SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES • NUMBER 1 

The Red Alga Polysiphonia 

(Rhodomelaceae) in the 

Northern Gulf of California 

George J. Hollenberg 

and James N. Norris 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1977 
ABSTRAC T 

Hollenberg, George J., and James N. Norris. The Red Alga Polysiphonia 

(Rhodomelaceae) in the Northern Gulf of California. Smithsonian Contributions 

to the Marine Sciences, number 1,21 pages, 10 figures, 1977.—Taxonomic studies 

of Polysiphonia show 14 species to be present in the northern Gulf of California, 

Mexico. One of these, P. sphaerocarpa var. cheloniae is described herein as a 

new variety, and is found growing exclusively on the green sea turtle, Chelonia 

mydas L. This and three other species are recorded for the first time in the Gulf 

of California. The distribution of six other species is extended. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: Seascape along the 

Atlantic coast. 

Library of Congress Cataloging in Publication Data 

Hollenberg, George J. 

The red alga Polysiphonia (Rhodomelaceae) in the northern Gulf of California. 

(Smithsonian contributions to the marine sciences ; no. 1) 

Bibliography: p. 

Supt. of Docs, no.: SI 1.29:37 

1. Polysiphonia. 2. Algae—Mexico—California, Gulf of. I. Norris, James N., joint author. 

II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to the marine 

sciences ; no. 1. 

QK569.R37H64 589'.41 76-608299 
Contents 

Page 

Introduction 1 

Key to the Species of Polysiphonia 2 

Polysiphonia confusa Hollenberg 2 

Polysiphonia flaccidissima Hollenberg 4 

Polysiphonia johnstonii Setchell Sc Gardner var. johnstonii 4 

Polysiphonia johnstonii var. concinna (Hollenberg) Hollenberg 7 

Polysiphonia masonii Setchell & Gardner 8 

Polysiphonia mollis J. D. Hooker & Harvey in Harvey 9 

Polysiphonia pacifica var. delicatula Hollenberg 10 

Polysiphonia paniculata Montagne 11 

Polysiphonia savatieri Hariot 12 

Polysiphonia scopulorum var. villum (J. G. Agardh) Hollenberg 14 

Polysiphonia simplex Hollenberg 14 

Polysiphonia sonorensis Hollenberg 16 

Polysiphonia sphaerocarpa var. cheloniae, new variety 16 

Polysiphonia species 17 

Literature Cited 19 

Index 21 

in 
The Red Alga Polysiphonia 

(Rhodomelaceae) in the 

Northern Gulf of California 

George J. Hollenberg 

and James N. Norris 

Introduction 

The earliest records of Polysiphonia Greville 

(1824:308) in the Gulf of California were provided 

by Setchell and Gardner (1924), who described three 

new species. Later, Dawson (1944), in his investigations on marine algae of the Gulf of California, 

recorded seven species, including a new one. A detailed study of the taxa of Polysiphonia from the 

Gulf of California and Pacific coast of Baja California was first presented by Hollenberg (1961). 

These three papers constitute the basis for our 

knowledge of Polysiphonia in the Gulf of California. 

Members of the genus Polysiphonia (Rhodomelaceae, Ceramiales) are distinguished by several important taxonomic characters: the number of pericentral cells; presence or absence of cortication; the 

manner of origin of rhizoids; the manner of origin 

of branches; the nature and position of trichoblasts; 

and the nature of spermatangial branchlets. These 

features have been pointed out in previous investigations by Hollenberg (1942a; 1942b; 1944; 1961; 

1968a). 

Herein we present our studies of recent intertidal 

and subtidal collections of Polysiphonia from the 

George J. Hollenberg, Department of Biology, University of 

Redlands, Redlands, California 92373. James N. Norris, Department of Botany, National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 20560. 

northern Gulf of California. These collections were 

obtained by the junior author during field expeditions in 1972-1974 on the Gulf coasts of Baja California and Sonora. In addition, the following collections were utilized during the course of this study: 

those of the senior author from Puerto Penasco, 

Sonora, in 1967; some additional material of the 

late Dr. E. Yale Dawson; and those of the junior 

author and Katina Bucher from the cruise of the 

R/V Dolphin to Las Islas de la Cintura (Midriff 

Islands) (Norris and Bucher, 1976). 

Collectors are designated as follows: JN for J. N. 

Norris; H for G. J. Hollenberg; D for E. Y. Dawson; 

and KB for Katina E. Bucher. The numbers cited 

correspond to the collector's field notebooks. Specimens are being deposited in the following herbaria: 

first set, U.S. National Herbarium, Smithsonian Institution (US); duplicate material to University of 

California, Berkeley (UC); Phycology Herbarium, 

University of Arizona (ARIZ); Universidad National Autonoma de Mexico (MEXU). Herbaria 

housing the holotypes are abbreviated in accordance 

with Holmgren and Keuken (1974). Geographical 

locales under "Specimens Studied" are listed from 

north to south. 

ACKNOWLEDGMENTS.—The Latin diagnosis was 

kindly provided by Dr. Hannah T. Croasdale 

(Dartmouth College). Field studies for collections 

were made possible through the support of the Na-
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

tional Science Foundation (research grant GB-28623) 

to Dr. M. Neushul and J. Norris (Marine Science 

Institute, University of California, Santa Barbara). 

The junior author also wishes to thank again the 

University of Arizona for use of their Laboratorio 

de Biologia Marina, Puerto Penasco, Sonora, while 

he served as resident Marine Biologist/Station Director of the facility. Collections by Norris and 

Bucher from Las Islas de la Cintura were made 

possible only through the generosity of Dr. William 

Fenical (Scripps Institution of Oceanography), Chief 

Scientist on the R/V Dolphin cruise of April 1974. 

Collections from the sea turtles were obtained by 

Dr. Richard S. Felger (Arizona-Sonora Desert Museum, Tucson, Arizona) and Edward W. Moser 

(Summer Institute of Linguistics, University of 

North Dakota). Further collections were given to 

Norris by Dr. Felger, Professor Richard Evans 

Schultes (Director, Botanical Museum of Harvard 

University), and Dr. Andrew T. Weil (Botanical 

Museum of Harvard University). Finally, we wish 

to thank Drs. Isabella A. Abbott, Robert W. 

Hoshaw, and Katina Bucher for reading the manuscript and offering suggestions for its improvement. 

10. 

11. 

12. 

13. 

Key to the Species of Polysiphonia 

THE SPECIES ARE ARRANGED ALPHABETICALLY IN THE TEXT 

Pericentral cells 4 (Section Oligosiphonia) 2 

Pericentral cells more than 4 (Section Polysiphonia) 11 

Trichoblasts or scar-cells in general, regularly 1 per segment 3 

Trichoblasts or scar-cells lacking or not regularly 1 per segment 9 

Branching dichotomous throughout P. masonii 

Branching lateral, not strictly dichotomous 4 

Growing on sea turtles P. sphaerocarpa var. cheloniae, new variety 

Not growing on sea turtles 5 

Branches arising from a primary branch of a trichoblast (axillary to a trichoblast) 

P. flaccidissima 

Branches arising from an entire trichoblast primordium 6 

Epiphytic, mostly less than 2 cm high P. savatieri 

Usually not epiphytic 7 

Segments in main branches mostly shorter than their diameter or sometimes as long 

P. simplex 

Segments in main branches mostly 1.5 times as long as their diameter or longer 8 

Trichoblasts usually well developed; spermatangial stichidia arising as a primary branch of 

a trichoblast P. mollis 

Trichoblasts very rudimentary; spermatangial stichidia arising from an entire trichoblast 

primordium Polysiphonia species 

Thalli delicate, mostly intertidal, commonly forming mats or furry patches 

P. scopulorum var. villum 

Thalli not as delicate, mostly subtidal, not forming mats or patches 10 

Thalli mostly less than 2 cm high; trichoblasts infrequent to rare ....P. pacifica var. delicatula 

Thalli up to 4 cm high, of a terracotta color; trichoblasts numerous, arising at irregular 

intervals P. sonorensis 

Pericentral cells 5-6 12 

Pericentral cells 8 or more 13 

Thalli occasionally epiphytic, mostly 5 cm or more high P. johnstonii var. johnstonii 

Thalli usually epiphytic, mostly less than 5 cm high P. johnstonii var. concinna 

Thalli mostly less than 2 cm high P. confusa 

Thalli mostly much larger, to 20 cm high P. paniculata 

Polysiphonia confusa Hollenberg 

FIGURE 1 

Polysiphonia confusa Hollenberg, 1961:350, pi. 1; fig. 5.— 

Dawson, Acelto, and Foldvik, 1964:87, pi. 33D.—Abbott and 

Hollenberg, 1976:696, fig. 643. 

