ever, been reported in New England by Hehre and Mathie- 
son (1970) and Taylor (1937, 1957). Kylin (1907, 1944) 
and Rosenvinge (1909), among others, have recorded them 
from Europe. Sexual stages remain unknown. 

Collins in Collins, Holden, and Setchell (1900, No. 741) 
described and distributed Chantransia virgatula f. tenuis- 
sima based on a collection of Monks from San Pedro, 
California. Later Drew (1928, p. 170, pl. 38, Figs. 26, 27) 
raised the taxon to specific rank (as Rhodochorton tenuis- 
simum (Collins) Drew), primarily on the bases of some- 
what narrower filaments, lack of hairs, and sessile and not 
opposite sporangia. 

The type collection of this species in FH as well as 
several isotypes (ADU, NY, WIS) have been examined and 
found to agree well with the types of Colaconema secun- 
data and of Callithamnion virgatulum. Type collection 
plants of all three taxa have the same characteristic de- 
velopment of the basal disc, which in some cases have 
produced proliferations. The type plants of Chantransia 
f. virgatula tenuissima examined also possess (in contrast 
to what Drew (1928) found in her material) unicellular 
hairs and do have some stalked monosporangia and some 
sporangia in an opposite arrangement. In agreement with 
Drew, most sporangia are sessile and scattered, Drew re- 
ported a parietal chromoplast, but this was not evident in 
the type material examined. Since there appear to be no 
other reliable criteria of separation, Chantransia virgatula 
f. tenuissima and Colaconema secundata are considered here 
to be conspecific as was suspected earlier (Woelkerling 
1971, p. 52). 


582 Rhodora [Vol. 75 


Acrochaetium subsimplex Levring (1953, p. 473, Figs. 
10A-D, 11) likewise is conspecific with Colaconema secun- 
data (see Woelkerling 1971, p. 51). 

A number of collections referred to Audouinella daviesi 
by Collins (see discussion under that taxon) have been 
examined and found to contain only (?) plants of Cola- 
conema secundata. If one assumes that these collections 
do not contain a mixture of two taxa (such a mixture was 
not found during the present study), Collins’ material has 
been misidentified and has been treated accordingly. 

Likewise one collection (NY, Holden collection No. 646) 
hitherto referred to Acrochaetium sagraeanum (see Collins 
1905, p. 231 and discussion of that taxon under “Species 
Excludendae") has been found to contain only plants of 
Colaconema secundata; the literature citations have been 
corrected accordingly. 

Two collections in NY (one from Massachusetts and one 
from New Jersey, the latter being distributed as specimen 
1696 in the Phycotheca Boreali Americana) hitherto re- 
ferred to Acrochaetium (= Chantransia) flexuosum by 
Collins (1906, 1911), Collins, Holden, and Setchell (1910), 
and Taylor (1937, 1957) have been examined and found 
to contain only young plants of Colaconema secundata. 
Parenchymatous discs typical of young plants of C. secun- 
data are abundant in both collections and in the largest 
plants these discs have become more or less obscured by 
subsequent cell division in prostrate system cells. Relatively 
few sporangia were present, but these fell within the range 
of dimensions found among other New England popula- 
tions. 

Specimens from Rhode Island referred by Taylor (1937, 
1957) to Acrochaetium flexuosum have not been available 
for examination and their status therefore remains in 
doubt. 


OTHER NEW ENGLAND RECORDS 


In addition to the species already discussed, several 
other audouinelloid taxa have been reported from New 


1973] Audouinella — Woelkerling 583 


England waters. As a result of the present investigation, 
the occurrence of four of these taxa, discussed below under 
“Collections Inquirendae", is surrounded by doubt. The 
taxonomic status of two other taxa, discussed below under 
“Species Inquirendae", is open to question, and the type 
collection of a seventh taxon, discussed below under 
“Species Excludendae", has been referred to the Chloro- 
phyta. 


Collections Inquirendae 


Acrochaetium attenuatum (Rosenvinge) Hamel 1927: 99; 
1928: 192. Edelstein et al. 1970: 625. Jao 1936: 244, pl. 
12, Fig. 1. Papenfuss 1945: 308. South 1970: 1. South 
and Cardinal 1970: 2079. Taylor 1937: 236, pl. 34, Fig. 1; 
1957: 222, pl. 34, Fig. 1. 

Chantransia attenuata Rosenvinge 1909: 106, Fig. 35. 
DeToni 1924: 56. 

Rhodochorton attenuatum (Rosenvinge) Nakamura 1944: 
103, Fig. 3. 

For detailed morphological accounts of this taxon, see 
Nakamura (1944) and Rosenvinge (1909). 

Type locality: Jelstrup, Limfjord, Denmark. 

Type: € (Rosenvinge 3883). 

Distribution: Northern Europe, Eastern Canada, Japan. 

Hosts: A variety of algae. 

Specimens examined: See below. 

This taxon has been recorded once in New England by 
Jao (1936) from Norton Point, Martha's Vineyard, Massa- 
chusetts and subsequently reported by Taylor (1937, 
1957). The collection upon which this record is based is 
apparently represented in MICH by a prepared slide bear- 
ing the number Woods Hole 280. (This same slide contains 
type collection specimens of two other taxa [Acrochaetium 
radiatum Jao and A. microfilum Jao] which are regarded 
here as conspecific with Colaconema humilis and Audout- 
nella microscopia respectively). 

The MICH slide has been examined during this study 
and found to contain two plants somewhat similar to those 


584 Rhodora [Vol. 75 


described by Jao (1936), but the material was too frag- 
mentary and too young to make any definite identification, 
although the specimens showed some features similar to 
young plants of Audouinella saviana. The taxonomic status 
of these plants therfore remains in doubt as does the occur- 
rence of A. attenuatum in New England waters. 

Likewise the record of Edelstein et al. (1970, p. 265) 
requires further investigation. 


Audouinella efflorescens (J. Agardh) Papenfuss 1945: 
326; 1947: 438. Edelstein and McLachlan 1968: 993, Figs. 
1-2. Taylor 1957: 225. 

Acrochaetium efflorescens (J. Agardh) Naegeli 1861: 
405. Hamel 1927: 103; 1928: 196. Taylor 1937: 236. 

Callithamnion efflorescens J. Agardh 1851: 15; 1876: 10. 

Chantransia efflorescens (J. Agardh) Kjellman 1875: 
14; 1883: 129. Kylin 1906: 113, Figs. 1-5; 1907: 119. 
Rosenvinge 1909: 134, Figs. 61-64. 

Grania efflorescens (J. Agardh) Kylin 1944: 26, Fig. 
24. 

Rhodochorton efflorescens (J. Agardh) Drew 1928: 151. 
Rosenvinge 1935: 7. 

For detailed morphological accounts of this taxon, see 
Kylin (1906) and Rosenvinge (1909). 

Type locality: Kattegat Channel, Sweden. 

Type: LD 35129. 

Distribution: Arctic, Eastern Canada, Northern Europe. 

Hosts: A variety of algae and invertebrates; also on 
rock. 


Specimens examined: Sweden: Kattegat, 13. VIII. 1833, ? (LD 
35129, type of Callithamnion efflorescens J. Agardh). 


This taxon has been recorded (as Chantransia efflores- 
cens) on Rhodymenia from Gay Head (Martha’s Vine- 
yard) Massachusetts by Farlow (1877, p. 245), but neither 
he nor anyone else apparently has made further reference 
to this collection. Moreover, no plants of Awdowinella ef- 
florescens from Gay Head on Rhodymenia could be located 
in FH or NY during this investigation, and this record of 


1973] Audouinella — Woelkerling 585 


the taxon in New England must therefore remain in doubt. 

Farlow (1881, p. 109) also records a Chantransia efflo- 
rescens Thuret from Gay Head on Cystoclonium. Whether 
this is the same specimen as the one discussed above is un- 
certain, but plants from the Cystoclonium collection, the 
major portion of which is in FH (a fragment cut from the 
sheet is in the Collins material in NY), have been examined 
and found to contain sexual plants of Auwdouinella daviesii 
rather than specimens of A. efflorescens. Carposporophytes 
of the two are easily distinguishable because carospores 
are only terminal in A. daviesii but occur in rows of 2-3 in 
A. efflorescens. The notes of Taylor (1937, p. 236; 1957, 
p. 225) refer to Farlow's material on Cystoclonium, which 
is regarded here to belong toA. daviesii (q.v. for further 
comments). 

Audouinella efflorescens is a distinctive species with its 
spirally twisted chromoplasts without pyrenoids and its 
seriate carpospores, and future study may show it to be a 
component of the New England flora. It apparently has 
recently been found in Nova Scotia (Edelstein and Mc- 
Lachlan 1968). 

Audouinella spetsbergense (Kjellman) comb. nov. Figs. 
61-63. 

Rhodochorton spetsbergense (Kjellman) Kjellman 1883: 
187. DeToni 1903: 1511. 

Thamnidium spetsbergense Kjellman 1875: 31, Figs. 
11-12. 

Rhodochorton | penicilliforme (Kjellman) Rosenvinge 
1894: 66, Fig. 9; 1923-24: 388, Figs. 325-327. Boergesen 
1902: 389, Fig. 60. Collins 1906a: 160. Drew 1928: 176. 
Edelstein et al. 1967: 200, Figs. 6, 28. Kylin 1907: 188; 
1925: 44, Figs. 26a-c; 1944: 28. Taylor 1987: 239; 1957: 
225. 

Rhodochorton mesocarpum (Carm.) Kjellman var. peni- 
cilliformis (Kjellman) Kjellman 1883: 187, pl. 16, Figs. 
6-7. Rosenvinge 1893: 792. 

Thamnidium mesocarpum (Carm.) Kleen f. penicilli- 
formis Kjellman 1875: 30; 1877: 25; 1877a: 23. 

Plants epiphytic, caespitose, up to 2.0 mm tall; original 


586 Rhodora [Vol. 75 


spore non-persistent. Prostrate system consisting of an 
irregularly discoid mass of pseudoparenchymatous fila- 
ments. Erect filaments sparingly to moderately and ir- 
regularly branched. Cells cylindrical, 8-13 „m wide and 
(10-) 20-35 (-50) um long, L/D (1-) 1.5-4 (4.5) ; each cell 
containing a number of discoid to irregularly shaped chro- 
moplasts without pyrenoids. Unicellular hairs not observed. 

Tetrasporangia ovoid, 25-35 um long and 16-25 „m wide, 
single or occasionally in pairs (rarely in groups of 3), 
sessile or on stalks, scattered over the erect filaments or 
occasionally terminal on lateral branches. 

Other reproductive structures not observed. 


Type locality: Fairhavn, Spitzbergen (79? 49' N.). 

Type: UPS. 

Distribution: In North America, from Washington state 
northwards on the Pacific coast and Rhode Island north- 
wards (?, see discussion) on the Atlantic Coast; also re- 
ported from northern Europe. 

Hosts: Various algae and hydroids. 

Specimens examined: SPITZBERGEN: Fairhavn Is. (79? 49' N.), 19. 
VIII. 1872, Kjellman (ups, type). Fairhavn Is. (79° 41' N.), 12. 
VIII. 1872, Kjellman (ups, type of R. penicilliforme (Kjellman) 
Rosenvinge). 


This taxon has been recorded (as Rhodochorton penicil- 
liforme) from Rhode Island (Collins 1906a, p. 160) and 
northern Massachusetts (Taylor 1937, p. 239; 1957, p. 225). 
Collins material has not been found either in FH or NY 
and the Massachusetts material has not been available for 
study. (The taxonomic description given above is based 
on data gathered from plants in the type collection; see 
Rosenvinge 1923-24 for a more complete account). More- 
over, no additional plants of this taxon have been col- 
lected during the present investigation and three collections 
in NHA examined have been found to contain only plants 
of other species. Thus, the occurrence of this taxon in 
New England waters is open to question, and further in- 
vestigation appears necessary. It has recently been re- 
ported from Nova Scotia (Edelstein et al. 1967), but did 


1973] Audouinella — Woelkerling 587 


not appear in a recent survey of Campobello Island, New 
Brunswick (Stone et al. 1970). 

Kjellman (1875) described two new taxa from collec- 
tions made on Fairhavn Island, Spitzbergen: Thamnidium 
spetsbergense Kjellman and T. mesocarpum (Carmichael) 
Kjellman forma pencilliformis Kjellman. The latter was 
given species status by Rosenvinge (1894, p. 66). The type 
specimens of both taxa, preserved on microscope slides in 
UPS, have been examined during this study and found to 
be identical in all respects including habit, cell dimensions, 
spore dimensions, and chromoplast morphology. Conse- 
qently, they are here considered conspecific, with the spe- 
cific epithet “spetsbergense” having nomenclatural priority. 
It has been referred to the genus Audowinella on the basis 
of Rosenvinge’s (1923-24, p. 388, Fig. 27) record of sperm- 
atangia. Culture studies may show that A. spetsbergense 
has a diphasic life cycle similar to that of A. purpurea (see 
West. 1969). 

The relationships of A. spetsbergense and A. purpurea 
require clarification. They differ from one another pri- 
marily in their habit (epibiotic vs. primarily saxicolous) 
and type of prostrate system (pseudo-parenchymatous vs. 
filamentous) and since none of the characters appear to be 
taxonomically reliable (see Woelkerling, 1971), the two 
entities may represent ecological variants of a single spe- 
cies. Because intermediate forms have not been found dur- 
ing this study, they are maintained as distinct taxa for the 
present, 


Colaconema polyides (Rosenvinge) Woelkerling 1971: 
19, Figs. 19, 27A. 

Acrochaetium polyides (Rosenvinge) Boergesen 1915: 
59. Kylin 1944: 26. Papenfuss 1945: 317. 

Chantransia polyides Rosenvinge 1909: 132, Figs. 59-60. 
Levring 1935a: 460, Fig. 2. 

For detailed morphological accounts of this taxon, see 
Rosenvinge (1909) and Woelkerling (1971). 

Type locality: Tonneberg Banke, Denmark. 

Type: C. 


