A Chronology of 

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Smithsonian Institution

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s m i t h s o n i a n c o n t r i b u t i o n s t o b o t a n y • n u m b e r 9 4

Marine Algae of the 

Northern Gulf of California:

Chlorophyta and 

Phaeophyceae

James N. Norris
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s m i t h s o n i a n c o n t r i b u t i o n s t o b o t a n y • n u m b e r 9 4

Marine Algae of the 

Northern Gulf of 

California: Chlorophyta 

and Phaeophyceae

James N. Norris

washington d.c.

2010
ABSTRACT

Norris, James N. Marine Algae of the Northern Gulf of California: Chlorophyta and Phaeophyceae. Smithsonian Contributions to Botany, number 94, x + 276 pages, 109 figures, 2010. — The present treatment 

constitutes a taxonomic study of the green and brown benthic marine algae currently known in the northern Gulf of California. In all, 133 species were found: 4 classes, 7 orders, 13 families, 20 genera, and 63 

species of Chlorophyta and 9 orders, 15 families, 26 genera, and 70 species of Phaeophyceae. The systematic account is a guide to the identification of marine algae from the northern Gulf of California, with 

the accepted name, keys, and descriptions of the orders, families, genera, and species. The date, place, and 

author(s) of valid publication of each taxon are cited. The current name for each species is given along 

with its basionym, type locality, synonyms, relevant taxonomic studies, habitat, and distribution in the 

Gulf of California and in the Pacific Ocean. Illustrations are provided for most species. A remarks section 

includes additional information on taxonomy, nomenclature, ecology, and/or other problems or facts of 

interest. In addition to reviewing the taxonomic phycological literature pertinent to the Gulf of California 

and Pacific México, new records and distribution extensions are given. Four new sections of Sargassum

subgen. Sargassum are proposed: S. sect. herporhizum E. Y. Dawson ex J. N. Norris, S. sect. johnstonii

E. Y. Dawson ex J. N. Norris, S. sect. lapazeanum E. Y. Dawson ex J. N. Norris, and S. sect. sinicola E. Y. 

Dawson ex J. N. Norris; five new combinations are made: Desmarestia munda subsp. mexicana (E. Y. 

Dawson) J. N. Norris, Epicladia condensata (Setchell et N. L. Gardner) J. N. Norris, Epicladia mexicana

(Setchell et N. L. Gardner) J. N. Norris, Hincksia bryantii (Setchell et N. L. Gardner) J. N. Norris, and 

Sargassum sinicola subsp. camouii J. N. Norris et Yensen. One new species, Sporochnus neushulii J. N. 

Norris, is described.

Cover images, left to right: Figures 55, 37, and 79A (details).

Published by Smithsonian Institution Scholarly Press

P.O. Box 37012, MRC 957

Washington, D.C. 20013-7012

www.scholarlypress.si.edu

Library of Congress Cataloging-in-Publication Data

Norris, James N.

Marine algae of the northern Gulf of California: Chlorophyta and Phaeophyceae / James N. Norris.

 p. cm. — (Smithsonian contributions to botany ; no. 94)

Includes bibliographical references and index.

1. Marine algae—Mexico—California, Gulf of. I. Title. II. Series: Smithsonian contributions to 

botany ; no. 94.

QK571.9.C35N67 2010

579.8'8091641—dc22

2009046617

ISSN: (print) 0081-024X; (online) 1938-2812

The paper used in this publication meets the minimum requirements of the American National Standard 

for Permanence of Paper for Printed Library Materials Z39.48–1992.
Contents

LIST OF FIGURES vii

INTRODUCTION 1

General Oceanographic Features of the Gulf of California 3

 General Features of the Marine Flora of the Northern 

Gulf of California 7

 Historical Review of Marine Botanical Exploration of the 

Gulf of California 13

Current Study 16

PART I: GREEN ALGAE 19

CHLOROPHYTA 19

ULVOPHYCEAE 20

Ctenocladales 21

 Ulvellaceae 21

 Entocladia Reinke 22

 Epicladia Reinke 25

 Stromatella Kornmann et Sahling 26

 Ulvella P. Crouan et H. Crouan 28

Phaeophilales 30

 Phaeophilaceae 30

 Phaeophila Hauck 30

Ulvales 31

 Ulvaceae 31

 Ulva Linnaeus 32

CLADOPHOROPHYCEAE 49

Cladophorales 49

 Cladophoraceae 50

 Chaetomorpha Kützing 50

 Cladophora Kützing 54

 Rhizoclonium Kützing 64
i v • 

Siphonocladales sensu stricto 67

 Boodleaceae 67

 Cladophoropsis Børgesen 68

 Phyllodictyon J. E. Gray 70

Siphonocladaceae 74

 Ernodesmis Børgesen 74

 Valoniaceae 75

 Valoniopsis Børgesen 75

BRYOPSIDOPHYCEAE 77

Bryopsidales 77

 Bryopsidineae 78

 Bryopsidaceae 78

 Bryopsis J. V. Lamouroux 79

 Derbesiaceae 82

 Derbesia Solier 82

 Halicystis-phase 85

 Codiaceae 85

 Codium Stackhouse 86

 Halimedineae 92

 Caulerpaceae 94

 Caulerpa J. V. Lamouroux 94

 Halimedaceae 98

 Halimeda J. V. Lamouroux 99

DASYCLADOPHYCEAE 101

Dasycladales 101

 Polyphysaceae 102

 Acetabularia J. V. Lamouroux 102

PART II: BROWN ALGAE 105

STRAMENOPILES 105

PHAEOPHYCEAE 106

Ishigeales 108

 Ishigeaceae 108

 Ishige Yendo 108

Sphacelariales sensu stricto  1

 Sphacelariaceae 111

 Sphacelaria Lyngbye in Hornemann 111

Dictyotales 117

 Dictyotaceae 117

 Dictyopteris J. V. Lamouroux 119

 Dictyota J. V. Lamouroux 122

 Padina Adanson 133

 Rugulopteryx De Clerck et Coppejans 141

 Spatoglossum Kützing 142

Desmarestiales 144

 Desmarestiaceae 144

 Desmarestia J. V. Lamouroux 145
n u m b e r 9 4 • v

Sporochnales 148

 Sporochnaceae 150

 Sporochnus C. Agardh 150

Ralfsiales* 155

 Ralfsiaceae 155

 Hapterophycus Setchell et N. L. Gardner in Setchell 156

 Ralfsia Berkeley in J. E. Smith et J. C. Sowerby 157

Cutleriales 161

 Cutleriaceae 162

 Cutleria Greville 162

Ectocarpales 165

 Acinetosporaceae 166

 Feldmannia Hamel 167

 Hincksia J. E. Gray 168

 Chnoosporaceae 171

 Chnoospora J. Agardh 172

 Chordariaceae 174

 Hecatonema Sauvageau 174

 Nemacystus Derbès et Solier 176

 Streblonema Derbès et Solier 177

 Ectocarpaceae 180

 Ectocarpus Lyngbye 180

 Pylaiellaceae 186

 Pylaiella Bory de Saint-Vincent 186

 Leathesiaceae** 187

 Leathesia S. F. Gray 187

 Scytosiphonaceae 189

 Colpomenia (Endlicher) Derbès et Solier 189

 Compsonema Kuckuck 194

 Hydroclathrus Bory de Saint-Vincent 195

 Rosenvingea Børgesen 196

Fucales 199

 Sargassaceae 200

 Sargassum C. Agardh 200

Excluded Records of Phaeophyceae 219

ACKNOWLEDGMENTS 221

APPENDIX 1: LIST OF NEW TAXA AND COMBINATIONS 225

APPENDIX 2: ENDEMIC TAXA OF THE GULF OF CALIFORNIA 

THAT OCCUR IN THE NORTHERN GULF 227

APPENDIX 3: ALGAL SPECIMENS ILLUSTRATED: SPECIES 

LOCALITY INFORMATION AND HERBARIUM 229

REFERENCES 235

INDEX OF SCIENTIFIC NAMES 271

* Contributed by Daniel León Álvarez and James N. Norris. 

** Contributed by Luis E. Aguilar-Rosas, Raúl Aguilar-Rosas, and James N. Norris.

Figures

1. Gemini V photograph of northern Gulf of California 2

2. Map of the northern Gulf of California, Mexico 4

3. Summer intertidal marine flora, Puerto Peñasco 8

4. Winter intertidal marine flora, Puerto Peñasco 9

5. Colpomenia sinuosa and Codium simulans 11

6. Intertidal zonation, low tide, Puerto Refugio, Isla Ángel 

de la Guarda 12

7. E. Yale Dawson (left) and Palmer T. Beaudette during 1959 

expedition to Pacific Mexico and Central America 15

8. E. Yale Dawson preparing for collecting dive during 1958 

cruise on M/V Stella Polaris; collecting subtidal algae in 

Costa Rica, 1959 15

9. Entocladia marchantiae, E. polysiphoniae, Entocladia sp., and 

E. viridis 24

10. Epicladia condensata and E. mexicana 26

11. Stromatella monostromatica epiphytic on Polysiphonia 27

12. Ulvella lens and U. setchellii epiphytes on Cladophora 29

13. Ulva anatomy: diagrammatic transections of Ulva sect. ulva

and Ulva sect. enteromorpha 31

14. Ulva californica habits, typical forms found in the 

northern Gulf 35

15. Ulva lactuca habit 36

16. Ulva lobata habit 37

17. Ulva nematoidea habits 38

18. Ulva rigida habit 40

19. Ulva acanthophora habit and close-up 42

20. Ulva clathrata habit 43

21. Ulva flexuosa lower thallus and close-up 45

22. Ulva intestinalis: cluster of tubular thalli 47

23. Ulva linza: three blades, characteristically unbranched 48

24. Chaetomorpha antennina and C. aerea 52

25. Chaetomorpha bangioides and C. antennina 53
v i i i • 

26. Cladophora albida and C. tiburonensis 56

27. Cladophora columbiana and C. macdougalii 57

28. Cladophora graminea 58

29. Cladophora hesperia 59

30. Cladophora microcladioides branching patterns 61

31. Rhizoclonium tortuosum, terminal portions of filaments 66

32. Cladophoropsis gracillima, drawings of type specimen 69

33. Phyllodictyon pulcherrimum axis with well-developed 

open reticulate fronds 71

34. Phyllodictyon robustum variation in habits 73

35. Ernodesmis verticillata, upper portion showing successive 

series of whorled branches 75

36. Valoniopsis hancockii and V. pachynema 77

37. Bryopsis pennata showing featherlike fronds 81

38. Derbesia marina filamentous sporophyte 84

39. Halicystis-phase globose gametophytes, epiphytic on 

articulated coralline Amphiroa 85

40. Codium amplivesiculatum, habit of large specimen 88

41. Utricles of Codium species: C. amplivesiculatum, 

C. brandegeei, C. simulans 89

42. Codium brandegeei, habit of “C. macdougalii-form” 90

43. Codium simulans habit 92

44. Codium sp. 1 specimen somewhat similar to “Codium 

cuneatum-form” of C. simulans 93

45. Caulerpa mexicana f. pectinata habit 95

46. Caulerpa sertularioides habits 97

47. Caulerpa vanbosseae habits 98

48. Halimeda discoidea habit, surface view and transection 100

49. Acetabularia caliculus cluster of thalli and gametangial rays 103

50. Ishige sinicola, habit variation and transection 110

51. Sphacelaria rigidula apical region and propagules 113

52. Sphacelaria californica and S. hancockii propagules 115

53. Sphacelaria tribuloides apical region and propagules 117

54. Northern Gulf of California Dictyotaceae: diagrammatic 

illustrations of apical region and transections of Dictyota, 

Rugulopteryx, Dictyopteris, Spatoglossum, and Padina 120

55. Dictyopteris undulata habit with prominent midrib 121

56. Dictyota concrescens habits of isotype and Gulf specimens 124

57. Dictyota coriacea habit 125

58. Dictyota crenulata habit and close-up showing characteristic 

dentate margins 127

59. Dictyota dichotoma var. intricata habit variations 129

60. Dictyota flabellata habit 130

61. Dictyota stolonifera thalli with basal stolons 131

62. Dictyota vivesii habit 132

63. Padina caulescens blades above long flat stipes 135

64. Padina crispata habit and upper, lower sides of blade 136

65. Padina durvillei, leathery flabellate blades 137

66. Padina gymnospora blades 139
n u m b e r 9 4 • i x

67. Padina mexicana var. mexicana blades forming rosettelike 

clusters 140

68. Rugulopteryx okamurae habit and transection 142

69. Spatoglossum lanceolatum, part of type specimen 143

70. Spatoglossum subflabellatum habit 144

71. Desmarestia filamentosa habit, specimen from the type 

locality 147

72. Desmarestia filamentosa, apex of branch and portion with 

developing opposite lateral branches 148

73. Desmarestia munda subsp. mexicana, main axis with lateral 

blades and primary blade with secondary blades 149

74. Sporochnus anomalus habit and branches with numerous 

stalked, tufted branchlets 151

75. Sporochnus anomalus branch with stalked receptacles and 

tufts of filaments 152

76. Sporochnus neushulii sp. nov. holotype: habit, tufted 

branchlets, and receptacles 154

77. Hapterophycus anastomosans holotype 157

78. Ralfsia hancockii, R. confusa, and R. pacifica 159

79. Cutleria hancockii gametophytes and sporophytes 164

80. Feldmannia irregularis, drawing of type specimen 167

81. Hincksia granulosa and H. bryantii 169

82. Hincksia mitchelliae filaments with sporangia 171

83. Chnoospora pacifica habit and close-ups 173

84. Hecatonema streblonematoides, upper portions of filaments 

with plurilocular gametangia 175

85. Nemacystus brandegeei habit, transection, and close-up 178

86. Streblonema transfixum, uniseriate filaments with terminal 

plurilocular structure 179

87. Ectocarpus acutus, filament portion showing barrel shape 

cells and acute apices 181

88. Ectocarpus commensalis, E. gonodioides, E. hancockii, and

E. sonorensis 183

89. Ectocarpus parvus filament with plurilocular sporangia 185

90. Leathesia nana, several thalli on Sargassum blade, habits, 

and transection 188

91. Colpomenia phaeodactyla and C. ramosa habits 191

92. Colpomenia sinuosa and C. tuberculata habits 192

93. Compsonema serpens, erect and prostrate filaments 195

94. Hydroclathrus clathratus habit 196

95. Rosenvingea floridana habit and portion showing opposite 

branching 198

96. Rosenvingea intricata, compressed tubular branches 

narrowing toward tips 199

97. Sargassum johnstonii branch and habit 203

98. Sargassum johnstonii f. gracile, habit of isotype specimen 204

99. Sargassum lapazeanum habit and close-up 205

100. Sargassum acinacifolium branch and isotype specimen 206

101. Sargassum macdougalii habit and upper portion of axis 208
x • 

102. Sargassum sonorense habit of young thallus 209

103. Sargassum sinicola subsp. sinicola habit 211

104. Sargassum sinicola subsp. sinicola habit 212

105. Comparison of Sargassum sinicola subsp. sinicola with 

S. sinicola subsp. camouii 214

106. Sargassum sinicola subsp. camouii habit, insert of conical 

holdfast 215

107. Sargassum herporhizum habit 216

108. Sargassum herporhizum thallus with basal rhizomes; 

upper portion of branch 217

109. Sargassum brandegeei habits 218
Marine Algae of the Northern Gulf of 

California: Chlorophyta and Phaeophyceae

INTRODUCTION

S

panish explorer Hernán Cortés described an island, “Baja California,” 

in a letter to the King of Spain (1524). Cortés’ account was based on reports of his ships’ captains who were exploring the Pacific Ocean (LeónPortillo, 1972). It was still considered an island 10 years later when 

Cortés himself sailed into the southern region and established a colony at Santa 

Cruz in 1535 (later renamed La Paz by Sebastián Viscaíno in 1596) (Lindsay, 

1983). In 1539 Cortés sent Francisco de Ulloa with three ships from Acapulco 

to explore northward along the west coast of Mexico (León-Portillo, 1995). 

