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 referred to C. ramosa Taylor. Although recorded from the Pacific coast of Baja 

California, C. ramosa has not yet been reported within the Gulf. Interrelationships of the species complex of Colpomenia with related genera such as Iyengaria Boergesen, Rosenvingea Boergesen, and Scytosiphon C. Agardh are discussed. 

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 , . , . ,. .. 

. . . . . . . . . . . . . . . . . . . . . . . 

... ....... 

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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 California, 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. According to Dawson (1944) the latter also included f. 

expansissirnu Setchell and Gardner. Under its various guises Colpomenia collectively forms a dominant element of the intertidal winter and spring 

flora of the Gulf of California. Field and microscopic studies, however, have led the present authors to conclude that the Colpomenia complex in 

the Gulf includes four distinctive species, separated 

by morphological, anatomical, and habitat differences. 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 NaMichael 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, California (GMS). Collection numbers of J. N. Norris 

carry the prefix JN- and refer to his field notebooks. Specimens were collected by him unless 

otherwise noted; those collected by Katina E. 

Bucher are noted as KB. 

ACKNOWLEDGMENTS.-we wish to thank Dr. Robert W. Hoshaw for the use of facilities while J. 

Norris was station director/resident marine biologist at Laboratorio de Biologia Marina of the University of Arizona, and for the loan of Dawson 

material (ARIZ). J. Norris is also grateful to Carl 

N. Hodges, Director, Environmental Research Laboratory, for the use of Laboratorio de Biologia 

Marina in Puerto Penasco, Sonora, Mexico. Support 

for field work in the Gulf of California was provided by a National Science Foundation grant NSFBMS-73-07000 (formerly GB-38623) awarded to J. 

Norris and hl. Neushul. It is a pleasure to acknowledge 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. Nicholson 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 Herbarium. Dr. Mark hl. Littler has generously shared 

unpublished productivity data for use in this paper. Dr. Hannah Croasdale kindly provided the 

Latin diagnosis. h1s. Alice Tangerini drew C. ~amosa 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 assistance 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 vesicular, becoming irregularly convoluted and expanded 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; paraphyses longer than plurilocular organs, to 55 pm 

in length; color golden brown; unilocular sporangia 

unknown. 

TYPE-LocALITY.-cadiz, Spain. 

HoLoTYPE.-Destroyed at Berlin-Dahlem (according 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.-NOvemberDISTRIBUTION.-COmmOn in tropical and subtropSPECIMENS 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, JN4929 (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 previously 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, California, 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. Discrete punctate sori of plurilocular organs are covered with a cuticle. On the other hand, C. peregrina is characterized by relatively thinner, smooth 

thalli, greenish in color upon pressing, with broad, 

confluent fertile zones,' lacking a cuticle, with plurilocular organs often composed of two rows of 

locules, and paraphyses shorter than the plurilocular 

'Nonconfluent sori were reported in C. peregrina in h'orway (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 populations of these trvo species. 
NUMBER 35 5 

organs. Sauvageau (1927) reported that the thalli 

of C. sinuosa are 420-560 pm thick and the plurilocular organs are 40 (-50) pm high, while thalli 

of C. peregrina are 140-280 pm thick and plurilocular 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 uniseriate and biseriate (or intermediate). Material illustrated by Okamura in Japan (1936, fig. 123) and 

Chamberlain for Gough Island in the South Atlantic (1965) showed a similar situation in C. sinuosa. 

Colpomenia sinuosa f. lacunosa was described by 

Taylor (1947) from Paita, Peru. Subsequent treatment 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” described 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 condition . . .” (1944:233). Unfortunately our opensheet form is not fertile, and lacking this, it seems 

judicious to merely call attention to this new material. Thus far only known from the Gulf of California, 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, JN4607 (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 vegetativus 70-88 (-110) pm crassus et compositus ex 

unico strato corticeo cellularum cuboidearum 

minorum, atque ex unico strato subcorticeo cellularum 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 between 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 colpomenioid 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 occurrence, approximately same height as plurilocular 

organs but at times shorter or taller; tufts of colorless 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.-DecemberSPECIMENS 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, Dawson12269 (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, JN5048 (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, midlittoral 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, JN3784 (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 California 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 believe that these two distinct entities were likewise 

passing under the name C. bullosa in Japan. Okamura (1936, fig. 124) presented the genuine C. bullosa, whereas the specimen presented in Chihara’s 

account (1970, pl. 19: fig. 3) resembles C. phaeodactyla. 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 distributions 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 “spinelike 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 coriaceous, 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, uniand biseriate, 5-7 locules high, occurring in extensive 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.; Pacific 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, littoral 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 elsewhere, we have concluded that Saunders’ C. tuberculata (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 plurilocular organs to be uniseriate, those observed in 

the Gulf of California specimens occasionally possessed biseriate portions as well. 

Tuberculate forms of C. sinuosa have been variously 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 tuberculate and crisp texture of C. tuberculata. Furthermore, 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 subdichotomously 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 Archipelago. 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); isotypes (MICH, US). 

Bahia San Luis Gonzaga to Bahia Tepoca, northern 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 .-JanuarySPECIMEZIS 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, Dawson 9836 (AHFH). Punta Malarrimo, Bahia Sebastian Vizcaino, 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, Dawson 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 Magdalena 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 Calamajue, 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 emphasize 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 resembling 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 plurilocular 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 reexamine 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 individually could be mistaken for enlarged forms of 

Scytosiphon lomentaria (Lyngbye) Link. One might 

question the placement of this species in Colpomenia, 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 hollow 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 different. 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 colpomenioid systems from which erect tubes will be produced 

as evaginations. 

The problem of distinguishing Zyengaria Boergesen (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 establishing Zyengaria (Boergesen, 1939) was the localization of growth to projections covering the thallus 

rather than the diffuse growth present in C. sinziosa. 

Although the surface of C. tuberculata is not covered 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 Colpomenia 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 character 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 distinguishing 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 tuberculata 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 plurilocular 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 bimonthly censuses, February to December 1974 (Norris, 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 coverage, 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. tuberculata 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 reOCT 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 individuals 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. Colpomenia tuberculata ranges throughout the intertidal zone from high to low tidal levels, growing 

on the platform, or epiphytic on other algae, particularly 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 different rates of carbon fixation. Colpomenia phaeodactyla 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. 
Literature Cited 

Agardh, C. A. 

1811. Part 11. Pages 17-26 in Dispositio algarum Sueciae. 

Lund. 

Algas Fucoideas Complectens. Volume 1 in Species, 

genera et ordines Algarum, seu descriptiones succinctac specierum, generum et ordinum, guibus 

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