588 Rhodora [Vol. 75 


Distribution: Denmark, South Australia, Tasmania. 
Hosts: Codium (Chlorophyta), Polyides (Rhodophyta). 
Specimens examined: See below. 


This taxon has been reported (as Acrochaetium polyides) 
from New Hampshire (Hehre and Mathieson 1970, p. 205) 
and Nova Scotia (Edelstein and McLachlan 1966, p. 38, 
Fig. 2; Edelstein et al. 1970, p. 625; South and Cardinal 
1970, p. 2078), apparently in a sterile condition. 

The New Hampshire collection and one Nova Scotia col- 
lection have been examined; in both cases the meagre 
amount of sterile Audouinella-like filaments present could 
not be identified to species with any certainty at all. The 
taxonomic affinity of these specimens and the occurrence 
of A. polyides in New England and Nova Scotia therefore 
remains doubtful. It seems likely that these collections have 
been referred to A. polyides solely on the basis of the type 
of host organism (see Edelstein and McLachlan 1969, 
p. 555), which, as noted above, is an unreliable criterion 
of systematic separation. 


Species Inquirendae 

Colaconema americana Jao 1936: 237, pl. 13, Fig. 8. 
Levring 1937: 94; 1953: 489. de Valera 1939: 3. Woel- 
kerling 1971: 42. 

Acrochaetium americanum (Jao) Papenfuss 1945: 312. 

Type locality: Gay Head, Martha’s Vineyard, Massachu- 
setts. 

Holotype: MICH, Woods Hole No. 272. 

Distribution: Type locality; Southern Australia. 

Hosts: Asparagopsis sp. 

The type collection of this species has not been available 
for examination. Two other collections (wJw 2693, wJw 
2699) of Asparagopsis containing an endophytic audouinel- 
loid alga have been examined and referred to Colaconema 
minima because of overall morphological agreement with 
that taxon. The cell and spore dimensions of plants in these 
collections are somewhat smaller than those reported by 
Jao (1936) for the type collection of C. americana (Table 


1973] Audouinella — Woelkerling 589 


6), and no greatly swollen cells were found, thus placing 
further doubt (see Woelkerling 1971, p. 42) upon the taxo- 
nomic significance of that character. Moreover, the mono- 
spores of the collections examined may or may not rest in 
cup-like bases (Fig. 74), thus casting doubt upon the taxo- 
nomic validity of that character also. Upon re-examination, 
plants in the type collection of C. americana will almost 
certainly be found to be conspecific with C. minima. 

The rather large number of audouinelloid taxa (includ- 
ing Colaconema minima and C. americana) whose thallus 
consists of a creeping prostrate system devoid of or bearing 
few celled erect filaments present more taxonomic prob- 
lems and difficulties with identification than any other 
group within the complex. Such taxa have often been de- 
scribed on the basis of single collections of one or a few 
plants and have been distinguished on minor differences in 
cell or spore dimensions, spore arrangement, branching 
and other morphological characters whose systematic value 
is dubious (see Woelkerling 1971). Another whole series 
of such taxa are based on differences in the species of host 
organism (see Baardseth 1941, p. 46), and host specificity 
is likewise of doubtful taxonomic value (Woelkerling 1971, 
DEED 

Until the type collections of all these audouinelloid algae 
can be critically compared and the taxa redefined (probably 
with considerable consolidation of present forms) on firmer 
morphological grounds, taxonomic procedure dictates a con- 
tinued recognition of a rather large assemblage of poorly 
known and rather confusing entities. 

Rhodochorton entozoicum (Reinsch In Giard) DeToni 
1903: 1514. 

Callithamnion entozoicum Reinsch In Giard 1890: 262. 

Type locality: Massachusetts. 


Type: BM. 
Distribution: Apparently known only from the type lo- 
cality. 


Hosts: Marine invertebrates. 
The type and only known New England collection of this 


590 Rhodora [Vol. 75 


taxon has not been available for examination. Reinsch 
(1879) first described and illustrated material of this taxon 
from sponges and bryozoans, but as noted by Collins 
(1883), Reinsch never assigned a name to it. Giard (1890), 
however, formally described the taxon under the specific 
epithet of *entozoicum," and this was subsequently re- 
corded by DeToni (1903). 

A re-examination of the type collection will almost cer- 
tainly show that Rhodochorton entozoicum is conspecific 
with Colaconema membranaceum. As noted by Collins 
(1883), DeToni (1903), and Giard (1890) himself, the 
two taxa are scarcely distinguishable from one another, 
and Hamel (1927, p. 59; 1928a, p. 152) regarded the two 
as conspecific. Hamel, however, gives no indication of hav- 
ing compared the type collections of the two taxa, and the 
present investigator prefers to defer final judgment until 
such a comparison can be made. Hamel also regards ma- 
terial of Giard (1890) from Wimeraux as belonging to C. 
membranacea. 


Species Excludendae 


Acrochaetium sagraeanum (Montagne) Bornet 1904: 
xxi — Cladophora sagraena Montagne 1856: 459. 


Bornet (1904) united under the name of Acrochaetium 
sagraeanum specimens from Cuba (Montagne’s type, which 
Bornet noted was sterile), Barbados (see Vickers 1905), 
California, and Connecticut (see Collins 1905). Later, 
Collins (1906) and Collins and Hervey (1917) referred 
specimens from Florida and Bermuda to this taxon. 

Montagne’s type has been re-examined during this study 
and found to belong to the Chlorophyta and probably to the 
genus Cladophora itself as originally suggested by Mon- 
tagne (1856). The specimens have a distinctive green 
color, lack pit connections and do not possess monospo- 
rangia or other reproductive organs, all of which strongly 
militate against placing Montagne's plants in the Awu- 
douinella complex as suggested by Bornet. 

New England plants referred to Acrochaetium sagraea- 


1973] Audouinella — Woelkerling 591 


num were originally collected by Holden (see Collins 1905, 
1906) and are deposited in FH and/or NY. Four of these 
(Holden collection number 17,701 [= Collins, Holden and 
Setchell 1895: 39], 708 and 757) contain plants of Au- 
douinella saviana. A fifth herbarium sheet (Holden collec- 
tion number 646) contains plants of Colaconema secundata. 
The sixth collection cited by Collins (1905), Holden 750, 
could not be located, and its taxonomic status remains in 
doubt. 

Plants in PC from California (which may represent frag- 
ments of vc 93071 or uc 789916 deposited at Berkeley) 
referred by Bornet to Acrochaetium sagraeanum have been 
examined and are referred here to Awudowinella daviesit. 
Likewise plants from Barbados (Bornet 1904, Vickers 
1905) are referred here to A. daviesii. Specimens from 
Bermuda (Collins and Hervey 1917); Collins, Holden and 
Setchell 1917, No. 2181), however, belong to A. saviana. 

Collins (1906, p. 192) cites a collection of Acrochaetium 
sagraeanum from Florida, but no specimens could be lo- 
cated in FH or NY, and its status remains uncertain. 

A number of literature references relating to one or more 
of the above collections can now be clarified. The name 
Acrochaetium sagraeanum (Montagne) Bornet as used by 
Boergeson (1915, p. 35), Collins (1906, p. 192), Hamel 
(1927, pp. 77, 99, Fig. 46; 1928, pp. 171, 193, Fig. 46), Jao 
(1936, p. 244), and Papenfuss (1945, p. 311) refers to the 
entire assemblage of collections described by Bornet (1904, 
p. XXI). DeToni (1924, p. 51) employed the name Chan- 
transia sagraeana (Montagne) DeToni for Bornet’s as- 
semblage. The collection upon which Hamel (1927, 1928) 
based his illustrations has not been determined. 

Vickers (1905, p. 60) relates to the Barbados collection 
while Collins (1905, p. 231), Collins, Holden and Setchell 
(1895, No. 39), Hylander (1928, pp. 158, 159), and Taylor 
(1937, p. 233; 1957, p. 220) pertain to the Connecticut 
material. The Bermuda collections are cited in Collins and 
Hervey (1917, p. 97) and Collins, Holden and Setchell 
(1917, No. 2181), Taylor (1960, p. 309) includes both the 


592 Rhodora [Vol. 75 


Bermuda and Barbados collections. Corrected citations for 
these literature references have been listed under the syn- 
onomy of Audouinella daviesii, A. saviana, and Colaconema 
secundata. 


EASTERN CANADIAN AND ARCTIC RECORDS 


According to South and Cardinal (1970), 28 species of 
audouinelloid algae have been recorded from the eastern 
Canadian coast from Cape Chidley, Labrador southwards 
to the New Brunswick-Maine border. 


In addition to those taxa listed from eastern Canada and 
the Arctic in the descriptive catalogues of Taylor (1957, 
1957), seven other taxa have been reported in this region, 
chiefly as a result of the Nova Scotia studies of Edelstein 
and McLachlan (1966, 1968, 1969) and of Edelstein et al. 
(1967, 1970). These include Acrochaetium bornetii Papen- 
fuss, A. endozoicum (Darbishire) Batters, A. inclusum 
(Levring) Papenfuss, A. kylinii Hamel, A. ralfsiae Boerg- 
esen, A. subtilissimum (Kuetzing) Hamel, and Kylinia 
cytophaga (Rosenvinge) Kylin. 

A number of these records appear highly questionable. 
Thus, for example, Edelstein and McLachlan (1969, p. 558) 
record Acrochaetium kylinii, A. ralfsiae, and A. subtilis- 
simum from Nova Scotia (the only records for this region) 
on the basis of some endophytic portions of sterile speci- 
mens with the comment *. .. we regard these species as 
preliminary records only." Other records of similar merit 
also occur (e.g. the Canadian record for A. polyides [Edel- 
stein and McLachlan 1966, p. 38] apparently is based on 
sterile material), and in other cases taxa have been identi- 
fied on the basis of type of host organism (Edelstein and 
MeLachlan 1969, p. 555) or similar criteria which are of 
dubious taxonomic significance. 

Until all of the specimens involved in publication records 
for the Cape Chidley, Labrador to New Brunswich-Maine 
border region can be critically re-examined along with 
other available material from the area (a task beyond the 


1973] Audouinella — Woelkerling 593 


scope of the present paper), the number and distribution 
of audouinelloid algae along these coasts must necessarily 
remain uncertain. South and Cardinal (1970, p. 2079, foot- 
note 3) also state that many of the species require further 
investigation. 

Only scattered records of audouinelloid algae north of 
Cape Chidley, Labrador have appeared in the literature and 
most of these are summarized in the papers of Kjellman 
(1883) and Lund (1959) and the catalogues of Taylor 
(1957, 1957). Until relevant herbarium specimens and 
additional field collections become available for study, the 
nature and distribution of audouinelloid algae in arctic 
regions will likewise remain uncertain. 


SUM MARY 


The systematics, morphology, ecology, and distribution of 
marine representatives of the Audouinella complex (Acro- 
chaetiaceae, Rhodophyta) in New England waters (includ- 
ing New York and New Jersey) have been studied, These 
investigations support the taxonomic proposals of Woelker- 
ling (1971) and have resulted in the recognition of two 
genera and eleven species to occur definitely in this region. 
Seven species, known to reproduce sexually are referred to 
Audouinella; the form genus Colaconema contains four New 
England representatives which are unknown in the sexual 
state. 

A number of published records based on misidentifica- 
tions have been corrected and in one instance, the type 
specimen of a taxon [Acrochaetium sagraeanum (Monta- 
gue) Bornet] has been excluded from the Rhodophyta. 
Records of questionable occurence, taxa whose status is in 
doubt, and the state of knowledge of marine audouinelloid 
algae in eastern Canada and the Arctic are considered 
briefly. 


ACKNOWLEDGMENTS 


Sincere thanks are due the curators of the algal collec- 


594 Rhodora [Vol. 75 


tions housed at the following institutions for the loaning of 
relevant herbarium specimens: Atlantic Regional Labora- 
tory, Halifax, Nova Scotia; the Botanical Museum and 
Herbarium, Copenhagen; the Botanical Museum, Lund; the 
Farlow Herbarium, Harvard University; the Herbarium 
Universitatis Florentinae, Firenze, Italy; the Institute of 
Systematic Botany, University of Uppsala; the National 
Museum of Canada; the National Museum of Wales, Car- 
diff, U.K.; the New York Botanical Gardens; the Museum 
National d'Histoire Naturelle, Laboratoire de Cryptogamie, 
Paris; the Rijksherbarium, Leiden, Netherlands; Rutgers 
University; Trinity College, Dublin, Ireland; the Univer- 
sity of Adelaide; the University of California; the Univer- 
sity of Hawaii; the University of Michigan; and the Uni- 
versity of New Hampshire. 

Thanks are also due to Dr. I. A. Abbott, Dr. E. J. Hehre, 
and Mr. R. Stone for the loan of specimens in their personal 
herbaria, to Miss Joan Conway for collecting several spe- 
cimens, and to Dr. R. T. Wilce and Dr. M. R. Carriker for 
arranging for use of facilities at the Marine Biological 
Laboratory in Woods Hole. 

This work was supported in part by National Science 
Foundation Grant GB-13250 to the Marine Biological Lab- 
oratory, Woods Hole, Massachusetts. Publication costs have 
been defrayed by the Norman C. Fassett Memorial Fund. 


ADDENDUM 


After this article had gone to press, Professor Peter S. 
Dixon provided me (personal communication) with some 
additional information about the typification of Audouinella 
daviesti and A. thuretii. 

According to Professor Dixon, the only specimen of Con- 
ferva daviesii in the Dillwyn collections at NMW comes 
from Swansea rather than from Bantry Bay, the type local- 
ity given by Dillwyn. A fragment of the NMW collection 
was sent to me on loan, and an examination of it indicated 
that it was material of C. daviesii. However, since this 
specimen apparently does not come from Bantry Bay, it 


1973] Audouinella — Woelkerling 595 


can at best be regarded only as authentic material rather 
than type material and the statements in the body of this 
paper should be modified accordingly. 