Polysiphonia inconspicua sensu Hollenberg, 1944:479 [not 

Polysiphonia inconspicua Reinsch, 1888:146]. 

DESCRIPTION.—Thalli 0.8-1.5(-3.0) cm high, with 

erect axes mostly assurgent from prostrate branches 

100-175 ^m in diameter, composed of segments 
NUMBER 1 

' 

100/urn- A lOOjum B 

: \ U 

FIGURE 1.—Polysiphonia confusa: A, upper portion of thallus (JN-5577); B, cystocarpic plant 

(JN-3835). 

about as long as the diameter or somewhat longer, 

attached by numerous unicellular rhizoids, arising 

1 to several per segment as cells cut off from the 

proximal ends of the pericentral cells, and frequently having digitate apices; erect branches 

60-150 ju,m in diameter, the segments 1.0-2.5 times 

as long; lateral branches arising exogenously in connection with trichoblasts at irregular intervals; pericentral cells 8-10, without cortication; trichoblasts 

to 1 mm long, with 1 or 2 dichotomies, arising 1 per 

segment in spiral sequence with 14 divergence, soon 

deciduous, leaving persistent scar-cells. 

Tetrasporangia, 60-80 /xm in diameter, in long 

spiral series in slightly swollen segments of ultimate 

and subultimate branches; cystocarps globular, to 

225 ju,m in diameter; spermatangial plants unknown. 

TYPE-LOCALITY.—Mid-intertidal rocks, Corona 

del Mar, Orange County, California. 

HOLOTYPE.—H-3285, 24 October 1942, in herbarium of G. J. Hollenberg (now at US). 

GUL F OF CALIFORNIA DISTRIBUTION.—Mid -

intertidal regions; epizoic on sand dollars, epiphytic 

on Codium and growing on hard substrate of buoy; 

Puerto Penasco to Desemboque de San Ignacio. 

PACIFIC COAST DISTRIBUTION.—U.S.A.: southern 

California. Mexico: northern Baja California. 

SPECIMENS STUDIED.—SONORA: JN-5577, sterile, growing on a 

sand dollar, Encope grandis, Bahia Cholla near Punta Pelicano, legit J. N. and K. B., 25 March 1974; JN-3835, cystocarpic, on NASA buoy, anchored off Laboratorio de Biologia 

Marina, Playa Estacion, Puerto Penasco, legit J. N., 22 

March 1973; JN-5158, growing on Codium sp., Desemboque 

de San Ignacio, legit R. E. Schultes, R. S. Felger, and A. T. 

Weil, 3 April 1974. 

REMARKS.—Previously known on the Pacific coast 

from southern California and northern Baja Cali-
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

fornia (Hollenberg, 1961:351), this taxon is now 

recorded in the Gulf of California. It is apparently 

rare in the northern Gulf; only three collections 

have come to our attention. Although none of these 

specimens is from the habitat described as characteristic of P. confusa Hollenberg, size and structural 

features leave little room for doubt as to the identity 

as given. 

Polysiphonia flaccidissima Hollenberg 

FIGURE 2E 

Polysiphonia flaccidissima Hollenberg, 1942b:783, figs. 8, 19; 

1961:351, pi. 2: fig. 2; 1968a:63, figs. 2A, 11.—Dawson, 

Acelto, and Foldvik, 1964:88, pi. 78B.—Abbott and Hollenberg, 1976:688, fig. 634. 

DESCRIPTION.—Epiphytic or saxicolous algae 10-

25 mm high, erect from creeping basal branches 

attached by numerous unicellular rhizoids, which 

are cut off as separate cells from the pericentral 

cells; main erect axes 70-80 ^,m in diameter, with 

segments mostly 1-2 times as long as the diameter; 

branches of several to many orders, arising exogenously from a primary branch of a trichoblast, 

mostly at intervals of 6 segments or less, narrowed 

gradually below and above, at first usually strict 

and curving toward the parent branch; pericentral 

cells 4, totally ecorticate, with thin hyaline walls; 

trichoblasts simple or mostly with a single dichotomy, tapering to slender apices, arising 1 per segment in spiral sequence, with 14 divergence, soon 

deciduous, leaving persistent scar-cells. 

Tetrasporangia 50-70 ^m in diameter, in spiral 

series in smaller branches, somewhat distending the 

segments; cystocarps globular; spermatangial stichidia arising as a primary branch of a trichoblast, 

with 1-2 celled sterile apices. 

TYPE-LOCALITY.—On a coralline alga, rocky 

point, Laguna Beach, Orange County, California. 

HOLOTYPE.—H-2269, 1 January 1938, in herbarium of G. J. Hollenberg (now at US). 

GULF OF CALIFORNIA DISTRIBUTION.—Inlertidal 

to subtidal, on rocks or occasionally epiphytic; 

Guaymas to Isla Espiritu Santo. 

PACIFIC COAST DISTRIBUTION.—U.S.A.: southern 

California; Hawaiian Islands. Mexico: northern 

Baja California; Revilla Gigedo Archipelago; Guerrero. Costa Rica. Panama. Peru. 

SPECIMENS STUDIED.—SONORA: JN-4267, tetrasporangial, and 

JN-4270, on Padina, from 1.5-6.1 m depth, Nueva Guaymas, 

legit Dale Evensen, 13 Oct 1973. 

REMARKS.—In the northern Gulf this species is 

apparently rare, restricted in distribution to Guaymas and vicinity, Sonora. Across the Gulf it has 

been recorded from Santa Rosalia, and in the 

southern Gulf from Puerto Escondido and Isla 

Espiritu Santo (Hollenberg, 1961:352). 

Polysiphonia johnstonii Setchell & Gardner 

var. johnstonii 

FIGURES 2B, 3 

Polysiphonia johnstonii Setchell & Gardner, 1924:767.— 

Hollenberg, 1944:475; 1961:357, pi. 2: figs. 4, 5.—Dawson, 

1944:329; 1949:237; 1951:53,57; 1954b:160; 1959:32; 1966:29. 

—Dawson, Neushul, and Wildman, 1960a:72; 1960b:26.— 

Norris, 1972:16.—Abbott and Hollenberg, 1976:699, fig. 649. 

Polysiphonia marchantae Setchell & Gardner, 1924:768, pi. 

49a. 

Polysiphonia sinicola Setchell & Gardner, 1924:769. 

Polysiphonia acuminata sensu Dawson, 1950:68; 1954b: 159 

[non Polysiphonia acuminata Gardner, 1927:100]. 

Polysiphonia richardsoni sensu Segi, 1951:253.—Dawson, 

1954b: 160 [in part, with reference to Gulf of California 

specimens cited by Setchell and Gardner, 1924:768 as 

P. marchantae only; non Polysiphonia richardsoni W. J. 

Hooker ex Harvey in W. J. Hooker, 1833:333]. 

DESCRIPTION.—Thalli relatively rigid, 5-12 cm 

high, dull red in color, nearly black on drying, 

attached by numerous unicellular rhizoids, which 

are cut off as separate cells from the pericentral 

cells of a limited prostrate base; primary erect 

branches several, relatively distinct, to 1 mm in 

diameter, repeatedly branched, the branches spirally 

arranged, arising at a narrow angle, which widens 

as the branch matures; ultimate branches usually 

relatively close and dense; segments in main 

branches 1.5 times as long as the diameter or 

shorter, with 5-6 pericentral cells, totally ecorticate; 

trichoblasts once or twice forked, arising 1 per segment in spiral sequence, with y5 or yG divergence, 

soon deciduous, leaving persistent scar-cells; branches 

replacing trichoblasts in origin at irregular intervals. 

Tetrasporangia l(-2) per segment, 70-90 ^m in 

diameter; cytocarps globular, 450-500 ^m in diam-
NUMBER 1 

50jjm 

I50jjm 

lOOjjm 

200um 

400pm 

FIGURE 2.—A, Polysiphonia johnstonii var. concinna, branch tip of a tetrasporic plant (after 

Hollenberg, 1944); B, P. johnstonii var. johnstonii, apical portion of an axis showing trichoblasts 

and characteristically incurved tips (after Hollenberg, 1961); c, P. sonorensis, branch tip of the 

holotype (after Hollenberg, 1961); D, P. masonii, upper portion of a tetrasporic plant (after 

Hollenberg, 1961); E, P. flaccidissima, branch tip (after Hollenberg, 1942b). 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

2 CM 

2CM 

FIGURE 3.—Polysiphonia johnstonii var. johnstonii: A, B, habits of two specimens showing the 

divergent manner of branching (JN-3201). 
NUMBER 1 

eter, nearly sessile; spermatangial stichidia arising 

as a primary branch of a trichoblast, 100-140 by 

30-40 /i,m in length and diameter, respectively. 