During the expedition Francisco de Ulloa became the first nonnative to circumnavigate what he called the “Mar de Cortés” (Sea of Cortez) (Lindsay, 1983), 

revealing Baja California to be a peninsula and not an island. The Gulf of California has also been called “Mar Bermejo” (Vermilion Sea) for the reddish color 

given by the muddy Río Colorado waters flowing into the upper Gulf; it was not 

named, as generally assumed, for the reddish color of the plankton blooms (van 

Andel and Shor, 1964).

A structurally complex trough reaching nearly oceanic depths, the Gulf of 

California is a large evaporation basin (Roden, 1964) lying between the arid, 

mountainous Baja California peninsula on the west and the dry Sonoran and 

Sinaloan coasts of the Mexican mainland on the east (Figure 1) (Álvarez-Borrego, 

1983, 2002). The Gulf of California is approximately 1400 km in length and 

narrow in width, varying from 150 to 200 km (average width is less than 166 

km), and has a surface area of 258,593 km2

 (=99,843 mi2) (Brusca et al., 2005).

The Gulf is composed of five physiographic zones (Lavín and Marinone, 

2003:figs. 1, 2): (Zone1) The entrance at the Gulf’s southern end, where the 

outer mouth extends from Cabo San Lucas (Baja California Sur) to Cabo Corrientes (Jalisco), is about 204 km wide, more than 3000 m deep, and in open 

communication with the eastern tropical Pacific Ocean. (2) The southern Gulf 

of California begins at the inner mouth from Cabo San Lucas to El Dorado 

(Sinaloa) and extends northward to the southern end of the Midriff Islands (Las 

Islas de la Cintura), and it has more oceanic conditions than zones 3 to 5 to the 

north. (3) The Midriff Islands have narrow channels and sill depths of 300–600 

m (note that Ballenas Channel (Canal de Ballenas) is extraordinarily deep, with 

James N. Norris, Department of Botany, National Museum of Natural History, Smithsonian 

Institution, Washington, D.C., USA; norrisj@

si.edu. Manuscript received 6 May 2008; accepted 31 December 2008.
2 • 

FIGURE 1. Northern Gulf of California. Gemini V photograph taken by astronauts Gordon Cooper and Charles Conrad, 21 August 1965 

(courtesy of NASA).
n u m b e r 9 4 • 3

depths to 1600 m, and the Guaymas Basin, a geologically 

active spreading center with depths of 2000 m and hydrothermal vents, may be considered oceanographic provinces themselves). (4) The northern Gulf of California has 

continental shelf sea characteristics. (5) The upper Gulf of 

California, north of 31°N to the Río Colorado Delta, is 

very shallow, with depths less than 30 m.

For the purpose of this study, the Gulf of California 

was divided into halves near latitude 28°N (Figure 2), separated south of the Islas de la Cintura (Midriff Islands), 

which includes the Gulf’s two largest islands, Isla Ángel 

de la Guarda and Isla Tiburón. The northern Gulf (Figures 1, 2), surveyed in this study, extends eastward from 

Bahía San Francisquito (Baja California) across the Gulf 

to Guaymas (Sonora) and northward to the Río Colorado. 

The area studied herein approximates the “Northern Gulf 

of California Faunal Region” of Brusca et al. (2005:fig. 

9.1) and covers more than 60,000 km2

 (=24,000 mi2). The 

southern Gulf, as referred to herein, extends from Bahía 

San Francisquito across the Gulf to Guaymas and southward to Cabo San Lucas (tip of Baja California Sur) and 

southeast to Cabo Corrientes (Jalisco).

The semi-enclosed Gulf is divided into a series of 

basins and trenches separated from one another by transverse ridges (Shepard, 1950; Rusnak et al., 1964; ÁlvarezBorrego, 1983). Allison (1964) reviewed the geology of 

the areas bordering the Gulf of California, and Lavín and 

Marinone (2003) provided an overview of its unique physical oceanographic features. Brusca (1980) summarized the 

tectonic origin of the Gulf, and more recent studies on the 

tectonic evolution of northwestern Mexico are in Johnson 

et al. (2003). Although earlier estimates of the age of the 

Gulf were up to 10 million years ago (Ma), the origin of 

the Gulf (“proto-Gulf”) began in the late Miocene about 

8 Ma, and by the early Pliocene, about 4 Ma, the Gulf was 

open to the Pacific Ocean (Carreño and Helenes, 2002).

The temperature-moderating effect of the Pacific 

Ocean upon the climate of the Gulf is greatly reduced by 

the series of mountain ranges that run the length of the 

Baja California peninsula (i.e., the Peninsular Ranges: 

Sierra Juárez, Sierra San Pedro Mártir, Tres Virgenes, 

Sierra de la Giganta, and Sierra de la Laguna). The mountains of Baja California, together with the Sierra Madre 

Occidental mountains of the mainland of western Mexico, 

isolate the Gulf’s atmosphere topographically, thus creating a semi-enclosed Gulf in a meteorological as well as an 

oceanic sense (Badan-Dangon et al., 1991; Álvarez-Borrego, 2002). As a result of these barriers, the climate along 

the Gulf coasts of Baja California and Sonora is more 

continental than oceanic (Roden, 1964), with greater 

extremes in temperature, hotter summers, and colder winters than on the Pacific (western) coast of the Baja California peninsula.

The effects of the surrounding deserts and their seasons influence ocean conditions in the northern Gulf and 

have a clear impact on its marine flora. For example, the 

harsh physical extremes in the upper Gulf are a naturally 

occurring seasonal disturbance that can induce algal mortality when conditions are beyond the physiological tolerance of a species. In the upper Gulf the benthic algal 

community is maintained in a subclimax condition by the 

strongly fluctuating environmental conditions (Littler and 

Littler, 1981).

General Oceanographic Feature

the Gulfof Cali

The oceanography of the Gulf of California has been 

described by Roden (1964), van Andel and Shor (1964), 

Maluf (1983), Álvarez-Borrego (1983, 2002), BadanDangon (1998), and Lavín and Marinone (2003). Several 

oceanographic features make the Gulf unique among the 

semi-enclosed seas of similar latitude and size, most notably the strong tidal mixing, some of which is close to deep 

stratification (Lavín and Marinone, 2003). The climate 

of the Gulf of California can be depicted as two seasons: 

a midlatitude winter and a subtropical summer (Mosiño 

and García, 1974). A summary of the general meteorological, hydrographic, and oceanographic features of this 

unique body of water is given herein.

Air Temperature: The mean annual air temperature range increases from about 6°C at Cabo Corrientes 

to 18°C at the Río Colorado Delta. During winter the air 

temperatures are higher near the southern entrance and 

decrease to their lowest at the Gulf’s northern end, while 

during summer the air temperatures shift to being lowest near the Gulf’s southern entrance and highest at the 

northern end. The differences in air temperature between 

the east (Gulf) and west (Pacific) coasts of the Baja California peninsula in summer are large, sometimes over 

10°C, while in winter the differences are small, about 2°C. 

Within the Gulf, air temperatures on its bordering coasts 

are usually the same at the same latitude. The water temperatures, are also generally equal to each other on both 

sides of the Gulf at the same latitude.

Precipitation: Generally, there is more rain on the 

east coast of the Gulf of California (Sonora and Sinaloa) 

than on its west coast (Baja California). With an annual 

rainfall of less than 10 cm, the northern Gulf is drier 

and more influenced by the surrounding desert than the 
4 • 

FIGURE 2. Map of the northern Gulf of California, Mexico (drawn by Alice R. Tangerini, National Museum of Natural History, Smithsonian 

Institution).
n u m b e r 9 4 • 5

southeastern part of the Gulf, where the annual rainfall 

can be greater than 100 cm. From Isla Ángel de la Guarda 

and Isla Tiburón northward, rains occur mostly during 

winter, while south of the Midriff Islands the most rain 

usually occurs from June to October. The amount of rain 

on land increases with altitude, the mountains of the Baja 

California peninsula having an average rainfall of 25–75 

cm/y. In contrast, the rainfall on the coasts is less than 10 

cm/y. However, month-to-month variations can be large, 

and year-to-year variations can change by as much as a 

factor of seven. The number of rainy days decreases from 

an average of 60 days/y at Cabo Corrientes (Jalisco) to 

only 5 days/y along the Baja California and Sonora coasts 

of the central Gulf.

Sediments: The chief source of waterborne sediments to the Gulf was the Río Colorado until the construction of Hoover Dam was completed in 1936. When the Río 

Colorado ran free, this river alone contributed 50% of the 

total sediment brought to the Gulf annually, or about 600 

m3

/s (mean annual). After the construction of Hoover Dam, 

sediment from the Río Colorado dramatically decreased to 

only an average of about 120 m3/s (ranging from about 

11 m3/s in October to 260 m3/s in January). Today, the 

combined diversion of waters by the Hoover Dam, Glenn 

Canyon Dam (completed in 1963), and other smaller dams 

in the United States and Mexico (e.g., Morelos Dam) has 

resulted in virtually no surface freshwater from the Río 

Colorado reaching the Gulf, except during winters with 

abundant rainfall or snowmelt or when water is released 

from the dams. Other rivers that contribute sediment and 

nutrients to the Gulf occur along its eastern coast: Río Concepción, Río Yaqui, and Río Mayo (Sonora); Río Fuerte, 

Río Sinaloa, Río Piaxtla, Río Presidio, and Río Baluarte 

(Sinaloa); and Río Grande de Santiago (Nayarit).

The sediment composition of these rivers is similar, 

being high in sulfate, sodium, and calcium ions, followed 

by bicarbonate, chlorides, and magnesium. Along with 

the waters of the Río Colorado, water from these ecologically important rivers has been increasingly put to use to 

support agriculture, ranching, and urban use by growing 

human populations, so that now little river water bearing sediments reaches the Gulf (Gilmartin and Revelante, 

1978; Álvarez-Borrego, 1983). Dispersion of sediments in 

the northern Gulf is now mostly cross-basinal rather than 

long-basinal, suggesting dispersion is mainly controlled by 

oceanic forcing rather than fluvial processes (Carriquiry 

et al., 2001).

The loss of both sediments and freshwater to the 

upper Gulf has resulted in the loss or degradation of 

wetland, lagoon, and estuarine environments. These are 

critical habitats for aquatic plants, birds, invertebrates, 

and spawning and nursery grounds for commercially 

important shrimp and fish, such as Totoaba macdonaldi Gilbert (Thomson et al., 2000). Habitat loss of their 

spawning and nursery grounds, along with overfishing and 

fishnet capture of juveniles (Guevara-Escamilla, 1973), 

and natural factors have had disastrous effects on totoaba 

populations, a Gulf-endemic species that once supported 

a commercially important fishery (Lercari and Chávez, 

2007). While it is assumed or suspected that the loss of 

river water with its sediments and nutrients has also had 

an effect on the benthic marine flora, there have been no 

quantitative ecological temporal and spatial studies on the 

impact to the marine intertidal and subtidal benthic communities of the upper or northern Gulf.

Evaporation: In the coastal areas of the Gulf the 

mean annual rate of evaporation is 200–250 cm/y, with 

a minimum in winter and a maximum during summer. 

Evaporation is less in the open waters; e.g., it has been 

estimated to be 128 cm/y in the southern Gulf. As water 

evaporates, residual salts are concentrated in the remaining seawater, and as freshwater input and precipitation 

are both extremely low in the northern Gulf, the salinity 

of northern Gulf waters increases.

Salinities: In general, salinities within the Gulf 

of California are 1.0–2.0‰ greater than at comparable 

latitudes outside the Gulf (Bray, 1988a, 1988b). In the 

northern Gulf, where evaporation exceeds precipitation, 

salinities range from 35.3‰ to 37.2‰ (Lavín et al., 1998). 

These increases are related to the Gulf’s geographical isolation from the Pacific Ocean and enhanced evaporation 

and higher water temperatures in the upper Gulf. Salinities 

greater than 36‰ are encountered locally (e.g., the Río 

Colorado Delta is now usually hypersaline, with salinities 

up to 39.0‰ in summer) and in semienclosed and shallow 

esteros (coastal lagoons) and bays of the northern Gulf 

(e.g., Bahía Cholla, Bahía de Adair, Bahía las Ánimas, and 

Bahía San Jorge) and in Bahía Concepción of the southern Gulf. In the upper Gulf, with no freshwater release 

from the Río Colorado, the salinities typically increase 

northwestward, ranging from 35.28‰ in October to over 

38.50‰ in July (Álvarez-Borrego et al., 1973; Lavín et al.,

1998; Álvarez-Borrego, 2002). In the southern third of the 

Gulf the offshore salinity, usually 34.6–35.8‰, is closer 

to that of oceanic waters. However, in the coastal areas 

of the southern Gulf it may be less than 34.6‰ during the 

rainy season, June to October.

Winds: The marked seasonal changes in the prevailing winds in the Gulf result from changes of atmospheric 

pressure in the vicinity of the Gulf and the channeling effect 
6 • 

of the mountain chains on both sides of the Gulf. Winds 

are northwesterly from November to May, typically with 

speeds of 8–12 m/s(meters per second). During the winter 

months, strong winds blow out of the north, sometimes for 

several days at a time, with wind speeds that gust 10–15 

m/s (Bray and Robles, 1991). Southeasterly winds predominate from June to October, with mean speed of about 5 

m/s (Badan-Dangon, 1998; Lavín and Marinone, 2003). 