Professor Dixon also informed me that the type material 
of Audouinella thuretii should be dated 5. IX. 1853 (see 
Bornet 1904), but that the earliest material he could find 
in PC is dated 1. IX. 1856. Portions of the latter collection 
have been examined during this study. Until the matter of 
the dates is further clarified, the 1856 material should be 
regarded as authentic rather than type, and statements in 
the body of this paper should be modified accordingly. 


596 Rhodora [Vol. 75 


Figs. 1-6. Audouinella alariae (Jonsson) Woelk. Figs. 1-3. Uni- 
cellular bases bearing a variable number of erect filaments. Figs. 4-5. 
Portions of densely branched erect filaments with numerous mono- 
sporangia. Fig. 6. Sporelinps. Figs. 1 3, 4: Ny (Hardhead Island, 
Penobscot Bay, Maine, VII. 1894, Collins [= Collins, Holden, and 
Setchell 1896a: 236]); Fig. 2: Ny (Hampton, New Hampshire, VII. 
1894, Collins); Fig. 5: WJw 2275 (Nubble Light, Maine, 19. VIII. 
1969, Hehre) ; Fig. 6: FH (Casco Bay, Maine, VII. 1939, Weatherill). 


1973] Audouinella — Woelkerling 597 


\ 
\ 
\\ 


l } 
\ / 
l ^ } 
\ 
l 


Figs. 7-9. Audouinella alariae (Jonsson) Woelk. Figs. 7-8. Habits 
of fairly small plants with mature monosporangia. Fig. 9. Portion 
of sparsely branched erect system with scattered monosporangia. 
Fig. 7: Ny (Hardhead Island, Penobscot Bay, Maine, VII. 1894, 
Collins [= Collins, Holden, and Setchell 1896a: 236]); Figs. 8, 9: 
wJw 2275 (Nubble Light, Maine, 19. VIII. 1969, Hehre). 


Figs. 10-19. Audouinella dasyae (Collins) Woelk. Figs. 10-12. 
Sporelings. Figs. 14-17. Variation in structure of prostrate system. 
Note central panduriform to pryriform cell which may become ob- 
secured in robust specimens (Fig. 16). Figs. 18-19. Chromoplasts in 
cells of erect filaments. Figs. 10-12: wyw 2853 (Old Silver Beach, 
West Falmouth, Massachusetts, 31. X. 1970, Woelkerling) ; Figs. 13, 
14: Edelstein 2563 (Malpeque Bay, Prince Edward Island, 5. VIII. 
1966, Edelstein); Figs. 15, 18, 19: wyw 3311 (Woods Hole, Massa- 
chusetts, 16. II. 1971, Woelkerling) ; Fig. 16: wyw 2977 (Old Silver 
Beach, West Falmouth, Massachusetts, 30. XII. 1970, Woelkerling) ; 
Fig. 17: wzw 2534 (Waquoit Bay [Harbor Entrance], Falmouth, 
Massachusetts, 27. IV. 1970, Woelkerling). 


1973] Audouinella — Woelkerling 599 


Figs. 20-31. Audouinella dasyae (Collins) Woelk. Figs. 20-21. 
Filaments of erect system with monosporangia. Figs. 22-23. Un- 
fertilized carpogonia. Figs. 24-25. Fertilized carpogonia. Note trans- 
verse division (Fig. 25). Figs. 26-28. Stages in development of car- 
posporophyte. Figs. 29-31. Spermatangia. Note varying arrange- 
ment, Fig. 20: wyw 3311 (Woods Hole, Massachusetts, 16. II. 1971, 
Woelkerling) ; Fig. 21: wyw 2820 (Woods Hole, Massachusetts, 13. 
X. 1970, Woelkerling) ; Figs. 22-23: FH (North Eastham, Massachu- 
setts, 9. VIII. 1959, Lamb a-174 [filed under the host, Dasya pedi- 
cellata]); Figs. 24-28, 30: wyw 1852 (West Yarmouth, Massachu- 
setts, 3. X. 1969, Woelkerling) ; Figs. 29, 31: wJw 2249 (West Yar- 
mouth, Massachusetts, 16. XI. 1969, Woelkerling). 


600 Rhodora [Vol. 75 


| 25um 
| (41-43) 


A 


Figs. 32-43. Audouinella daviesii (Dillwyn) Woelk. Figs. 32-35. 
Filaments of erect system bearing monosporangia. Note clustered 
arrangement of sporangia and variation in cell size. Fig. 36. Chro- 
moplasts in cells of erect filaments. Figs. 37-40. Spermatangia and 
unfertilized earpogonia. Figs. 41-42. Stages in carposporophyte de- 
velopment. Fig. 43. Tetrasporangia. Fig. 32: wsJw 2719 (Woods Hole, 
Massachusetts, 17. VII. 1970, Woelkerling); Fig. 33: wiw 2978 (Old 
Silver Beach, West Falmouth, Massachusetts, 30. XII. 1970, Woelker- 
ling); Fig. 34: waw 2536 (Waquoit Harbor, Falmouth, Massachu- 
setts, 27. IV. 1970, Woelkerling) ; Fig. 35: wyw 3302 (Woods Hole, 
Massachusetts, 29. I. 1971, Woelkerling) ; Fig. 36: wyw 2332 (Woods 
Hole, Massachusetts, 4. II. 1970, Woelkerling) ; Figs. 37-42: waw 2678 
(Woods Hole, Massachusetts, 2. VII. 1970, Woelkerling) ; Fig. 45: 
wJw 2198 (Montauk Point, New York, 20. I. 1970, Woelkerling). 


1973] Audouinella — Woelkerling 601 


Figs. 44-45. Audouinella unifila (Jao) Woelk. Habit of two plants 
MICH (Norton Point, Martha’s Vineyard, 3. VIII. 1934, Jao, holo 
type). 

Figs. 46-51. Audouinella microscopica (Naegeli In Kuetzing) 
Woelk. Habit of monosporangial plants. Figs. 46-47: MiCH (Norton 
Point, Martha's Vineyard, 3. VIII. 1934, Jao, type of Acrochaetium 
microfium Jao); Figs. 48-51: MICH (Black Rock, Sconticut Neck. 
New Bedford, Massachusetts, 25. VII. 1934, Jao, type of Aerochae- 
tium compactum Jao). 

Figs. 52-55: Audouinella purpurea (Lightfoot) Woelk. Fig. 52 
Portion of erect system bearing carposporophytes. Figs. 53-55. Chro- 
moplasts in cells of erect filaments. Figs. 52-55. wJw 3297 (Sand- 
wich, Massachusetts, 22. I. 1971, Woelkerling). 


602 Rhodora [Vol. 75 


N 
J} 
Jj 

/ 


Figs. 56-60. Audouinella saviana (Meneghini) Woelk. Figs. 56- 
57. Prostrate systems. Fig. 58. Chromoplasts in cells of erect fila- 
ments. Figs. 59-60. Erect filaments bearing monosporangia (Fig. 59) 
and tetrasporangia (Fig. 60). Figs. 56-59: wJw 2753 (Woods Hole, 
Massachusetts, 21. VII. 1970, Wilce); Fig. 60: ww 3818 (Sandwich, 
Massachusetts , 13. X. 1970, Woelkerling). 


1973] Audouinella — Woelkerling 603 


oy [2oum 
| 61,63) 
lOum 
(62) 


4O0gm 
| (64,65) 


Figs. 61-63. Audouinella spetsbergense (Kjellman) Woelk. Figs. 
61, 63. Erect filaments bearing tetrasporangia. Fig. 62. Chromo- 
plasts in cells of erect filaments. Fig. 61: ups (Fairhavn Island, 
Spitzbergen, 19. VIII. 1872, Kjellman, type of A. spetsbergense) ; 
Figs. 62, 63: urs (Fairhavn Island, Spitzbergen, 12. VIII. 1872, 
Kjellman, type of Rhodochorton penecilliforme). 

Figs. 64-65. Caloconema membranacea (Magnus) Woelk. Habit 
of portions of two plants. Note tetrasporangia (Fig. 65). Figs. 64- 
65: FH (Revere Beach, Massachusetts, 29. V. 1886, Collins, from 
Hauck and Richter 1888: 154). 


604 Rhodora [Vol. 75 


Figs. 66-73. Colaconema humilis (Rosenvinge) Woelk. Fig. 66. 
Sporelings. Figs. 67-73. Habit of monosporangial plants. Figs. 66- 
69: MICH (Norton Point, Martha's Vineyard, Massachusetts, 3. VII. 
1934, Jao, type of Acrochaetium radiatum Jao); Figs. 70, 72, 73: 
Edelstein 1382 (Ketch Harbor, Nova Scotia, 31. V. 1965, Edelstein) ; 
Fig. 71: wsw 1838 (Woods Hole, Massachusetts, 26. VI. 1969, 
Conway). 


1973] Audouinella — Woelkerling 605 


Figs. 74-76. Colaconema minima (Collins) Woelk, Habit of mono- 
sporangial plants. Note variation in shape of monosporangia and in 
cells subtending them, Fig. 74: wsw 2699 (Sandwich, Massachusetts, 
30. VI. 1970, Woelkerling) ; Fig. 75: FH (Robinson's Hole, Elizabeth 
Islands, Massachusetts, VIII. 1907, Collins, type [= Collins, Holden. 
and Setchell 1908: 1493]); Fig. 76: C (Mollegrund, Denmark, 8. 
VIII. 1899, Rosenvinge, type of Chantransia emergens Rosenvinge). 


606 Rhodora [Vol. 75 
NE ° @ 


f) 
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EY \ dp d " /| 


\ A 
u AN \ \ N Ü / 
\\\ \ š li 
r YR | Wi 
Ta0um NN ' | ) / 
(77-81) | ' N 
N j + EN \ ; "n 
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Ma + Y \\ | / \\ \ 1 
| EI 20ym « | V A I N | [i 82 | 
qe 1 \ Ny PY i 
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Figs. 77-83. Colaconema secundata (Lyngbye) Woelk. Fig. 77. 
Sporelings showing characteristic parenchymatous grouping of cells. 
Figs. 78-81. Prostrate system of mature plants. Fig. 82. Habit of 
virgate plant. Fig. 88. Chromoplasts in cells of erect filaments. 
Fig. 77: wyw 2316 (Woods Hole, Massachusetts, 4. II. 1970, Woelker- 
ling); Figs. 78-79, 82: wyw 2276 (West Falmouth, Massachusetts, 
6. I. 1970, Woelkerling) ; Figs. 80-81: wyw 2476 (Nantucket Center, 
Massachusetts, 14. IV. 1970, Woelkerling) ; Fig. 88: wyw 2808 ( Peni- 
kese Island, Massachusetts, 1. X. 1970, Woelkerling). 


1973] Audouinella — Woelkerling 607 


Table 1. Present status of audouinelloid taxa listed by 
Giard (1890), Hehre and Mathieson (1970), and Tayloi 
(1957), (Author citations for taxa are recorded in the taxo- 
nomic treatments.) 


Name of Taxon in 


Older Literature Present Status 
Acrochaetium Conspecific with Audouinella daviesii 
alcyonidae 

A. amphiroae Conspecific with A. daviesii 

A. attenuatum See "Collections Inquirendae" 

A. dasyae Audouinella dasyae 

A. daviesii Audouinella daviesii 

A. emergens Conspecific with Colaconema minima 

A. flexuosum All New England collections referred to 
Colaconema secundata 

A. intermedium Conspecific with Audouinella dasyae 

A. microfilum Conspecific with Audouinella 
microscopica 

A. minimum Colaconema minima 

A. polyides See “Collections Inquirendae” 

A. radiatum Conspecific wıth Colaconema humilis 

A. sagraeanum See “Species Excludendae" 

A. thuretii Conspecific with Audouinella saviana 

A. zosterae Conspecific with Audouinella dasyae 


608 


Audouinella 
efflorescens 


A. membranacea 


Colaconema 
americana 


Conchocelis 
rosea 


Kylinia alariae 


K. compacta 


K. hallandica 


K. moniliformis 


K. secundata 
K. unifila 


K. virgatula 


Rhodochorton 
entozoicum 


R. penicilliforme 


R. purpurea 


Rhodora [Vol. 75 
Table 1 — Continued 


See “Collections Inquirendae” 


Colaconema membranacea 


See “Species Inquirendae” 


Referred to Bangiophycidae; not 
discussed in this account. 


Audouinella alariae 


Conspecific with Audouinella 
microscopica 

Not recorded for New England and not 
discussed in this account 

Conspecific with Audouinella 
microscopica 

Colaconema secundata 


Audouinella unifila 


Conspecific with Colaconema secundata 


See “Species Inquirendae" 


See “Collections Inquirendae" under 
Audouinella spetsbergense 


Audouinella purpurea 


609 


Audouinella — Woelkerling 


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610 Rhodora [Vol. 75 


A. dasyae A. intermedium A. zosterae 
Height 3mm 2mm 3mm 
Cell 6-12 (-16) um 8-10 um 6.4-9.6 um 
diameter [8-11 um] [6-11 um] 


Cell length 15-70 (-90) pm (26-) 32-64 um 32-70 um 
[15-70 um] [21-75 um] 


L/D 2-7 (-10) — 
[2-7.5] [2-7] 
Monospore 16-24 um 19-25 um 22-31 um 
length [16-20 um | [18-24 um | 
Monosopore 9-12 (-16) „m 9-13 um 6.5-9.5 um 
diameter [9-12 (-15) um] [8-11 um] 


Table 3. Data on populations of A. dasyae complex. 
Note: Data on A. dasyae is a composite of all New Eng- 
land collections examined. Data for A. intermedium and 
A. zosterae in brackets is based on personal studies of the 
types; data not in brackets is that reported by Jao (1936). 


A. thuretii A. saviana A. saviana 
(type) (type) (New England) 
Height up to3 mm up to 3 mm up to 4 mm 


Cell 
diameter — 7-10 (-13) „m 7-12um . (7-) S-12(-14) „m 
Cell 


length 20-45 im 20-40 um 20-60 um 
L/D 2-5 1.5-4 2-6 (-8) 
Monospore 

length 15-20 „m 13-21 m 18-27 um 
Monospore 

diameter 9-12 m 8-12 um 10-15 pm 


Table 4. Data on populations of the Audouinella saviana 
complex. The first two columns represent data on type 
colleetion plants; the third is a composite of data on New 
England populations. 