TYPE-LOCALITY.—On Gracilaria sp., Isla San 

Esteban, Gulf of California, Mexico. 

HOLOTYPE.—Ivan M. Johnston 110, April 1921 

(CAS 1361, now at UC). 

GULF OF CALIFORNIA DISTRIBUTION.—Intertidal 

and subtidal, on rocks or epiphytic; Puerto Penasco 

to La Paz. 

PACIFIC COAST DISTRIBUTION.—Santa Catalina 

Island, California; northern Baja California; Sinaloa. 

SPECIMENS STUDIED.—SONORA: JN-3102, rocky intertidal, 

Punta Pelicano, 30 Jun 1972; JN-3260, on Sargassum sp. on 

rock platform with sand areas, Playa Hermosa, Puerto Penasco, 

legit J. N. and K. B., 26 Jul 1972; JN-4040, spermatangial, 

intertidal rock platform, Playa Hermosa, Puerto Penasco, legit 

J. N. and K. B., 3 May 1973; JN-5150, beach drift after strong 

winds, Desemboque de San Ignacio, legit R. E. Schultes, R. S. 

Felger, and A. T. Weil, 3 Apr 1974. BAJA CALIFORNIA DEL 

NORTE: JN-5462, 1-7 m depth, Punta la Gringa, Bahfa de 

Los Angeles, legit J. N. and K. B., 28 Apr 1974; JN-3029, 

5 m depth, Punta la Gringa, Bahia de los Angeles, 22 May 

1972; JN-3201, and JN-3209 (a slender form), estuary, Bahia 

de las Animas, legit J. N. and G. Boehlert, 23 May 1972. LAS 

ISLAS DE LA CINTURA: JN-5280, tetrasporangial and cystocarpic, 

10 m depth, W side of Roca Blanca, Puerto Refugio, Isla 

Angel de la Guarda, legit J. N., J. Paul, and K. Robertson, 

21 Apr 1974; JN-5314, intertidal to 10 m depth, Puerto 

Refugio, Isla Angel de la Guarda, legit J. N. and K. B., 

21 Apr 1974; JN-5369, 5-12 m depth off small islet, Puerto 

Refugio, Isla Angel de la Guarda, 21 Apr 1974; JN-5762, 

intertidal, rocky shore NE side of Puerto Refugio, Isla Angel 

de la Guarda, 23 Apr 1974; JN-5483, 3-10 m depth, S end of 

Isla Estanque, legit J. N. and K. B., 27 Apr 1974; JN-5508 and 

JN-5616A, 3-12 m depth, S end of Isla Estanque, legit J. N. 

and K. B., 27 Apr 1974; JN-5561, tetrasporangial, 3-8 m depth, 

rocky shore SE end of Isla San Esteban, 25 Apr 1974; JN-5528, 

cystocarpic, 3-5 m depth, off SE end of Isla San Esteban, 

26 Apr 1974; JN-5707, epiphytic on Laurencia sinicola, and 

JN-5709, 3-8 m depth, SE end of Isla San Esteban, 25 Apr 

1974; JN-5518, 1-5 m depth, SE end of Isla San Esteban, legit 

K. B., 26 Apr 1974. 

REMARKS.—One of the most frequently encountered species of Polysiphonia in the northern Gulf, 

its distribution is now extended northward to 

Puerto Penasco from Puerto Libertad (Hollenberg, 

1961:358). In the northern Gulf, specimens are 

usually epiphytic and commonly sterile. 

Polysiphonia johnstonii var. concinna 

(Hollenberg) Hollenberg 

FIGURES 2A, 4D 

Polysiphonia concinna Hollenberg, 1944:474, fig. 10.—Dawson, 

1951:53, 56; 1954b: 159; 1959:32. 

Polysiphonia johnstonii var. concinna (Hollenberg) Hollenberg, 1961:358, pi. 3: fig. 4.—Abbott and Hollenberg, 1976: 

699, fig. 650. 

DESCRIPTION.—This variety is similar to P. johntonii var. johnstonii in detailed features, but 

smaller and shorter in dimensions throughout. 

TYPE-LOCALITY.—Epiphytic on other algae, near 

Scripps Institution of Oceanography, La Jolla, San 

Diego County, California. 

HOLOTYPE.—H-2015, December 1936, in herbarium of G. J. Hollenberg (now at US). 

GUL F OF CALIFORNIA DISTRIBUTION.—Mid -

intertidal to subtidal, epiphytic on various algae, 

occasionally on rocks; Puerto Penasco to Bahia 

Agua Verde. 

PACIFIC COAST DISTRIBUTION.—Southern California; northern Baja California; Sinaloa. 

SPECIMENS STUDIED.—SONORA: H 67-64, spermatangial, cystocarpic, on Sargassum sp. in drift, Playa Tucson, Punta Pelicano, near Puerto Penasco, 8 Apr 1967; H 67-70, on Sargassum 

sp. in drift, Playa Estaci6n, Puerto Penasco, 8 Apr 1967. BAJA 

CALIFORNIA DEL NORTE: JN-4667, and JN-4668 spermatangial, 

and JN-4669 cystocarpic, the latter two on Eucheuma uncinatum, and JN-4670, cystocarpic, on Codium sp., 3-5 m depth 

N side of Puerto Calamajue, legit J. N. and K. B., 28 Mar 

1973; JN-3011, cystocarpic, Islas de los Gemelos, Bahia de 

los Angeles, 21 May 1972; JN-4752, on Padina, Isla de los 

Gemelos, Bahia de los Angeles, 22 Dec 1972; JN-3238, spermatangial, on Codium sp., Bahia San Francisquito, 24 May 1972. 

LAS ISLAS DE LA CINTURA: JN-5804, cystocarpic, 3-6 m depth, off 

W side of Roca Blanca, Puerto Refugio, Isla Angel de la 

Guarda, 22 Apr 1974; JN-5616B, 1-10 m depth off S end of 

Isla Estanque at the S end of Isla Angel de la Guarda, legit 

J. N. and K. B., 27 Apr 1974; JN-5520b, on Codium sp., 1-5 m 

depth, SE end, off Isla San Esteban, legit K. B., 26 Apr 1974. 

REMARKS.—Previously encountered off Isla San 

Jorge in the upper Gulf (Hollenberg, 1961:358), it 

was not surprising to find P. johnstonii var. concinna in Puerto Penasco and vicinity. Numbe r JN3011, cited above, is placed in this variety with some 

hesitation, since the branches frequently arise in 

pairs one segment apart. Also the procarpial 

branches are numerous and present an unusual 

appearance. They seemingly arise from relatively 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 4.—A-C, Polysiphonia sphaerocarpa var. cheloniae (JN-4765): A, the upper branching and 

a young cystrocarp; B, a nearly mature cystocarp; c, two spermatangial stichidia and the related 

trichoblast of one. D, P. johnstonii var. concinna, several young cystocarps with seemingly arrested 

development (JN-3011). 

large trichoblast primordia (scar-cells) and remain 

prominent and undeveloped, at least for the most 

part. 

Polysiphonia masonii Setchell & Gardner 

FIGURE 2D 

Polysiphonia masonii Setchell & Gardner, 1930:160.—Hollenberg, 1942b:783; 1961:358, pi. 6: figs. 2, 3.—Dawson, 

1954b: 160. 

DESCRIPTION.—Reddish brown epiphytes 2-3 cm 

high, flaccid, attached by a bundle of short unicellular rhizoids with digitate apices, arising from 

the lower segments of the erect primary axis; main 

axis 350-400 ^m in diameter below, repeatedly and 

regularly branched dichotomously, composed of 

segments mostly as long as the diameter; branches 

nearly as large as the main axis at the point of 

insertion, arising from an entire trichoblast primordium; pericentral cells 4, ecorticate; trichoblasts 

to 480 ^m long, with 2 or 3 dichotomies, tapering 

to very slender apices, arising 1 per segment in 

spiral sequence with 14 divergence, soon deciduous, 

leaving persistent scar-cells. 

Tetrasporangia in spiral series in the ultimate 

branches, 85-95 /an in diameter, not much distending the segments; cystocarps subspherical nearly 
NUMBER 1 

sessile, 300-325 ^m in diameter; spermatangial 

stichidia arising from a primary branch of a trichoblast, broadly fusiform. 

TYPE-LOCALITY.—Epiphytic on Zostera, Isla Guadalupe, off the Pacific Coast of Baja California. 

HOLOTYPE.—H. L. Mason 86, April 1925 (CAS 

173618, now at UC). 