During summer, the hot desert winds strongly influence the 

coastal marine systems of the upper Gulf.

In the northern Gulf, winds can be extremely variable 

and, as any fisherman or sailor of these waters knows, often 

unpredictable. Near the coast the land and sea breeze system prevails, with diurnal wind changes larger than annual 

ones. Sea breezes reach a maximum velocity in early afternoons and usually calm down after sunset. Land breezes 

are generally weaker, but mountain and valley winds predominate in the northern mountainous coasts.

Storms: Northwest gales occur in the northern 

Gulf from December to February and are particularly 

strong in the Ballenas Channel, between the mountainous Isla Ángel de la Guarda and the Gulf coast of Baja 

California. The southern Gulf has violent, short-duration 

storms, called chubascos, during the rainy season of June 

through October. Some of these may occasionally move 

northward.

Large tropical storms and hurricanes are generated in the eastern North Pacific off the southern coast 

of mainland Mexico, from June to November (mainly in 

September and October). These can either continue west 

or, although not often (Serra, 1971), turn north to northeast and enter the Gulf (Álvarez-Borrego, 1983), where 

they usually dissipate along the Sinaloa and Sonora coasts 

before reaching Isla Tiburón and Isla Ángel de la Guarda. 

However, those that do proceed up the Gulf, occasionally 

as far north as Arizona, can cause much damage and can 

be devastating to humans, but no short-term or long-term 

studies have been done on their ecological effects on the 

Gulf’s marine communities (Álvarez-Borrego, 1983).

Wave Action: The seasonal winds in the Gulf of 

California drive ocean swells that influence the wave intensity. Wave action is characteristically stronger on the north 

sides of the islands and on coastal peninsulas. In general, 

wave action is much reduced on the Gulf of California 

coasts when compared to outer coasts of Pacific Mexico, 

except during large storms, such as chubascos, tropical 

storms, and hurricanes, when waves can be high and damaging, especially when in combination with high tides.

Water Circulation: Beier (1997) developed a model 

that showed a seasonal reversing gyre that forms in the 

northern Gulf (Lavín et al., 1997). Typically, in the northern 

Gulf there is an anticyclonic (clockwise) circulation during 

winter and a cyclonic (counterclockwise) circulation during 

summer (Lepley et al., 1975; see Álvarez-Borrego, 2002:fig. 

3.5), with two transitional periods (Carrillo and PalaciosHernández, 2002). However, there are continuous changes 

throughout the year, and it is not always clearly one or 

the other (Álvarez-Borrego, 2002). The water masses of the 

Gulf of California have been reviewed by Bray and Robles 

(1991) and Badan-Dangon (1998).

Tides: Tides are a mixture of semidiurnal (twice a 

day) and diurnal (once a day) components. In the regions 

of the Gulf’s entrance and in the upper Gulf and, to a 

lesser extent, in the southern Gulf and the northern Gulf, 

the semidiurnal tidal components are dominant, larger 

than the diurnal components. In the central Gulf, around 

Santa Rosalía, the diurnal tidal components dominate 

over the semi-diurnal components (Marinone and Lavín, 

1997). The tidal range increases gradually from the Gulf 

entrance to Isla Tiburón and Isla Ángel de la Guarda, and 

then increases rapidly northward to the Río Colorado.

The lowest tides of the year are from February to 

April, and the highest are from July to September. The time 

of the daily high and low tides gets later as one proceeds 

north from the Gulf’s entrance to the upper Gulf. The time 

difference is approximately 5.5 hours for high water and 

6.0 hours for low water, resulting in high water at one 

end of the Gulf while it is low at the other (Roden, 1964). 

These tidal changes contribute to high water velocities in 

the central Gulf area, with the strongest tidal currents in 

the narrow channels between the large Midriff Islands and 

the adjacent coasts (e.g., Canal de Ballenas and Canal de 

Infiernillo; Figure 2). The speed of the currents is variable, 

dependent on the stage of the moon and prevailing winds. 

Current speeds increase from 3 cm/s at the Gulf’s mouth to 

60 cm/s at the head of the Gulf (Álvarez-Borrego, 2002:fig. 

3.4). Tidal currents are strong in the narrow passages to 

shallow bays and estuaries and are especially notable in 

the narrow channels between the Midriff Islands and Gulf 

coasts, with speeds reported up to 3 m/s (=6 knots) in the 

Ballenas Channel (Álvarez-Sánchez et al., 1984).

The semidiurnal tidal amplitude in the region of the 

Gulf entrance is moderate, about 30 cm. In contrast, the 

tidal amplitude increases dramatically northward. During spring tides, vertical amplitude is greater than 4 m in 

the Midriff Islands, up to 7 m in the northern Gulf, and 

even more pronounced in the upper Gulf, greater than 9 

m (Álvarez-Borrego, 2002). The difference between the 

semidiurnal tide and diurnal tide is also striking (ÁlvarezBorrego, 2002). The amplitude of the diurnal tide slowly 
n u m b e r 9 4 • 7

increases northward to the upper Gulf to about twice that 

of the mouth of the Gulf.

The northern Gulf has some of the greatest tides in 

the world (Brusca et al., 2005). The large vertical amplitude covers and exposes vast areas and is especially pronounced at low tides in the shallow, low-angle sloping, 

mudflats, sandy beaches, and rocky reefs, tide pools, terraces, and platforms.

Currents: At the Gulf entrance (Cabo San Lucas 

to Cabo Corrientes) the surface current is variable. In winter, north of Cabo Corrientes, the wind-driven southward 

currents characterize surface circulation. During summer 

a northward current flowing along the mainland coast 

of western Mexico enters the Gulf at the central to eastern regions of the mouth (entrance) and continues north, 

while a southward current flows along the Gulf coast of 

Baja California (Roden, 1958; Granados-Gallegos and 

Schwartzlose, 1974). Tidal currents predominate north of 

Isla Tiburón. In the narrows between the Midriff Islands 

and adjacent coasts, they are often accompanied by rips 

and eddies,where currents of 1–3 m/s have been measured. 

The largest interannual variations in current patterns in 

the Gulf of California are due to the ENSO (El Niño–

Southern Oscillation) phenomenon (Bray and Robles, 

1991; Lavín and Marinone, 2003).

Net outflow of surface currents at the Gulf entrance is 

–10 cm/s in February, and net inflow is +21 cm/s in August. 

However, compensation must take place or sea level 

changes would take place. How that compensation occurs 

is not fully understood, but there are three hypotheses: (1) 

opposite flow at different levels (depths), (2) opposite flow 

along shores, or (3) a combination of both processes.

Sea Temperature: From April to September , the 

Gulf seawater is warmer than that of Pacific Baja California at 10 m depths, while from October to March their sea 

temperatures at 10 m are about the same.

Within the Gulf, diurnal fluctuations are greater at the 

surface and become more pronounced in the northern portions of the Gulf. The annual range of seawater temperatures 

is about 15°C at Río Colorado, in comparison to about 9°C 

at the Gulf’s southern entrance. In the northern Gulf there 

are marked seasonal changes in coastal seawater temperatures. They are lowest in winter, 8oC–12oC, whereas in summer they are significantly higher, 30oC or more. The coldest 

sea surface temperatures occur where mixing is strongest, 

e.g., Canal de Ballenas-Canal de Salsipuedes (Martínez-Díaz 

de León et al., 2006; Marinone, 2007).

Vertical Distribution of Water Properties: The 

northern Gulf is relatively shallow; it has large seasonal 

and year-to-year variations of temperature and salinity in 

the upper 150 m. The strong tidal components contributing to the vertical mixing, some of it close to deep stratification, make the Gulf unique from other semi-enclosed 

seas (Lavín and Marinone, 2003).

The thermocline in winter is almost isothermal, 

whereas in late summer, it is strongest, with the water temperature difference between the surface and 150 m depth 

approximately 14°C. During spring, salinities are higher 

along the Gulf’s Baja California peninsula side than off 

the coast of Sonora, while in summer the highest salinities 

are in the central Gulf. Oxygen at the surface is 5.5 ml/L 

in April but decreases to 4.5 ml/L in August because of the 

increase in water temperature. At 150 m there is a slight 

salinity increase from spring to summer, despite the temperature increase and strong thermocline (Roden 1964). 

There may be a convective overturn in the upper Gulf (i.e., 

surface density exceeds the density of subsurface layers). 

This is possible because the density increases with decreasing temperature and increasing salinity, and because the 

lowest temperatures and highest salinities occur near the 

coastal regions of the upper Gulf.

Upwelling and Downwelling: Upwelling in the 

Gulf of California occurs along the Mexican mainland 

coast (e.g., Lluch-Cota, 2000) at the same time downwelling is occurring along the Baja California Gulf coast 

and vice versa. The dominant effects of atmospheric 

pressure, wind directions, and temperature cause this 

phenomenon.

The prevailing winds in the Gulf produce upwelling in 

some areas. In summer, southeasterly winds lead to upwelling and cooler temperatures along the Gulf side of the Baja 

California peninsula. The low temperatures along the lee 

side (east coast) of Isla Tiburón in winter are related to 

upwelling, caused by the prevailing northwesterly winds.

General Features f the Marine Flo

the Northern Guf of Ca

The most conspicuous feature of the marine flora of the 

northern Gulf, especially to those familiar with the marine 

flora of the Pacific coast of North America, is the relatively 

smaller size of the algae present. This general impression 

is due mostly to the absence of large brown kelps (Laminariales) and Fucales other than Sargassum. Although the 

marine flora of the northern Gulf is rich in species (over 

430 species recorded), the brown algal species of Sargassum (Fucales) and those of Padina (Dictyotales) compose 

the greatest portion of the biomass of fleshy macroalgae.

Largely influenced by its desert surroundings, the 

uniqueness of the northern Gulf is due, in part, to its 
8 • 

FIGURE 3. Summer marine flora. Intertidal platform with characteristic yellow-brown hue of summer flora, Playa Estación, Puerto Peñasco, 

Sonora, Mexico (July 1973).
n u m b e r 9 4 • 9

FIGURE 4. Winter marine flora. Intertidal platform with the dark brown color of winter flora, Playa Estación, Puerto Peñasco, Sonora, Mexico 

(February 1973).
1 0 • 

continental rather than oceanic climate, with wide air 

temperature variation between summer, 31.2°C–32.8°C 

(88°F–91°F), and winter, 10.0°C–11.1°C (50°F–52°F). 

The intertidal regions of the northern Gulf are subject to 

extreme tidal exposure during the low tide series: up to 4 

m vertical amplitude in the Midriff Islands, as much as 7 

m in Puerto Peñasco, and up to 10 m at the Río Colorado 

Delta. This tidal exposure subjects the intertidal algae and 

sea grasses to severe desiccation, high light intensity (solar 

burn), temperature extremes, and low surf action. The 

northern Gulf has a distinctive and diverse flora because of 

these unique variations of temperatures, tides, and water 

circulation, along with seasonally alternating desert conditions of winter cold and summer heat, which operate 

on a great diversity of protected and exposed shore environments, from estuaries, esteros (lagoons), shallow bays, 

intertidal mudflats, sand/rock, coquina limestone, caliche 

shell-hash, and rocky tidal platforms (Figures 3, 4), sandy 

and rocky shores (Figures 5B, 6), and subtidal reefs and 

sand plains.

While the benthic marine flora of the northern Gulf 

of California is largely subtropical/warm temperate, 

there are also some tropical and temperate elements. A 

high percentage, 19% of the 133 species of northern Gulf 

Chlorophyta and Phaeophyceae, are endemic species (see 

Appendix 2) (about 11% of the greens, 7 of the 63 species, 

and about 27% of the browns, 19 of the 70 species).

Dawson (1941, 1960c) observed that some of the 

prominent benthic algae of the northern Gulf of California 

are species also known in Japan. For example, the Japanese 

brown algae Dictyopteris undulata and Dictyota coriacea

were possibly introduced as viable fragments in ballast 

water of Japanese fishing boats operating in the northern 

Gulf of California in the early 1900s (Dawson, 1960c, as 

“Pachydictyon coriaceum”). Ballast waters from Japan 

would have been dumped into Gulf waters, most likely as 

the catch of shrimp or fish was onloaded. Other species of 

marine algae, such as Gracilaria textorii and Lomentaria 

hakodatensis, with a similar Gulf of California–Japan distribution may have also been unintentionally transported 

by Japanese fishing boats as viable fragments, spores or 

zygotes, and subsequently introduced into the Gulf. It is 

also conceivable that these same boats may have inadvertently transported some Gulf algal species back to Japan 

along with their catch. Without any significant collections 

of the Gulf’s benthic macroalgae prior to 1921 (Setchell 

and Gardner, 1924a; Dawson, 1960c; Norris, 1976b), it 

is difficult to discern if this was the case. However, with 

current molecular analyses and systematic, phylogenetic, 

and phylogeographic studies (e.g., Gavio and Fredericq, 

2002; Gurgel et al., 2004; De Clerck et al., 2006), it is now 

possible to test this hypothesis.

In the upper Gulf there is a strong seasonal turnover 

in species and their relative abundance, resulting in a pronounced alternation of a summer flora with a winter flora. 

At low tide, these seasonal changes are obvious to a novice 

observer who returns to the same locality at different seasons. An example of the marked contrast between the seasons is seen on the tidal platform at Playa Estación, Puerto 

Peñasco, Sonora (Figure 2). The marine flora of this region 

in summer (July) has a yellow-brown appearance (Figure 

3). This color hue is caused by the turf-forming community of the mid intertidal zone, due mostly to Chondrophycus paniculatus (= Laurencia paniculata), which grows 

in association with Porolithon sonorense (a crustose coralline that may be responsible as a cementing agent for 

the rock/shell hash material which comprises the tidal 

platform). In contrast, during winter (February), the tidal 

platform takes on its characteristic dark-brown color (Figure 4) because of the conspicuous ephemeral and annual 

brown algal species that grow, mostly epiphytically on 

and among the Chondrophycus. These are predominately 

the large, coarse, and wart-like Colpomenia tuberculata, 

the more delicate and smooth, but often convoluted, C. 

sinuosa (Figure 5A), the brown finger-like hollow sacs of 

C. phaeodactyla; and the fan-shaped Cutleria hancockii.

Although usually not growing on the top of the Playa 

Estación platform, other perennials, such as species of 

Sargassum and Padina, also contribute to the dark-brown 

winter coloration. Occurring in the numerous tide pools 

(such as the large Cumpleaños Tide Pool, Playa Estación), 

tidal runoff areas, and crevices throughout the platform, 

these brown algae can obtain their largest sizes in winter 

to spring in these habitats. Some of the intertidal macroalgae can persist through the summer in these less-exposed 

habitats off the platform top.

