SMITHSONIAN CONTRIBUTIONS TO BOTANY 0 NUMBER 35 
The Genus Colpomenia 
Derbes et Solier (Phaeophyta) 
in the Gulf of California 
Michael 3. Wynne 
and James N. Norris 
SMITHSONIAN INSTlTUTION PRESS 
City of Washington 
1976 
ABSTRACT 
Wynne, Michael J., and James N. Norris. The Genus Colpomenia Derbks et 
Solier (Phaeophyta) in the Gulf of California. Smithsonian Contributions to 
Botany, number 35, 18 pages, 11 figures, 1976.-Four species of the brown algal 
genus Colpomenza Derbes et Solier (Scytosiphonales) are recognized as occurring 
within the Gulf of California: C. sinuosa (Roth) Derbes et Solier, C. tziberculata 
Saunders, C. ramosa Taylor, and C. phaeodartyla, new species. Some of the 
specimens interpreted by Dawson (1944) as Rosenvingea intricata are now re- 
ferred to C. ramosa Taylor. Although recorded from the Pacific coast of Baja 
California, C. ramosa has not yet been reported within the Gulf. Interrelation- 
ships of the species complex of Colpomenia with related genera such as Iyen- 
garia Boergesen, Rosenvingea Boergesen, and Scytosiphon C. Agardh are dis- 
cussed. 
OFFICIAL PUBLICATION DATE is handstam ed in a limited number of initial copies and is recorded 
in the Institution?s annual report, Srnitisonian Year. SERIES COVER DESIGN: Leaf clearing from the 
katsura tree Cercidphyllum japonicum Siebold and Zuccarini. 
Library of Congress Cataloging in Publication Data 
Wynne, Michael James. 
The genus Colpomenia Derbes et Solier (Phaeophyta) in the Gulf of California. 
(Smithsonian contributions to botany ; no. 35) 
Bibliography: p. 
1. Colpomenia. 2. Marine algae-California, Gulf of. I. Norris, James N., joint author. 11. 
QKl.Sz747 no. 35 [QK569.S36] 581?.08s [589?.45] 76-8392 
Title, 111. Series: Smithsonian Institution. Smithsonian contributions to botany ; no. 35. 
Contents 
Introduction . , , , , , , , . . , . . , . . , . . . . . . 
..(.............. ,........ 
Key to Colpomenia Species in the Gulf of California . . . . . . . . . . . . . . . 
2. Colpomenia phaeodactyla, new species , 
4. Colpomenia ramosa Taylor . . . . . . . . . . . , . , . . . . . . . . , . . . . . . . . 
Discussion , ................... , .,......... ..................... 
Ecological Observations . . . . . . . . . . . . . . . , . . . . . . . . . . 
Literature Cited ,.. ...,.. ........... ,,.,...., ...... ..... .. 
1. Colpomenia sinuosa (Roth) Derbes et Solier . . . . . . . . .... 
3. Colpomenia tuberculata Saunders , . , . ,. .. 
. . . . . . . . . . . . . . . . . . . . . . . 
... ....... 
Page 
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111 

The Genus Colpomenia 
Derbes et Solier (Phaeophyta) 
in the Gulf of California 
Michael J. Wynne 
and James N. Norris 
Introduction 
Three distinctive morphological forms of Cob 
pomenia Derbes et Solier within the Gulf of Cali- 
fornia, Mexico, have been recognized by previous 
authors (Setchell and Gardner, 1924; Dawson, 1944, 
1966) as representing separate forms of a single 
variable species, C. sinuosa (Roth) Derbes et Solier: 
f. sinuosa, f. deformans Setchell and Gardner, and 
f. tuberculata (Saunders) Setchell and Gardner. Ac- 
cording to Dawson (1944) the latter also included f. 
expansissirnu Setchell and Gardner. Under its var- 
ious guises Colpomenia collectively forms a domi- 
nant element of the intertidal winter and spring 
flora of the Gulf of California. Field and micro- 
scopic studies, however, have led the present au- 
thors to conclude that the Colpomenia complex in 
the Gulf includes four distinctive species, separated 
by morphological, anatomical, and habitat differ- 
ences. The variable appearance of these species of 
Colpomenia raises problems of distinguishing them 
from related genera of the family Scytosiphonaceae. 
This paper, then, analyzes certain of the generic 
delimitations within the family. 
Specimens studied are deposited in the herbaria 
of the following institutions: United States Na- 
Michael J. Wynne, Department of Botany, The University of 
Texas at Austin, Texas 78712. James N. Norris, Department 
of Botany, National Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 20560. 
1 
tional Herbarium, Smithsonian Institution (US); 
University of Michigan (MICH); University of 
California, Berkeley (UC); University of Texas at 
Austin (TEX); University of Arizona, Tucson 
(ARIZ); Universidad Autonoma de Mexico 
(MEXU); Allan Hancock Foundation Herbarium, 
University of Southern California (AHFH); and the 
Gilbert Morgan Smith Herbarium, Hopkins Marine 
Station of Stanford University, Pacific Grove, Cali- 
fornia (GMS). Collection numbers of J. N. Norris 
carry the prefix JN- and refer to his field note- 
books. Specimens were collected by him unless 
otherwise noted; those collected by Katina E. 
Bucher are noted as KB. 
ACKNOWLEDGMENTS.-we wish to thank Dr. Rob- 
ert W. Hoshaw for the use of facilities while J. 
Norris was station director/resident marine biolo- 
gist at Laboratorio de Biologia Marina of the Uni- 
versity of Arizona, and for the loan of Dawson 
material (ARIZ). J. Norris is also grateful to Carl 
N. Hodges, Director, Environmental Research Lab- 
oratory, for the use of Laboratorio de Biologia 
Marina in Puerto Penasco, Sonora, Mexico. Support 
for field work in the Gulf of California was pro- 
vided by a National Science Foundation grant NSF- 
BMS-73-07000 (formerly GB-38623) awarded to J. 
Norris and hl. Neushul. It is a pleasure to acknowl- 
edge the support and encouragement that Dr. 
Michael Neushul has continuously provided. We 
also appreciate the critical reading and comments 
of Dr. Harold Robinson, Finally thanks are due 
2 
Drs. M. 51, Littler, F. R. Fosberg, and B. Simpson 
for their comments on the ecological observations. 
We wish to acknowledge the following for kindly 
loaning specimens: Dr. I. A. Abbott (GMS); Dr. 
TV. R. Taylor (MICH and TVRT); Dr. N. L. Nich- 
olson and Llr. R. B. Setzer (AHFH); and Dr. C. T. 
Rogerson (KY). We thank Dr. P. C. Silva for 
searching for the type of C. tuberculata Saunders 
in the University of California, Berkeley Herbar- 
ium. Dr. Mark hl. Littler has generously shared 
unpublished productivity data for use in this pa- 
per. Dr. Hannah Croasdale kindly provided the 
Latin diagnosis. h1s. Alice Tangerini drew C. ~a- 
mosa and prepared the bar graphs. 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
For the opportunity to participate on the R/V 
Dolphin cruise to the northern Gulf of California, 
JN thanks Dr. William Fenical, Chief Scientist, 
Scripps Institution of Oceanography. He is also 
grateful to the scuba diving and collecting assist- 
ance of Katina Bucher, Dr. Howard Sleeper, ;\lark 
Helvey, George Boehlert, and David Moore. Thanks 
are due to Drs. Richard S. Felger, Mark 51. Littler, 
Richard Evans Schultes, Andrew T. TVeil, and 
Sicholas P. Yensen, Diane Littler, James Woessner, 
and William Sliheeler for generously sharing their 
collections. Finally, we thank Katina Bucher for 
her help throughout this study and assistance in 
processing the specimens studied. 