GULF OF CALIFORNIA DISTRIBUTION.—Intertidal, 

Puerto Penasco. 

PACIFIC COAST DISTRIBUTION.—Isla Guadalupe. 

SPECIMEN STUDIED.—SONORA: H 67-60, tetrasporangial, intertidal, Playa Estaci6n, Puerto Penasco, 8 Apr 1967. 

REMARKS.—Now reported for the first time in the 

Gulf of California, this seems to be the only record 

of the species other than those recorded from Isla 

Guadalupe (Hollenberg, 1961:359). Rarely encountered, it is known only from the present Gulf 

collection; three others, including the type specimen, from Isla Guadalupe. 

Polysiphonia mollis J. D. Hooker & Harvey 

in Harvey 

FIGURE 5B 

Polysiphonia mollis J. D. Hooker & Harvey in Harvey, 

1847:43.—Dawson, 1957:8; 1966:29.—Dawson, Neushul, and 

Wildman, 1960a:72.—Hollenberg, 1961:359, pi. 4: fig. 2.— 

Abbott and Hollenberg, 1976:688, fig. 636. 

Polysiphonia eastwoodae Setchell & Gardner, 1930:161.—Hollenberg, 1948:160.—Dawson, 1954b: 160. 

Polysiphonia tongatensis var.? sensu Setchell & Gardner, 

1930:160 [non Polysiphonia tongatensis Harvey in Kuetzing, 

1864:14]. 

Polysiphonia snyderae Kylin, 1941:35, pi. 12: fig. 34.—Hollenberg, 1942a:784, fig. 9.—Dawson, 1944:330; 1951:56; 

1954b:160. 

Polysiphonia snyderae var. heteromorpha Hollenberg, 1942b: 

784. 

Polysiphonia snyderae var. intricata Hollenberg, 1942b:785.— 

Dawson, 1954b: 161. 

Polysiphonia tongatensis sensu Segi, 1951:207 [in part, with 

reference to Revilla Gigedo Islands' material of Setchell 

and Gardner, 1930:160 only; non Polysiphonia tongatensis 

Harvey in Kuetzing, 1864:14]. 

DESCRIPTION.—Thalli dull reddish brown, mostly 

5-12 cm high from a discoid base or mostly assurgent from prostrate branches of limited extent, 

attached by unicellular rhizoids with frequently 

digitate apices, arising 1-2(-5) per segment from 

the proximal ends of the pericentral cells, from 

which they are cut off as separate cells by a curving 

wall; erect branches 300-400 ^m in diameter below, 

commonly 60-180 ^m in diameter in upper parts; 

segments in main branches mostly 1-2 times as long 

as the diameter, but sometimes considerably longer; 

main erect branches richly branched pseudodichotomously above, mostly naked below; pericentral 

cell 4, ecorticate; trichoblasts 1 per segment in 

spiral sequence, with 14 divergence, soon deciduous, leaving persistent scar-cells; branches arising 

at irregular intervals, commonly 6-10 segments 

apart, a branch replacing a trichoblast in the spiral 

sequence. 

Tetrasporangia 60-70 ^m in diameter, in spiral 

series in the ultimate and subultimate branches; 

cystocarps abundant, ovoid to nearly globular, 300-

350 //in in diameter; spermatangial stichidia comprising a primary branch of a trichoblast. 

TYPE-LOCALITY.—Epiphytic on other algae, Tasmania. 

HOLOTYPE.—Collection of R. Gunn labeled 

"original sp. . . parasitical on large algae" in Harvey Herbarium (TCD). 

GULF OF CALIFORNIA DISTRIBUTION.—Usually 

intertidal, epiphytic, or growing on rock, wood, 

shells, etc.; El Tornillal to Bahia San Lucas. 

PACIFIC COAST DISTRIBUTION.—Southern British 

Columbia to northern Baja California; Sinaloa. 

SPECIMENS STUDIED.—SONORA: JN-3326, adrift, inlet, El Tornillal, legit D. L. Lindquist; H 67-1.5, legit T. Scalione, Playa 

Estacidn, Puerto Penasco, 28 Jan 1967; D27210 (US), rocky 

reef, Playa Tucson, N side of Punta Pelicano, vicinity of 

Puerto Penasco, legit E. Y. Dawson, 6 Apr 1966. BAJA CALIFORNIA DEL NORTE: JN 2998a, spermatangial, from 7.6 m depth, 

Isla la Ventana, Bahia de los Angeles, 21 May 1972; JN-3265a, 

growing on "pen shell," Pinna rugosa, 3-4.5 m depth, Bahia 

San Francisquito, 24 May 1972; JN-3226, a slender form, 

growing on shells, Bahia San Francisquito, 24 May 1972. 

REMARKS.—The recent collection from El Tornillal represents a slight northward extension from 

Puerto Penasco, Sonora (Dawson, 1966:29). This 

species is widely distributed in temperate and subtropical regions. It is most frequently encountered 

in sheltered waters. 

In collection JN-3265a the segments in the main 

axes are shorter than the typical length for the 

species. A more unusual feature in JN-2998a is the 

fact that the spermatangial stichidia arise from the 

entire trichoblast primordium. In both JN-2998a 
10 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

50pm 

lOO^m 

FIGURE 5.—A, Polysiphonia paniculata, branch tip (after Hollenberg, 1961); B, Polysiphonia 

mollis, branch tip (after Hollenberg, 1961). 

and JN-3265a the trichoblasts are very rudimentary 

or are represented by scar-cells only. Furthermore, 

rhizoids in JN-3265a are cut off as separate cells 

from the pericentral cells. The latter feature does 

not normally occur in species such as P. paciftca 

in which spermatangial stichidia arise from the 

entire trichoblast primordium. More material representing these two entities is needed for confident 

identification. 

Polysiphonia paciftca var. delicatula Hollenberg 

FIGURE 6B 

Polysiphonia paciftca var. delicatula Hollenberg, 1942b:778, 

1961:362.—Smith, 1944:360.—Abbott and Hollenberg, 1976: 

690. 

DESCRIPTION.—Usually medium red, 0.5-2.0 cm 

high from prostrate branches 50-110 ujri in diameter, arising endogenously from prostrate branches, 
NUMBER 1 11 

90pm B 

40pm 

FIGURE 6.—A, Polysiphonia sphaerocarpa var. cheloniae, branch apices of a spermatangial plant 

(JN-4765); B, P. paciftca var. delicatula, branch apices (JN-4848). 

and composed of segments 1.0-2.0 (-3.0) times as 

long; branches short and mostly with blunt apices; 

pericentral cells 4, ecorticate; trichoblasts lacking 

or exceedingly rare. 

Tetrasporangia 50-60 ^m in diameter, in straight 

series; cystocarps slightly urceolate; spermatangial 

stichidia slender, 140-200 ixm in length by 30-40 

^m in diameter, on very short pedicels, leaving 

persistant scar-cells when shed. 

TYPE-LOCALITY.—On wharf piling, low tide level, 

Municipal Wharf, Monterey, Monterey County, 

California. 

HOLOTYPE.—H 2888, 20 July 1939, in herbarium 

of G. J. Hollenberg (now at US). 

GULF OF CALIFORNIA DISTRIBUTION.—Epiphytic 

on various algae or growing on rocks, low intertidal to subtidal; Puerto Penasco to Guaymas; 

Bahia San Lucas. 

PACIFIC COAST DISTRIBUTION.—Central California; 

Chiapas; Galapagos Archipelago. 

SPECIMENS STUDIED.—SONORA: H 67-53, on coralline algae, 

Playa Estaci6n, Puerto Penasco, 25 Mar 1967; JN-4848a, 

epiphytic on Gelidium johnstonii (JN-4827), 3.3 m depth, on 

rock reef, Cabo Lobos, vicinity of Puerto Libertad, legit J. N. 

and K. B., 17 Nov 1973. 

REMARKS.—Seemingly rare in the Gulf of California, its distribution is now extended northward 

to Puerto Penasco from Bahia Guaymas (Hollenberg, 1961: 362). This species is generally found on 

rocks, wood, or epiphytic in deep or quiet waters. 

Polysiphonia paniculata Montagne 

FIGURES 4A, 5A, 7 

Polysiphonia paniculata Montagne, 1842:254.—Kuetzing, 1863: 

15, pi. 45c-e.—Howe, 1914:142-—Hollenberg, 1944:480, fig. 7; 

1961:362, pi. 7: fig. 1.—Dawson, Neushul, and Wildman, 

1960b: 26.—Dawson, Acelto and Foldvik, 1964:89, pis. 78A, 

79.—Dawson, 1954b: 160; 1966:29.—Smith, 1969:115 [725].-— 

Abbott and Hollenberg, 1976:701, fig. 652. 
12 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Polysiphonia californica Harvey, 1853:48.—Kylin, 1941:36.— 

Dawson, 1944:332.—Smith, 1944:362, pi. 93: fig. 1. 