Midriff Island Environments: There are more than 

900 islands and islets in the Gulf of California (Carreño 

and Helenes, 2002; Brusca et al., 2005). The largest are 

the Midriff Islands (Islas de la Cintura) of the central Gulf 

(Figure 2), where large bathymetric gradients in combination with currents and tidal forces cause great volumes of 

water to accelerate in the narrow passages between the 

islands and the nearby coasts. Around the Midriff Islands, 

strong mixing and turbulence occur year round, advecting 

deep cool water with high nutrient concentrations to upper 

water layers, boosting biological productivity (Douglas et 

al., 2007). Isla Tiburón, the largest of the Gulf’s islands, is 

separated from mainland Sonora (Bahía Sargento to Bahía 

Kino) by the Infiernillo Channel (Canal de Infiernillo), and 
n u m b e r 9 4 • 1 1

FIGURE 5. A. Colpomenia sinuosa, low intertidal platform, Playa Estación, Puerto Peñasco, Sonora (February 1974). B. Codium simulans, 

intertidal rock-sand flat at low tide, El Desemboque, Sonora, Mexico (April 1973).
1 2 • 

FIGURE 6. Intertidal zonation, low tide at Puerto Refugio, Isla Ángel de la Guarda, Mexico. High intertidal zone dominated by green algae, 

particularly Ulva species; mid to low intertidal zone dominated by large, dark brown Sargassum johnstonii, with scattered individuals of goldenbrown species of Colpomenia (R/V Dolphin cruise, April 1974).
n u m b e r 9 4 • 1 3

the second largest, Isla Ángel de la Guarda is separated 

from the Gulf coast of Baja California (Punta Bluff to 

Bahía de Los Ángeles), by the Ballenas Channel (Figure 

2). The Ballenas Channel is characterized by strong deepwater currents near deep basins (800–1000 m depths) that 

create conditions resembling persistent upwelling (López 

et al., 2006) and provide sustained nutrient enrichment to 

the shallow upper water layers.

At Puerto Refugio, a protected bay on the north end 

of Isla Ángel de la Guarda (Figure 2), these conditions support abundant seaweed populations that along with other 

primary producers sustain an associated wealth of animal 

populations (e.g., invertebrates, fish, and marine mammals and bird and sea lion rookeries). The algal communities at Puerto Refugio are diverse and luxuriant (Figure 

6), with some of the Gulf’s macroalgal species reaching 

their largest size on the Midriff Islands. While most of the 

northern Gulf’s marine flora is generally considered to be 

warm temperate to subtropical, the cooler, more stable 

subtidal water temperatures associated with the Ballenas 

Channel (López et al., 2006) influence the species composition of the marine flora of the Midriff Islands. For 

example, subtidal brown algal species of Desmarestia and 

Sporochnus, both with cool-temperate to cold-temperate 

affinities, have been found in the Gulf of California only 

in the Midriff Islands (Dawson, 1941, 1944; Norris and 

Bucher, 1976). Deep subtidal populations of Desmarestia

and Sporochnus, although different species than those of 

the northern Gulf, have also been reported in the tropical 

northwest Pacific, off Necker Island (Northwest Hawaiian 

Islands [NWHI], Papaha ̄naumokua ̄kea Marine National 

Monument) by Abbott and Huisman (2003b, 2004).

Graham et al. (2007) presented a predictive model for 

kelp refugia in the tropics by identifying habitats where 

kelp (Phaeophyceae) could survive. While there is no evidence of kelps (i.e., Laminariales) in the northern Gulf, 

certain subtidal habitats of the Midriff Islands host populations of cold-water large brown algae, such as Desmarestia, that are often associated with kelps. Whether these 

are relic or pioneering populations, their island subtidal 

habitats are unique within the Gulf due to their stable 

cool, nutrient-rich waters and may serve as refugia (sensu 

Graham et al., 2007) for these ecologically and geographically isolated brown algae.

Historical Reviw of Marine Botanical Ex

January 1870 marked the collection of the first known 

scientific specimen of a marine alga from the Gulf of California when Wurdemannia miniata (Rhodophyta) was 

obtained by Edward Palmer (McVaugh, 1956; Beaty, 1964; 

Lindsay, 1983), a professional plant collector and an early 

ethnobotanist, on the shore of Isla Carmen, off Loreto 

(Baja California Sur) (Dawson, 1944; Norris, 1976b). 

The next known algal collections, consisting mostly of 

specimens of Sargassum, came from the third expedition 

of the California Academy of Sciences (San Francisco) 

to the Gulf in 1890 (Dawson, 1944; Lindsay and Engstrand, 2002) and were collected by Townshend S. Brandegee (Setchell, 1928) and Walter E. Bryant (Fisher, 1905; 

Godínez-Ortega, 2008). M. Paul Hariot (1895) published 

the earliest account of some Gulf of California algae on 

the basis of a small collection of marine (mostly corallines) 

and freshwater algae obtained by Léon Diguet, a French 

chemical engineer who became interested in the natural 

history and ethnology of Baja California while employed 

at the Boleo mines in Santa Rosalía between 1888 and 

1892 (Lindsay, 1955, 1983). His collections were of such 

interest to the Museum d’Histoire Naturelle (Paris) that he 

returned to Baja California as a field investigator and collector for the Paris museum from 1893 to 1913 (GodínezOrtega, 2008). During his exploration, Diguet spent some 

time in La Paz and on Isla Espiritu Santo investigating the 

pearl fisheries then under French concession and used their 

facilities at La Paz to study the marine fauna.

The first marine algal collections from the northern 

Gulf were made in 1904 by Daniel T. MacDougal (Humphrey, 1961), while he collected for the New York Botanical Garden on a boat trip from Yuma, Arizona, to San 

Felipe, Baja California. Shortly thereafter, drift material 

was gathered in the southern Gulf at La Paz by Gastón L. 

Vives in 1911, which together with the San Felipe collections of MacDougal, were the basis for the first detailed 

study of several Gulf algae by Marshall A. Howe (1911; 

Setchell, 1938). Both of these collectors had species named 

for them by Howe (1911), i.e., Cladophora macdougalii

and Dictyota vivesii and Gracilaria vivesii.

The expedition of the California Academy of Sciences 

to the Gulf of California in 1921 yielded extensive collections during the over 2914.5 km (= 1811 miles) covered by 

the schooner Silver Gate (Slevin, 1923). The marine algal 

specimens were collected by the botanist aboard, Ivan 

M. Johnston (Thomas, 1969), who was then a student 

of W. A. Setchell. These specimens comprised the largest 

part of the monograph of Gulf of California algae (1924a) 

by William A. Setchell (Drouet, 1943; Campbell, 1945; 

Papenfuss, 1976) and Nathaniel L. Gardner (Setchell, 

1937c; Papenfuss, 1976). For Johnston’s efforts, Setchell 

and Gardner (1924a) named nine species for him, including Sargassum johnstonii. Their study also incorporated 
1 4 • 

the 1917 collection of Dr. and Mrs. Daisy R. Marchant 

(Godínez-Ortega, 2008), and the earlier materials of T. 

S. Brandegee and W. E. Bryant, with several new species 

named after those collectors, e.g., Pringsheimia marchantiae, Gonodia marchantiae, Codium brandegeei, Sargassum brandegeei, and Ectocarpus bryantii. In all, 144 

species and varieties of red, brown, and green algae were 

reported, 111 of which were described as new taxa (Setchell and Gardner, 1924a). In a separate publication, Setchell and Gardner (1924b) described another new brown 

alga from the Gulf, Nemacystus brandegeei.

Collections of D. T. MacDougal in 1923 from Puerto 

Libertad, Sonora, resulted in two new species being 

described by Gardner (1927), and later two more, Codium 

macdougalii and Sargassum macdougalii, were described 

by E. Yale Dawson (1944). The 1932 Templeton Crocker 

Expedition of the California Academy of Sciences on the 

yacht Zaca from Isla Guadalupe to the Galápagos yielded 

several specimens from the southern end of the Gulf. The 

algae, collected mostly by John Thomas Howell (Daniel et 

al., 1994), were included in the studies of Setchell (1937b) 

and Dawson (1944). Waldo L. Schmitt (National Museum 

of Natural History, Smithsonian Institution (Chace, 1978) 

obtained algae from Baja California during the Allan Hancock Pacific Expedition of 1933 and, from his dredging 

of the cape region of southern Baja California during the 

Presidential Cruise of 1938. Both collections were later 

treated by William R. Taylor (1939, 1945, respectively; 

Wynne, 1991). The Field Museum (Chicago) Expedition to 

Sonora, made by Francis Drouet (Forest, 1983) and Donald Richards in the winter of 1939, yielded numerous algal 

collections, including many blue-greens, from the Sonoran 

coast. Other upper Gulf specimens were obtained in the 

spring of 1941 by John Poindexter at Puerto Peñasco, 

Sonora. Edward F. Ricketts (Astro, 1976, 2001; Rodger, 

2006) made some algal and sea grass collections from 

Puerto Refugio, Isla Ángel de la Guarda, in April 1940 

(for an excellent account of this Sea of Cortez trip and 

natural history, particularly of the marine fauna, see Steinbeck and Ricketts, 1941; Steinbeck, 2001). During the fall 

of 1940, J. Wyatt Durham brought in some algae while 

aboard the schooner E. W. Scripps on a research expedition of the Geological Society of America and Scripps 

Institution of Oceanography (Anderson et al., 1950).

E. Yale Dawson studied the Gulf of California 

marine algae for his doctoral dissertation (University of 

California, Berkeley, 1942; published 1944) under the 

joint direction of Lee Bonar (UC Berkeley) and Gilbert M. 

Smith (Stanford University; Silva, 1960; Wiggins, 1962). 

All of the previous Gulf of California collections mentioned above, except the earliest of Diguet (Hariot, 1895) 

and those of Schmitt (Taylor, 1939, 1945), were studied 

by Dawson. While a student at UC Berkeley, his phycological interests were encouraged early on and mentored 

by William “Uncle Bill” Setchell. As a boy of 14, Dawson was first introduced to plants of the desert and coast 

during a trip to Baja California with his father in 1933, 

and his interest in cacti (e.g., Setchell and Dawson, 1941) 

and seaweeds persisted throughout his life (Hawkes, 

2005). Dawson’s first sea voyage to the Gulf was aboard 

the M/Y Velero III expedition of the Allan Hancock 

Foundation (University of Southern California) in 1940 

(Garth, 1967), allowing him to be the first marine botanist to visit and collect marine algae of the Gulf of California (Dawson, 1941). After serving in the U.S. Army 

Air Corps during World War II (1943–1945), Dawson, 

as a fellow of the John Simon Guggenheim Foundation 

in 1946–1947, made several overland collecting trips in 

Baja California and along the Mexican mainland from 

Punta Peñasco (Sonora) to Tapachula (Chiapas) and 

obtained additional collections from the Midriff Islands 

while aboard the boat Guanero of Guanos y Fertilizantes 

de México. Beginning in 1948 and continuing to 1954, 

Dawson participated in five other expeditions aboard 

the M/Y Velero IV of the Allan Hancock Foundation. 

While Research Director and Vice President of the Beaudette Foundation (Santa Ynez, California) and, later, as 

Curator of Botany (Los Angeles County Museum), Dawson made other cruises (1957–1964) on the M/Y Stella 

Polaris (Figure 7) that resulted in further knowledge of 

the marine algae of the southern Gulf (Dawson, 1959a). 

Having learned to scuba dive, he became the first marine 

botanist to dive and explore the subtidal waters of the 

Gulf of California and Central America (Figure 8A, B). 

Several friends and biologists, including Carl L. Hubbs, 

Bibiano F. Osorio-Tafall, J. Laurens Barnard, Michael 

Neushul Jr., Robert D. Wildman, Wheeler J. North, and 

Robert W. Hoshaw, made or shared collections with him. 

Among the scuba divers he dove with in Mexico were 

pioneering underwater photographer Don Ollis (Beaudette Foundation, Santa Barbara) and Michael Neushul, 

Robert D. Wildman, Richard H. Rosenblatt, James R. 

Stewart (Scripps Institution of Oceanography), and John 

Bleck (Santa Barbara). Acknowledging their interest and 

appreciating their help, Dawson named species for them, 

e.g., Myriogramme osorioi, Botryocladia neushulii, Kallymenia bleckii, Fauchea hoshawsii, Lejolisia hoshawii, 

and Antithamnion hubbsii.
n u m b e r 9 4 • 1 5

In all, E. Yale Dawson made more than 20 expeditions 

into the Gulf of California from 1940 until his untimely 

death in 1966, with over 50,000 specimens collected 

from 130 different stations during those years (Dawson’s 

field notebooks are deposited in the Phycology Library 

of the Algal Collection of the U.S. National Herbarium, 

National Museum of Natural History, Smithsonian Institution, where he was curator from 1965 to 1966). Dawson’s published accounts of marine algae are numerous, 

with taxonomic, floristic, and distributional studies on the 

algae from the Gulf of California and the Pacific Coast of 

Mexico (see Abbott, 1966; Zaneveld, 1966; Garth, 1967; 

Silva, 1967; Norris, 1976b). Among his major contributions to Mexican phycology, resulting from more than 

25 years of research, are “Marine Algae of the Gulf of 

California” (Dawson, 1944) and eight monographs,: 

“Marine Red Algae of Pacific Mexico” (Dawson, 1953a, 

1954c, 1960b, 1961a, 1962a, 1963a, 1963b). Of the studies published after his death, two concerned Gulf marine 

algae: an annotated list of marine algae of Puerto Peñasco, 

Sonora (Dawson, 1966a), and new species and distribution records (Dawson, 1966b).

Contributions to our knowledge of the marine algae of 

the Gulf of California from 1965 to 1986 were furthered 

FIGURE 7. E. Yale Dawson (left) with Palmer T. Beaudette (right) 

during an expedition to Pacific Mexico and Central America in 

1959; note the M/V Stella Polaris anchored in the background.

FIGURE 8. E. Yale Dawson (A) preparing for algal collecting dives 

using scuba during the 1958 cruise of the M/V Stella Polaris and (B) 

diving and collecting subtidal algae in 1959 in Costa Rica. (Photos 

by Don Ollis, formerly of Beaudette Foundation for Biological Research [now defunct]; photos now part of Dawson papers collection 

at NMNH Department of Botany).
1 6 • 

by the collections of Robert W. Hoshaw (University Arizona, Tucson) and his students, particularly, Arthur E. 