Key to CoZpomenia Species in the Gulf of California 
1. Thalli consisting of clustered finger-like sacs aiising from a common prostrate holdfast; 
wall of sacs thin, delicate, usuall? less than 100 pm thick 2. C. Phaeodactyla 
Thalli not consisting of such finger-like sacs, thalli ranging from spherical, balloon-like 
shapes to irregularly shaped, prostrate or sessile expanses; thallus wall thicker, firmer, 
2 
2. Thalli with irregular lobed branches, forming a spreading adherent clump 4. C. rarnosa 
Thalli smooth or tuberculate but lacking definite branches, forming sessile expanses or 
3. Thalli with tuberculate, !\arty surface, of a crisp, rigid texture: thickness exceeding 0.5 mm, 
3. C. tuberculata 
Thalli with smooth or conloluted surface, hut nexer tuberculate or rigid; thickness 0.3-0.5 
1. C. sinuosa 
exceeding 200 pm in thickness 
globose sacs 3 
often about 1.0 mm; plurilocular organs up to 25 pm long 
mm; plurilocular organs 40 pm or more long 
1. Colpomenia sinuosa (Roth) Derbks et Solier 
FIGURES 1, 2a,b, lla 
Ulva smuosa Roth, 1806:327, pl. 12a-c. 
CoIpornenia sinuosa (Roth) Derbes et Solier, 1856:11, pl. 22: 
figs. 18-20.--Saunders, 1898: 164, pl. 32: figs. 7,8.-Setchell 
and Gardner, 1925:539, pl. 45: figs. 82-86 [with reference 
to Gulf of California specimens only].-Dawson, 1944232; 
1959319; 1966:lO. [Cf. Misra, 1966:115-6, for synonymy.] 
DESCRIPTIoN.-Thalli (Figure 1) globular or vesic- 
ular, becoming irregularly convoluted and ex- 
panded with age, reaching a diameter of 14 cm; 
wall 300-500 pm thick, composed of a surface layer 
of small pigmented cuboidal cells (Figure 2a) and 
4-6 layers of gradually larger, irregularly shaped 
subcortical and medullary cells; plurilocular organs 
(Figure 2b) uniseriate (to partially biseriate), about 
40 pm in length, clustered in fairly discrete sori 
often around an invaginated tuft of hairs; pa- 
raphyses longer than plurilocular organs, to 55 pm 
in length; color golden brown; unilocular sporangia 
unknown. 
TYPE-LocALITY.-cadiz, Spain. 
HoLoTYPE.-Destroyed at Berlin-Dahlem (ac- 
cording to Dawson et al., 1964). 
rocks and epiphytic on various algae, subtidal (1 3- 
16 m depths) to midlittoral levels; throughout the 
Gulf of California from Puerto Peiiasco to Bahia 
Agua Verde. 
April. 
ical seas throughout the world. 
GULF OF CALIFORNIA DIsTRIsUTIoN.-Growing On 
GULF OF CALIFORNIA SEASONALITY.-NOvember- 
DISTRIBUTION.-COmmOn in tropical and subtrop- 
SPECIMENS EXAMIXED.--GUI~ of California. Sonora: Punta 
Pelicano, vicinity of Puerto Penasco, 17 Mar 1973, JN-3811 
(UC); 19 Apr 1973, JN-3966 (MEXU); 9 Mar 1974, J9-4982 
(US), (leg. JN and KB); Feb 1965, Darison 27558 (ARIZ). 
Playa Arenosa, vicinity of Puerto Pefiasco, 26 Dec 1972, 
Wynne 3754 (MICH, TEX); 17 Apr 1973, JK-3911 (MICH). 
3 
NUMBER 35 
Playa Hermosa, Puerto Penasco, littoral rock platform and 
sand areas, 2 Feb 1973, JS-3702 (MICH). Playa Estacion, 
Puerto Penasco, littoral rock platform and tide pools, Feb 
1968, (TEX), (leg. R. Hoshaw); 25 Dec 1972, Wynne 3733 
(MICH, TEX); 22 Jan 1973, JN-3621 (US); 25 Nov 1972, 
JN-3663 (UC), (leg. JN and KB); 4 Feb 1973, JN-3746 
(MICH); 4 Mar 1973, JN-3783 (US), JN-3786 (TEX), JS-3801 
(MEXC), (leg. JN and KB); 3 Feb 1973, JN-4717 (UC); 18 
Jan 1974, J3-4760 (US), (leg. Jh' and KB); 9 Feb 1974, JN- 
4929 (UC); 5 Feb 1974, JN-4952 (US) and JN-4956 (ARIZ, 
MICH, US); 1 Mar 1974, JN-5013 (ARIZ, MICH); 24 Mar 
1974, JN-5108 (US), (leg. J. Woessner and W. Wheeler). N 
side of Punta Lobos, licinity of Puerto de Lobos, 3 m depth, 
sand covered rock, 17 Feb 1973, JN-6035 (US), (leg. M. 
Helvey). IV side of Punta Robinson, vicinity of Puerto 
Libertad, 4.5 m depth, rock reef, 17 Nov 1973, JN-4824 
(UC), (leg. JN and KB). Punta Cirio, S of Puerto Libertad, 
epiphytic on Sargassum stipes, W side of cobe, 18 Nov 1973, 
JN-4913 (TEX), (leg. JN and KB). Desemboque de San 
Ignacio, drift, 3 Apr 1974, JN-5140 (TEX), (leg. R. S. Felger, 
R. E. Schultes, and A. T. Weil). Isla San Pedro Nolasco, 
epiphytic on Sargassum stipe, NE side of cove, 20 Mar 1974, 
JS-5206 (GMS), (leg. N. P. Yensen). Baja California: Isla 
Willard, Bahia San Luis Gonzaga, littoral rocks, 20 Apr 
1974, JN-5400 (MEXU, US), (leg. JN and KB). Islas de la 
Cintura: 3 m depth, E end of Isla Mejia, Puerto Refugio, 
23 Apr 1974, JN-5681 (AHFH), leg. JN and KB). Isla Angel 
de la Guarda, low littoral rocks, NE shore of Puerto Refugio, 
23 Apr 1974, JS-5787 (MICH, US). 
REMARKS.-BlaCkler (1964) differentiated the two 
FIGURE ].-Habits of Colpomenia sznuosa: a,b, from Playa 
Arenosa, Puerto Penasco (Wynne-3754). 
' 50rm ' 
a 
x 
FIGURE 2.-Transections of Colpomenia sinuosa: a, a sterile thallus from Playa Estacion, Puerto 
PeAasco, (JN-4952); b, fertile plant showing the uniseriate plurilocular organs and unicellular 
paraphyses, also from Playa Estacibn (Wynne-3733). 
4 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 3.-Open expanded-sheet form of Colpomenia sinuosa from 13-16 meter depth, off Isla la 
Ventana, Bahia de Los Angeles UN-2995). 
species of Colpomenia (C. sinuosa (Roth) Derbks et 
Solier 1856 and C. peregrina (Sauvageau) Hamel 
1937) occurring on the west coast of North America, 
which had been confused in the literature (Setchell 
and Gardner, 1925; Smith, 1944). They had pre- 
viously been recognized as distinct in Europe by 
Sauvageau (1927). Apparently it is only in the 
southern California region that the ranges of these 
two species overlap; C. peiegiina occurs from 
Amchitka Island in the Aleutians to La Jolla, Cali- 
fornia, while C. sinuosa ranges southward from 
Point Concepcion including the Channel Islands 
(Blackler, 1964). 