DESCRIPTION.—Thalli densely tufted, soft, dark 

brownish red, mostly saxicolous, 10-25 cm high, 

arising from densely matted prostrate branches, attached by numerous unicellular rhizoids, with usually digitate apices, arising 2 or 3 per segment from 

the proximal end of the pericentral cells, from 

which they are cut off as separate cells; erect 

branches arising endogenously from the prostrate 

branches at close intervals, loosely branched to relatively naked below, densely clothed with branches 

above; branches of all orders exogenous, the ultimate branches commonly densely plumose, narrowed at the base; pericentral cells mostly 10-12, 

but sometimes as many as 14 in older parts or as 

few as 8 in younger parts, totally ecorticate; main 

axes 300-430 jxm in diameter and composed of segments 2.0-2.5(-4.0) times as long; trichoblasts to 

800 ju,m long, simple or mostly with 1 or 2 dichotomies, arising 1 per segment in spiral sequence with 

14 divergence, tardily deciduous, leaving relatively 

Mm 4 

FIGURE 7.—Polysiphonia paniculata, tetrasporic specimen (JN4552) from the mud/sand flats of Bahia Cholla (vicinity of 

Puerto Penasco), Sonora. 

large scar-cells; branches arising in association with 

trichoblasts. 

Tetrasporangia 80-100 ju.m in diameter, more or 

less spiraling in the ultimate and subultimate 

branches; cystocarps globular-ovoid, somewhat truncate, 350-400 fim in diameter; spermatangial stichidia 130-250 ,^m by 50-70 ^m without sterile 

apex at maturity; thalli adhering to paper when 

dried. 

TYPE-LOCALITY.—Peru. 

HOLOTYPE.—Collection by Gaudichaud labeled 

"in fronde Ulvae nematoideae, ad littora Peruviana" 

in Montagne Herbarium (PC). 

GULF OF CALIFORNIA DISTRIBUTION.—Midintertidal to subtidal, usually in protected bays and 

estuaries, Puerto Penasco to Guaymas. 

PACIFIC COAST DISTRIBUTION.—British Columbia 

to northern Baja California. Peru. 

SPECIMENS STUDIED.—SONORA: D 27254, slender form, mud 

flats, Bahia Cholla, 6 Apr 1966; JN-4551, cystocarpic, sand 

flats, 1 m depth, Bahia Cholla, 8 Dec 1973; JN-4552, tetrasporic, growing on living sand dollar (Encope grandis), sand 

flats, Bahia Cholla, legit J. N. and J. Kudenov, 8 Dec 1973; 

JN-4993, sand flats, Bahia Cholla, legit J. N. and K. B., 

9 Mar 1974; JN-3697, and JN-3713, both tetrasporangial, intertidal rock platform, W of Playa Hermosa, Puerto Penasco, 

2 Feb 1973; JN-4718, tetrasporangial, attached to sea shells, 

midway between Isla Tiburon and W of Campo Viboras, 

Sonora, legit G. L. Kooyman and E. Senate, 19 Dec 1973; 

JN-4773.5, spermatangial, collected along with P. sphaerocarpa 

var. chelonia (JN-4773), in the Infiernillo Channel between 

Campo Ona and Viboras, Sonora, legit R. S. Felger and 

E. Moser from a green sea turtle ($ Chelonia mydas) that 

was harpooned by Guadalupe Lopez, a Seri Indian, 21 Jan 

1974. 

REMARKS.—This species apparently occurs only in 

the northern Gulf. Here it has been found mostly 

in calm waters of estuaries, mud flats, protected 

bays, and coastal areas. It is also one of the marine 

algae known to be epizoic on the green sea turtle, 

Chelonia mydas L. in the Gulf (see Felger, Cliffton, 

and Regal, 1976; Norris, unpublished data). 

Polysiphonia savatieri Hariot 

FIGURE 8A 

Polysiphonia savatieri Hariot, 1891:226.—Segi, 1951:202, fig. 

lOa-n, pi. 3: fig. 8.—Dawson, 1954b: 160.—Hollenberg, 1961: 

363, pi. 7: fig. 4; 1968a:77, figs. 37, 38.—Abbott and Hollenberg, 1976:692, fig. 639. 

Polysiphonia minutisima Hollenberg, 1942b:781, fig. 21. 
NUMBER ] 13 

FIGURE 8.—A, Polysiphonia savatieri, upper portion of a tetrasporic thallus (JN-4195); B, P. 

scopulorum var. villum, entire thallus (JN-5098A); note the dichotomous branched trichoblast. 

DESCRIPTION.—Thalli epiphytic, dull reddish 

brown, mostly 3-10 mm high from a tuft or cushion 

of rhizoids, usually penetrating the host; rhizoids 

up to 1 mm long, cut off by a curving wall from 

the proximal end of the pericentral cells; main axes 

indistinct, commonly assurgent from the basal attachment, u p to 148 ^m in diameter; branches of 

several orders, exogenous, not associated with trichoblasts in origin, arising at variable intervals; 

pericentral cells 4, ecorticate; trichoblasts relatively 

short, 1 per segment in spiral sequence, with 14 

divergence, soon deciduous, leaving persistent scarcells. 

Tetrasporangia in short spiral series, u p to 80 /j,m 

in diameter, somewhat distending the segments; 

cystocarps slightly urceolate or mostly globular at 

maturity, 225-290 ^m in diameter; spermatangial 

stichidia about 135 fjjn. long and 40 ju.m in diameter, 

comprising one primary branch of a trichoblast, 

without sterile apex. 

TYPE-LOCALITY.—Epiphytic on large algae, Yokosuka, Japan. 

HOLOTYPE.—A collection by Dr. Savatier, about 

1885 (PC). 

GUL F OF CALIFORNIA DISTRIBUTION.—Mid -

intertidal to shallow subtidal, epiphytic on various 

algae, growing on rocks or in tide pools; Puerto 

Penasco to Bahia de Las Animas. 

PACIFIC COAST DISTRIBUTION.—Central California 

to northern Baja California; Isla Guadalupe. Japan. 

Western tropical Pacific. 

SPECIMENS STUDIED.—SONORA: JN-5030, on Neoagardhiella 

baileyii, Punta Pelicano, legit J. N., K. B., and D. Moore, 

17 Mar 1974; JN-3182, tetrasporangial, on intertidal rocks, 

W of Casa Garcia, Playa Estacion, Puerto Penasco, legit J. N. 

and K. B., 11 Jul 1972; JN-3545, cystocarpic, Cumpleafios tide 
14 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

pool, Playa Estacion, Puerto Penasco, legit J. N. and K. B., 

8 Sep 1972; JN-4537, tetrasporangial, Cumpleanos tide pool, 

Playa Estacion, Puerto Penasco, legit J. N. and K. B., 9 Nov 

1973; JN-4195, tetrasporangial, and JN-4197, cystocarpic, on 

intertidal platform and tide pools, Playa Estacion, Puerto 

Penasco, 1 Jun 1973; H 67-14, low intertidal, Playa Estaci6n, 

Puerto Penasco, 25 Feb 1967; JN-3878, spermatangial, on 

Codium sp. in front of Laboratorio de Biologia Marina, Playa 

Estacion, Puerto Penasco, 14 Apr 1973; JN-4266, cystocarpic, 

6 m depth, Neuva Guaymas, legit D. Evanson, 13 Oct 1973. 

BAJA CALIFORNIA DEL NORTE: JN-3126, epiphytic on Sargassum, 

9 m depth, Rocas Consag, legit D. L. Lindquist, 2 Jun 1972; 

JN-3281, spermatangial, on Codium sp., Puertecitos, legit J. N. 

and G. Boehlert, 17 May 1972; JN-4448, Isla la Ventana, 

Bahia de Los Angeles, 27 Jul 1973; JN-4442, on Dictyota sp., 

rocky intertidal, Isla la Ventana, Bahia de Los Angeles, 27 Jul 

1973; JN-3216b, tetrasporangial, on Gracilaria sp., Bahia de 

Las Animas, legit J. N. and G. Boehlert, 23 May 1972. 

REMARKS.—This species is infrequently encountered throughout its Pacific coast distribution (Hollenberg, 1961:364). It may now be added to the 

northern Gulf of California marine flora, where it 

was rather frequent among the recent collections. 

Polysiphonia scopulorum var. villum 

(J. G. Agardh) Hollenberg 

FIGURE 8B 

Polysiphonia scopulorum var. villum {J. G. Agardh) Hollenberg, 1968a:81, fig. 7A.—Abbott and Hollenberg, 1976:692, 

figs. 640, 641. 