611 


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612 Rhodora [Vol. 75 


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dophyceae 1. K. Danske Vidensk. Selsk. Skr. (Afd. 7, Raekke) 

7(1): 1-151. 


1923-1924. The marine algae of Denmark. 1. 
Rhodophyceae 3. K. danske Videnske Selsk. Skr. (Afd. 7, Raekke) 
7(3): 285-487. 


1935. Distribution of the Rhodophyceae in the 
Danish waters. K. danske Vidensk. Selsk. Skr. (Afd. 9, Raekke) 
6(2): 1-44. 

RorH, A. G. 1797-1806. Catalecta Botanica quibus Plantae Novae 
et minus Cognitae Describuntur atque Illustrantur. Leipzig. 
ScHIFFNER, V. 1916. Studien ueber Algen des Adraitischen Meeres. 
Wiss. Meeresunters. (Abt. Helgoland) (N. F.) 11(2): 127-198. 

1931. Neue und bemerkenswerte Meeresalgen. Hed- 
wigia 71: 139-205. 

SCHMITZ, F. and P. FALKENBERG. 1897. Rhodomelaceae. In A. Eng- 
ler und K. Prantl, *Die Naturlichen Pflanzen familien." 1(2). 
Leipzig. 

Stropot, S. 1876. Le Balbiania investiens étude organogénique et 
physiologique. An». Sci. Nat. (Bot.) (6), 3: 146-174. 

SoutH, G. R. 1970. Checklist of marine algae from Newfoundland, 
Labrador, and the French Islands of St. Pierre and Miquelon. 
Mar. Sci. Res. Lab. (Memorial University of Newfoundland) 
Technical Report No. 2. iv + 20 pp. 

, and A. CARDINAL. 1970. A checklist of marine algae 
of Eastern Canada. Can. J. Bot. 48: 20771-2095. 

STONE, R. A., E. J. HEumE, J. M. Conway, and A. C. MATHIESON. 
1970. A preliminary checklist of the marine algae of Campa- 
bello Island, New Brunswick, Canada. Rhodora 72: 313-338. 

SUNDENE, O. 1953. The algal vegetation of Oslofjord. Norske 
Vidensk. Oslo. I. Mat. — Naturw. Klasse 1953(2) : 1-244. 


1973] Audouinella — Woelkerling 621 


TAYLOR, W. R. 1928. The marine algae of Florida with special 
reference to the Dry Tortugas. Pap. Tortugas Lab. 25: 1-219. 

1937. Marine Algae of the Northeastern Coast of 
North America. IX + 427. Ann Arbor, Michigan. 

—— Th F 1941. Notes on the marine algae of Texas. Pap. 
Mich. Acad. Sci. 26: 69-79. 

—————. 1957. Marine Algae of the Northeastern Coast of 
North America. 2nd. Ed. Ann Arbor. 

1960. Marine Algae of the Eastern Tropical and 
Subtropical Coasts of the Americas. Ann Arbor. 

TURTON, W. 1806. A General System of Nature. 3. London. 

VALERA, M. DE 1939. Some new or critical algae from Galway Bay, 
Ireland. K. fysiogr. Sallsk. Lund. Forh. 9(9) : 1-14. 

VICKERS, A. 1905. Liste des algues marine de la Barbade. Ann. 
Sci. Nat. (Bot.) (9) 1: 45-66. 

WEBER VAN Bosse, A. 1921. Liste des algues du Siboga. II. Rho- 
dophyceae. Part I. Protoflorideae, Nemalionales, Cryptonemiales. 
Siboga-Exped. Monogr. No. 59B, 187-310, Plates 8-10. 

WEST, J. A. 1967. The life histories of Rhodochortom purpureum 
and Rhodochorton tenue in culture. J. phycol. 3(Suppl.), 11. 

1968. Morphology and reproduction of the red alga 
Acrochaetium pectinatum in culture. J. phycol. 4: 89-99, 

1969. The life histories of Rhodochorton purpureum 
and R. tenue in culture. J. phycol. 5: 12-21. 

— 1970. A monoecious isolate of Rhodochorton pur- 

pureum, J. phycol. 6: 368-370. 

1970a. The Life history of Rhodochorton concrescens 
in culture. Br. phycol. J. 5: 179-186. 

WHELDEN, R. M. 1947. Algae. pp. 13-127. In Polunin, N. ed. 
Botany of the Eastern Canadian Arctic. Part II. Thallophyta 
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WOELKERLING, W. J. 1970. Acrochaetium botryocarpum (Harvey) 
J. Ag. in Southern Australia. Br. phycol. J. 5(2) : 159-171. 

1971. Morphology and taxonomy of the Au- 

douinella complex (Rhodophyta) in Southern Australia, Aust. 

J. Bot., Suppl. Ser. Suppl. 1. 91 pp. 

1972. Studies on the Audouinella microscop- 

ica (Naeg.) Woelk. complex (Rhodophyta). Rhodora 74: 85-96. 

1973. The  Audouinella complex (Rhodo- 
phyta) in the Western Sargarso Sea. Rhodora: 75: 78-101. 

YAMADA, Y. 1944. Notes on some Japanese algae. 10. Sci. Pap. 
Inst. algol. Res. Hokkaido Univ. 3: 11-25. 


DEPARTMENT OF BOTANY 
UNIVERSITY OF WISCONSIN 
MADISON, WISCONSIN 53706 


BOOK REVIEW 


THE GENUS LESQUERELLA (CRUCIFERAE) 
IN NORTH AMERICA' 


It is quite satisfying and even reassuring in these times 
to encounter a substantial taxonomic work unencumbered 
by flights of fancy. It seems commendable for the authors 
to state on page 33, “We see little to be gained by putting 
our own speculations concerning a detailed phylogeny into 
print.” The gift of imagination is good if rightly used but 
it is no substitute for hard work and relentless probing 
after facts which are both so well exemplified in the attrac- 
tive, informative and impressive volume under review. 

In their introduction the authors explain that, for more 
than thirty years, collecting and field studies of Lesquerella 
have been carried on by the senior author. They state that 
this treatment includes sixty-nine species and twenty-nine 
infraspecific taxa but excludes some dozen species of South 
America. It is obviously too much to have asked of the 
present authors that they studied these exotic species with 
anything like the detail and care given to the North Amer- 
ican areas, their study embracing the examination of 9000 
sheets, the study of wild populations, the experimental cul- 
ture of some taxa, the determination of chromosome num- 
bers and the examination of type material. 

Following the introduction is a discussion of the limits 
of the genus, understandably an important consideration in 
the Cruciferae. The extent of their coverage of Lesquerella 
is indicated by merely listing the remaining topics treated 
in the first thirty-five pages, breeding system, interspecific 
hybridization, chemical information of systematic value, 
the type species and subdivisions of the genus, chromosome 
numbers, pollen, trichomes, taxonomic characters and evo- 
lution within the genus. 


‘Rollins, Reed C. and Elizabeth A. Shaw. The Genus Lesquerella 
(Cruciferae) in North America. pub. date 6 June 1973. Harvard 
University Press, Cambridge, Massachusetts. X pp 1-288, illustra., 
$18.00. 


622 


1973] Book Review 623 


It is of interest to note that an aneuploid relationship 
was found to occur frequently between species of Lesque- 
rella. Also in some cases infraspecific polyploidy was found 
to occur accompanied by evolutionary changes and some- 
times not. The following statement by Rollins and Shaw on 
p. 12 would seem to suggest a reasonable taxonomic hand- 
ling of this kind of problem, *If no significant and readily 
detectable morphological physiological or ecological changes 
have followed the event of chromosome complement mul- 
tiplication within a species then there is little justification 
for insisting that the taxonomy merely reflect the ployploid 
picture." 

Scanning electron microscope photographs strikingly il- 
lusrate the types of pollen and details of the variety of 
trichomes in Lesquerella. Indeed the book throughout is 
well illustrated with readily accessible plates, figures and 
maps. Particularly in the main body of the treatment, 
beginning on p. 36, and dealing with the taxonomy of the 
group, it is refreshing to note the uncrowded handling of 
taxa and the convenient juxtaposition of illustrations, de- 
scriptive material and discussions. 

It is of particular importance to call attention to the fact 
that many new combinations and a number of new species 
are published by Rollins and Shaw in this work. 

The use by Rollins and Shaw of two important infraspe- 
cific categories, those of subspecies and variety seems to 
work out realistically though the authors don't seem to 
explain in all cases what their reasons are for preferring 
one or the other rank. 

There are unresolved questions raised by the authors 
but they seem in this treatment to have done all they might 
have done with Lesquerella at this time. 


ALBION R. HODGDON 
DEPARTMENT OF BOTANY & PLANT PATHOLOGY 
UNIVERSITY OF NEW HAMPSHIRE 

DURHAM, NEW HAMPSHIRE 03824 


INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA 


Manuscripts must be double-spaced or preferably triple- 
spaced (not on corrasable 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 indicate the style of type through the use of capitals 
or underscoring, partieularly in the citations of specimens, 
except that the names of species and genera may be under- 
lined 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 System- 
atics and Ecology of Poison-Ivy and the Poison-Oaks," 
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu- 
larly with reference to the indentation of keys and syno- 
nyms. Papers of a floristic nature should follow, as far as 
possible, the format of "Contribution to the Fungus Flora 
of Northeastern North America. V.," H. E. Bigelow & M. E. 
Barr, Rhodora 71: 177-203. 1969. For bibliographic cita- 
tions, a recommended list of standard journal abbreviations 
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot. 
Club 85: 277-300. 1958. 


Volume 75. No. 804. including pages 493-644, was issued December 31. 1975 


624 


Dodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
ALBION REED HODGDON, Editor-in-Chief 


ROLLA MILTON TRYON 
RADCLIFFE BARNES PIKE 
STEPHEN ALAN SPONGBERG 
GERALD JOSEPH GASTONY 
ALFRED LINN BOGLE 
RICHARD EDWIN WEAVER 3 


> Associate Editors 


VOLUME 75 


1973 


Che Nem England Botanical Club, Hue. 


Botanical Museum, Oxford St., Cambridge, Mass. 02138 


INDEX TO VOLUME 75 


New scientific names and combinations are printed in bold face type 


Abies balsamea 314 

Acacia tortuosa 416 

Acalypha setosa 393 

Acerates hirtella 398 

Achillea millefolium 321 

Acnida cannabina 57 

Acorus calamus 54, 57, 61, 316 

Acrochaetium alariae 541, 543- 
545; aleyonidii 273, 551, 554- 
556, 607; var. cylindricum 551, 
554; americanum 588; amphi- 
roae 551, 556, 607; antillarum 
92; attenuatum 583, 607; av- 
rainvillae 89, 563; bornetii 88, 
557, 592; catenulatum 86, 557; 
collopodum 86, 557; compactum 
86, 557; crassipes 86, 557; das- 
yae 278, 545, 602; daviesii 81, 
550, 577, 602; dufourii 84, 85, 
97; efflorescens 584; effusum 
92; emergens 572, 607; endo- 
phytieum 92, 574; endozoicum 
592; flexuosum 273, 532, 577, 


582, 602; hallandicum 82; hu- 
mile 569; hypneae 563; inclu- 
sum 592; infestans 89; inter- 


medium 545, 548, 549, 607; 
kylinii 574, 592; Levringii 87; 
luxurians 94, 97, 575; micro- 
filum 87, 558, 583, 607; micro- 
seopieum 86, 557; minimum 
572, 607; moniliforme 87, 558; 
Nemalionis 89; opetigenum 
549; pallens 563; polyides 587, 
588, 607; radiatum 273, 569, 
571,.583, 604, 607; ralfsiae 592; 
rhipidandra 541; robustum 549; 
sagraeanum 532, 551, 556, 590, 
591, 593, 607; sagraeanum 560, 


562, 577, 582; sargassi 84, 85, 
97; savianum 560; secundatum 
94, 575; subseriatum 545, 548; 
subsimplex 575, 582; subtilissi- 
mum 592; tenuissimum 576; 
thuretii 88, 551, 556, 560, 562, 
563, 607; trifilum 272, 273; uni- 
filum 563; unipes 549; vir- 
gatula 96, 97; virgatulum 94, 
273, 576; f. luxurians 273, 576; 
f. tenuissimum 576; zosterae 
545, 548, 549, 607 

Actinastrum 266 

Agardhiella tenera 274 

Ageratum conyzoides 422; ssp. 
conyzoides 422; ssp. latifolium 
422; latifolium 422 

Agmenellum thermale 282; quad- 
ruplicatum 282 

Agropyron repens, f. aristatum 
314; f. pilosum 314; f. trichor- 
rachis 314 

Agrostis perennans 315; scabra 
315; stolonifera, var. major 
315; var. palustris 315; tenuis, 
f. aristata 315 

Alabama and Middle Tennessee, 
Some Notes on the Flora of the 
Southern States, Particularly 
366 

Alaria esculenta 543 

Alaska, New Record for the Yel- 


low Lady's Slipper Orchid, 
Cypripedium calceolus L. ssp. 
parviflorum (Salisb.) Hult. 
from 491 


Alaskan Species of Erigeron L. 
(Compositae), A New 116 


627 


628 


Algae, Benthie, and Vascular 
Plants of the Lower Merrimack 
River and Adjacent Shoreline 
52; of Virginia and Maryland, 
Including the Chesapeake Bay 
Area, the Marine 258 


Allium cernuum 385; stellatum 
384 

Alnus crispa, var. mollis 317; 
rugosa, var. americana 317 


Alopecurus pratensis 315 

Alysicarpus vaginalis 390 

Ambrosia artimisiifolia, var. ela- 
tior 321; bidentata 407 

Amellus asper, var. canescens 
192; var. glabriusculus 194; 
var. normalis 192 

Ammophila breviligulata 56, 315 

Amphicarpon floridanum 374; 
muhlenbergianum 374 

Amphipleura rutilans 59, 60 

Amsonia rigida 398 

Anabaena inaequalis 292; sphae- 
rica 292; torulosa 292 

Anacystis cyanea 282; dimidiata 
282; marina 283; montana, f. 
minor 283; f. montana 283; 
thermalis, f. thermalis 283 