According to Sauvageau (1927) and Blackler 
(1964, 1967), C. sinuosa is characterized by fairly 
firm, thick, often deeply folded thalli. With 
pressing, these are yellowish-brown in color. Dis- 
crete punctate sori of plurilocular organs are cov- 
ered with a cuticle. On the other hand, C. pere- 
grina is characterized by relatively thinner, smooth 
thalli, greenish in color upon pressing, with broad, 
confluent fertile zones,' lacking a cuticle, with pluri- 
locular organs often composed of two rows of 
locules, and paraphyses shorter than the plurilocular 
'Nonconfluent sori were reported in C. peregrina in h'or- 
way (Lund, 1949), whereas Clayton (1975) has presented 
statistical evidence that the shape of the sori and presence 
or absence of a cuticle over the plurilocular organs are the 
only criteria for effectively distinguishing Australian popula- 
tions of these trvo species. 
NUMBER 35 
5 
organs. Sauvageau (1927) reported that the thalli 
of C. sinuosa are 420-560 pm thick and the pluri- 
locular organs are 40 (-50) pm high, while thalli 
of C. peregrina are 140-280 pm thick and pluri- 
locular organs are about 20 pm high, The Gulf of 
California material agrees in large measure with 
the combination of traits of C. sinuosa, except that 
the plurilocular organs (Figure 2b) are both uniseri- 
ate and biseriate (or intermediate). Material illus- 
trated by Okamura in Japan (1936, fig. 123) and 
Chamberlain for Gough Island in the South Atlan- 
tic (1965) showed a similar situation in C. sinuosa. 
Colpomenia sinuosa f. lacunosa was described by 
Taylor (1947) from Paita, Peru. Subsequent treat- 
ment by Dawson et al. (1964) placed this taxon into 
synonymy with C. sinuosa. Since the present paper 
records C. tuberculata from Peru, we considered it 
desirable to examine the type material of C. sinuosa 
f. lacunosa. We have inspected the holotype (US) 
and concur with the conclusion of Dawson et al. 
(1964) that it should be placed with C. sinuosa. The 
thickness of the wall is in the range of C. sinuosa 
rather than C. tuberculata. 
Open expanded blades (Figure 3) that appear 
to be a form of C. sinuosa were recently collected 
during subtidal surveys. Apparently a deep water 
form, they were found growing attached at 13-16 m 
depths. These sheets were smooth, fairly thin, and 
measured up to 25 cm broad. Our specimens 
lacked the ?minute, spine-like projections? de- 
scribed by Setchell and Gardner (1924:726; 1925: 
541) for their C. sinuosa f. expanissima. Their form 
was found floating in Bahia de San Francisquito, 
whereas the recent collections were attached and 
subtidal. Dawson regarded the former taxon as a 
form of C. sinuosa f. tuberculata, which ?achieved 
extensive development in a detached floating con- 
dition . . .? (1944:233). Unfortunately our open- 
sheet form is not fertile, and lacking this, it seems 
judicious to merely call attention to this new ma- 
terial. Thus far only known from the Gulf of Cali- 
fornia, for now we refer our specimens to C. 
sinuosa. The following collections are cited: Sonora: 
Ensenada de San Francisco, vicinity of Puerto San 
Carlos, 24 Mar 1949, Dawson 7231 (US). Baja 
California: Puerto Calamajue, 23 Mar 1973, JN- 
4607 (US), (leg. JN and KB); Punta La Gringa, 
Bahia de Los Angeles, 8 m, 28 Apr 1974, JN-6103b 
(US), (leg. JN); Isla la Ventana, Bahia de Los 
Angeles, 13-16 m, 21 May 1972, JN-2995 (US, UC), 
(leg. JN and G. Boehlert); Bahia Salinas, Isla 
Carmen, 4-14 m, 20 Mar 1965, Dawson 6986 (US); 
Puerto Escondido, 4-8 m, 18 Mar 1949, Dawson 
7144 (US). Las Islas de la Cintura: Isla San Esteban, 
6.5-39 m, 26-27 Jan 1960, Dawson 21547 (US), (leg. 
Calif. Dept. Fish and Game). 
2. Colponaenia phaeoductyla, new species 
FIGURES 4, 5a,b, llc 
C. sinuosa f. deformans of Setchell and Gardner, 1924:726, pl. 
19: figs. 61,62; 1925:542 [in part, with reference to the Gulf 
of California specimens only].-Daw~son, 1944:233; 1966: 11 .- 
Wynne, 1972:137, fig. 1; 1973:141 [not C. sinuosa f. 
deformans Setchell and Gardner 1903:242]. 
Colpomenia bullosa of Norris 1972:6 [not C. bullosa 
(Saunders) Yarnada 1948:6]. 
Thalli e fasciculis saccorum cavorum longorum, 
qui plerumque ab 8 ad 15 cm long. (raro usque ad 
25 cm long.) atque ab 0.8 ad 1.5 cm lat. (raro usque 
ad 5.5 cm lat.) variant, constantes, e basi adhaerente 
colponemioidea, c. 1-2 diam., enascentes; sacci ad 
basim attenuati, cavi, autem, per longitudinem 
suam, a 6 ad 12 e uno basi enascentes, interdum, 
autem, paucior vel numerosior (25-30); saccus vege- 
tativus 70-88 (-110) pm crassus et compositus ex 
unico strato corticeo cellularum cuboidearum 
minorum, atque ex unico strato subcorticeo cel- 
lularum paululorum maiorum, atque e zona 
interiore 2-3 cellulis crassis, cellularum maiorum 
sine colore; sacci textura tenues delicatique, cum 
emersi omnino flaccidi; organa plurilocularia 28-30 
pm longa, uni- atque biseriata, soros densos amplos 
super saccos erectos efficientia; paraphyses rarae, 
quasi eaedem altitudine ac organa plurilocularia, 
interdum, autem, breviores altioresve; penicilli 
pilorum sine colore rari; thalli aureo-brunnei, 
fusciores cum fertiles; chloroplastum unicum in 
cellulis omnibus corticeis; sporangia unilocularia 
ignota. 
DEsCRIPTIoN.-Thalli (Figure 4) consisting of 
clusters of long, hollow sacs, typically ranging be- 
tween 8 and 15 cm in length (rarely up to 25 cm 
in length) and 0.8-1.5 cm in width (rarely up to 
5.5 cm in width), arising from an adherent col- 
pomenioid base of about 1-2 cm in diameter; sacs 
tapered toward the base yet hollow throughout 
their length, with (1-) 6-12 (-30) sacs arising from 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 4.-Holotype of Colpomenia phaeodactyla, Playa Estacion, in front of Laboratorio de 
Biologia Marina, Puerto Pefiasco, Sonora (JN-3629). 
a single base; vegetative sac 70-88 (-110) pm thick 
and composed (Figure 5a) of a single cortical layer 
of smaller cuboidal cells, a single subcortical layer 
of slightly larger cells, and an inner zone 2-3 cells 
thick of larger colorless cells; sacs are thin and 
delicate in texture, completely flaccid when 
emersed; plurilocular organs (Figure 5b) 28-38 pm 
long, uni- and biseriate, forming dense, extensive 
sori over the erect sacs; paraphyses of rare occur- 
rence, approximately same height as plurilocular 
organs but at times shorter or taller; tufts of color- 
less hairs of rare occurrence; color of thalli golden 
brown, darker when fertile; cortical cells each with 
one chloroplast; unilocular sporangia unknown. 