Polysiphonia villum J. G. Agardh, 1863:941. 

Lophosiphonia villum (J. G. Agardh) Setchell & Gardner, 

1903:329.—Hollenberg, 1942a:535.—Dawson, 1944:332, pi. 48: 

figs. 1-6; 1951:53; 1954b:161; 1963:421, pi. 169: figs. 1-3.— 

Taylor, 1945:304. 

Lophosiphonia scopulorum sensu Dawson, 1959:36 [non 

Lophosiphonia scopulorum (Harvey) Womersley, 1950:188]. 

DESCRIPTION.—Brownish red thalli mostly 5-10 

mm high from prostrate branches attached by unicellular rhizoids arising from the center of the pericentral cells, from which they are not cut off by 

cross walls; erect branches unbranched or mostly 

very sparingly branched, mostly 40-60 ^m in diameter, composed of segments 1.0—1.5 times as long as 

the diameter, arising at irregular intervals in a 

strictly endogenous manner; lateral branches exogenous or endogenous, independent of trichoblasts, 

which they replace; pericentral cells 4, ecorticate; 

trichoblasts mostly infrequent to wanting, with 

1 or 2 dichotomies, 250-480 ju,m long, soon deciduous. 

Tetrasporangia 50-60 ^m in diameter, in straight 

series, somewhat distending the segments; cystocarps ovoid, 150-190 ^m in diameter; spermatangial 

stichidia cylindrical, without sterile apices, arising 

from an entire trichoblast primordium. 

TYPE-LOCALITY.—"Ad littus americae tropica," 

probably on the Pacific coast of Mexico. 

HOLOTYPE.—A collection by Liebmann in the 

Agardh Herbarium (LD). 

GULF OF CALIFORNIA DISTRIBUTION.—Intertidal 

rocks, Puerto Penasco; Isla Angel de la Guarda; 

Isla Partida; Isla Turner; Isla Estanque. 

PACIFIC COAST DISTRIBUTION.—British Columbia, 

Canada, to Costa Rica. Widely distributed in the 

northern Pacific and western Atlantic oceans. 

SPECIMEN STUDIED.—SONORA: JN-5098a, collected with P. 

simplex (JN-5098b), midtidal rocks, Playa Estacion, Puerto 

Penasco, 24 Mar 1974. 

REMARKS.—This species is found growing alone 

or with other algae, often forming a continuous 

turf on intertidal rocks. Dawson (1944:332; 1954b: 

161) reported this taxon from three islands of Las 

Islas de la Cintura under the name of Lophosiphonia villum (J. G. Agardh) Setchell & Gardner. Our 

new collection, moves the known distribution northward to Puerto Penasco from Puerto Refugio, Isla 

Angel de la Guarda (Dawson, 1944:333, 1963:422). 

T h e reasons for placing this species in Polysiphonia rather than in Lophosiphonia are given by 

Hollenberg (1968a: 56). In this connection, the criteria listed by Hollenberg indicate that Lophisiphonia mexicana described by Dawson (1944:333) 

cannot be properly assigned to that genus. Further 

collections of "L. mexicana" are needed before a 

definitive disposition of the taxon can be made. 

Polysiphonia simplex Hollenberg 

FIGURE 9 

Polysiphonia simplex Hollenberg, 1942b:782, fig. 18; 1961:364, 

pi. 5: fig. 1.—Dawson, 1944:331; 1951:53; 1954a:6; I954b:160; 

1966:29.—Abbott and Hollenberg, 1976:694, fig. 641. 

Polysiphonia ferulacea sensu Segi, 1951:209 [in part, with reference to Pacific North American specimens cited by Hollenberg, 1942b:782 only; not Polysiphonia ferulacea Suhr in 

J. G. Agardh, 1863:980]. 

DESCRIPTION.—Thalli medium to dark brownish 

red, drying nearly black; usually saxicolous, commonly forming dense mats often of considerable 

extent, alone or mixed with other algae, or occa-
NUMBER 1 15 

FIGURE 9.—Polysiphonia simplex: A, upper portion of the thallus showing the delicate trichoblasts 

(JN-5098B); B, upper portion of a cystocarpic thallus (JN-5038). 

sionally epiphytic; with creeping basal branches 

250—360 ju,m in diameter, composed of short segments, and attached by numerous unicellular 

rhizoids, which often have digitate apieces and are 

always cut off as separate cells from the proximal 

end of the pericentral cells; erect branches l-3(-7) 

cm high and 160-250 ^m in diameter near the base, 

arising in an exogenous assurgent manner; main 

axes usually distinct but not prominent, sparingly 

branched; branches exogenous, radially directed, 

of several orders; trichoblasts delicate, to 730 /*m 

long with 1 or 2 dichotomies, 1 per segment in 

spiral sequence with 14 divergence, soon deciduous, 

leaving persistent scar-cells; branches replacing 

trichoblasts in the spiral (arising from an entire 

trichoblast primordium). 

Tetrasporangia up to 70 ju,m in diameter, in spiral 

series in the ultimate branches, more or less distending the segments; cystocarps ovoid-globular, 300-

350 ixm in diameter, spermatangial stichidia 100-

170 /xm long, 35-40 ju,m in diameter, comprising one 

primary branch of a trichoblast; without sterile 

apex. 

TYPE-LOCALITY.—On rocks, Laguna Beach, 

Orange County, California. 

HOLOTYPE.—H 2115, 14 May 1937, in herbarium 

of G. J. Hollenberg (now at US; isotype at GMS). 

GULF OF CALIFORNIA DISTRIBUTION.—Intertidal, 

growing alone or with other turf algae forming 

mats; Puerto Penasco to Cabeza Ballena. 

PACIFIC COAST DISTRIBUTION.—Southern California to southern Mexico. Costa Rica. 

SPECIMENS STUDIED.—SONORA: D 27173, Bahia Cholla, mudflat, 5 Apr 1966; H 67-64, tetrasporangial, cystocarpic, Playa 

Tucson, Punta Pelicano, 9 Apr 1967; JN-5038, cystocarpic, low 

intertidal, Punta Pelicano, 17 Mar 1974; JN-5098b, midintertidal rock platform, Playa Estacion, Puerto Penasco, 

legit J. N. and K. B., 24 Mar 1974. 
16 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

REMARKS.—This species is reported by Dawson 

(1944:331) and Hollenberg (1961:364) from a number of Gulf localities. With the recent collections 

from Puerto Penasco, we now know P. simplex to 

be widely distributed throughout the Gulf. 

Polysiphonia sonorensis Hollenberg 

FIGURE 2C 

Polysiphonia sonorensis Hollenberg, 1942b:779, figs. 5, 20.— 

Dawson, 1944:330; 1954b:161.—Hollenberg, 1961:365, pi. 5: 

fig. 2. 

DESCRIPTION.—Densely tufted algae of a terracotta color, not strongly adhering to paper on drying, 

4—5 cm high from prostrate creeping branches 120-

175 ju,m in diameter attached by unicellular rhizoids, 

which arise as outgrowths of the center of the pericentral cells, from which they are not cut off as 

separate cells; erect branches arising mostly endogenously or sometimes exogenously from the prostrate brances, 70-100 ^m in diameter, composed of 

segments mostly 1.5-2.0 times as long as the diameter; branching dense, the branches arising exogenously at irregular intervals of mostly 1—20 segments and mostly at an acute angle with the parent 

branch; main axes indistinct; branches replacing 

trichoblasts; trichoblasts numerous, delicate, to 1 mm 

long, arising at irregular intervals, with 1 or 2 

dichotomies, soon deciduous, leaving inconspicuous 

scar-cells. 

Reproductive material has not been found. 

TYPE-LOCALITY.—Afloat along the southeastern 

shore of Bahia Empalme, Sonora, Gulf of California. 

HOLOTYPE.—Francis Drouet and Donald Richards 3426a (F); isotype: herbarium of G. J. Hollenberg (now at GMS). 

GULF OF CALIFORNIA DISTRIBUTION.—Intertidal; 

Bahia Empalme to Guaymas. 

SPECIMENS STUDIED.—SONORA: in addition to the type, there 

is one other collection (Frances Drouet and Donald Richards3144) from rocks between tide marks on NW shore of harbor 

at Guaymas. 

REMARKS.—Known only from two specimens, this 

species is apparently endemic to the Gulf. It was 

not found in the recent collections. 