Anaphalis margaritacea, var. in- 
tercedens 321; var. subalpina 
321 

Andropogon arctatus 874; capil- 
lipes 875; divergens 375; hirti- 
florus 375; littoralis 375; niveus 
375; perangustatus 376; terna- 
rius 374 

Anthoxanthum odoratum 315 

Antigonon leptopus 386 

Antithamnion cruciatum 276 

Aquatic and Wetland Plants of 
the Southwestern United States 
(Review) 114 

Aralia hispida 319 

Arenaria lateriflora 317; 
loides, var. robusta 317 

Arethusa bulbosa 316 


pep- 


Rhodora 


[Vol. 75 


Aristida tuberculosa 371 

Aronia melanocarpa 390 

Artemisia ludoviciana 409; stel- 
leriana 56, 511 

Arthrocladia villosa 564 

Arthrospira brevis 284 

Asclepias aceratoides 399; hir- 
tella 398; quadrifolia 160; to- 
mentosa 399; verticillata 135 

Ascocyclus orbicularis 269 

Ascophyllum nodosum 58, 271; f. 
scorpioides 271 

Asparagopsis hamifera 273, 2774 

Asperococcus echinatus 270 

Asplenium trichomanes 167 

Aster acuminatus 1-4, 6-9, 11, 12, 
15, 17-19, 21, 22, 27, 30, 321; 
Blakei 1-4, 6-9, 11-16, 17, 19-22; 
Chemical, Cytological and Ge- 
netic Evidence for the Hybridi- 
zation of 1; X Blakei 321; 
chapmanii 405; Chromosome 
Numbers in 26; ciliolatus 27, 
30; concinnus 26, 27, 30; cordi- 
folius 27, 30; dumosus 166; 
eryngiifolius | 405; foliaceus, 
var. arcuans 321; gracilis 26, 
27, 30; johannensis 321; junci- 
formis 27, 30; lateriflorus 27. 
30; lowerianus 28, 30; macro- 
phyllus 28, 30; nemoralis 1-4, 
6-9, 11, 12, 15, 17-22, 321; x 
acuminatus 13-15; novi belgii 
56; patens 28, 31; pilosus 28, 
31; puniceus 28, 31; sagitti- 
folius 28, 31; simplex 28, 31; 
spectabilis 29, 31, 405; subu- 
latus 57; umbellatus 29, 31, 
321; undulatus 26, 29, 31 

Astragalus atlasovi 304, 305; po- 
laris 304; A Range Extension 
and a New Synonym for 304; 
tennesseensis 135, 136, 158 

Athanasia hastata 184, 185 

Athyrium filix-femina, var. mi- 
chauxii, f. rubellum 314 


1973] 


Atriplex glabriuscula 314; patula 
53; var. hastata 56, 106-110, 
317; Variation in the Seed Size 
of 106 

Audouinella alariae 534, 541, 
543-545, 550, 555, 596 fig., 597 
fig., 608, 609; attenuatum 584; 
australis 564, 565; Complex 
(Acrochaetiaceae, Rhodophyta) 
in the Northeastern United 
States, The Morphology and 
Systematics of the 529; Com- 
plex (Rhodophyta) in the 
Western Sargasso Sea, The 78; 
dasyae 534, 545, 547-550, 598, 
599, 607, 610; daviesii 79, 81, 
82, 533, 550, 553-556, 563, 582, 
585, 591, 592; efflorescens 557, 
584, 585, 608; hallandica 79, 80, 
82, 83, 85, 97; hermanni 535; 
intermedium 610; membranacea 
566, 608; microscopica 79, 80, 86, 
88, 534, 557, 559, 565, 583, 601, 
607, 608; purpurea 530, 534, 
536, 539-41, 568, 587, 601, 608; 
saviana 534, 560-63, 584, 591, 
592, 602, 607, 610; spetsber- 
gense 585, 587, 603, 608; thu- 
retii 79, 81, 88, 89, 560, 610; 
unifila 534, 563, 565, 601; zos- 
terae 610 

Averett, John E., Biosystematic 
Study of Chamaesaracha (Sol- 
anaceae) 325 

Avicennia germinans 102-105; 
Salt Concentrations in Ground 
Waters Beneath Rhizophora 
mangle and 102 

Azalea austrina 396 

Azorella selago 241 


Bahama Flora Since Britton and 
Mllspaugh, Additions to the 411 

Bangia ciliaris 272; fuscopur- 
purea 272 

Baptisia australis 159 


Index to Volume 75 


629 


Barkley, T. M. 
R. 

Barneby, R. C., A Range Exten- 
sion and a New Synonym for 
Astragalus polaris, Benth. 304 

Baskin, Carol C. see Baskin Jerry 
M. 

Baskin, Jerry M. and Carol C. 
Baskin, The Past and Present 
Geographica] Distribution of 
Petalostemon foliosus and 
Notes on Its Ecology 132 

Betula papyrifera, var. cordifolia 
316 

Bidens cernua 321, 408; frondosa 
321; nivea 170, 171, 184; tri- 
partita 408 

Blidingia minima 58 

Bocconia frutescens 415 

Boehmeria nivea 386 

Book reviews 114, 140, 323, 622 

Bostrychia radicans 280; f. mon- 
iliforme 280; rivularis 280 

Botrychium biternatum 366; luna- 
rioides 366 

Boufford, David E., New or Note- 
worthy New Hampshire Plants 
158 

Bougainvillea glabra 415 

Bowler, Peter A. and Phillip W. 
Rundel, The Status of Rama- 
lina subampliata (Nyl.) Fink 
in North America 306 

Bowley, Donovan R., The En- 
vironment of Schistostega pen- 
nati (Hedw.) Hook & Tayl.: 
New Vermont Stations 149 

Brachiaria extensa 372; platy- 
phylla 372 

Bryopsis hypnoides 267; plumosa 
327 

Bulbostylis capillaris 378 

Byssus purpurea 536 


see Kowal, Robert 


Cacalia diversifolia 409; tuberosa 
136 


630 Rhodora [Vol. 75 


Caesalpina divergens 416 

Cakile edentula 318 

Calamagrostis canadensis, var. 
canadensis 315 

Calea aspera 171, 192 

Callicarpa americana 420 

Callithamnion baileyi 276; bys- 
soides 276; corymbosum 276; 
daviesii 81, 550, 555, 577; var. 
secundatum 94, 575; efflorescens 
584; entozoicum 569, 589; lux- 
urians 94, 96, 575; membrana- 
ceum 566, microscopicum 86, 
557; purpureum 536; rothii 
537; savianus 560, 563, secun- 
data 94, 575; virgatulum 94, 
545, 550, 576, 581 

Callitriche verna 319 

Caloglossa lepricurii 279 

Calopogon tuberosus 316 

Calothrix confervicola 292; crus- 
taceae 292; pulvinata 293 

Canna flaccida 385 

Capsella bursa-pastoris 318 

Cardamine bulbosa 388; douglas- 
sii 388; pratensis 160 

Carex abscondita 251; aestivalis 
251, 253; alata 251, 253; al- 
bolutescens 250, 254; albursina 
250, 253; amphibola 250; annec- 
tens 250; arenaria 252; argy- 
rantha 250, 253; artitecta 250; 
atlantica 250; austro-carolini- 
ana 380; baileyi 251, 254; bar- 
rattii 251, 253; Bigelowii 156; 
biltmoreana 249, 252, 253; 
blanda 250; brevior 249, 253; 
bromoides 250, 254; brunnescens 
249, 253; var. sphaerostachya 
315; bullata 251, 253; bushii 
250, 253; buxbaumii 249, 254, 
255; canescens 249, 254; var. 
disjuncta 315; var. subloliacea 
515; capitata 155, 156; carey- 
ana 250, 253; caroliniana 250; 
cephalantha 315; cephalophora 


250; chapmannii 249, 252, 255; 
cherokeensis 249, 251, 253; col- 
linsii 251, 253; communis 250, 
253, 380; comosa 250, 253; com- 
planata 250; conjuncta 250, 
253; conoidea 316; crawei 381; 
crebriflora 251; crinita 250; 
cristatella 250, 253;  crus- 
corvi 251, 253; dasycarpa 
381; debilis 250; decomposita 
251, 253;  dichotoma 269; 
digitalis 250; divisa 252; ebur- 
nea 250, 253; emmonsii 250, 
253; extensa 249, 252; festu- 
cacea 250; flaccosperma 251; 
folliculata 250, 254; frankii 
250; gigantea 251, 253; glau- 
cescens 251, 253; gracilescens 
250; gracillima 251; granularis 
250; grayi 251, 253; grisea 250; 
hitcheockiana 251, 253; hormo- 
thodes 315; howei 250, 255; 
hyalinolepis 251, 253; hystri- 
cina 250, 253; interior 250, 2553; 
intumescens 250, 252; jamesii 
250, 254; joori 251, 253; kobo- 
mugi 252; laevivaginata 250; 
lasiocarpa 249, 254, 255; laxi- 
culmis 251; laxiflora 250; leav- 
enworthii 250; leptonervia 251, 
253; leptalea 250, 254, 316; 
louisianica 251, 253; lupuli- 
formis 250, 253; lupulina 250; 
lurida 249, 250; microdonta 
381; muhlenbergii 250; muri- 
cata 249, 253; nigromarginata 
250; normalis 251; novi-angliae 
316; oligocarpa 250, 254; oxy- 
lepis 251, 253; paleacea 316; 
pallescens 249, 253; var. neo- 
gaea 316; paupercula, var, ir- 
rigua 316; pedunculata 249; 
pensylvanica 249; physorhyn- 
cha 250; plantaginea 167, 251, 
253; platyphylla 251; polymor- 
pha 249, 251, 253; prasina 251, 


1973] 


254; reniformis 251, 253; retro- 
flexa 250; rosea 250; rostrata 
249, 253; salina 56; scabrata 
251, 254; scoparia 250; scorsa 
251, 254; shortiana 251, 253; 
sparganoides 250, 254; spicata 
252; squarrosa 251; stipata 250, 
315; striatula 250; stricta 250; 
styloflexa 250; suberecta 251, 
253; swanii 251, 254; tenera 
251; tenax 381; tetanica 249, 
251, 253; torta 251, 254; tribu- 
loides 250, 315; triehocarpa 249, 
251, 253; trisperma 251, 253, 
315; typhina 250, 253; umbel- 
lata 250, 253; validior 381; 
venusta 251, 253; vestita 251, 
253; virescens 251, 254; viri- 
dula 316; vulpinoidea 250; wal- 
teriana 251, 253; willdenowii 
250, 254; woodii 249, 251, 253 

Carices of Virginia, Phytogeog- 
raphy of the 248 

Carum carvi 319 

Carya alba 38; cordiformis 34, 
36, 38, 41, 48; glabra 34, 36, 
37, 44; var. odorata 34; glabra- 
ovalis 44, 46, 47, 49; ovalis 34, 
36, 37, 42, 44; ovata 34, 36, 40, 
41, 45, 49; tomentosa 34, 37, 38, 
48 

Cassandra calyculata, var. angus- 
tifolia 320 

Cassia deeringiana 390; fascicu- 
lata 136; nicitans 158 

Cedrela odorata 417 

Cenchrus gracillimus 412 

Ceramium daviesii 81, 550; dia- 
phanum 276; fastigiatum 276; 
rubriforme 277; rubrum 277; 
secundatum 94, 575; strictum 
278; violaceum 537 

Cerastium vulgatum 317 

Chaetomorpha area 266; 
266 

Chamaesaracha 


linum 


boliviensis 364; 


Index to Volume 75 


651 


coniodes 326, 328, 329, 334, 339, 
841, 355, 357, 359, 860, 362; 
coronopus 326-328, 334, 337-342, 
345, 349; crenata 326, 328, 
331, 334, 338, 351-354; echinata 
364; edwardsiana 328, 334, 339, 
341, 344, 345, 347-349; grandi- 
flora 364; heterophylla 364; 
japonica 364; nana 364; pallida 
326, 328, 334, 339, 345-349, 355; 
physaloides 364; potosina 364; 
savatieri 364; sinensis 364; 
(Solanaceae), Biosystematic 
Study of 325; sordida 326-328, 
334, 339, 347, 353, 355, 356, 
358; villosa 328, 334, 338, 349, 
350, 351, 353, 354; watanabei 
364 

Champia parvula 275 

Chantransia alariae 541; attenu- 
ata 583; catenulata 86, 557; 
collopoda 86, 557; corymbifera 
88, 551, 556, 563; crassipes 86, 
558; dasyae 545; daviesii 81, 


550, 577; dufouri 84, 85; 
efflorescens 584; var. thur- 
etii 88, 55, 556, 560, 563; 


emergens 92, 572, 574, 605, 611; 
endozoica 92; flexuosum 532, 
577, 582; hallandica 84; hu- 
milis 569; infestans 91; lux- 
urians 94, 575; macounii 549; 
microscopica 86, 557; minima 
572; moniliformis 87, 558; par- 
vula 85; polyides 587; rhipi- 
dandra 541, 544, 609; sagrae- 
ana 551, 560, 577, 591; sargassi 
84; saviana 560; secundatum 
94, 541, 545, 558, 575, 580; 
tenuissima 576; thuretii 88, 560, 
562; unilateralis 544; virgatula 
94, 541, 545, 560, 576, 580; var. 
luxurians 576; f. tenuissima 
576, 581 

Chasmanthium nitidum 369 

Chenopodium album 317 


632 


Chesapeake Bay Area, The Ma- 
rine Algae of Virginia and 
Maryland, Including the 258 

Chimaphila maculata 162-164; 
(L.) Pursh in Maine and New 
Hampshire 162 

Chinnappa, C. C. see Gill, L. S. 