TYPE-LOCALITY.-Playa Estacion, Puerto Pefiasco, 
Sonora, Gulf of California, Mexico. 
HOLOTYPE.-]. Nor?% 3629 (US), 22 Jan 1973, 
Playa Estacion, Puerto Penasco, Sonora, Mexico; 
isotype (MICH). 
GULF OF CALIFORNIA DIsTRIsuTI0N.-Growing on 
rocks, middle to low littoral and occasionally upper 
littoral, in Gulf of California from Puerto Pefiasco 
south to Isla Partida. 
May. 
DIsTRIBLJrIoN.-~Orthern Gulf of California, and 
Pacific Baja California, Mexico; Pacific Coast of 
Costa Rica; southern and central Japan. 
GULF OF CALIFORNIA SEASONALITY.-December- 
SPECIMENS ExiarlNED.-Pacific Coast of Baja Calzfornia. 
Punta Thurloe, San Bartolome, intertidal, Mar 1934, Taylor 
34-611 (MICH, US): 3 Apr 1955, Dawson-13320 (US). Bahia 
Tortuga, Sulphur Is., intertidal, 12 Feb 1954, Dawson- 
12269 (US). Gulf of California. Sonora: Punta Pelicano, 
vicinity of Puerto Penasco, littoral granite rocks, 17 Mar 
1973, JN-3813 (TEX); 3 Mar 73, (US), (leg. hl. and D. Littler); 
19 Apr 1973, JN-3968 (MEXU); 9 Mar 1974, JN-4980 (US), 
(leg. JN and KB). Punta Pelicano, SE side, 17 Mar 1974, JN- 
5048 (TEX, US, MEXU), (leg. JN, KB, and D. Moore); 8 
Apr 1966, Dawson 27389 (US); Playa Arenosa, Feb 1965, 
NUMBER 35 
7 
I 
50ym 
I 
a 
I 
50rm 
1 
,1, 
b 
FIGURE 5.-Transections of Colpomenia phaeodactyla from Punta Pelicano, Puerto Penasco 
(JN4980): u, the vegetative sac of a sterile thallus; b, uniseriate and biseriate plurilocular 
organs and unicellular paraphyses of a fertile thallus. 
Dawson 27557 (ARIZ, IJS); Playa Hermosa, Puerto Pefiasco, 
littoral rock platform, 2 Feb 1973, JN-3700 (US); 19 Jan 
1973, JN-4061 (UC); Playa Estacih, Puerto Penasco, mid- 
littoral rock platform and tide pools, Feb 1968, (leg. R. 
Hoshaw), (TEX); 25 Dec 1972, Wynne 3732 (TEX, MICH, 
UC, US); 22 Jan 1973, JN-3629 (US); 3 Feb 1973, JN-3727 
(MICH); 4 Feb 1973, JN-3748 (MICH): 4 Mar 1973, JN- 
3784 (ARIZ); 18 Jan 1974, JN-4759 (UC, US), (leg. JN and 
KB); 5 Feb 1974, JN-4950 (MICH, UC, US), (leg. JN and KB); 
24 Mar 1974, JN-5100 (US), (leg. JN and KB). Playa Estacibn, 
Puerto Penasco, on NASA buoy anchored offshore, 22 Mar 
1973, JN-3834 (US). Cab0 Tepoca, Puerto de Lobos, littoral 
rocks, 16 Mar 1973, JN-6020 (AHFH). Puerto de Lobos, 
littoral rocks, 17 Feb 1973, JN-6045 (MEXU), (leg. JN and 
M. Helvey). NW of Punta Robinson, vicinity of Puerto 
Libertad, rock outcropping, 17 Feb 1973, JN-5937 (MICH). 
Bahia Kino, 2 Jan 1973, Wynne 3857 (MICH, TEX). Baja 
California: Campo El Heurfanito, 20 miles S of Puertecitos, 
2 Jan 1972, (US), (leg. M. and D. Littler). Punta Bufeo, N of 
Arroyo San Luis, 25 Feb 1975, (US), (leg. M. and D. Littler). 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
Punta Willard, Bahia San Luis Gonzaga, Jan 1940, Dawson 
308 (MICH). N side of Puerto Calamajue, 28 Mar 1973, 
JN-4620 (MICH) and JN-4707 (UC), (leg. JN and KB). 
Punta La Gringa, Bahia de Los Angeles, 2.4-6.1 m depth, 22 
May 1972, JN-3071 (UC), and on littoral rocks, 28 Apr 1974, 
JN-5431 (MICH, UC, US), (leg. JN and KB). Islas de la 
Cintura: Isla Mejia, Puerto Refugio, Isla Angel de la 
Guarda, littoral rocks, 23 Apr 1974, JN-5840 (GMS), (leg. 
JN and KB). NW of rock window on shore, Puerto 
Refugio, Isla Angel de la Guarda, littoral rocks, 21 Apr 
1974, JN-5345 (ARIZ), (leg. JN and KB). Puerto Refugio, 
Isla Angel de la Guarda, littoral rocks, 20 Apr 1974, 
JN-5422 (US), (leg. KB). NE shore of Puerto Rerugio, Isla 
Angel de la Guarda. littoral rocks, 23 Apr 1974, JN-5735 
(AHFH) and JN-5769 (MICH, US), (leg. JN and KB). Isla 
Turner, off Isla Tiburon, Jan 1940, Dawson 96 (MICH). Isla 
San Esteban, Feb 1940, Daicson 435 (MICH). SE end of Isla 
San Esteban, littoral to 3 m depth, 25 Apr 1974, JN-5537 
(MICH, UC, US) and JN-5726 (US). Costa Rica. Port Parker, 
Salinas Bay, Mar 1939, Taylor 39-76 (MICH). Japan. Nabeta, 
Shimoda, Shizuoka, Feb 1974, (MICH, TEX), (leg. M. 
Chihara). 
REMARKS.-It was earlier pointed out by Wynne 
(1972) that specimens identified as Colpomenia 
sinuosa f. deformans (sensu Setchell and Gardner, 
1924; Dawson, 1944, 1966) from the Gulf of Cali- 
fornia are quite distinct from those of the Pacific 
Coast of North America (Setchell and Gardner, 
1903, 1925). The correct name for the Pacific Coast 
C. sinuosa f. deformans Setchell and Gardner (1903) 
is Colpomenia bzillosa (Saunders) Yamada (1948) 
(Smith, 1969). Evidence supporting the recognition 
of C. phaeodacfyla as a separate species includes the 
delicate nature of the sacs and their clustered aspect 
from a common basal system. This is in contrast to 
the usually solitary condition and firmer texture of 
the balloon-like sacs of C. bullosa. 
An examination of Japanese works led us to be- 
lieve that these two distinct entities were likewise 
passing under the name C. bullosa in Japan. Oka- 
mura (1936, fig. 124) presented the genuine C. bul- 
losa, whereas the specimen presented in Chihara?s 
account (1970, pl. 19: fig. 3) resembles C. phaeodac- 
tyla. An examination of samples of the latter kindly 
provided by Dr. RI. Chihara confirmed this belief. 
Furthermore, Chihara (pers. comm.) has informed 
us that while C. bullosa occurs in northern Japan, 
including Hokkaido, Sanriku, and the Kurile 
Islands, where the Oyashio Current is predominant, 
the new species, C. phaeodactyla, occurs in southern 
and central Japan. Thus, C. phaeodactyla would 
fall in the category of algal species whose distribu- 
tions are known in both the Gulf of California and 
southern Japan (Dawson, 1960; Hommersand, 
1972). 