Dennis, Nichols P. Yensen, and Richard M. McCourt. 

Marine flowering plants (sea grasses) were recorded from 

the Sonoran coast by Steinbeck and Ricketts (1941), Dawson (1960c), and Norris (1985c). The ecological and natural history knowledge of the Seri Indians (Comcáac) has 

contributed to our understanding the plants and animals 

of coastal Sonora; e.g., their use of the eel grass seed has 

added to our knowledge of distribution and ecology of 

Zostera marina in the Gulf (Felger and Moser, 1973), sea 

turtle temporal distribution (Felger et al., 1976), and Seri 

ethnobotany (Felger and Moser, 1985; Norris, 1985).

Professor Laura Huerta-Múzquiz (Escuela Nacional 

de Ciencias Biológicas, Instituto Politécnico Nacional; 

Calderón de Rzedowski and Rzedowski-Rotter, 2000) 

began studying Gulf and Baja California algae in the late 

1950s (e.g., Huerta-Múzquiz, 1960), and her students, 

more recently Catalina Mendoza-Gonzalez and Luz Elena 

Mateo-Cid, are among those who have continued to make 

contributions to our knowledge of Gulf of California and 

Pacific Mexico algae. In the 1980s, Professor Marta M. 

Ortega (Universidad Nacional Autónoma de México) and 

her students also worked on algae of Sonora and Sinaloa, 

although focusing primarily on marine algae of the Gulf of 

Mexico and the Caribbean Sea (e.g., Ortega et al., 2001).

The 1980s also marked the beginning of phycological 

research centered at Universidad Autónoma de Baja California, Ensenada, with Luis E. Aguilar-Rosas and Raúl 

Aguilar-Rosas and marine botanists Marcos A. AguilarRosas, Isaí Pacheco-Ruíz, and José A. Zertuche-González, 

along with their students, all making numerous contributions based on their collections and studies of the marine 

algae of Baja California and the Gulf of California (see 

References).

Jorge González-González (Universidad Nacional 

Autónoma de México) and his students, including Daniel León-Alvarez, Hilda León-Tejera, and Elisa ServièreZaragoza, have been adding to our knowledge of Gulf 

of California and Pacific Mexico marine algae with their 

studies, including their useful catalogue of benthic marine 

algae of Mexico (González-González et al., 1996). More 

recently, Jose Espinoza-Ávalos and Rafael RiosemaRodriguez (Universidad Autónoma de Baja California 

Sur), Francisco F. Pedroche (Universidad Autónoma Metropolitana, Unidad Iztapalapa), and others studying the 

Gulf marine algae have continued to further our knowledge of the floristics, taxonomy, and ecology of Gulf algae. 

Today, the marine botanists of Mexico represent an active 

and productive era in Gulf of California phycology (see, 

for example, their numerous publications on Gulf greens 

and browns among the References).

Current Stu

Although my visits to Baja California and the Gulf 

of California via vehicle began in 1963, my first trip as 

a student of marine biology was by sea in 1969 on the 

cruise of the M/Y Makrele (Moss Landing Marine Laboratories of the California State Universities) to the northern Gulf of California (Norris, 1973). During 1972–1975, 

while a graduate student at the University of California, 

Santa Barbara, it was my pleasure to be in residence as 

Station Director of Laboratorio de Biología Marina of 

the University of Arizona and Universidad de Sonora in 

Puerto Peñasco, supported and encouraged by Robert W. 

Hoshaw, Donald A. Thomson, and John R. Hendrickson 

(University of Arizona). Field explorations on the Sonora 

and Baja California coasts were made possible through 

support from National Science Foundation grants (BMS73-07000-A01 and BMS-75-13960) and USDC NOAA04-5-158-20. The Islas de la Cintura were surveyed during 

the 1974 cruise of the R/V Dolphin (Scripps Institution of 

Oceanography) to the northern Gulf of California (Norris 

and Bucher, 1976, 1977). From 1972 to 1979, selected 

localities on the Baja California and Sonora coasts (Figure 2) of the northern Gulf were sampled several times to 

assess seasonal changes of the marine algae. The algal collections from these field trips form the basis for the present 

taxonomic study and were supplemented by the appreciable collections of E. Y. Dawson (US Alg. Coll.; AHFH, 

now UC [see below for list of definitions for abbreviations 

used in this paper]) and those of Ivan Johnston and others that were included in the studies of W. A. Setchell and 

N. L. Gardner (CAS, now UC; US Alg. Coll.).

Specimens were obtained by intertidal collecting and 

subtidally with use of scuba by me and Katina E. Bucher; 

collection numbers designated by the prefix JN- refer to 

our field notebooks (Algal Collection, National Museum 

of Natural History, Smithsonian Institution). Methods 

for field handling of algal collections and their subsequent 

preservation, mounting as herbarium specimens, and preparation for light microscope study are as given by Tsuda 

and Abbott (1985). Photomicrographs were made using an 

Olympus Q-5 digital camera mounted on a Zeiss Universal microscope; all images used herein were prepared using 

Adobe Photoshop PS-3 on a Macintosh G-5 computer.

Specimens studied or examined for this work are deposited in collections at the institutions listed below, which are 

abbreviated for subsequent mention in the text. Herbarium 
n u m b e r 9 4 • 1 7

abbreviations herein follow the Index Herbariorum (Holmgren et al., 1990; Holmgren and Holmgren, 1998, continuously updated, http://sweetgum.nybg.org/ih/).

AHFH Herbarium of the Allan Hancock Foundation, University of Southern California, 

Los Angeles; Algae Collection to LAM 

now UC

CAS Herbarium of the California Academy of 

Sciences, San Francisco; now UC

CMMEX Algae Herbarium of the Universidad 

Autónoma de Baja California, Ensenada, 

Mexico

LAM Herbarium Los Angeles County Museum, 

Los Angeles; Algae Collection now UC

LD Botanical Museum, Lund, Sweden

MICH Herbarium of the University of Michigan, Ann Arbor

PC Muséum National d’Histoire Naturelle, 

Laboratoire de Cryptogamie, Paris

UC Herbarium of the University of California, Berkeley

US Alg. Coll. Algal Collection of the U.S. National Herbarium, National Museum of Natural 

History, Smithsonian Institution, Washington, D.C.

Several botanical and phycological references were 

invaluable throughout this study, notably the second edition of Taxonomic Literature (referred to as TL-2) by 

Stafleu and Cowan (1976–1988) and its supplement volumes (Stafleu and Mennega, 1992–2000; Dorr and Nicolson, 2008), Stafleu (1972) on dates of publication of the 

extensive volumes of Engler and Prantl, Stearn’s (2004) 

Botanical Latin, and the thorough AlgaeBase (http://

www.algaebase.org) developed, updated, and maintained 

by Michael and Wendy Guiry (Guiry and Guiry, 2008). 

The citation of the authors of taxa names follows Brummitt and Powell (1992).

Place and locality names of the land-based collection 

sites and ship-based cruise stations that are used herein 

are from the U.S. National Oceanic and Atmospheric 

Administration (NOAA) charts 21008, 21120, 21124, 

21141, 21161, 21181, 21182, and 21014, Carta de Farod 

e Hidrografía de México F.H. 600, and the boating and 

cruising sea guides to Baja California and the Gulf of California by Lewis (1971), Williams (1996), Cunningham 

(2004, 2006a, 2006b), and Rains (2006).

It is hoped this publication will encourage algal taxonomic and phylogenetic research as it makes available a 

taxonomic baseline for further biodiversity and marine 

botanical and ecological investigations in the northern 

Gulf of California. Accurate identification of the marine 

algae is essential for conservation studies and ecosystem 

management of this unique region. The biological uniqueness of the northern Gulf of California is today internationally recognized, and conservation of its ecosystems 

biodiversity is hopefully provided by the several marine 

biosphere reserves, marine protected areas (for natural 

resources or flora and fauna), marine sanctuaries, and 

national marine parks in the Gulf of California (e.g., Case 

et al., 2002; Cartron et al., 2005a, 2005b; Danemann and 

Ezcurra, 2008).

CHLOROPHYTA

Chlorophyta Pascher, 1914:158

The Chlorophyta (green algae) are an extremely large, diverse group with 

wide variation in vegetative morphologies, ranging from unicellular to structurally complex species. Members of this group are found in freshwater (including 

snow and ice), estuarine, marine, terrestrial, and subaerial (e.g., on trees, walls, 

sloth fur, and polar bear hairs, etc.) habitats. In marine environments, they 

range from microscopic phytoplankton to the benthic microalgae and minute 

to large macroalgae of intertidal and subtidal habitats, down to 210 m depths 

(Littler et al., 1985, 1986; Norris and Olsen, 1991).

The color of these algae varies greatly, from dark green to pale or yellow 

green, because of the dominance of the green photosynthetic pigments, chlorophylls a and b, over the various accessory pigments, such as the carotenes (alpha, 

beta, and gamma) and xanthophylls (lutein, antheraxanthin, neoxanthin, violaxanthin, and zeaxanthin). While most are noncalcified, some marine genera 

are calcified. Cells are either uninucleate or multinucleate, with cellulose found 

in the cell walls in all but a few unicellular species. Chloroplasts vary in size, 

shape, and number per cell but always have two encircling membranes without 

a periplastid endoplasmic reticulum and often contain one or more pyrenoids 

(storage reserves of starch). Thylakoids (structures inside chloroplasts where 

light-dependent reactions of photosynthesis occur) are singular or stacked. Cell 

division occurs with a phragmoplast (cytoskeletal structure instrumental in 

deposition of new cell wall material).

Vegetative reproduction occurs by cell division and fragmentation. Reproduction can be by non-motile aplanospores and sometimes resting spores (akinetes). Swimming cells have two flagella or multiples of two, with a stellate structure 

linking nine pairs of microtubules at the basal transition region. Zoospores are 

usually biflagellate or quadriflagellate, with acronematic (whiplash) type of flagella. Parthenogenesis is also reported in a few species. Sexual reproduction may 

be isogamous or anisogamous, usually with biflagellate gametes, or oogamous.

Remarks. The most recent classification system considers the Chlorophyta to be within the Chloroplastida Adl et al. (2005; Lewis and McCourt, 

PART I: 

GREEN ALGAE
2 0 • 

2004, as “Chlorobionta C. Jeffery 1982”) as members of 

the Archaeplastida; the latter is one of the seven clusters 

that represent the basic groups (or traditional kingdoms) 

of the eukaryotes (Adl et al., 2005).

The benthic green macroalgae found in the Gulf of 

California are in four classes: Ulvophyceae, Cladophorophyceae, Bryopsidophyceae, and Dasycladophyceae.

Key to the Classes of Chlorophyta in the Nort

1a. Algae unicellular or multicellular; cells uninucleate or multinucleate; chloroplasts with pyrenoids; no cytoplasmic streaming within cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1b. Algae coenocytic; a single large multinucleate cell or simple to branched tubes or siphons; chloroplasts with or without 

pyrenoids; with cytoplasmic streaming (observable in living specimens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2a. Cells uninucleate, containing only a single parietal chloroplast with 1 to several pyrenoids. . . . . .

2b. Cells multinucleate, with numerous chloroplasts; in many (but not all) there is a single pyrenoid per chloroplast 

. . . . . . . . . 

3a. Algae siphonous and radially symmetrical, with a central axis that bears lateral branches in whorls; chloroplasts without 

pyrenoids  . . . . . . . .

3b. Algae of various filamentous or vesiculate forms (not radially symmetrical); numerous chloroplasts with or without 

pyrenoids  . . . . . . . .

Ulvophyce

Ulvophyceae Mattox et K. D. Stewart, 1984:41, 66, nom. 

invalid. [see McNeill et al., 2006:Art. 36.2; Pröschold 

and Leliaert, 2007:129]; O’Kelly and Floyd, 1984:123.

Species range from individual to colonial unicellular 

forms to the macrothalli of variable morphologies that 

include (1) multicellular filamentous forms, (2) multicellular monostromatic or distromatic sheets, or (3) multicellular or siphonous forms, mostly of cushion-like clumps 

of entangled or compacted filament-like tubes. Members 

are predominately marine or estuarine and may be simple 

and unbranched or branched. Each cell has a single cupshaped or band-shaped chloroplast that contains one to 

several pyrenoids. Cell division occurs by the furrowing of 

the persistent, closed, centric mitotic spindle.

Life histories, where known, involve an isomorphic or 

heteromorphic alternation of generations. Mobile reproductive cells, occurring in most members, have one or two 

pairs of flagella without mastigonemes (flagellar “hairs”). 

Each flagellum has basal bodies with four microtubular 

roots arranged in a cruciate pattern and smaller rootlets of 

two sizes that alternate between two or more microtubules.

One of the orders, the Ulotrichales Borzi (1895) is present 

in the southern Gulf, based on the report of Ulothrix Kützing 

(1843) by Rocha-Ramírez and Siqueiros-Beltrones (1991).

Three of the orders, the Ctenocladales, Phaeophilales, and Ulvales, are represented in the northern Gulf of 

California.

Key to the Orders of the Ulvophyceae In the 

1a. Algae macroscopic, and either biseriate filaments; or tubular (1 cell layer); or membranaceous and partially tubular and 

partially bladelike (1–2 cell layers); or membranaceous blades throughout (1–4 cells layers)  . . . 

1b. Algae microscopic or minute, uniseriate branched filaments that may be erect or prostrate or prostrate with erect axes; 

uniseriate filaments either remain free and not adjoined throughout or, in some, are partially or entirely consolidated into 

pseudoparenchyma (1 cell thick at margins and 1–4 cells thick centrally) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Algae microscopic to macroscopic, but minute; mostly endophytic or epiphytic on cells of other algae; uniseriate filaments 

free or partially or completely consolidated into pseudoparenchyma, 1–4 cells thick  . . . . . . 

2b. Algae microscopic, uniseriate filaments; mostly endolithic in dead shells, coral rubble, or crustose corallines; 1 cell thick 

throughout, and further separated by zoosporangia formation, and zoospore ultrastructure  . . . . . . 
n u m b e r 9 4 • 2 1

Ctenoc

“Ctenocladales” Silva, 1982:147.

Ctenocladales P. C. Silva in Silva, Meñez et Moe, 1987:131.

Algae are uniseriate, branched filaments, which may 

be (1) entirely erect, (2) entirely prostrate, or (3) a combination of both with an attached prostrate portion from 

which erect branching filaments develop. Prostrate filaments may form a pseudoparenchyma. Cells are uninucleate, with a single chloroplast and usually one or more 

pyrenoids. Unicellular colorless hairs or multicellular setae 

(hair-like) branches are present in some species.