Chiogenes hispidula 320 

Chlorella 265 

Chondria baileyana 280; sedifolia 
280; tenuissima 280 

Chondrus crispus 275 

Choris floridana 370 

Chromastrum alariae 541; col- 
Iopodum 86, 557; compactum 
86, 557; crassipes 87, 558; hal- 
landieum 84; humile 569; in- 
festans 91; microscopicum 86, 
557; moniliforme 87, 558; rhip- 
idandra 541; secundatum 94, 
575; unifilum 564; virgatulum 
94, 576 

Chromosome  Pairing in  Obli- 
gately Apogamous Ferns; Pel- 
laea atropurpurea and Pellaea 
glabella, var. glabella 122 

Cnroococeus turgidus 283 


Chrysanthemum leucanthemum, 
var, pinnatifidum 321 
Cienfuegosia heterophylla 418; 


yucatanensis 418 

Cinna arundinacea 315; latifolia 
315 

Circaea alpina 319 

Cirsium arvense 321; vulgare 321 

Citrus aurantium 417 

Cladophora albida 266; crystal- 
lina 266; expansa 266; fascicu- 
laris 266; flexuosa 267; var. 
densa 267; gracilis 267; magda- 
lenae 267; rupestris 267; sag- 
raena 590 

Clematis dioscoreifolia 158, 159; 
var. robusta 158 

Clerodendrum speciosissimum 421 

Closterium gracile 261 


Rhodora 


[Vol. 75 


Coeccochloris elabens 283; penio- 
cystis 283; stagnina 283 

Coccothrinax argentata 414; in- 
aguensis 414 

Coelopleurum lucidum 319 

Colaconema americana 574, 588, 
589, 608, 611; bonnemaisoniae 
94, 566; humilis 534, 569-571, 
583, 604, 607; infestans 79, 80, 
89, 90 fig., 92, 94; membranacea 
534, 556, 568, 569, 590, 603, 608; 
minima 534, 572-574, 588, 589, 
605, 607, 611; nakamurai 564; 
polyides 587; porphyrae 92 

Colaconema secundata 79, 81, 93- 
97, 532, 533, 555, 556, 575, 579, 
581, 582, 591, 592, 606-608; 
tenuissima 576 

Collinsonia canadensis 400 

Colobanthus kerguelensis 240 

Colt, L. C., Jr. and Steven Swartz 
An Unusual Substrate for Po- 
lysiphonia paniculata Montagne 
488 

Colubrina asiatica, var. asiatica 
418 

Conchocelis rosea 608 

Conferva daviesii 81, 550; purpu- 
rea 536; rothii 537; rupestris 
95; violacea 537 

Conopholis americana 167 

Convolvulus sepium 320; f. mala- 
chophyllus 320 

Conyza bonariensis 406 

Coptis groenlandica 318 

Corculum leptopus 386 

Cornus amomum, ssp. microcarpa 
396; canadensis 319; purpusii 
396 

Cotula plumosa 241 

Crotalaria lanceolata 391 

Croton alabamensis 371; discolor 
417; linearis 417 

Cuphaea carthaginensis 393 

Cyathea weatherbyana 144 

Cymophyllum fraseri 252, 255 


Cynanchum graminifoium 420 
Cyperus aristata 413; dentatus 
379; difformis 379; globosus 
414; globulosus 414 
Cypripedium calceolus 491; ssp. 
parviflorum 491; From Alaska, 
New Record for the Yellow 
Lady's Slipper Orchid 491; gut- 
tatum 491; passerinum 491 
Cystopteris fragilis 239 


Danthonia spicata 315 

Dasya pedicillata 279 

Delphinium virescens 136 

Dennstaedtia punctilobula 315 

Dentaria laciniata 158, 160 

Deschampsia flexuosa 315, 371 

Desmanthus illinoensis 136 

Desmotrichum undulatum 270 

Dianthus prolifer 387 

Diarina festucoides 369 

Diarrhena americana 369 

Dicentra canadensis 158, 161 

Dichothrix penicillata 292 

Dichromena colorata 379; 
folia 380 

Dictyosiphon foeniculaceus 270 

Digitalis lutea 159 

Diodia hirsuta 403 

Dioscorea bulbifera 414 

Diplotaxis muralis 388 

Distichlis spicata 57 

Drosera rotundifolia 318 

Dryopteris cristata 314; Goldiana 
167; spinulosa, var. americana 
314 

Drysdale, Frank R. Variations 
of Seed Size in Atriplex patula, 
var. hastata (L.) Gray 106 


lati- 


Eastman, Lesley M. Some New 
Locals in the Maine Flora 166; 
see also Hodgdon, A. R. 

Echinocephalum latifolium 
180 | 

Ectocarpus confervoides 58, 260; 


173, 


Index to Volume 75 


633 


elachistaeformis 268; penicil- 
latus 268; siliculosis 268 

Eichornia crassipes 486, 487; sub- 
ovata 478 

Elachistea fucicola 58, 62, 269 

Eleocharis acicularis 57; elliptica 
315; flavescens 377; geniculata 
414; halophila 315; melano- 
carpa 376; olivacea 377; rob- 
binsii 377; smallii, var. major 
315; tricostata 377; wolfii 378 

Elephantopus cuneifolius 184 

Elymus arenarius, var. villosa 
315; virginicus 315 

Emilia javanica 423; sonchifolia 
423 

Empetrum nigrum 319 

Enteromorpha erecta 58, 60, 62, 
265; clathrata 262; compressa 
262; groenlandica 58, 60; intes- 
tinalis 58, 263; lingulata 263; 
linza 58, 263; marginata 264; 
micrococca 264; minima 264; 
plumosa 264; prolifera 264 

Entocladia viridis 261; wittrockii 
262 

Entophysalis conferta 284; deusta 
284 

Epilobium angustifolium 319; cil- 
iatum 3894; glandulosum, var. 
adenocaulon 319 

Epipactis Helleborine 166 

Equisetum arvense, var. boreale 
313; sylvaticum, var. paucira- 
mosum 313 

Eragrostis amabilis 369; ciliaris 
413; var. laxa 412; excelsa 412; 
urbaniana 413 

Erigeron L. (Compositae), A 
New Alaskan Species of 116 

Erigeron grandiflorus 116, 118, 
119; Hultenii 116-119; hyper- 
boreus 118; Muirii 118; myri- 
onactis 406; peregrinus 118, 
119; repens 406; simplex 118; 
uniflorus, ssp. eriocephalus 118 


634 


Eriocaulon lineare 369, 382; tex- 
ense 383 

Eriophorum angustifolium 315; 
spissum 315; virginicum 315 

Erythroclada subintegra 272 

Erythrotrichia rhizoides 272 

Euphrasia americana 320; randii 
320 


Faasen, Paul Van Chromosome 
Numbers in Asters 26 

Fantz, Paul R. see Lewis, Walter 
H. 

Faust, W. Z and S. B. Jones, Jr. 
The Systematic Value of Tri- 
chome Complements in a North 
American Group of Vernonia 
(Compositae) 517 

Festuca myuros 369; rubra, var. 
rubra 314 

Filipendula ulmaria 318 

Fimbristylis annua 414; autumn- 
alis 414; perpusilla 378, 379; 
puberula 368 

Flaveria bidentis 422; trinervia 
422 

Forestiera ligustrina 135 

Fosliella farinosa 274; lejoisii 274 

Fothergilla major 389 

Fragaria virginiana 318 

Fralick, Richard A. see Mathie- 
son, A. C. 

Fremyella grisea 293 

Fucus vesiculosis 58, 262, 269, 
270; var. sphaerocarpus 271; 
var. spiralis 58, 60 

Fuirena pumila 376 


Gaillardia prlchella 422 

Galapagos Islands, Flora of (Re- 
view) 140 

Galeopsis tetrahit, var. bifida 320 

Galium trifidum 321 

Gelidium crinale 274 

Geranium robertianum 392 

Geum laciniatum 390 


Rhodora 


[Vol. 75 


Giffordia duchassaiginiana 268 
Gilia achilleaefolia, ssp. achilleae- 
folia 235; ssp. multicaulis 235 
Gill, L. S. and C. C. Chinnappa 

A Note on the Karyology 
of  Haplopappus spinulosus 
(Pursch) DC., ssp. typicus Hall 
220 
Gillis, Wm. T., Richard A. How- 
ard and George R. Proctor 
Additions to the Bahama Flora 
Since Britton and Millspaugh 
411 
Glandularia drummondii 400 
Glaux maritima 320 
Gnaphalium uliginosum 321 
Gomontia polyrhiza 265 
Gomphosphaeria | aponina 
lacustris 284 
Goniotrichum alsidii 271 
Goodyera repens, var. opnioides 
316 
Gracilaria 
cosa 275 
Grand Manan, New Brunswick, 
The Vascular Flora of Kent 
Island 311 
Grania efflorescens 584 
Griffithsia tenuis 278 
Grinnelia americana 262, 280 
Guilandina divergens 416 
Gutierrezia dracunculoides 404 
Gyrotheca tinctoria 385 


285; 


foliifera 274; verru- 


Habeneria Hookeri 166; obtusata 
316 

Haplopappus spinulosus, ssp. co- 
tula 221; ssp. spinulosa 224; 
ss. typicus 220, 221, 225; A 
Note on the Karyology of 220 

Harms, Vernon L. New Record 
for the Yellow Lady’s Slipper 
Orchid, Cypripedium calceolus 
L. ssp. parviflorum  (Salisb.) 
Hult. From Alaska 491 

Harrisia undata 419 


1973] 


Harvill, A. M., Jr. Phytogeog- 
raphy of the Carices of Vir- 
ginia 248 

Haynes, Robert R. and James L. 
Reveal A Re-evaluation of Po- 
tomageton  fibrillosus Fern. 
(Potomagetonaceae) 75 

Hedeoma drummondii 400 

Hedyotis nigricans, var. filifolia 
421 

Helenium autumnale 136 

Heracleum maximum 319 

Heteranthera limosa 486, 487 

Hibiscus acetosella 418 

Hickories in New England, The 
Northern Limits of the Distri- 
bution of 34 

Hieracium aurantiacum 322; flo- 
ribundum 322; pilosella 322; 
pratense 322; umbellatum 159, 
161; vulgatum 159 

Hildenbrandia prototypus 59, 61 

Hill, Michael L. and O. M. Rogers 
Chemical, Cytological and Gen- 
etic Evidence for the Hybrid 
Origin of Aster Blakei (Por- 
ter) House 1 

Hippurus vulgaris 319 


Hodgdon, A. R. Flora of the 
Galapagos Islands (Review) 
140; The Genus Lesquerella 


(Cruciferae) in North America 
(Review) 622; and Lesley M. 
Eastman Chimaphila maculata 
(L.) Pursh. in Maine and New 
Hampshire 162; see also Mc- 
Clair, John; Steele, F. L. 

Homalobus amblyodon 304 

Hordeum jubatum 314 

Howard, Richard A. A Partial 
History of the New England 
Botanical Club 493; see also 
Gillis, Wm. T. 

Hudsonia tomentosa 56 

Hydrocoleum holdenii 285 

Hydrocotyle umbellata 420 


Index to Volume 75 


635 


Hypericum virginicum, var. fra- 
seri 319 

Hypnea musciformis 274 

Hypochaeris glabra 409 

Hypoxis juncea 385 

Hyptis mutabilis 482 


Impatiens capensis, f. immaculata 
319 

Ipomoea muricata 399; nil 420 

Iris versicolor 316 

Isnardia spathulata 394 

Isoetes butleri 367, 368; melano- 
poda 367, 368 

Isotria medeoloides 166, 167 

Iva annua 407; ciliata 407; mi- 
erocephala 406 


Jasminum fluminense 420 

Jones, S. B., Jr. see Faust, W. Z. 

Juncus brachycephalus 384; brev- 
icaudatus 316; bufonius 316; 
var. halophilus 316; effusus, 
var. compactum 316; filiformis 
316; gerardi 57, 316; nodatus 
384; roemerianus 414; tenuis 
316 

Juniperus 
pressa 314 


communis, var. de- 


Kalmia angustifolia 320 

Kerguelen Islands, Palynological 
Evidence for the Late Glacial 
Occurrence of Pringlea and 
Lyallia on 239 

Kiang, Y. T. Floral Structure, 
Hybridization and Evolutionary 
Relationship of Two Species of 
Mimulus 225 

Kowal, Robert R. and T. M. 
Barkley Senecio anonymous 
Wood, An Earlier Name for 
Senecio smallii Britton 211 

Kral, Robert Some Notes on the 
Flora of the Southern States, 


636 


Particularly Alabama and Mid- 
dle Tennessee 366 

Kylinia alariae 541, 608; austra- 
lis 564; catenulata 86, 557; 
collopoda 86, 557; compacta 86, 
557, 608; crassipes 87, 558; cy- 
tophaga 592; dufourii 84; hal- 
landica 84, 608; humile 569; 
infestans 91; microscopica 86, 
557; moniliformis 87, 558, 608; 
rhipidandra 541; secundata 94, 
575, 608; sargassi 84; unifila 
564, 608; virgatula 94, 576, 
608; f. luxurians 576 


Lachnanthes caroliniana 382, 385 

Lachnocaulon digynum 383; eng- 
leri 383; minus 369, 384 

Laciniaria chapmanii 404 

Lactuca saligna 410 

Laguncularia racemosa 103, 105 

Laminaria digitata 58; saccha- 
rina 59 

Larix laricina 314 

Lathyrus japonicus 319 

Leathesia difformis 269 

Ledum groenlandicum 320 

Leontodon autumnalis, var. pra- 
tensis 321 

Lepidium filicaule 416; virgini- 
cum 416 

Leptilon bonariense 406 

Leptochloa fascicularis 370; uni- 
nervia 370 


Lesquerella (Cruciferae) in 


North America, The Genus 
(Review) 622 
Leticula resupinata 403 
Leucophysalis grandiflora 332, 
364; heterophylla 364; nana 


364 
Lewis, Walter H. and Paul R. 
Fantz Tribal Classification of 
Triosteum (Caprifoliaceae) 120 
Liagorophila endophytica 574 
Liatris chapmanii 404 