Colpomenia sinuosa f. deformans was recorded, 
with a query, by Taylor (1945) from Isla Isabela in 
the Galapagos Archipelago and has remained in 
the literature (Silva, 1966). However, this material 
has been examined (W. R. Taylor #I42 (MICH, 
US)) and is not C. phaeodactyla. It appears to be 
eroded specimens of C. sinuosa. Our new species, 
C. phaeodactyla, would resemble C. mollis Taylor 
(1945) from Colombia in the delicate nature of the 
wall, but the latter species is irregularly divided 
with branch endings truncate and bearing ?spine- 
like projections? (Taylor, 1945:85). 
3. Colpomenia tuberculata Saunders 
FICURFS 6, 7a,b llb 
Colpomenia tuberculata Saunders, 1898:164, pl. 32: figs. 1-3. 
Colpomenia sinuosa f. tuberculata Setchell and Gardner, 
1903:242; 1924:725; 1925:541.-Dawson, 1944:233; 1949:234, 
242; 1959:19: 1966:11,--Norris, 1972:6. 
DEScRrPTIoIL?.-Thalli (Figure 6) sessile, hollow, 
rigid, crisp, hemispherical to irregularly shaped, 
flattened expanses, covered with blunt tubercles 
1-10 mm high and 1-10 mm across; surface coria- 
ceous, deeply convoluted, and folded; 0.6-1.5 mm 
thick; cortex (Figure 7a) of 4-5 cuboidal cells, inner 
layer of 4-8 increasingly larger colorless cells; 
plurilocular organs (Figure 7b) 16-23 pm long, uni- 
and biseriate, 5-7 locules high, occurring in exten- 
sive soral regions over entire thallus; paraphyses 
present, slightly shorter than plurilocular organs; 
hairs arising in tufts from deep invaginations 
(Figure 9), especially in soral areas; color olive 
brown; unilocular sporangia unknown. 
TYPE-LOCALITY.-Near San Pedro, California. 
HoLoTwE.-unknown (see ?Remarks?). 
GULF OF CALIFORNIA DISTRIBUTION.-Growing On 
rocks and occasionally epiphytic on other algae, 
mid-littoral to upper-littoral; Puerto Penasco to 
San Jose del Cabo. 
June. 
DIsTRIBUTION.-SOUthern California, U.S.A.; Pa- 
cific Baja California, Gulf of California, and 
Sinaloa, Mexico; and Peru. 
GULF OF CALIFORNIA SEAsONALITY.-DeCember- 
NUMBER 35 
9 
FIGURE 6.-Habit of Colpomenia tuberculata, from Bahia Cholla, vicinity of Puerto Petiasco, 
Sonora (Dawson-27203). 
SPECIMENS EXAMlNED.-CalifOrnia. Los Angeles County: San 
Pedro, Feb 1932, (AHFH), (leg. Templeton). San Diego 
County: Pacific Beach, Nov 1900, (NY), (leg. Saunders). 
La Jolla, Nov 1900, (Phycotheca Boreali-Americana in AHFH 
and TEX), (leg. E. Snyder); JuI 1946, Dawson 2445-2446 
(mixed with C. sinuosa) (AHFH). Pacific Coast of Baja 
California. N of Bahia de Todos Santos, Oct 1948, Dawson 
5207 (AHFH). Bahia San Quintin, Nov 1949, Dawson 8720 
(AHFH). Punta Maria, Apr 1946, Dawson 1542 (AHFH). 
Miller?s Landing, Bahia Viscaino, Apr 1951, Dawson 1366 
(AHFH). Isla Piedras, Laguna Ojo de Liebre (Scammon?s 
Lagoon), Apr 1946, Dawson 2521 (AHFH). Punta Mallarimo, 
Bahia Viscaino, Apr 1951, Dawson 9902 (AHFH). Bahia 
Asuncion, Apr 1950, Dawson 9162 (AHFH). Punta Abreojos, 
Apr 1950, Dawson 9479 (AHFH). Bahia Magdalena, Apr 
1946, Dawson 7875 (AHFH). Punta Entrada, Isla Magdalena, 
May 1950, Dawson 9258 (MICH, AHFH). Gulf of California. 
Sonora: Bahia Cholla, Apr 1968, B. Gittins 4823 (MICH); 
Apr 1966, Dawson 27203 (ARIZ). Punta Pelicano, vicinity of 
Puerto Penasco, littoral rocks, 17 Mar 1973, JN-3812 (ARIZ); 
19 Apr 1973, JN-3967 (MEXU); 9 Mar 74, JN-4983 (US, 
MEXU). Punta Pelicano, SE, vicinity of Puerto Petiasco, 17 
Mar 1974, JN-5027 (US), (leg. JN, KB, and D. Moore). 
Playa Hermosa, Puerto Penasco, littoral platform, 2 Feb 
1973, JN-3701 (US): 3 Mar 1973, JN-4015 (MICH), (leg. 
JN and KB). Punta Penasco, Feb 1940, Dawson 351 (AHFH); 
Mar 1942, Poindexter (AHFH). Playa Estacion, Puerto 
Penasco, littoral rock platform and tide pools, May 1965, 
Dawson 27522 (ARIZ); 3 Feb 1973, JN-3729 (MICH) and 
JN-3747 (ARIZ); 4 Mar 1973, JN-3785 (UC) and JN-3800 
(US), (leg. JN and KB); 18 Mar 1973, JN-3855 (TEX); 20 
May 1973, JN-4047 (US); 18 Jan 1974, JN-4761 (ARIZ); 
9 Feb 1974, JN-4928 (US), (leg. JN and KB); 5 Feb 1974, 
JN-4951 (US), (leg. JN and KB); 6 Feb 1974, JN-4954 (TEX) 
and JN-4957 (US), (leg. JN and KB); Feb 1968, (TEX), 
(leg. R. Hoshaw); 24 Mar 74, JN-5073 (US, UC). Playa 
Estacion, Puerto Penasco, vicinity of transect line, 4 Jun 1974, 
JN-5626 (AHFH). Playa Arenosa, 26 Dec 1972, Wynne 3748 
(MICH). Cabo Tepoca, vicinity of Puerto de Lobos, littoral 
rocks, 16 and 17 Feb 1973, JN-5950 (US), JN-5965 (GMS), 
and JN-5977 (AHFH), (leg. JN and M. Helvey); Feb 1946, 
Dawson 824 (AHFH); Feb 1940, Dawson 384 (AHFH). Punta 
Lobos, Puerto de Lobos, tide channel, sand covered rocks, 
16 Feb 1973, JN-5989 (US), (leg. M. Helvey). NW side, 
Punta Robinson, Puerto Libertad, 19 Feb 73, JN-5927 
(MICH, ARIZ). Puerto Libertad, Mar 1946, Dawson 693 
10 
ShIITHSONIAN CONTRIBUTIONS TO BOTANY 
I 
i 50pm 
a b 
FIGURE 7.-Transections of Colpomenia tuberculata, from Playa Estacion, Prlerto PeAasco, Sonora 
(JN-5073): a, sterile thallus with unicellular paraphyseq; b, fertile thallus rvith plurilocular 
organs and unicellular paraphyses (note the long true hairs arising from a depression in the 
cortex). 
(AHFH). Desemboque de San Ignacio, drift, 3 Apr 1974, 
JS-5139 (MEXU, TEX, UC, US), (leg. R. S. Felger, R. E. 