Polysiphonia sphaerocarpa var. cheloniae, 

new variety 

FIGURES 4A-C, 6A 

DESCRIPTION.—Plantae ad 3.5 cm altae, a crista 

basali rhizoideorum per rhizoidea e ramis assurgentibus suppletorum, ortae; rhizoidea per muros 

curvatos a finibus proximalibus cellulorum pericentralium ut cellulae discretae abscissa; cellulae 

pericentrales 4; ramificatio pseudodichotoma; rami 

principales ad 300 fjja. crassi, e segmentis 1.0-2.5 

(-3.0) plo longioribus quam lata conpositi; segmentis in ramis iunioribus turgidulis; trichoblastae 

ad 400 jam longae, una in unoquoque segmento, 

1-3 dichotomias habentes, ad apices delicatos 

attenuatae; rami e primordio integro trichoblastae 

intervalis fere 5-12 segmentorum enascentes, basim 

versus angusti. 

Tetrasporangia ignota; cystocarpi globosi, ad 370 

lira, diametro; stichidia spermatangialia 160-175 

fxxa. longa, 40-48 ^m diametro, cylindrica aut fusiformia, fere sine cacumine sterili, ut furca primaria 

trichoblastae enascentia. 

Thalli to 3.5 cm high, from a basal tuft of rhizoids supplemented by rhizoids from assurgent 

branches; rhizoids cut off as separate cells from the 

proximal end of the pericentral cells; pericentral 

cells 4, ecorticate; branching pseudodichotomous; 

main branches to 300 jitn in diameter, composed of 

segments 1.0-2.5(-3.0) times as long as the diameter; 

branches arising from an entire trichoblast primordium in intervals of mostly 5-12 segments, narrowed toward the base; segments in younger 

branches turgid; trichoblasts one per segment to 

400 /j.m long, with 1-3 dichotomies, tapering to 

delicate apices. 

Tetrasporangia unknown; cystocarps globular, to 

370 ^m in diameter; spermatangial stichidia 160-175 

/j.m long, 40-48 ^m in diameter, cylindrical to fusiform, mostly without sterile apices at maturity, 

arising as a primary fork of a trichoblast. 

TYPE-LOCALITY.—On green sea turtle, Chelonia 

mydas L., in Canal de Infiernillo between Campo 

Ona and Campo Viboras, Sonora, opposite the east 

coast of Isla Tibur6n. Gulf of California. 

HOLOTYPE.—J. Norris-4801, cystocarpic (US). 

GULF OF CALIFORNIA DISTRIBUTION.—Canal de 

Infiernillo, between Sonora and Isla Tiburon. 
NUMBER 1 17 

FIGURE 10.—Polysiphonia species: A, upper portion of a spermatangial thallus (JN-2998); B, branch 

tip, showing the spermatangial stichidia, which originates from an entire trichoblast primordium 

(JN-2998); c, upper portion, showing the usual absence of trichoblasts (JN-3265); D, upper portion 

of a tetrasporangial thallus, showing the developing tetrasporangia initials in the characteristic 

spiral sequence (JN-3265). 
18 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

SPECIMENS STUDIED.—SONORA: all collections of this alga were 

made 21 Jan 1974 in the Infiernillo Channel between Campo 

Ona and Campo Viboras, Sonora (opposite the east coast of 

Isla Tiburon). They were collected by R. S. Felger and E. W. 

Moser from medium size to young adult green sea turtles 

(Chelonia mydas), which were harpooned by Guadalupe Lopez, 

a Seri Indian. Five collections were made: JN-4801, cystocarpic, 

holotype (US) was taken from the carapace of female turtle 

number 4, harpooned at approximately 9 m depth; JN-4765, 

partly spermatangial, was from the dorsal surface of the head 

and flippers of another female turtle number 2, harpooned at 

approximately 7.6 m deptli; JN-4773, sterile, was collected 

from the carapace of female turtle number 1, taken from 

approximately 9 m depth; JN-4777, immature cystocarpic, was 

from the carapace of turtle number 1; and JN-4794, sterile, 

was from the dorsal surface of the head and front flippers of 

female turtle number 1. 

REMARKS.—This alga seems very close in its general features to P. sphaerocarpa Boergesen (1918: 

271) and especially to var. sphaerocarpa as described 

by Hollenberg (1968a:90) from the central and 

western Pacific Ocean. It differs from that variety in 

the substratum (i.e., the green sea turtle), and the 

consequent habitat. It is also somewhat taller than 

previously described specimens of that taxon. 

At least two other species of Polysiphonia have 

been described as growing on sea turtles. Polysiphonia tsudana, collected from a species of sea 

turtle (Chelonia) taken near Laysan Island, Hawaii, 

was described by Hollenberg (1968b:205). It is 

smaller than the Gulf of California taxon, with 

branches much smaller in diameter and having 

shorter segments. Furthermore, the Laysan specimens of P. tsudana had a variable number of pericentral cells and the spermatangial stichidia arise 

from the entire trichoblast primordium, rather than 

from a primary branch as in P. spaerocarpa var. 

cheloniae from the northern Gulf. Polysiphonia 

carettia, also described by Hollenberg (1971:15), 

was found growing on the carapace of a Loggerhead sea turtle (Caretta caretta L.), taken at the 

west end of Santa Catalina Island, off the coast of 

southern California. This alga is likewise smaller 

than the northern Gulf of California specimens. It 

differs, furthermore, in that the branches of the 

California P. carettia are all or mostly all cicatrigenous in origin, arising from the scar-cells. Finally the 

rhizoids are cut off as separate cells in the Gulf of 

California specimens, but remain in open connection with the pericentral cells in the case of the 

California species. 

Polysiphonia species 

FIGURE 10 

DESCRIPTION.—Thalli to 3 cm high, repeatedly 

pseudodichotomously branched at irregular intervals of mostly 8-10 segments; main axes to 320 ^m 

in diameter below, composed of segments 1.0-1.5 

times as long as the diameter; pericentral cells 4, 

ecorticate; rhizoids not observed; trichoblasts very 

rudimentary or absent, scar-cells 1 per segment in 

spiral sequence. 

Tetrasporangia in spiral series in the smaller 

branches; cystocarps not seen; spermatangial stichidia arising from an entire trichoblast primordium. 

SPECIMENS STUDIED.—BAJA CALIFORNIA DEL NORTE: two collections are assigned to this entity: JN-2998b, spermatangial, 

from 7.6 m depth, Isla la Ventana, Bahia de Los Angeles, 

21 May 1972; and JN-3265b, tetrasporangial, growing on the 

"pen shell" (Pinna rugosa), 3-5 m depth, Bahia San Francisquito, 24 May 1972. 

REMARKS.—The manner of origin of spermatangial stichidia is very much like that in P. paciftca and P. scopulorum, but the presence of a scarcell or a trichoblast on each segment is not a feature 

normally associated with the origin of spermatangial stichidia from an entire trichoblast primordium as in the last named species. Also there 

are differences in the arrangement of the tetrasporangia. In P. paciftca and P. scopulorum they 

are in a straight series, whereas in our Gulf of California collections they are arranged in a spiral 

series (Figure 10D). It seems probable that these 

specimens represent an undescribed species. Further 

collections are needed to determine their taxonomic 

status. 
Literature Cited 

Abbott, I. A., and G. J. Hollenberg 

1976. Marine Algae of California, xii + [2] + 827 pages. 

Stanford: Stanford University Press. 

Agardh, J. G. 

1863. Species, genera et ordines Algarum, seu descriptiones 

succinctae specierum generum et ordinum, quibus 

algarum regnum constituitur. Volume 2, part 3, pages 

701-1291. Lund: G. W. K. Gleerup. 

Boergesen, F. 

1918. The Marine Algae of the Danish West Indies, III: 

Rhodophyceae. Danske Botanisk Arkiv, 3:241-304. 

Dawson, E. Y. 

1944. The Marine Algae of the Gulf of California. Allan 

Hancock Pacific Expeditions, 3(10): 189-454. 

1949. Resultados preliminares de un reconocimiento de las 

algas marinas de la costa Paciftca de Mexico. Revista 

de la Sociedad Mexicana de Historia Natural, 9(3/4): 

215-254. 

1950. A Note on the Vegetation of a New Coastal Upwelling Area of Baja California. Journal of Marine Research, 9(2): 66-68. 

1951. A Further Study of Upwelling and Associated Vegetation along Pacific Baja California, Mexico. Journal 

of Marine Research, 10:39-58. 

1954a. The Marine Flora of Isla San Benedicto Following 

the Volcanic Eruption of 1952-1953. Allan Hancock 

Foundation Publications Occasional Paper, 16:1-13. 

1954b. Resumen de las investigaciones recientes sobre algas 

marinas de la costa Paciftca de Mexico, con una 

sinopsis de la literatura, sinonimia y distribucion de 

las especies descritas. Revista de la Sociedad Mexicana 

de Historia Natural, 13:97-197 + i-x [reprint of 1953 

with corrections, index, pagination, and addenda]. 