Asexual reproduction is by fragmentation or by akinetes. Sporangia are formed within vegetative cells and 

produce aplanospores, or biflagellate, triflagellate, or 

quadriflagellate zoospores. Sexual reproduction has not 

been confirmed for most species. Where reported, sexual 

reproduction, may be isogamous or anisogamous, with 

biflagellate gametes formed within enlarged vegetative 

cells (gametangia). Sporangia and gametangia may form a 

pore, papilla, or neck.

Remarks. The taxonomy of the members of 

the Ctenocladales and Chaetophorales group has been 

largely based on culture and ultrastructural studies, and 

the ordinal and family placement of their members has 

been variously interpreted. The reliability of certain taxonomic characters has been questioned, and taxonomic 

placement of some genera is still in a state of flux (e.g., 

Silva et al., 1996b). For example, the Ulvellaceae is placed 

in the Ctenocladales herein (following, e.g., Silva et al., 

1996b; Abbott and Huisman, 2004; Pedroche et al., 

2005), but others consider the family in the Ulvales (e.g., 

Schneider and Searles, 1991; Kraft, 2007). At present, 

their taxonomy and phylogenetic relationships are somewhat problematic and in need of further elucidation.

There is one family in the northern Gulf of California.

Ulvel

Ulvellaceae Schmidle, 1899:57, emend. O’Kelly and G. L. 

Floyd, 1983:162.

Algae are usually microscopic, prostrate, and composed of filaments that are freely branched or laterally 

coherent or coalesced to from a pseudoparenchyma, and 

may be, endophytic or epiphytic on other algae or marine 

plants, or grow on other marine organisms. Cells have a 

lateral, band-shaped or platelike chloroplast and are with 

or without pyrenoids. Colorless setae or hair-like projections from the cell wall are present in some genera and 

absent in others.

Sporangia and gametangia are developed in vegetative cells and become larger than vegetative cells. Asexual 

reproduction is usually by biflagellate or quadriflagellate 

zoospores or aplanospores. Sexual reproduction where 

known, is usually by biflagellate isogametes or anisogametes but is in need of investigation for many species.

Remarks. Members are found in brackish and 

marine environments. Species are based on the growth form 

of the filaments (i.e., if the filaments are consolidated into 

pseudoparenchyma versus not consolidated and free), differences in cell size and number of pyrenoids, and habitat and 

host (Nielsen, 1984; Ricker, 1987; Kraft, 2000). O’Kelly 

and Yarish (1981b) suggested detailed features of sporangial 

development may be more reliable taxonomic characters.

Four genera are known in the northern Gulf of 

California.

Key to the Genera of Ulvellaceae in the

1a. Microscopic, prostrate filaments spreading, remaining free of each other throughout (not coalescing into a pseudoparenchyma) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Entocladia (in part)

1b. Microscopic, disc-shaped to spreading; prostrate filaments either entirely coalesced into a pseudoparenchyma, or partially 

coalescing in varying degrees with some filaments free.

2a. Crusts initially a single cell layer, developing to 2 to 4 cell layers (polystromatic) centrally and a single cell layer (monostromatic) outward  . . . 

2b. Crust or sheet-like, a single cell layer thick (monostromatic) throughout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Filaments originating from a central point; forming a more or less circular disc, with or without filaments extending from 

margin  

3b. Filaments not originating from a central point, not circular, but forming a crust with irregular outer margins .

4a. Filaments with setae (hair-like structures without nuclei; not always present in field-collected material) 

. . . . . . . . . . .

4b. Filaments without “hairs” or setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epicladia
2 2 • 

Entocladia Reinke

Entocladia Reinke, 1879:476.

Algae are microscopic prostrate filaments, usually epiphytic on or endophytic within the cell walls of various 

larger algae, sea grasses, or other aquatic vascular plants. 

Filaments are either unconsolidated and irregularly branched 

and free; or may be compacted in varying degrees into a 

pseudoparenchyma. Cells are quadrate to elongated, with a 

single nucleus and chloroplast that contains one to several 

pyrenoids. Hairs or setae-like structures are present (observable in culture, but not always seen in field collections).

Life histories, where known, are with isomorphic sporophytes and gametophytes. Asexual reproduction is by 

quadriflagellate zoospores, formed by sequential division 

in cells near the center of the thallus. Sexual reproduction 

is by biflagellate isogametes formed in the same manner as 

zoospores.

Remarks. Entocladia and Epicladia are recognized herein; the two usually separated by growth form 

and if the cells of the filaments have hairs or setae (Entocladia) or lack them (Epicladia) (see Nielsen, 1980, 1984). 

This generic separation is problematic; e.g., Kraft (2000) 

noted that hairs were rarely (if at all) found in field collections, but they have been shown to occur under culture 

conditions for some species.

Four species of Entocladia and two of Epicladia are 

known in the northern Gulf of California.

5a. Filaments forming pseudoparenchymatous, sheet-like thallus, with outer noncoalesced to loosely coalesced filaments; percurrent axes sometimes visible as primary and lateral branches develop; cells with 1(–3) pyrenoids . . . . . . . . Stromatella

5b. Filaments loosely to tightly coalescing in some portions, with outer short to long filaments remaining free of each other; 

cells with 1–3 (–7) pyrenoids  . . . . . . . . . .

Key to the Species of Entocladia and Epicladia (Ulvellaceae) in 

1a. Prostrate filamentous thallus disc-shaped (more or less circular) in surface view. 

1b. Prostrate filamentous thallus not disc-shaped in surface view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Filaments free of each other (not consolidated into pseudoparenchyma); cells mostly elongated, some irregular in shape, 

3–5 μm wide, 10–40 μm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Entocladia polysiphoniae

2b. Filaments mostly free, sometimes with portions consolidated to varying degrees to form a pseudoparenchyma  

3a. Filaments mostly free, with portions aggregated (i.e., not closely consolidated); cells variable in shape, squarish to rectilinear, most 6–8 μm wide, 6–20 (-35) μm long; 1–3 pyrenoids per cell . . . . . . . . . . . . . . . . . . . . . . . . . . . Entocladia viridis

3b. Filaments extensively consolidated, spreading and covering host cells; margins with extending filaments; 3–7 pyrenoids per 

cell  . . . . . . . 

4a. Disc somewhat circular; inner cells isodiametric, with few to several central cells empty; outer cells narrower, much longer; 

margin border more or less even. . . . . . . . . . . .

4b. Disc irregular in outline, with filaments extending from the margins  

5a. Disc-shaped with only a few, short marginal filaments; central cells 12–16 μm wide; marginal cells 5–9 μm in diameter . .

. . . . . . . . . . .

5b. Discoid, with fringelike margin of numerous longer filaments; central cells 15–18 μm in diameter; marginal cells 5–7 μm in 

diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epicladia mexicana

Entocladia marchantiae

(Setchell et N. L. Gardner) Cri

FIGURE 9A

Pringsheimia marchantiae Setchell et N. L. Gardner, 

1924a:720 [as “P. marchantae”], pl. 12: fig. 8; Dawson, 

1944:205; González-González et al., 1996:138.

Entocladia marchantiae (Setchell et N. L. Gardner) Cribb, 

1995:23, pl. 1: fig. 5; Pedroche et al., 2005:18.

Acrochaete marchantiae (Setchell et N. L. Gardner) 

Nielsen et McLachan, 1986b:516, figs. 3–14.

Pringsheimiella marchantiae (Setchell et N. L. Gardner) 

Schmidt et Petrak in Schmidt, 1935:29 [as “P. marchantae”]; Dawson, 1961b:372; Nielsen and McLachan, 

1985:511; González-González et al., 1996:138.
n u m b e r 9 4 • 2 3

Algae microscopic, more or less discoid, 100–120

(–170) μm in diameter; monostromatic pseudoparenchyma 

with few to several empty central cells (released spores?); 

outer marginal filaments of 1–7 cells, unbranched or 

sparsely, alternately branched; some outer filaments free. 

Cells with a parietal chloroplast and 1(–2) pyrenoids. Central cells isodiametric, (8–)10–20 μm wide. Outer marginal cells narrower and longer (more or less rectangular), 

(2–)3–5 μm wide and (4–)5–10 μm long. Hairs not observed; 

reported in culture studies (Nielsen and McLachan, 1986b).

Habitat. Epiphytic on species of Laurenci

and Gracilaria textorii; low intertidal to shallow subtidal.

Distribution. Gulf of California: Isla Coronado (Isla Smith), off Bahía de Los Ángeles; La Paz

Type Locality. Epiphytic on Laurencia, La

Paz, Baja California Sur, Gulf of California.

Remarks. Entocladia marchantiae from Isla 

Coronado, represents a new record for the northern Gulf 

(JN-4411; US Alg. Coll. slide 4257). Regarding the species epithet, Nielsen and McLachan (1986b) thought it was 

named only for Mrs. Marchant and corrected the spelling 

to “marchantiae.” However, Setchell and Gardner (1924a) 

are rather vague in their species dedication, citing only the 

collection number (“Marchant no. 68x”), then in their 

introduction “a collection made by Dr. and Mrs. Marchant 

in 1917” (p. 696), and later acknowledging “our indebtedness to Dr. and Mrs. Marchant...for collections of marine 

algae” (p. 697). If it can be shown that it was their intention to name the species for both Dr. and Mrs. Marchant, 

then the species name would be correctable to “marchantiorum” (McNeill et al., 2006:Art. 60C.1).

Entocladia polysiphoniae 

Setchell et N. L. Gardn

FIGURE 9B

Entocladia polysiphoniae Setchell et N. L. Gardner, 

1924a:718, pl. 13: fig. 18; Dawson, 1944:204; Dawson, 

1961b:371; Noda, 1968:56, fig. 4; Nielsen, 1972:265; 

Konno and Noda, 1980:27, fig. 2; Espinoza-Ávalos, 

1993:333; González-González et al., 1996:137; Yoshida, 1998:18; Kraft, 2000:518, fig. 3F–H; Pedroche et 

al., 2005:18; Kraft, 2007:31, fig. 11A–E.

Entocladia viridis sensu Mendoza-González and MateoCid, 1999:45, figs. 37, 38 [non Entocladia viridis Reinke, 1879:476].

Algae endophytic, microscopic; composed of irregularly branched filaments, mostly 3–5 μm in width, that 

remain free of each other (not consolidated into a pseudoparenchyma) within cells of the host, and covering an 

area up to 1 mm in diameter. Cells irregularly variable in 

shape, mostly elongated, 10–40 μm in length and 3–5(–9) 

μm in width; parietal chloroplast with a single pyrenoid.

Sporangia (?) intercalary in filaments, up to 24 μm in 

diameter, numerous, formed at irregular intervals.

Habitat. Endophytic in Polysiphonia

Distribution. Gulf of California: El Desemboque to Guaymas; La Paz. Pacific coast: Oaxaca; Japan

Type Locality. Endophytic in cell walls of

Polysiphonia marchantiae (now P. johnstonii), Guaymas, 

Sonora, Gulf of California.

Remarks. In the Gulf, Entocladia polysiphoniae is only known from the type specimen and a few other 

collections. It is distinguished by filaments that remain free 

of each other (never consolidated) and branches that can 

develop, often at right angles, from the middle or toward 

the ends of intercalary cells. Entocladia polysiphoniae has 

also been reported from Japan (Noda, 1968, 1975; Konno 

and Noda, 1980; Yoshida, 1998) and from Lord Howe 

Island, Australia (Kraft, 2000, 2007). Further collections 

of Gulf E. polysiphoniae will be helpful in accessing its life 

history, taxonomic, and phylogenetic status.

Entocladia sp.

FIGURE 9C

Algae microscopic, cells of filaments coalescing forming a spreading, monostromatic pseudoparenchyma; up to 

2 mm across; sometimes completely surrounding cell of 

host Cladophora; cells of outer marginal filaments mostly 

coherent or occasionally with some outermost filaments 

extending distally up the host cell and remaining free. 

Cells with a parietal chloroplast and (2–)3–7 pyrenoids; 

cells mostly irregularly squarish or polygonal, (8–)10–16 

μm in diameter; cells of outer filaments more or less rectangular, elongated, rectilinear, 15–30 μm long and 8–12 

μm wide. Hairs not observed in Gulf specimen.

Habitat. Endophytic within lower cells of

Cladophora; shallow subtidal.

Distribution. Gulf of California: Islas de Los

Gemelos (west island), vicinity of Bahía de Los Ángeles.

Remarks. The northern Gulf specimen of 

Entocladia sp. A (JN-4492; US Alg. Coll. slide 4154) is 

apparently an unknown species. It is somewhat similar to 

Entocladia robustum Cribb (1995; = Endoderma viride

f. major Børgesen, 1920) in cell shape and size and the 

number of pyrenoids per cell. It is also remotely similar to 
2 4 • 

FIGURE 9. Species of Entocladia. A. Entocladia marchantiae: Surface view, disc of consolidated cells, isodiametric in interior (some central cells 

empty), and narrower, elongated cells toward margin; epiphytic on Gracilaria textorii (JN-4411b). B. Entocladia polysiphoniae: Free filaments, 

branching mostly from midregions of intercalary cells; epiphytic on Polysiphonia (JN-5190). C. Entocladia sp. A: Filaments aggregated into 

loose pseudoparenchyma; several pyrenoids per cell; covering cell wall of host Cladophora (JN-4492b). D, E. Entocladia viridis: Free filaments 

extending from aggregated cells; growing horizontally across a basal cell of Chaetomorpha aerea (JN-5180b).
n u m b e r 9 4 • 2 5

Entocladia velutina Cribb (1995) in cell shape and number 

of pyrenoids per cell, but the cell sizes are smaller in the Gulf 

Entocladia sp. A, and the hair cells described in the Australian type (Heron Island, Queensland) were not observed.

Entocladia viridis Reinke

FIGURE 9D,E

Entocladia viridis Reinke, 1879:476, pl. VI: figs. 6–9; 

Setchell and Gardner, 1920b:289; Smith, 1944:35; 

Dawson, 1957a:2; Dawson, 1961b:372; Dawson and 

Neushul, 1966:174; Abbott and Hollenberg, 1976:64, 

fig. 18; O’Kelly and Yarish, 1981a:549, figs. 1–4; 

O’Kelly and Yarish, 1981b:33, figs. 1–12; O’Kelly and 

Floyd, 1983:153, figs. 1–29; Huerta-Múzquiz -Muzquiz 

and Mendoza-González, 1985:42; Ortega et al., 

1987:77, pl. 13: fig. 52; Santelices and Abbott, 1987:5; 

Cribb, 1995:25, pl. 5: figs.1–3; González-González et 

al., 1996:286; León-Tejera and González-González, 

2000:327 [as “E. aff. viridis”]; Kraft, 2000:517, fig. 