Schultes, and A. T. Weil). Isla Alcatraz, Bahia Kino, Feb 
1946, Dawson 1059 (AHFH). Bahia Kino, 1 Jan 1973, Wynne 
3843 (MICH) and 2 Jan 1973, Wynne 3858 (MICH). Bahia 
Sail Carlos, near Guaymas, 31 Dec 1972, Wynne 3813 (MICH, 
TEX, UC). Baja California: Playa Blanca, N of San Felipe, 
littoral rocks, 15 May 1972, JN-2961 (UC). Punta Estrella, 
S of San Felipe, 15 May 1972, JS-3311 (US), (leg. JN and 
G. Boehlert). Punta ?(Villard, Bahia San Luis Gonzaga, lit- 
toral to 6.1 m depth, 20 May 1974, JN-5410 (MICH), (leg. 
JS and KB) and JN-5382 (US). Puerto Calamajue, N side, 
23 Mar 1973, 5x4618 (MICH, US), (leg. JN and KB). Bahia 
de 10s Angeles, Apr 1946, Dawson 1308 (AHFH). Islas de 
10s Gemelos, Bahia de 10s Angeles, 21 May 1952, JN-3012 
(UC). La Mona, Bahia de 10s Angeles, littoral rocks, 20 
May 1972, JN-2983 (ARIZ), (leg. JN and G. Boehlert). Bahia 
San Francisquito, 24 May 1972, JN-3219 (US) and JN-3272 
(US). San Marcia1 Reef, S of Bahia Agua L?erde, Mar 1937, 
Daivson (AHFH). San Jose del Cabo, Feb 1940, Dawson 636 
(AHFH). Islas de la Cintura: Isla Mejia, Puerto Refugio, 
littoral rocks, 23 Apr 1974, JN-5838 (AHFH), (leg. JS and 
KB). Puerto Refugio, Isla Angel de la Guarda, Jan 1940, 
Dawson 220 (AHFH). Puerto Refugio, littoral rocks, NE 
shore, Isla Angel de la Guarda, 23 Apr 1974, JN-5770 
(MICH). Isla Estanque, Feb 1940, Dawson 422 (AHFH). Isla 
Partida, Feb 1946, Dawson 987 (AHFH). Isla Turner, off Isla 
Tiburon, Jan 1940, Dawon 90 (AHFH). Sinaloa: Mazatlan, 
Apr 1952, Daiison 10822 (AHFH). Peru. Telegraph Point, 
13 Apr 1968, I. A. Abbott 4586 (GMS), (leg. D. P. Abbott). 
REMARKS.-/lfter careful scrutiny of ?tuberculate? 
material from both the Gulf of California and else- 
where, we have concluded that Saunders? C. tuber- 
culata (1898) should be revived as a legitimate 
species, not merely a form of C. sinuosa, to which 
rank it had been relegated by Setchell and Gardner 
(1903, 1924, 1925) and Dawson (1944, 1966). We 
have been unsuccessful in locating the holotype of 
C. tuberculata. Although some collections made by 
Saunders are now housed in the herbaria of the 
NUMBER 35 
Netv York Botanical Garden, the University of 
California, Berkeley, and Farlow Herbarium, they 
did not include any specimen of C. tuberculata 
from San Pedro, California, the type-locality. It is 
possible that the specimen was deposited in the 
Herbarium of the California Academy of Sciences, 
San Francisco, California, and could have been lost 
in the 1906 San Francisco earthquake. In any case, 
the Gulf of California material is consistent with 
Saunders? description of this species in regard to 
thickness, averaging 0.6 to 0.9 mm. The average 
length of plurilocular organs measured slightly less 
than the figures given by Saunders, but the ranges 
overlap. Although Saunders recorded the pluri- 
locular organs to be uniseriate, those observed in 
the Gulf of California specimens occasionally pos- 
sessed biseriate portions as well. 
Tuberculate forms of C. sinuosa have been vari- 
ously reported in the literature (e.g., Dawson, 1954; 
Joly, 1957; Chamberlain, 1965), but what often 
seems to be represented are much divided, gnarled 
11 
forms of C. .~inuosa rather than the distinctly tuber- 
culate and crisp texture of C. tuberculata. Further- 
more, it is likely that contorted forms of C. 
peregrina have been mistakenly recorded as C. 
tuberculata from Alaska (Dawson, 1961). 
4. Colpomenia ramosa Taylor 
FIGURE 8a,b 
Colpomenia ramosa Taylor, 1945:84, pl. 6: fig. 2.-Dawson, 
Rosenvingea intricata of Dawson, 1944:233, pl. 52; fig. 1 
1949:228; 1951:52; 1952:431: 1954a:117; 1961:394. 
[in part]. 
DEscRIprIoN.-Thalli (Figure 8) forming adherent 
clumps to 4 cm broad, to 2 cm tall, crisp, with 
several areas of attachment, irregularly subdichot- 
omously to polychotomously branched, the branches 
in congested, closely set vertically directed series 
from broader (to 8 mm) basal portions; the terminal 
divisions small, cylindrical, 1-2 mm across, 2-3 mm 
b 
FIGURE 8.-Colpomenia ramosa from Puerto Calamajue, Baja California del Norte: a, portions 
from a clumped thallus (JN-4688); b, detail showing the cylindrical to slightly compressed 
nature of the branches (JN-4697). 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
long, with rounded ends; becoming hollow, the 
wall 200-400 pm thick (to 500 pm thick in type 
material), 6-8 cell layers thick; small-celled cortex 
progressing into larger cells inward; plurilocular 
sporangia in sori, uniseriate, 10-12 locules, 35-40 
pm long. 
A collection from Academy Bay, Santa Cruz 
Island, Dawson 26219 (US), now extends the known 
distribution southward to the Galapagos Archipe- 
lago. Previously it had been reported from Isla 
Cedros to Costa Rica (Dawson, 1954a; Dawson, 
1961). 
TYPE-LmALI?rY.-Bahia Sur, Isla Cedros, Pacific 
Baja California, Mexico. 
HOLOTYPE.-W. R. Taylor 34-651 (AHFH); iso- 
types (MICH, US). 
Bahia San Luis Gonzaga to Bahia Tepoca, north- 
ern Gulf; Isla Espiritu Santo, southern Gulf. 
March. 
DIsTRrBUTroK.-Pacific coast of Baja California, 
and Gulf of California, Mexico; Pacific coast of 
Costa Rica; and the Galapagos Islands. 
GULF OF CALIFORKIA DIsTRIBUTIoN.-~ccasional, 
GULF OF CALIFORNIA SEASONALITY .-January- 
SPECIMEZIS EXA~IINED.--P~C~~~C Coast of Baja California. 
Punta Santa Rosalia, 13 Apr 1946, Darvson 1422 (AHFH), 
Dawson 1514 (AHFH); 9 Oct 1946, Dawson 2760 (AHFH, 
US); 10 Oct 1946, Darvson 2898 (AHFH). Miller?s Landing, 
Bahia Sebastian I?izcaino, 12 Apr 1946, Dawson 1365 (AHFH). 
Bahia Sur, Isla Cedros, 10 Mar 1934, W. R. Taylor 34-651 
(WRT); 5 Mar 1949, DaTvson 6550 (US); 19 Apr 1951, Daw- 
son 9836 (AHFH). Punta Malarrimo, Bahia Sebastian Viz- 
caino, 16 Apr 1951, Daivson 10016 (AHFH). Punta San 
Eugenio, 1 Nov 1951, Dawson 10351 (AHFH). Bahia San 
Bartolome (Bahia Tortugas), 11 Feb 1954, Daivson 12267 
(US). Bahia Asuncion, 28 Apr 1950, Da~vson 9163 (AHFH). 