Rhodora 


[Vol. 75 


Ligusticum scothicum 319 

Limonium carolinianum 53, 56; 
nashii 320 

Lindernia crustacea 403 

Linnaea borealis, var. americana 
321 

Linum westii 392 

Listera cordata 316 

Lobelia Kalmii 160 

Lomentaria baileyana 276 

Lonchocarpus sericeus 416 

Lonicera canadensis 321 

Lowden, Richard M. Revision of 
the Genus Pontederia L. 426 

Ludwigia spathulata 394; sphae- 
rocarpa 395; suffruticosa 395 

Luzula multiflora 316 

Lyallia kerguelensis 241, 243; on 
Kerguelen Islands, Palynologi- 
eal Evidence for the Late Gla- 
cial Occurrence of Pringlea and 
239 

Lycopodium alopecuroides 368; 
annotinum, var. acrifolium 313; 
clavatum 144, 256; lucidulum 
256; obscurum, f. dendroideum 
313; prostratum 368 

Lycopus uniflorus 320; virginicus 
320 

Lyngbya 60; aerugineo-coerula 
285; aestuarii 285; confervoides 
285; gracilis 285; lutea 285; 
semiplena 286 

Lysimachia gramineum 397; lan- 
ceolata 398; quadrifolia 398; 
terrestris 320 

Lythrum salicaria 56 


Maianthemum canadense 316 

Maine Flora, Some New Locals 
in the 166 

Maine and New Hampshire, Chi- 
maphila maculata (L.) Pursh 
in 162 

Manisuris rugosa 374; tubercu- 
losa 374 


1973] 


Manning, Wayne E. The North- 
ern Limits of the Distribution 
of Hickories in New England 
34 

Marshallia mohrii 368, 408 

Maryland, Including the Chesa- 
peake Bay Area, The Marine 
Algae of Virginia and 258 

Mastigocoleus 293 

Mathieson, A. C. and Richard A. 
Fralick Benthic Algae and 
Vascular Plants of the Lower 
Merrimack River and Adjacent 
Shoreline 52 

Matricaria maritima, var. agres- 
tis 321; matricarioides 321 

Maytenus buxifolia 418; phyllan- 
thoides 418 

McCain, John, R. B. Pike and A. 
R. Hodgdon The Vascular 
Flora of Kent Island, Grand 
Manan, New Brunswick 311 

Medicago minima 391; orbicu- 
laris 391 

Melanthera amethystina 192; an- 
gustifolia 172, 174, 176-179, 
181, 183, 187, 197-199, 202, 206; 
var. subhastata 198; aspera 
169, 171, 177, 180, 181, 188, 190, 
192, 206; var. aspera 174, 176, 
177, 182, 184, 187, 188, 190, 192, 
193; var. glabriuscula 171, 173, 
174, 176, 177, 181, 182, 184, 187, 
194, 195, 207; brevifolia 194; 
buchii 192; calcicola 194; can- 
escens 192; carpenteri 184; 
(Compositae), A Revision of 
North American and Caribbean 


169; confusa 194; corymbosa 
192; crenata 194; deltoidea 
192; fruticosa 206; hastata 


171, 184, 186, 188; ssp. cuben- 
sis 193; var. cubensis 192; var. 
lobata 184; var. pandurata 184; 
var. parvifolia 200; hastifolia 
206; lanceolata 198, 199; ligu- 


Index to Volume 75 


637 


lata 172, 174, 176, 177, 180, 183, 
187, 199, 203-205; linearis 177, 
198; Linnaei 192; lobata 184; 
microphylla 198;  molliuscula 
192; montana 193; nivea 170, 
171, 179, aay 1716177, 180) 
183-188, 199, 205-207; oxycar- 
pha 184; oxylepis 184, 188; 
panduriformis 171, 184; parvi- 
ceps 206, 207; parvifolia 172, 
174, 176-180, 182, 187, 199-201; 
purpurascens 198, 199; radiata 
179, 200; trilobata 184; urtici- 
folia 192 

Melica nitens 371 

Meliotus alba 416 

Melosira 59 

Menyanthes trifoliata, var. minor 
320 

Merismopedia 60; glauca 284 

Merremia tuberosa 420 

Merrimack River and Adjacent 
Shoreline, Benthic Marine Al- 
gae and Vascular Plants of the 
Lower 52 

Mertensia maritima 320 

Mesadenia diversifolia 409 

Metastelma graminifolium 420 

Miconia Robinsoniana 144 


Microcoleus chthonoplastes 286; 
lyngbyaceus 287; tenerrimus 
287; vaginatus 287 

Micromenia bahamensis 421; 


brownei 421 

Mimulus, Floral Structure, Hy- 
bridization and Evolutionary 
Relationship of Two Species of 
225; guttatus 225-237; nasutus 
225-237 

Mirabilis longiflora 235; 
ginea 161 

Monanthochloé littoralis 413 

Monarda clinopodia 482 

Moneses uniflora 319 

Monochoria hastata 431 


nycta- 


638 


Monostroma  leptodermum 262; 
oxyspermum 58, 60, 262 

Monotropa hypopithys 320; uni- 
flora 319 

Morinda citrifolia 421 

Morrow, Loring and Norton H. 
Nickerson Salt Concentrations 
in Ground Waters Beneath 
Rhizophora mangle and Avicen- 
nia germinans 102 

Mosla dianthera 401 

Muhlenbergia frondosa 372; gla- 
briflora 371 

Myrica gale 316 

Myrionema strangulans 269 

Myriotrichia subcorymbosa 269 


Myriophyllum  exalbescens 395; 
heterophyllum 115; pinnatum 
115 


Najas marina 412 

Narukila cordata 454; var. lanci- 
folia 454, 562 

Nemalion multifidum 273 

New Brunswick, The Vascular 
Flora of Kent Island, Grand 
Manan 311 

New England, The Northern 
Limits of the Distribution of 
Hickories in 34 

New Hampshire, Chimaphila ma- 
culata (L.) Pursh in Maine and 
162; Plants, New or Note- 
worthy 158; Two Interesting 
Plants on Mt. Cardigan, 
Orange, 155 

Nickerson, Norton H. and Joseph 
W. Tripp Floral Dimorphism in 
Rachicallis americana (Jacq.) 
Hitch. (Saltwater Bush) 111; 
see also Morrow, Levring 

Nodularia harveyana 292 

Nuphar luteum 68; microphyllum 
65-71, 73; Some Morphological 
Aids in Distinguishing From 
Similar Aquaties 65; X rubro- 


Rhodora 


[Vol. 75 


discum 66-68, 70, 73; variega- 
tum 66-70 
Nymphoides cordata 65, 70-73 


Oenothera perennis 319 

Onoclea sensibilis 313 

Oscillatoria 60; amphibia 287; 
brevis, var. neapolitana 287; 
laetevirens 287; lutea 288; 
margaritifera 288; nigro-viridis 
288; princeps 288; retzii 288; 
salinarum 288; splendida 289; 
submembranaceae 289; subuli- 
formis 289 

Osmunda cinnamomea 313 

Ott, Franklyn D. The Marine 

Algae of Virginia and Mary- 
land, Including the Chesapeake 
Bay Area 258 

Oxalis montana 319 

Oxopolis filiformis 420 


Panax quinquefolium 167 

Panicum adspersum 373; amarum 
373; capillare 136; echinulatum 
413; linearifolium 373, 374; 
malacophyllum 373; rigidulum 
166; webberianum 373; werneri 
374 

Parks, James C. A Revision of 
North American and Caribbean 
Melanthera (Compositae) 169 

Paspalum bakeri 413; laxum 413, 
minus 372; molle 413; propin- 
quum 413; urvillei 413; vagina- 
tum 372 

Pediastrum boryanum 266; du- 
plex 266; simplex 266 


Pellaea atropurpurea 122-125, 
127-130: And Pellaea glabella, 
var. glabella, | Chromosome 


Pairing in Obligately Apoga- 
mous Ferns 122; glabella 122; 
var. glabella 123, 124, 126-130; 
var. occidentalis 129 
Percursaria percursa 264 


1973] 


Pernettya Howellii 142 

Persicarria bicornis 386 

Petalonia fascia 59, 62, 270 

Petalostemon foliosus 132-138; 
The Past and Present Geogra- 
phical Distribution of, and 
Notes on Its Ecology 132 

Philactis Liebmannii 206 

Phleum pratense 315 

Phlox divaricata 397; pilosa, var. 
ozarkana 400 

Phoenix dactylifera 414 

Phormidium . persicinum 289; 
retzii 289; submembranaceum 
289; unciniatum 289; weissi 289 

Physaliastrum japonicum 364; 
savatieri 364 

Physalis lobata 333; 364; longi- 
folia 402; sinensis 364; subgla- 
brata 402; wrightii 333 

Phytolacca purpurascens 415 

Picea glauca 314; mariana 314; 
rubens 3 

Pike, R. B, see McCain, John 

Pilaiella littoralis 59 

Pinus elliotti 177 

Piriqueta caroliniana 419; tomen- 
tosa 419; viridis 419 

Pisonia floribunda 143 

Plantago hookeriana 403; juncoi- 
des 56; var. decipiens 320; var. 
glauca 321; major 320; oligan- 
thos 56, 321: patagonia 403; 
virginica 421: wrightiana 403 

Plectonema  calothricoides 293; 
golenkinianum 293: tenebrans 
293 

Poa annua 314: compressa 314; 
Cookii 240; kerguelensis 240; 
palustris 314: pratensis 314 

Poinsettia cyathophora 417, 418; 
heterophylla 418 

Polanisia dodecandra 389 

Polygonella fimbriata 386 

Polygonum allocarpum 317; am- 
phibium 317: erectum 386; 


Index to Volume 75 


639 


hydropiper 57; hydropiperoides 
317; lapathifolium 317; longi- 
stylum 386 

Polysiphonia denudata 280; fibril- 
losa 59, 60; harveyi 281; ni- 
grescens 281;  novae-angliae 
282; paniculata 488, 490; An 
Unusual Substrate For 488; 
subtillissima 282; urceolata 
282 

Pontederia angustifolia 452-454; 
brasiliensis 473; caerulea 453; 
cordata 426-429, 431, 433, 435, 
443, 445, 4471, 448, 450, 452-454, 
460; f. albiflora 454; f. angusti- 
folia 453, 454, 462; f. bernardi 
455; f. brasiliensis 453, 454; 
f. latifolia 454; f. sagittata 
467; f. taenia 455; var. albiflora 
453; var. angustifolia 452; var. 
cordata 430, 436, 442, 446, 450- 
452, 456-458, 460, 464, 466, 484, 
486, 487; var. lanceolata 462; 
var. lancifolia 436, 445, 440, 
450, 454, 4577, 458, 460, 461, 464, 
467, 486, 487; var. ovalis 430, 
445, 446, 450, 461, 464, 466, 486, 
487; var. parviflora 471; var. 
sagittata 467; var. typica 453, 
462; cordifolia 473; eriantha 
473; hastata 427-429, 431; hete- 
rantherimorpha 462; L., Revi- 
sion of the Genus 426: lagoén- 
sis 478; lanceolata 462; f. bra- 
siliensis 454; f. ovalis 466; f. 
trullifolia 454; var. vichadensis 
466; lancifolia 460-462; nym- 
phaeifolia 453; oblonga 462; 
ovalis 461, 466; ovata 427, 428; 
parviflora 430, 432, 434, 435, 
437, 439, 441, 445, 446, 450, 
469, 471-473, 484, 486, 487; 
reniformis 473; rotundifolia 
430, 432-435, 438, 439, 441-443, 
445, 446, 450, 473, 474, 476, 
480, 484, 486, 487; var. nym- 


640 


phaeifolia 454; sagittata 430, 
434, 435, 437, 445-447, 450, 460, 
467-469, 478, 484, 487; schom- 
burgkiana 478; subovata 435, 
438, 443, 449, 450, 478-480 

Porphyra leucosticta 59, 60, 62, 
272; miniata 272; umbilicalis 
59, 272 

Porphyrosiphon 
splendidus 290 

Potamogeton fibrillosus 75, 76; 
A Re-evaluation of 75; foliosus 
75, 76; var. fibrillosus 76, 77; 
var. foliosus 77 

Potentilla anserina 318; egedei, 
var. groenlandica 57; norvegica 
318, 390; tridentata 155 

Psidium galapageium 143 

Pseudendoclonium marinum 58, 
61, 262 

Pteridium aquilinum 144 

Pteris vittata 412 

Ptilimnium costatum 395 

Ptilota serrata 59, 60, 62 

Pucciniellia paupercula, var. alas- 
kana 314 

Punctaria latifolia 270; plantagi- 
nea 270 

Pylaiella littoralis 268 

Pyrus americana 318; decora 318; 
floribunda 318; malus 318 

Pringle Herbarium, The 297 

Pringlea and Lyallia on Kergue- 
len Islands, Palynological Evi- 
dence for the Late Glacial Oc- 
currence of 239 

Pringlea antiscorbutica 241, 242 

Proctor, George R. see Gillis, 
Wm. T. 