3D,E; Abbott and Huisman, 2003a:283, fig. 13; Abbott and Huisman, 2004:43, fig. 4A; Pedroche et al., 

2005:18; Mateo-Cid et al., 2006:58; Kraft, 2007:30, 

fig. 10A–D.

Endoderma viride (Reinke) Lagerheim, 1883:74; Collins, 

1909b:279; Børgesen, 1920:416, figs. 397–399.

Endoderma viride (Reinke) De Toni, 1889:209.

Acrochaete viridis (Reinke) R. Nielsen, 1979:442, fig. 

2A–L; Nielsen and McLachan, 1986a:809; MendozaGonzález et al., 1994:111.

Phaeophila viridis (Reinke) Burrows in Parke and Dixon, 

1976:568.

Algae minute epiphytes, of irregularly branched filaments, mostly 5–8(–10) μm in width, usually remaining free (not coalesced; or slightly coalescing to varying 

degrees) on cell walls of host. Cells variable in shape, inner 

cells squarish to rectilinear, and outer cells elongate, 5–8

(–10) μm in diameter and (5–)10–20(–30) μm long. Cells 

contain a parietal chloroplast with up to 3 pyrenoids.

Habitat. Growing on outer cell walls of Acetabularia, Chaetomorpha, and probably other algae; intertidal to shallow subtidal

Distribution. Gulf of California: Puerto

Peñasco; Estero de Bacorehuis (Laguna Agiabampo) to 

Bahía Balandra (north of La Paz); Playa Los Cerritos (vicinity of Mazatlán), Sinaloa. Pacific coast: British Columbia 

to southern California; Oaxaca; Hawaiian Islands; Rapa 

Nui (Easter Island); Galápagos Islands.

Type Locality. Napoli (Naples), Italy

Remarks. Entocladia viridis may be a species 

complex of more than one taxon (Kylin, 1938; Kraft, 

2000). O’Kelly and Yarish (1981b) also noted Entocladia 

viridis is a polyphyletic assemblage and that specimens 

from California reported by Setchell and Gardner (1920b) 

as “E. viridis” are possibly another species. Reports of 

Entocladia viridis in the Gulf of California (HuertaMúzquiz and Mendoza-González, 1985; Ortega et al., 

1987; Mateo-Cid et al., 2006) are probably this species 

in the sense of Smith (1944) and Abbott and Hollenberg 

(1976).

Epicladia Reinke

Epicladia Reinke, 1889b:31.

Algae are microscopic, uniseriate branched filaments, 

usually growing as endophytes or epiphytes on various 

algae and sea grasses or epizoic on invertebrates; also on 

shells. Most species are prostrate and composed of either 

irregularly branched filaments from the center portion or 

of cells compacted into pseudoparenchyma, usually with 

short to long filaments extending from the margin. Hairs 

and setae are not present. Vegetative cells are uninucleate 

with a single parietal, platelike or band-shaped chloroplast and one or more pyrenoids.

Sporangia are formed by sequential division in vegetative cells near the center of the thallus; pyriform-shaped, 

biflagellate or quadriflagellate zoospores exit through a 

papilla or short tube. Sexual reproduction is by biflagellate 

isogametes formed in a similar manner as the zoospores.

Remarks. Although Epicladia has been considered a synonym of Entocladia by O’Kelly and Yarish 

(1981a, 1981b), Nielsen (1980, 1984) concluded the genera were distinct, with species referred to Epicladia differing primarily from those of Entocladia by the absence of 

hairs or seta in both nature and in culture.

There are two species of Epicladia known from the 

Gulf of California (see “Key to the species of Entocladia

and Epicladia”).

Epicladia condensata (Setchell et N. L. Gardne

J. N. Norris, comb. nov.

FIGURE 10A,B

Entocladia condensata Setchell et N. L. Gardner, 

1924a:718, pl. 12: figs. 4, 5; Dawson, 1944:204; Dawson, 1961b:371; Espinoza-Ávalos, 1993:333; González-
2 6 • 

González et al., 1996:137; Pedroche et al., 2005:17; 

Pacheco-Ruíz et al., 2008:201.

Algae disc-shaped, with compacted cells in the center and a few short filaments extending outward from the 

margin. Hairs or setae absent. Vegetative cells in center, 

angular, 12–16 μm in diameter in surface view; cells of 

short marginal filaments, 7–9 μm in diameter.

Sporangia (?) slightly larger than central vegetative cells.

Habitat. Endophytic within the cell wall of

utricles of various species of Codium; intertidal.

Distribution. Gulf of California: Puerto

Libertad; Bahía San Francisquito; Isla San Esteban.

Type Locality. In utricles of Codium; Bahía

San Francisquito, Baja California, Gulf of California.

Remarks. Apparently a Gulf endemic, Epicladia condensata is found in the utricles of Codium. 

Similarily, Acrochaete codicola (Setchell et N. L. Gardner) O’Kelly (in Gabrielson et al., 2006; basionym: Entocladia codicola Setchell et N. L. Gardner, 1920a, 1920b) 

from the Pacific coast is found in the utricles of Codium 

fragile (Suringar) Hariot (1889). Epicladia condensata is 

disc-shaped, with an irregular margin, and composed of a 

much larger central pseudoparenchyma of cells (12–16 μm 

in diameter) that is more compacted throughout and has 

only a few short marginal filaments of 1–2 angular cells 

(7–9 μm in diameter) (Setchell and Gardner, 1924a), thus 

differing from A. codicola, which has a smaller central 

pseudoparenchyma and numerous, free (not adjoined), 

much longer, branched filaments that are up to at least 

six, long, cylindrical cells (3–4 μm in diameter, 7.5–21 μm 

in length) (see Smith, 1944; Abbott and Hollenberg, 1976; 

both as “Entocladia codicola”).

Epicladia mexicana (Setchell et N. L. Gardne

J. N. Norris, comb. nov.

FIGURE 10C

Entocladia mexicana Setchell et N. L. Gardner, 1924a:719, 

pl. 19: fig. 57; Dawson, 1944:205; Dawson, 1961b:371; 

Espinoza-Ávalos, 1993:333; González-González et al., 

1996:137; Pedroche et al., 2005:18; Pacheco-Ruíz et al., 

2008:201.

Algae endophytic, disc-shaped, 350–450 μm in diameter; pseudoparenchymatous in center, and with numerous 

free filaments around the margin. Cells of pseudoparenchymatous center of thallus 15–18 μm in diameter; cells of marginal filaments, 7.5–17.5 μm long and 5–7 μm in diameter.

Sporangia (?) numerous, scattered, up to 20 μm in 

diameter.

Habitat. Endophytic within Chaetomorpha

antennina and Valoniopsis hancockii; mid to low intertidal.

Distribution. Gulf of California: Puerto

Refugio, Isla Ángel de la Guarda; La Paz.

Type Locality. Endophytic in cell wall

of Chaetomorpha; La Paz, Baja California Sur, Gulf of 

California.

Remarks. Although not well known, Epicladia 

mexicana is apparently an endemic species in the Gulf of 

California.

Stromatella Kornmann et Sahling

Stromatella Kornmann et Sahling, 1985:223.

Stromatella Kornmann et Sahling, 1983:10, nom. inval. 

[note that lacking their selection of a species as the geneFIGURE 10. Species of Epicladia. A, B. Epicladia condensata: A. 

Surface view, endophytic on upper portion of a Codium utricle (after 

Setchell and Gardner, 1924a: pl. 12, fig. 4). B. Longitudinal section 

of a Codium utricle showing E. condensata within the utricle cell 

wall (after Setchell and Gardner, 1924a: pl. 12, fig. 5). C. Epicladia 

mexicana: Surface view, pseudoparenchymatous center with scattered sporangia, and free filaments extending from around the margin (after Setchell and Gardner, 1924a: pl. 19, fig. 57).
n u m b e r 9 4 • 2 7

ritype, the first use of the generic name was invalid (see 

McNeill et al., 2006:Art. 37.1); Stromatella was later 

validly published by Kornmann and Sahling (1985)].

Algae are microscopic filaments, forming monostromatic crusts that grow as prostrate epiphytes on various 

algae. Cells of the filaments coalesce to form a single-layer 

pseudoparenchyma with laterally branched, marginal filaments that are loosely aggregated (not coalescing). Cells 

have a single chloroplast with one pyrenoid.

Reproduction is by the conversion of vegetative cells 

into sporangia and the subsequent division into eight zoospores. Sporangia develop progressively from the interior 

of the pseudoparenchyma outward toward the margin.

Remarks. Although superficially similar to 

Ulvella, Stromatella differs in vegetative anatomy (monostromatic throughout) and reproductive characters (Kornmann 

and Sahling, 1983). Cells of Stromatella have a chloroplast 

with one pyrenoid, whereas in Ulvella there are 0–3 pyrenoids per cell. The taxonomic status of Stromatella was subsequently corroborated by studies of Nielsen (1988b).

There is one species in the northern Gulf of California.

Stromatella monostromatica (P. J. L. Dangea

Kornmann et Sahling

FIGURE 11A,B

Ulvella monostromatica P. J. L. Dangeard, 1965:45, pl. I: 

figs. 8–13, pl. III: fig. 1.

Stromatella monostromatica (P. J. L. Dangeard) Kornmann et Sahling, 1985:223 [footnote]; Nielsen, 

1988a:255, fig. 22a–h; Nielsen, 1988b:427, figs. 1–17; 

Kraft, 2000:516, fig. 3B,C; Abbott and Huisman, 

2003a:283, fig. 12; Abbott and Huisman, 2004:43, fig. 

4B; Kraft, 2007:23, fig. 7A–C.

Stromatella monostromatica (P. J. L. Dangeard) Kornmann et Sahling comb. inval., 1983:10, fig. 4.

Algae epiphytic, monostromatic crusts. Interior of 

compact angular cells, 8–12 μm long and 6–8 μm in width, 

forming a pseudoparenchyma; with filaments at margins 

loose and unconsolidated, laterally branched, of narrow, 

elongated cells, 6–15 μm long and 4–5 μm in width. Cells 

with a single pyrenoid.

Habitat. Epiphytic on Polysiphonia, Bryopsis, and probably other algae; intertidal to shallow

subtidal.

Distribution. Gulf of California: Isla Coronado (Isla Smith) and Isla la Ventana, Bahía de Los Ángeles. Pacific coast: Washington; Hawaiian Islands

Type Locality. Collioure, Pyrénées-Orientales, southern France

Remarks. Apparently a highly variable species 

(Nielsen, 1988a, 1988b), Stromatella monostromatica has 

been reported on many different hosts and from a wide 

range of habitats and geographical localities. The collections from Isla la Ventana (JN-4442b; US Alg. Coll. slide 

4160) and Isla Coronado (JN-4401; US Alg. Coll. slide 

4142), off Bahía de Los Ángeles, are the first records for 

FIGURE 11. Stromatella monostromatica, epiphytic on Polysiphonia (JN-4442b): A. Aggregation of cells forming monostromatic pseudoparenchyma; margins irregular with branching peripherally extending filaments. B. Filaments consolidate forming a single-layer pseudoparenchyma spreading over host; percurrent axes barely discernible.
2 8 • 

the Gulf of California and are similar to thalli reported 

from Hawaii (Abbott and Huisman, 2003a, 2004) and 

Australia (Kraft, 2000, 2007).

Ulvella P. Crouan et H. Cr

Ulvella P. Crouan et H. Crouan, 1859:288.

Algae are microscopic to minute, prostrate, more or 

less circular discs, usually less than 2 mm in diameter, and 

attached directly (without rhizoids) to algae, sea grasses, 

or other substratum. Initially, a monostromatic disc of 

rounded to angular cells in the center with outward radiating, bifurcate filaments. As development continues, the 

central portion of the disc becomes 2–4 cell layers thick, 

while the outer portion remains monostromatic. Cells are 

uninucleate and have a parietal chloroplast with (0–)1–3 

pyrenoids. Hairs are usually not present in field collections 

but may develop under certain culture conditions (Nielsen, 

1977; Notoya, 1983).

Reproduction by quadriflagellate zoopores has been 

reported (Nielsen, 1977).

Remarks. The genera distinction of Ulvella 

and Pseudulvella Wille (1910) has long been considered 

problematic (e.g., Dangeard, 1931), with the nature of the 

zoospores (quadriflagellate in Ulvella and biflagellate in 

Pseudulvella) the only apparent character separating them 

(Papenfuss, 1962; Yarish, 1975). The two were merged 

into a single genus by Nielsen (1977), but later she noted 

that not all of the species of Pseudulvella belong in Ulvella

(Nielsen in Silva et al., 1996a:725). Recent studies of the 

generitype, Pseudulvella americana (Snow) Wille (1910), 

have shown the two to be distinct genera, belonging not 

only to different families but in different classes of green 

algae (Sanchez-Puerta et al., 2006). The taxonomic status of the species of Ulvella in the Gulf of California, as 

well as those on the Pacific coast (Abbott and Hollenberg, 

1976), needs to be critically studied.

Two species of Ulvella are now known in the northern 

Gulf of California.

Key to the species of Ulvella in the nort

1a. Central cells mostly rounded; marginal cells 6–10 μm wide, and <20 μm in length . . . . . . . . . . . . . . . . . . . . . . . . . U. lens

1b. Central cells mostly elongate; marginal cells thinner and longer, 3–5 μm wide, usually >20 μm in length  . . . U. set

Ulvella lens P. Crouan et H. Cr

Figur

Ulvella lens P. Crouan et H. Crouan, 1859:288, pl. 22: fig. 

E; Nielsen, 1977:1, figs. 1–14; Kornmann and Sahling, 

1983:16, Notoya, 1983:9, fig.1; Burrows, 1991:130; 

Hoffman and Santelices, 1997:42, fig. 2 (1–2); Yoshida, 1998:19; Kraft, 2000:514, figs. 2D–G; Abbott 

and Huisman, 2003a:283, fig. 9; Abbott and Huisman, 

2004:44, fig. 4C; Kraft, 2007:25, fig. 8A–D.

Algae minute prostrate epiphytes, disc-shaped, usually 

less than 500 μm in diameter; initially monostromatic, then 

becoming polystromatic, up to 2–3 cell layers, in the central portion; outer portions of 1 cell layer. Cells of central 

portion, rounded to angular, (6–)10–12 μm in diameter; 

outer cells and margin cells, elongate and narrower, 10–18 

μm in length, 6–10 μm in width; a few marginal cells bifid 

(more readily seen in smaller, developing specimens). Cells 

either lacking pyrenoids or with a single pyrenoid.