Punta Abreojos, 30 Apr 1950, Daivson 9478 (AHFH, MICH, 
US). Punta Pequefia, Bahia San Juanito, 1 May 1950, Daw- 
son 9218 (AHFH, MICH, US). Isla Santa Magdalena, 21 
Aug 1946, Daivson 7876 (AHFH); 6 Apr 1955, Dawson 13406 
(US). Punta Hughes, Isla Santa Magdalena, 4 May 1950, 
Dawson 9333 (AHFH). Isla Santa Margarita on Bahia Mag- 
dalena side, 9 Mar 1949, Dawson 6613 (US). Bahia Almejas, 
7 mi KW of Cab0 Tosco, Isla Santa Margarita, 24 Apr 
1955, Da?ivson 13445 (US). Gulf of California. Sonora: Bahia 
Tepoca, 4 Feb 1940, Damon 385 (AHFH); 19 Feb 1946, 
Datvson 825 (AHFH): 17 Feb 1973, 5 m depth, JN-6059, 
(US, UC). Baja California: Isla Willard, Bahia San Luis 
Gonzaga, 30 Jan 1940, Dawon 306 (AHFH). Puerto Cala- 
majue, 7 m, 28 Mar 1973, JN-4688 (US), and 6 m, JS-4697, 
(MICH). Bahia San Gabriel, Isla Espiritu Santo, 14 Feb 
1940, Dawson 606 (AHFH). Pacific Coast of Costa Rica. Port 
Parker, near Salinas Bay, 24, 25 Mar 1938, FV. R. Taylor 
39-76 (AHFH, US, WRT). Galapagos Archipelago. Academy 
Bay, Santa Cruz Island, 24 Jan 1964, Darvson 26219 (US). 
REMARKS.-U~ to the present C. ramosa Taylor 
was recorded from Pacific Baja California (Taylor, 
1945; Dawson, 1961) and Rosenwingea intricata (J. 
Agardh) Boergesen was recorded from within the 
Gulf of California (Dawson, 1944). Yet both algae, 
although placed in two different genera of the 
same family, present very similar forms and empha- 
size the tenuous nature of the distinction between 
these two genera. After examining many specimens, 
we have concluded that differences do exist to 
distinguish two species and that although C. ramosa 
is more frequently encountered, there is evidence 
that R. intricata also occurs both in the Gulf of 
California and on the Pacific coast of Mexico. The 
specimen presented by Dawson (1944, pl. 52: fig. 1) 
as ?Rosenwingea inricata? conforms to Colpomenia 
mmosa. One of his collections cited (Dawson 424) 
is identifiable as R. intricata; the others cited 
(Dawson 1944:234) are herein referred to C. ramosa. 
The four collections from the Gulf and Pacific 
Mexico we examined in this study and identified as 
Rosenwingea intricata (J. Agardh) Boergesen 
(1914:26) are the following: Gulf of California, 
Baja California: Punta Bufeo, N of Arroyo San 
Luis, in drift, 25 Feb 1975, (US), (leg. hl. and D. 
Littler); Lagoon, Isla Estanque (off S end of Isla 
Angel de la Guarda) 4 Feb 1940, Dawson 424 
(AHFH); Punta Colorada, Bahia de la Paz, 20 May 
1965, Php-1711 (US), (leg. 0. Holquin and J. 
Vazquez). Guerrero: Acapulco, inner bay, 3 Feb 
1947, Dawson 3891 (AHFH, US). 
Taylor (1945) described C. ramosa as forming a 
broad, crisp clump, with multiple attachments and 
with irregular subdichotomous to polychotomous 
branches. Occasionally this species has thalli re- 
sembling those of R. intricata. Thalli of the latter 
taxon are usually erect with loose branching and 
attached by a basal disk (Taylor, 1928, 1960; 
Boergesen, 1914; Earle, 1969), but specimens 
tending toward a subrepent habit also occur. One 
distinction is that specimens of C. ramosa are 
basically prostrate expanses, coarsely branched and 
often with short peglike branches, approaching 
more closely the adherent forms expressed in the 
genus Colpomenia (especially C. tuberculata) in 
contrast to the erect forms typical of Rosenvingea. 
Another difference is that the wall of C. ramosa 
NUMBER 35 
13 
is much thicker than that of R. intricata. According 
to Taylor (1945), the wall of C. ramosa is 6-8 cell 
layers and about 400-500 pm thick, and the pluri- 
locular organs are uniseriate. Our observations 
confirm the presence of uniseriate plurilocular 
organs, up to 12 locules in length, which are in 
fairly extensive sori. 
Although the type material of C. ramosa and 
other collections from Pacific Baja have relatively 
thick walls, specimens from within the Gulf of 
California generally have thinner walls, often in 
the range of 200-300 pm. The wall of R. intricata 
is much less thick, however, being only 3-4 cell 
layers and less than 100 pm thick (Vickers and 
Shaw, 1908, as Striaria attenuata; Srinivasan, 1960, 
1969). Srinivasan (1960) illustrated plurilocular 
organs in R. intricata as biseriate, and we have 
seen similar short, ovate biseriate plurilocular 
organs in R. intricata collected in Florida and 
Hawaii (Cape Haze Marine Laboratory, Sarasota, 
S. Earle 66-303 (US); Waikiki Beach, Honolulu, 
Oahu, 0. Ravanko, 8 Jun 1967 (MICH)). Branching 
in C. ramosa is highly irregular, often with branches 
arising in groups (polychotomously) randomly over 
the surface, whereas in R. intricata branching is 
typically divaricate. The texture of C. ramosa 
ranges from crisp to stiff, while that of R. intricata 
is flaccid. Finally, the sori of C. ramosa are fairly 
extensive and visible to the naked eye, but those of 
R. intricata are discrete and small. 
Discussion 
The assemblage of Colpomenia species within 
the Gulf of California has stimulated us to re- 
examine the generic delimitations usually employed 
for the family Scytosiphonaceae. The species of 
Colpomenia discussed in this paper demonstrate 
that the criteria used in distinguishing Colpomenia 
from related genera, Scytosiphon C. Agardh (181 l), 
Zyengaria Boergesen (1939), and Rosenvingea 
Boergesen (1914), are often blurred. For example, 
the cylindrical sacs of C. phaeodactyla taken indi- 
vidually could be mistaken for enlarged forms of 
Scytosiphon lomentaria (Lyngbye) Link. One might 
question the placement of this species in Colpo- 
menia, which is regarded as having spherical or 
globose thalli in contrast to the tubular thalli of 
Scytosiphon. 
We concur with Dawson (1944), who posed this 
same question, concluding that C. phaeodactyla (as 
C. sinuosa f. dejormans of Dawson) has a very 
different development from that of Scytosiphon. 
Ontogenetically, the sacs of the former arise as 
hollow evaginations that push outward from the 
colpomenioid prostrate basal system: they are hol- 
low from their earliest formation and are hollow 
even at the juncture with the basal system which 
produces them. The tubes of Scytosiphon, on the 
other hand, develop as uniseriate filaments 
(Tatewaki, 1966; Wynne, 1969: Luning and Dring, 
1973), that undergo a series of transverse arid 
longitudinal divisions resulting in a hollow tube 
but remaining solid at the base. Thus, the initial 
form observed in these two genera is quite differ- 
ent. Wynne (1972) has shown in culturing C. 
phaeodactyla that the young germlings are spherical 
(conforming to its assignment to Colpomenia). From 
field studies we have also seen that the young stages 
of C. phaeodactyla consist of spreading colpomeni- 
oid systems from which erect tubes will be produced 
as evaginations. 