1957. Notes on Eastern Pacific Insular Marine Algae. Los 

Angeles County Museum Contributions in Science, 

8:1-8. 

1959. Marine Algae from the 1958 Cruise of the Stella 

Polaris in the Gulf of California. Los Angeles County 

Museum Contributions in Science, 27:1-39. 

1963. Marine Red Algae of Pacific Mexico, Part 8, Ceramiales: Dasyaceae, Rhodomelaceae. Nova Hedwigia, 

6:401-481. 

1966. Marine Algae in the Vicinity of Puerto Penasco, 

Sonora, Mexico. Gulf of California Field Guide Series, l:iii + 57. Tucson: University of Arizona. 

Dawson, E. Y., C. Acleto O., and N. Foldvik 

1964. The Seaweeds of Peru. Beihefte zur Nova Hedwigia, 

13: [5] + 111 pages. 

Dawson, E. Y., M. Neushul, and R. D. Wildman 

1960a. Seaweeds Associated with Kelp Beds along Southern 

California and Northwestern Mexico. Pacific Naturalist, 1(14):1-81. 

1960b. New Records of Sublittoral Marine Plants from 

Pacific Baja California. Pacific Naturalist, 1(19): 1-30. 

Felger, R. S., K. Cliffton, and P. J. Regal 

1976. Winter Dormancy in Sea Turtles: Independent Discovery and Exploitation in the Gulf of California 

by Two Local Cultures. Science, 191:283-285. 

Gardner, N. L. 

1927. New Rhodophyceae from the Pacific Coast of North 

America, VI. University of California Publications, 

Botany, 14:99-138. 

Greville, R. K. 

1824. Flora Edinensis. lxxxi -f 478 pages. Edinburgh: W. 

Blackwood. 

Hariot, P. 

1891. Liste des algues marines reportees de Yokusuka 

(Japon) par M. le Dr. Savatier. Memoires de la Societe 

des Sciences Naturelles et Mathematiques de Cherbourg, 27:211-230. 

Harvey, W. H. 

1847. Nereis Australis, or Algae of the Southern Ocean. 

[2] + viii + 124 pages. London: Reeve Brothers. 

1853. Nereis Boreali-Americana; or, Contributions towards 

a History of the Marine Algae of the Atlantic and 

Pacific Coasts of North America, Part II: Rhodospermeae. 258 pages. Washington, D. C: Smithsonian 

Institution. 

Hollenberg, G. J. 

1942a. Phycological Notes, I. Bulletin of the Torrey Botanical Club, 69:528-538. 

1942b. An Account of the Species of Polysiphonia on the 

Pacific Coast of North America, I: Oligosiphonia. 

American Journal of Botany, 29:772-785. 

1944. An Account of the Species of Polysiphonia on the 

Pacific Coast of North America, II: Polysiphonia. 

American Journal of Botany, 31:474-483. 

1948. Notes on Pacific Coast Marine Algae. Madrofio, 

1948:155-162. 

1961. Marine Red Algae of Pacific Mexico, Part 5: The 

Genus Polysiphonia. Pacific Naturalist, 2(6):345-375. 

1968a. An Account of the Species of Polysiphonia of the 

Central and Western Tropical Pacific Ocean, I: Oligosiphonia. Pacific Science, 22(1): 56-98. 

1968b. An Account of the Species of the Red Alga Polysiphonia of the Central and Western Tropical Pacific 

Ocean, II: Polysiphonia. Pacific Science, 22(2): 198-

207. 

1971. Phycological Notes, V: New Species of Marine Algae 

from California. Phycologia, 10(1): 11-16. 

19 
20 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Holmgren, P. K., and W. Keuken 

1974. Index Herbariorum, Part I: The Herbaria of the 

World. Sixth edition, vii + 397 pages. Utrecht, 

Netherlands: Oosthoek, Scheltema & Holkema. 

Hooker, W. J. 

1833. Mosses, Hepatiae, Lichens, Characeae and Algae. 

Part I of volume 5 (Class XXIV, Cryptogamia) in 

The English Flora of Sir James Edward Smith. 

x + 4 + 432 pages. London: Longman, Rees, Orme, 

Brown, Green & Longman. [Listed under "J. E. 

Smith" in most libraries.] 

Howe, M. A. 

1914. The Marine Algae of Peru. Memoirs of the Torrey 

Botanical Club, 15:1-185. 

Kuetzing, F. T. 

1863. Tablulae Phycologicae. Volume 13, [i] + 31 pages. 

Nordhausen. 

1864. Tabulae Phycologicae. Volume 14, [i] + 35 pages. 

Nordhausen. 

Kylin, H. 

1941. Californische Rhodophyceen. Lunds Universitets 

Arsskrift, N. F. Avd. 2, 37(2)1-51; and Kunglig Fysiograftska Sallskapets Handlingar, new series, 52(2): 1-51. 

Montagne, C. 

1842. Troisieme centurie de plantes cellulaires exotiques 

nouvelle, part 2. Annales des Sciences Naturelles, 

Botanique, series 2, 18:241-282. 

Norris, J. N. 

1972. Marine Algae from the 1969 Cruise of "Makrele" to 

the Northern Part of the Gulf of California. Boletin 

de la Sociedad Botanica de Mexico, 32:1-30. 

Norris, J. N., and K. E. Bucher 

1976. New Records of Marine Algae from the 1974 R/V 

Dolphin Cruise to the Gulf of California. Smithsonian 

Contributions to Botany, 34:1-22. 

Reinsch, P. F. 

1888. Species et genera nova Algarum ex insula Georgia 

Australi. Berichte der Deutschen Botanischen Geselischaft, 6:144-156. 

Segi, T. 

1951. Systematic Study of the Genus Polysiphonia from 

Japan and Its Vicinity. Journal of the Faculty of 

Fisheries, Prefectural University of Mie, 1(2): 169-272. 

Setchell, W. A., and N. L. Gardner 

1903. Algae of Northwestern America. University of California Publications, Botany, 1:165-418. 

1924. The Marine Algae: Expedition of the California 

Academy of Sciences to the Gulf of California in 

1921. Proceedings of the California Academy of Sciences, fourth series, 12:695-949. 

1930. Marine Algae of the Revillagigedo Islands Expedition in 1925. Proceedings of the California Academy 

of Sciences, fourth series, 19:109-215. 

Smith, G. M. 

1944. Marine Algae of the Monterey Peninsula, California. 

ix + 622 pages. Stanford: Stanford University Press. 

1969. Marine Algae of the Monterey Peninsula, California. 

Second edition (incorporating the 1966 Supplement 

by G. J. Hollenberg and I. A. Abbott), x + 752 pages. 

Stanford: Stanford University Press. 

Taylor, W. R. 

1945. Pacific Marine Algae of the Allan Hancock Expeditions to the Galapagos Islands. Allan Hancock Pacific 

Expeditions, 12: iv 4- 528 pages. 

Womersley, H. B. S. 

1950. The Marine Algae of Kangaroo Island, III: List of 

Species, 1. Transactions of the Royal Society of 

Southern Australia, 73(2): 137-197. 

•fr u.= . GOVERNMENT PRINTING OFFICE: 1977 O 227-134 
Index 

(Synonyms and principle entries in italics; new variety in SMALL CAPITALS) 

Lophosiphonia, 14 

mexicana, 14 

scopulorum, 14 

villum, 14 

Polysiphonia, 2 (key) 

acuminata, 4 

californica, 12 

concinna, 7 

confusa, 2, 3 (fig.) 

eastwoodae, 9 

ferulacea, 14 

flaccidissima, 4, 5 (fig) 

inconspicua, 2 

johnstonii var. concinna, 5 (fig), 7, 8 (fig.) 

johnstonii var. johnstonii, 4, 5 (fig.), 6 (fig.) 

marchantae, 4 

masonii, 5 (fig.), 5 

minutisima, 12 

mollis, 9, 10 (fig.) 

paciftca var. delicatula, 10, 11 (fig) 

paniculata, 10 (fig.), 11, 12 (fig.) 

richardsoni, 4 

savatieri, 12, 13 (fig.) 

scopulorum var. villum, 13 (fig), 14 

simplex, 14, 15 (fig.) 

sinicola, 4 

snyderae, 9 

snyderae var. heteromorpha, 9 

snyderae var. intricata, 9 

sonorensis, 5 (fig.), -76 

species, 77, 18 (fig.) 

sphaerocarpa var. CHELONIAE, 8 (fig), 11 (fig), 16 

tongatensis, 9 

tongatensis var. ?, 9 

villum, 14 

21