Protoderma marinum 262 


notarisii 289; 


Rachicallis americana 111-113; 
(Saltwater Bush), Floral Di- 
morphism in 111 

Ralfsia verrucosa 59 


Ramalina calicaris, f. pyrifera 


Rhodora 


[Vol. 75 
307, 308; var. subamplicata 
306, 307; f. subpapillosa 307, 
308; canariensis 307; fasti- 
giata, var. subampliata 306, 


307; subampliata 306, 307, 309; 
(Nyl) Fink in North America, 
The Status of 306 

Ramium niveum 386 

Ranunculus 57; acris 318; cymba- 
laria 318; Mosleyi 240 

Ratibida columnaris 407; columni- 
fera 407 

Reimarochloa oligostachya 372 

Reussia grazielae 474; lagoénsis 
478; rotundifolia 474; subovata 
478; triflora 449, 473, 478 

Reveal, James L. see Robert R. 
Haynes 

Reviews 114, 140, 323, 622 

Rheum rhaponticum 317 

Rhexia mariana 394; var. mari- 
ana 394; salicifolia 369, 394; 
virginica 394 

Rhinanthes crista-galli, var. fal- 
lax 320 


Rhizoclonium kockianum 267; 
riparium 58, 269; tortuosum 
267 


Rhizophora mangle 102-105; And 
Avicennia germinans, Salt Con- 
centrations in Ground Waters 
Beneath 102 

Rhodochorton amphiroae 551, 
555; attenuatum 583; catenula- 
tum 86, 551; concrescens 568; 
daviesii 81, 551; efflorescens 
584; emergens 572; endophyti- 
cum 574; entozoicum 589, 590, 
698; hallandicum 84; humile 
569; infestans 91; intermedium 
536; islandicum 536, 540; mem- 
branaceum 91, 94, 566; meso- 
carpum, var. penicilliformis 
585; microscopicum 86, 557; 
moniliforme 87, 558; parasiti- 
cum 536, 540; penetrale 92; 


1973] 


penicilliforme 585, 586, 603, 
608; purpureum 530, 536, 540, 
608; rhipidandra 541; Rothii 
540; spetsbergense 585; sub- 
simplex 576; tenue 537, 540; 
tenuissimum 576, 581; thuretii 
88, 560; virgatulum 94, 577 

Rhododendron  austrinum 396, 
397; calendulaceum 396; cana- 
dense 320; canescens 397; 
maximum 163 

Rhus aromatica 135 

Rhyncheletrum repens 413; ro- 
seum 413 

Rhynchospora capillacea 161; fer- 
naldii 380; perplexa 368; plei- 
antha 380 

Ribes glandulosum 318; hirtellum 
318; lacustre 318 

Rigby, Sylvia Jane Chromosome 
Pairing in Obligately Apoga- 
mous Ferns; Pellaea atropur- 
purea and Pellaea glabella, var. 
glabella 122 

Rivularia nitida 293 

Rogers, O. M. see Hill, L. Michael 

Rorippa islandica, var, fernaldi- 
ana 318; var. hispida 318 

Rosa rugosa 318 

Rotala ramosier 422 

Rubus chamaemorus 318; hispidus 
318; idaeus, var. canadensis 318 

Rudbeckia hirta 136, 407; mollis 
407; triloba 136 

Rumex acetosella 317; domesticus 
317; fenestratus 317; orbicula- 
tus 317; pallidus 317 

Rundel, Phillip W. see Bowler, 
Peter A. 

Ruppia maritima 142 


Sabatia difformis 398; paniculata 
398; quadrangula 398; stellaris 
420 

Sagina procumbens 317 

Sagittaria isoetiformis 368 


Index to Volume 75 


641 


Salicornia europaea 53, 54, 56, 
317 

Salix amygdaloides 
rigida 385 

Sanguisorba canadensis 256 

Saracha coronopus 339; potosina 
364 

Sargasso Sea, The Audouinella 
Complex (Rhodophyta) in the 
Western 78 

Sargassum fluitans, 79, 95; hy- 
strix 271; natans 79, 84, 89, 95, 
271 

Sarracenia purpurea 318 

Satureja rigida 421 

Saxifraga rivularis 153, 154: In 
the White Mountains, New 
Hampshire, A New Station For 
153 


158, 161; 


Scenedesmus acuminatus 266; 
opaliensis 266; quadricauda 266 
Schistostega osmundacea 149; 


pennati 149; The Environment 
of, New Vermont Stations 149 

Schizothrix arenaria 290; calci- 
cola 290; tenerrima 290 

Schoenolirion croceum 368 

Schofield, Eileen K. see Young, 
Steven B. 

Scirpus americanus 315; 
cinctus 315; atrovirens 315; 
cyperinus 315: debilis 376; 
maritimus 54, 61; var. fernaldi 
57; purshianus 376; validus 54, 
57, 61 

Scutellaria epilobiifolia 320; race- 
mosa 401 

Seytosiphon lomentaria 59, 270 

Sedum roseum 318; telephium, f. 
purpureum 318 

Senecio anonymus 211-214, 216- 
218; An Earlier Name For 
Senecio smallii Britton 211; 
aureus, var. angustifolius 217; 
confusus 423;  earlei 219; 
smallii 211, 212, 214, 216, 217; 


atro- 


642 


f. tristis 219; sylvaticus 321; 
tomentosus 211, 214, 216-218; 
vulgaris 321 

Sesuvium microphyllum 415 

Setaria geniculata 413; faberi 
372 

Seymour, Frank C. The Pringle 
Herbarium 297 

Sida elliottii 393 

Silene gallica 387 

Sisyrinchium exile 415; micran- 
thum 415; montanum, var. 
crebrum 316 

Sium suave 57 

Smilax dominguensis 414; hava- 
nensis 414 


Smilacina stellata, var. crassa 
316; trifolia 316 
Solanum eleagnifolium 421; 


chamaesarachidium 364; coni- 
odes 359; coronopus 339 

Solidago elliotti 405; graminifolia, 
var. nuttallii 321; macrophylla 
321; odora 160; rugosa, var. 
villosa 321; sempervirens 53, 
54, 56, 321, 423; sempervirens, 
var. mexicana 423; uliginosa 
405 

Sonchus asper 322 

Sorghum halepense 413 

Sparganium americanum 314 

Spartina alterniflora 53, 54, 56, 
61, 315; patens 53, 54, 56, 61; 
spartinae 370 

Spathelia bahamensis 417 

Spergularia canadensis 317; ma- 
rina 57, 317; rubra 317 

Spermacoce aspera 422 

Spermolepis inermis 395; patens 
395 

Spiraea latifolia 318 

Spiranthes lucida 158, 160; ro- 
manzoffiana 316 

Spirogyra 261 

Spirulina major 


290; subsalsa 


Rhodora 


[Vol. 75 


291; f. oceanica 291; tenerrima 
291 

Sphacelaria fusca 268 

Spondias purpurea 418 

Spongberg, Stephen A. A New 
Alaskan Species of Erigeron L. 
(Compositae) 116 

Spongomorpha arcta 58, 60, 62 

Sporobolus vaginiflorus 135 

Spyridia filamentosa 278 

Stachys floridana 401; hispida 401 

Stanford, Jack W. Umaldy T. 
Waterfall (1910-1971) 146 

Staurastrum 261 

Steele, F. L. and A. R. Hodgdon 
Two Interesting Plants on Mt. 
Cardigan, Orange, New Hamp- 
shire 155 

Steironema gramineum 397, quad- 
riflorum 398 


Stellaria calycantha, var. iso- 
phyla 317; graminea 317; 
humifusa 317; media 317 
Stenandrium carolinae 422 
Stenophyllum capillaris 378 
Sterk, Fern Frank see Van 


Fassen, Paul 

Stilophora rhizoides 269 

Streptopus amplexifolius, var. 
americanus 316 

Striaria attenuata 269 

Stylosanthes calcicola 416; tuber- 
culata 416 

Suaeda americana 317; maritima 
317 

Svida microcarpa 396 

Swartz, Steven see Colt, L. C.. 
Jr. 

Symphoriocarpos orbiculatus 135 

Symploca atlantica 291 


Talinum triangulare 415 
Tanacetum vulgare 321 
Taraxacum erythrospermum 410; 
laevigatum 410: officinale 322 
Tennessee, Some Notes on the 


1973] 


Flora of the Southern States, 
Particularly Alabama and Mid- 
dle 366 

Terminalia muelleri 419 

Thalietrum debile 387; dioicum 
388; macrostylum 388; mirabile 
387; polygamum 318; texanum 
388 

Thamnidium intermedium 536, 
540; mesocarpum, f. penicilli- 
formis 585, 587; rothii 537; 
spetsbergense 585, 587 

Thelypteris palustris, var. pubes- 
cens 314 

Thermopsis villosa 159 

Thysanella fimbriata 386 

Tiffney, Wesley N., Jr. A New 
Station for Saxifraga rivularis 
L. in the White Mountains, 
New Hampshire 153 

Tofieldia glutinosa 160 

Trailliella intricata 273, 279 

Trentopohlia daviesii 81, 551, 577; 
purpurea 536; virgatula 95, 
577; var. secundata 577 

Trichodesmium thiebautii 291 

Trientalis borealis 320 

Trifolium angustifolium — 391; 
agrarium 319; pratense, var. 
sativum 318; repens 319; vesi- 
culosum 391 

Triglochin maritima 56, 
palustre 314 

Triosteum (Caprifoliaceae), Tri- 
bal Classification of 120: perfo- 
liatum 121 

Trismeria trifoliata 412 

Tryon, Rolla Aquatic and Wet- 
land Plants of the Southwest- 
ern United States (Review) 
114 

Tunica prolifera 387 

Typha latifolia 57, 314 


314; 


Ulothrix ^ endosporangia 261; 
flacca 58, 261; subflaccida 261 


Index to Volume 75 


643 


Ulva lactuca 58, 63, 364; var. 
latissima 265; var. rigida 265 

Uniola nitida 369 

Unisema acutifolia 453; cordata 
454; f. angustifolia 454; f. 
latifolia 454; deltifolia 452; 
heterophylla 453; var. lanceo- 
lata 453; var. stenocardia 453; 
lancifolia 462; f. trullifolia 
454; latifolia 452; var. albiflora 
453; var. elatior 453; var. pal- 
lida 453; var. undulata 453; 
media 452; var. albiflora 452, 
454; mucronata 452; obliquata 
452; obtusifolia 452; orbiculata 
473; peduncularis 453; var. 
parviflora 453; purshiana 459: 
rotundifolia 453; sagitata 452 

Urospora penicilliformis 58 

Urtica procera 317; viridis 317 

Utricularia resupinata 403 


Vaccinium angustifolium, var. 
laevifolium 320; oxycoccus 320; 
var. ovalifolium 320; vitis- 


idaea, var. minus 320 

Valeriana pauciflora 404 

Van Fassen, Paul and Fern Frank 
Sterk Chromosome Numbers 
in Aster 26 

Vaucheria 59-62 

Verbena bipinnatifida 400 

Vermont, New Stations (and) 
The Environment of Schistos- 
tega pennati (Hedw.) Hook & 
Tayl. 149 

Vernonia acaulis 519, 528, 525; 
angustifolia 519, 523-525, ar- 
kansana 519, 523-525; baldwinii 
525; ssp. baldwinii 519, 520, 
523, 526, 527; ssp. interior 519, 
520, 523, 526; blodgettii 519, 
523, 525, 526; The Systematic 
Value of Trichome Comple- 
ments in a North American 
Group of 517; ervendbergii 519, 


644 
523-525; fasciculata 519, 522- 
525; flaccidifolia 519, 522, 523, 
525; gigantea 519, 523-525; 
glauca 519, 525; greggii 
519, 523-525; interior 525; 


larsenii 519, 522, 523, 525; let- 
termannii 519, 523, 525; lindhei- 
meri 519, 522, 523, 525; mar- 
ginata 519, 520, 522, 523, 525; 
missurica 519, 520, 523, 525, 
526; noveboracensis 519, 523- 
525; pulchella 519, 523, 525; 
shaffneri 519, 523-525; texana 
519, 523-525 

Veronica anagallis-aquatica 482 

Vicia cracca 319 

Viguirea porteri 408 

Viola cucullata 319; pallens 319; 
septentrionalis 319 

Virginia and Maryland, Including 
the Chesapeake Bay Area, The 
Marine Algae of 258 

Virginia, Phytogeography of the 
Carices of 248 

Vitis munsoniana 393 


Warea amplexifolia 389; cunei- 
folia 389; sessilifolia 389 

Waterfall, Umaldy T. (1910-1971) 
146 

Wicks, Johonet C. Some Morpho- 
logical Aids in Distinguishing 


Rhodora 


[Vol. 75 


Nuphar microphyllum from 
Similar Aquatics 65 

Withania sordida 355 

Woelkerling, Wm. J. The Audou- 
inella Complex (Rhodophyta) 
in the Western Sargasso Sea 
78; The Morphology and Sys- 
tematics of the Audouinella 
Complex (Acrochaetiaceae, Rho- 
dophyta) in the Northeastern 
United States 529 


Xanthocephalum dracunculoides 
404 
Xyris difformis, var. floridana 


381; fimbriata 381, 382; iridi- 
folia 382; jupicai 382; longi- 
sepala 369, 382; platylepis 
382; smalliana 382 


Young, Steven B. and Eileen K. 
Schofield Palynological Evi- 
dence for the Late Glacial Oc- 
currence of Pringlea and Lyal- 
lia on Kerguelen Islands 239 

Youngia japonica 423 


Zanthoxylem bifoliatum 417 

Zizania aquatica 54, 57, 61 

Zostera marina 488; var. steno- 
phylla 314 


Ln 


1282 


STATEMENT OF OWNERSHIP, MANAGEMENT AND 
CIRCULATION 


Title of Publication: RHODORA 
Date of Filing: October 22, 1973 
Frequency of Issue: Quarterly 


Location of Known Office of Publication: 
Botanical Museum, Harvard University, Oxford Street, 
Cambridge, Mass. 02138 


Location of the Headquarters of the Publishers: 
Botanical Museum, Harvard University, Oxford Street, 
Cambridge, Mass. 02138 


Publisher: 
New England Botanical Club, Inc., Botanical Museum, 
Harvard University, Cambridge, Mass. 02138 


Editor: 
Dr. Albion Hodgdon, Dept. Botany, 
University of New Hampshire, N.H. 03824 


Managing Editor: 
Dr. Herman R. Sweet, Botanical Museum, 
Harvard University Cambridge, Mass. 02138 


Owner: New England Botanical Club, Inc., Botanical Museum, Oxford 
Street, Cambridge, Mass. 02138 


No stockholders 
Known Bondholders, Mortgagees, and other Security Holders: None 


Average No. Copies Single issue 

each issue during nearest to 

preceding 12 months filing date 
Total No. copies printed 850 850 

Paid Circulation 

Sales through dealers, etc. none none 
Mail subscriptions 782 792 
Total paid circulation 782 792 
Free distribution 12 12 
Total distribution 794 804 


I certify that the statements made by me above are correct and 
complete. Herman R. Sweet, Business Manager and Treasurer.