The problem of distinguishing Zyengaria Boer- 
gesen (1939) from Colpomenia has been discussed 
earlier (Wynne, 1972). The only species of this 
genus, I. stellata (Boerg.) Boergesen was described 
as Rosenvingea stellata Boergesen (1928) and later 
transferred to Colpomenia (Boergesen, 1930). 
Boergesen's primary distinguishing trait in estab- 
lishing Zyengaria (Boergesen, 1939) was the locali- 
zation of growth to projections covering the thallus 
rather than the diffuse growth present in C. sinziosa. 
Although the surface of C. tuberculata is not cov- 
ered with the densely clustered projections that 
occur in I. stellata, there would seem to be a 
difference in degree rather than in kind, and we 
would suggest that Zyengaria be merged with Col- 
pomenia and that C. stellata (Boerg.) Boergesen 
(1930) be reinstated. Chamberlain (1965) also 
referred to the dubious nature of Zyengaria, but 
states that another apparent difference from 
Colpomenia is the absence of paraphyses among the 
plurilocular organs in Zyengaria. The latter char- 
acter is of questionable value, and in other genera 
of the family, such as Scytosiphon (cf. Wynne, 
1969), there is variation from one species to the 
next. 
A third difficult distinction is that between the 
genera Colpomenia and Rosenvingea. The primary 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
characteristic for separating Rosenvingea from the 
remaining Scytosiphonaceae is its branching nature. 
Yet branching Colpomenias have been described 
as C. ramosa, C. mollis Taylor (1945), and C. 
nainativensis Durairatnam (1962). Difficulties arise 
in distinguishing taxa such as C. ramosa from R. 
intricata (J. Agardh) Boergesen and in distin- 
guishing C. nainativensis from R. nhatrangensis 
Dawson (195413). We will focus our attention on 
the former pair of species. Rosenuingea contains 
species with typically erect, tubular, branching 
thalli. Specimens of R. intl-icata are usually erect 
with loose branching and attached by a basal disk 
(Taylor, 1928, 1960; Boergesen, 1914; Earle, 1969, 
figs. 111, 112), but sometimes specimens tending 
toward a subrepent condition occur. On the other 
hand, specimens of C. ramosa are basically prostrate 
expanses, coarsely branched, approaching more 
closely the adherent forms expressed in the genus 
Colpomenia (e.g., C. tuberculata) than the erect 
forms typical of Rosenvingea. Other differences 
might be cited to distinguish C. ramosa from R. 
intricata. The wall of C. l-amosa is 6-8 cell layers 
about 400-500 pm thick, and, according to Taylor 
(1945), the plurilocular organs are uniseriate. The 
wall of R. intricata is much less thick, being only 
FIGURE 9.-Intertidal Tegion of Playa Estacion (rn fiont of Laboratorio de Biologia Marina), 
Puerto Penasco: a, high to middle tidal levels in the spring dominated by Colpomenia tuber- 
culata and Lauremia paniculata, with a ferc plants of C. phaeodactyla also present (April 1974) 
(note the transect line running between permanent markers); b, loirermost intertidal region, 
dominated at this season by Colpomenia sinuosa and Sargassurn herporhizum (February 1974). 
NUMBER 35 
15 
3-4 cell layers and less than 100 pm thick (Vickers 
and Shaw, 1908, pl. xxiv, as Striaria attenuata; 
Srinivasan, 1960). Srinivasan (1960) illustrated pluri- 
locular organs to be biseriate. So in conclusion, we 
are of the opinion that forms of these two genera 
closely approach each other in their morphologies, 
but Colpomenia includes forms which range from 
spherical to saccate to prostrate expanses, while 
Rosenuingea includes freely branched, primarily 
erect forms. 
Ecological Observations 
Studies on the intertidal biotic community at 
Playa Estacion, Puerto Penasco, Sonora, (Figure 
9a,b) have shown marked seasonal changes in algal 
species and their population sizes. Data from bi- 
monthly censuses, February to December 1974 (Nor- 
ris, unpublished), were obtained from a 100 meter 
line transect placed perpendicular to the shoreline. 
Frequency, number of individuals, and percent 
cover were recorded on each census for seasonal 
comparisons. Seasonal differences in the population 
sizes of the three Colpomenia species present (C. 
phaeodactyla, C. tuberculata, and C. sinuosa) are 
summarized in Figure 10. 
SEAsoNALiTY.-Plants of each species begin to 
appear in November. By December populations of 
C. phaeodactyla (Figure 1 lc) and C. sinuosa (Figure 
1 la) are larger than those of C. tuberculata (Figure 
llb). In February the populations of the three 
species are similar in numbers of individuals and 
percent coverage. Combining the percent coverage 
figures for all three species shows that during 
February Colpomenia species covered 19% of the 
entire intertidal transect. With such a high cover- 
age, it intuitively seems that biomass would also be 
high (not measured, since it would have disturbed 
the sample transect), and the species of Colpomenia 
would play an important ecological role from 
February to April ( 12.5y0 coverage). Populations of 
C. phaeodactyla and C. sinuosa dropped by April: 
in June they appeared absent. In contrast, C. tuber- 
culata reached its peak in April, and during that 
month it was the most abundant organism present. 
No species of Colpomenia were observed on the 
study line in August or October. 
DISTRIBUTlON.-CO~pOmeniU phaeodactyla is re- 
OCT DEC FEE APR JUN AUG 
- n n I, 280+ 
OCT DEC FEE APR JUN AUG 
C. phaeodactyla C. tuberculata C. sinuasa 
FIGURE 10.-Seasonal comparison of the numbers of indi- 
viduals and coverage of the species of Colpornenia on the 
intertidal transect at Playa Estacibn, Puerto PeAasco during 
1974. 
stricted mostly to the middle and low intertidal 
range, growing on top of the platform (caliche 
capped coarse sandstone with shell materials) and 
occurring only occasionally in tide pools. Col- 
pomenia tuberculata ranges throughout the inter- 
tidal zone from high to low tidal levels, growing 
on the platform, or epiphytic on other algae, par- 
ticularly Laurencia paniculata. Colpomenia sinuosa 
is almost entirely found in the low tidal range 
(sometimes at higher levels, but then usually in 
tide pools) and extended into the shallow subtidal. 
While occasionally attached to rocks, it is mostly 
epiphytic on other algae including Sargassum. 
PRoDucTIv1TY.-Measurements for two species of 
Colpomenia growing sympatrically, reveal differ- 
ent rates of carbon fixation. Colpomenia phaeodac- 
tyla has a mean net rate (& one S.D.) of 2.58 k 
16 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 11 .-Habitat photographs of Colpornenia species from Playa Estacion, Puerto Penasco 
(February 1974): a, C. sinuosa; b, C. tuberculata; c. C. phaeodactyla. 
0.08 mg C fixed/gram dry weight/hour, and in for each species at Punta Bufeo (just north of 
contrast, C. tuberciilnta sliows a mean rate of 1.10 Punta Willard, Bahia San Luis Gonzaga), Baja 
f 0.04 mg C fixed/gram dry weight/hour. These California del Norte, 25 Feb 1975. Light varied 
data were obtained by Dr. hl. Littler (pers. comm.) between 1200-1700 pE/m2/sec (PAR) and 52,500- 
during a high tide period using three light and two 66,000 lux during the 10:30 am to 2:50 pm period 
dark bottle replicates (each containing 3-5 thalli) of in situ incubation. 
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