Sporangia not observed in Gulf specimens (sporangia, 

slightly larger than vegetative cells, develop from central 

cells; zoospores released through a hole in top of sporangium [Nielsen, 1977:1, fig. 12]).

Habitat. Epiphytic on Cladophora and probably on other algae as well and possibly epizoic on mollusk shells; intertidal to shallow subtidal

Distribution. Gulf of California: Puerto

Refugio, Isla Ángel de la Guarda. Pacific coast: Japan; 

Hawaiian Islands; Chile.

Type Locality. Rade de Brest, Finistère,

Atlantic coast of France (Burrows, 1991:130).

Remarks. Ulvella lens is now reported for the 

Gulf of California (EYD-271; US Alg. Coll. slide 4165) 

but is probably more common in the Gulf than the current record indicates. Thalli of northern Gulf U. lens are 

relatively small in diameter but have been reported up to 

2 mm in diameter elsewhere. Although similar to U. setchellii, U. lens differs in having rounded to angular central 

cells and shorter outer cells.

The Pacific coast Ulvella applanata (Setchell et N. L. 

Gardner) G. R. South et Tittley (1986; = Pseudulvella 

applanata Setchell et N. L. Gardner, 1920a; Abbott 

and Hollenberg, 1976) has been considered to be a synonym of U. lens (see Guiry and Guiry, 2008); however, 
n u m b e r 9 4 • 2 9

the taxonomic relationship of these two taxa needs to be 

reinvestigated.

Ulvella setchellii P. J. L. Dange

Figur

Ulvella setchellii Dangeard, 1931:317, figs. 1D,1E, pl. 1; 

Feldmann, 1937:188, fig. 9C; Smith, 1944:38; Hollenberg, 1948:155; Dawson, 1961b:372; Abbott and Hollenberg, 1976:59, fig. 9; Nielsen, 1977:3, figs. 15–21; 

Stewart and Stewart, 1984:140; Stewart, 1991:25; 

González-González et al., 1996:140; Cribb, 1995:28, 

pl. 6: fig. 5, pl. 7: fig. 7; Kraft, 2000:514, fig. 3A; Abbott 

and Huisman, 2003a:283, fig. 10; Abbott and Huisman, 2004:45, fig. 4D; Pedroche et al., 2005:18; Kraft, 

2007:27, fig. 9A–E.

Ulvella lens sensu Setchell and Gardner, 1920b:295, pl. 33; 

Setchell and Gardner, 1930:138; Dawson, 1946b:169; 

Mendoza-González et al., 1994:111 [non Ulvella lens P. 

Crouan et H. Crouan 1859:288].

Algae minute epiphytes, circular to irregularly discshaped, 30–300 μm in diameter; initially monostromatic, 

later becoming polystromatic of 2–4 cell layers in central 

portion; outer portions of 1 cell layer. Cells of central 

portion angular to slightly elongate, mostly 5–8 μm in 

diameter. Marginal cells of Gulf specimens, long and narrow, 12–27 μm long and 3–5 μm wide, most terminating 

with divided tips (appearing Y-shaped, with one side usually longer than the other). Cells uninucleate, chloroplast 

with 2–3 pyrenoids.

Sporangia not observed in Gulf specimens (central 

cells become sporangia and develop conical protuberances 

that function as exit tubes for zoospores [Nielsen, 1977:4, 

fig. 16]).

Habitat. Epiphytic on Cladophora and probably on other algae as well; intertidal

Distribution. Gulf of California: Bahía Agua

Dulce (Bahía Tecomate), Isla Tiburón; Bahía Kino; Playa 

Los Cerritos (vicinity Mazatlán), Sinaloa. Pacific coast: 

British Columbia to Isla Guadalupe (Baja California).

Syntype Localities. Roscoff, Finistère,

France (on Polysiphonia); Croisic, Loire Inférieure, France 

(on Phyllophora); and Pacific Grove, Monterey County, 

California (on Laurenica).

Remarks. Specimens of Ulvella setchellii from 

Bahía Agua Dulce, Isla Tiburón (US Alg. Coll. slide 892), 

epiphytic on Cladophora (EYD-46-980), and Bahía Kino 

(US Alg. Coll. slide 4166), also epiphytic on a Cladophora 

(EYD-2928), represent the first records for the northern 

Gulf of California. Although the cell diameters of Gulf U. 

setchellii and U. setchellii from California (e.g., Abbott 

and Hollenberg, 1976) are in general agreement, the outer 

FIGURE 12. Species of Ulvella. A. Ulvella lens: Immature specimen, nearly circular with some bifurcate cells at disc margin; epiphytic on 

Cladophora (EYD-271). B. Ulvella setchellii: Disc with elongate cells outward from central region and long, narrow, commonly bifurcate marginal cells; epiphytic on Cladophora microcladioides (EYD-46-980).
3 0 • 

marginal cells differ in length; Gulf U. setchellii are not as 

long (only up to 27 μm) compared to those from California (up to 50 μm in length). This may be a reflection of a 

smaller, immature state of the Gulf specimens.

Phaeo

Phaeophilales Chappell, O’Kelly, L. W. Wilcox, et G. L. 

Floyd, 1990:516.

Algae are microscopic, multicellular, and filamentous. 

Vegetative cells contain a single nucleus.

The order is defined on developmental and ultrastructural characters. Reproductive development: initially the 

zoosporangia become multinucleate, and then the zoospores are formed by simultaneous cleavage (O’Kelly and 

Yarish, 1981a). Zoospore ultrastructure: reproduction is 

by scaleless, quadriflagellate zoospores that are formed in 

the zoosporangia. The anterior-most pair of flagellar basal 

bodies in the zoospores has simple, convex terminal caps.

Remarks. On the basis of the ontogeny of the 

zoosporangia and ultrastructural characters of the zoospores of the generitype Phaeophila dendroides, Chappell et al. (1990) removed the genus from the Ulvellaceae

and established a new family and order. Although the 

Phaeophilales and Phaeophilaceae are recognized herein, 

their status is problematic. O’Kelly et al. (2004) noted that 

while their molecular phylogenetic reconstructions supported six lineages, including a Phaeophila lineage, it may 

be premature to interpret their higher level classification, 

including the proposed order and family of Chappell et al.

(1990).

Phaeo

Phaeophilaceae Chappell, O’Kelly, L. W. Wilcox, et G. L. 

Floyd, 1990:516.

Algae are branched uniseriate filaments, with the 

characteristics of the order.

Remarks. The taxonomic status of the family 

is unresolved (O’Kelly et al., 2004; see also Remarks for 

Phaeophilales).

A monotypic family, its single genus is represented in 

the northern Gulf of California.

Phaeophila Hau

Phaeophila Hauck, 1876:56.

Algae minute, endophytic or epiphytic, and composed of branching uniseriate filaments, with cells that 

bear hairs (without a septa separating them from the cell). 

Cells contain parietal chloroplasts with few to several 

pyrenoids.

Zoosporangia can be similar in size to the vegetative cells or slightly larger and are often terminal on the 

branches. The zoospores are quadriflagellate.

Remarks. Although Nielsen (1972) recognized 11 species in Phaeophila (some of which have been 

subsequently transferred to other genera [Nielsen, 1983, 

1988a; Nielsen et al., 1995]), most have considered it to 

be a monotypic genus (e.g., O’Kelly and Yarish, 1981a; 

Chappell et al., 1990; Kitayama and Yoshida, 1992).

More recently, O’Kelly et al. (2004) noted it is unclear 

whether any species other than P. dendroides actually 

belong in Phaeophila. For example, results from culture studies by Moewus (1949) of Ectochaete ramulosa

L. Moewus (1949; = Phaeophila ramulosa (L. Moewus) 

Nielsen) and by Yarish (1975) of Endoderma vagans

Børgesen (1920; = P. vagans (Børgesen) Nielsen) suggest 

both are not Phaeophila but belong to Acrochaete sensu 

Nielsen (1979).

There is one species in the northern Gulf of California.

Phaeophila dendroides

(P. Crouan et H. Crouan) Batt

Ochlochaete dendroides P. Crouan et H. Crouan, 

1852:Vol. 3, No. 346; Crouan and Crouan, 1867:128, 

pl. 8.

Phaeophila dendroides (P. Crouan et H. Crouan) Batters, 

1902:13; Taylor, 1945:47; van den Hoek, 1958:212, 

fig. 6d–f; Dawson, 1961b:371; Nielsen, 1972:255–

257, 261, figs. 2, 3, pl. 3a–f; O’Kelly and Yarish, 

1981a:549, figs. 1, 15–27; Garbary et al., 1985:43; 

Mendoza-González and Mateo-Cid, 1986:419; Nielsen, 

1987:435, figs. 9C–E, 10A–P; Kitayama and Yoshida, 

1992:47, figs. 1–12; Mendoza-González and MateoCid, 1992:23; Servière-Zaragoza et al., 1993:482; 

Cribb, 1995:21, pl. 1: fig. 4; González-González et al., 

1996:288, 372; Yoshida, 1998:21, fig. 1-1F; O’Kelly et 

al., 2004:789–798; Wysor, 2004:214; Pedroche et al., 

2005:19; Mateo-Cid et al., 2006:58; Tribollet et al., 

2006: 292–303; Servière-Zaragoza et al., 2007:7.

Phaeophila engleri Reinke, 1889a:86; Thivy, 1943:245, 

pls. 1, 2; Taylor, 1945:46; Dawson, 1961b:371; 

Earle, 1972:85; León-Tejera and González-González, 

1993:498; González-González et al., 1996:372; Mendoza-González and Mateo-Cid, 1998:24.
n u m b e r 9 4 • 3 1

Minute algae of uniseriate filaments, irregularly alternate or oppositely branched; endophytic or epiphytic, 

endoepilithic or penetrating shells or dead coral. Cells 

elongated, cylindrical, (5–)7–25 μm in diameter and 

15–55 μm long; some may be swollen (subglobose) up to 

25 μm in diameter; with parietal chloroplast with 1–13 

pyrenoids; and wavy setae (hairs), up to 400 μm long and 

(1–)2–5 μm in diameter.

Sporangia 10–40 μm in diameter, with an emission 

tube, 3.5–7.7 μm in diameter and up to 4 times longer 

than diameter of sporangium.

Habitat. In shells, coral rubble, and crustose

corallines; intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas. Pacific coast: British Columbia to 

Washington; Isla Clarión (Islas Revillagigedo); Isla María 

Magdalena (Islas Tres Marías), Nayarit to Oaxaca; Panamá; Hawaiian Islands; Galápagos Islands.

Type Locality. Brest, Finistère, France

Remarks. Phaeophila dendroides is usually 

reported to be within mollusk or barnacle shells and 

crustose corallines. However in the northern Gulf it was 

reported as epiphytic on non-calcareous algae, i.e., on 

“Laurencia” (Mendoza-González and Mateo-Cid, 1986) 

and Chaetomorpha (Mateo-Cid et al., 2006).

Ulva

Ulvales Blackman et Tansley, 1902:136.

Algae are multicellular and consist of biseriate filaments or monostromatic or distromatic thalli that are 

tubular or compressed or form partly to completely flattened blades. Growth is by intercalary cell division. Cells 

are uninucleate, with a single laminate or cup-shaped 

chloroplast and one or more pyrenoids.

Life histories are generally isomorphic with phases of 

morphologically similar asexual and sexual thalli, but a 

few species are known to be heteromorphic, with dissimilar gametophytes and sporophytes. Asexual reproduction 

is by bi- or quadriflagellate zoospores, which are formed 

within cells. Many can also reproduce vegetatively by 

fragmentation. Sexual reproduction may be isogamous, 

anisogamous, or oogamous.

One family is well represented in the northern Gulf of 

California.

Ulva

Ulvaceae Lamouroux ex Dumortier, 1822:71.

Most members of the Ulvaceae are marine species that 

are membranous and exhibit high morphological variability within species, often overlapping between species. They 

may be (1) flat, distromatic (two cells thick throughout; 

Figure 13A), and entire, divided, or branched, with smooth, 

ruffled, or puckered surfaces; (2) tubular to compressed, 

monostromatic (one cell thick throughout; Figure 13B), 

and simple to extensively branched; or (3) with both a flattened portion (two cell layers) and tubular margins and/or 

a tubular basal portion (both one cell layer) (Figure 13C). 

Members are attached to the substratum by a basal disc 

or rhizoidal outgrowths from basal cells of blade margins.

Asexual reproduction is by biflagellate or quadriflagellate zoospores. For most, where the life history is 

known, there is an alternation of isomorphic gametophytes and sporophytes. Sometimes a single phase, either 

gametophytic or sporophytic, is repeated without alternating phases, and for a few the life history is entirely sporophytic. Gametophytes produce biflagellate isogametes or 

anisogametes.

Remarks. Ulva is a cosmopolitan genus 

and widespread in distribution throughout the Gulf of

FIGURE 13. Ulva anatomy, diagrammatic transections: A. Ulva 

sect. ulva: Flat blade of two cell layers throughout. B, C. Ulva sect. 

enteromorpha: B. Cylindrical, hollow, and one cell layer throughout. C. Flattened blade of two cell layers, with hollow, one-cell-layer 

margins. (drawn by Alice R. Tangerini, NMNH)
3 2 • 

California. The reports of Blidingia minima (Nägeli ex 

Kützing) Kylin (1947b) from La Paz (Huerta-Múzquiz 

and Mendoza-González, 1985; Rodríguez-Morales and 

Siqueiros-Beltrones, 1999, both as “E. micrococca”), 

Pacific Baja California (Huerta-Múzquiz, 1978, as “E. 

minima and E. micrococca”), and Oaxaca (HuertaMúzquiz and Tirado-Lizárraga, 1970, as “E. micrococca”) 

and B. marginata (J. Agardh) P. J. L. Dangeard ex Bliding 

(1963) from Isla San Benedicto (Islas Revillagigedo), Oaxaca, and Chiapas (Huerta-Múzquiz and Garza-Barrientos, 

1975, as “E. marginata”) establish another genus, Blidingia Kylin (1947b), in the southern Gulf and Pacific Mexico.

One genus, Ulva, is known in the northern Gulf of 

California.

Ulva Linnae

Ulva Linnaeus, 1753:1163.

Enteromorpha Link in Nees, 1820:5.

Chloropelta Tanner, 1980:129.

Although members are of two basic forms, they can 

exhibit widely variable morphologies. One group, Ulva

sect. ulva, includes flattened blades of two layers (distromatic) of closely adherent cells with gelatinous walls of 

varying thickness (Figure 13A). These blades range from 

narrowly lanceolate to broadly expanded forms and may 

be entire, irregularly lobed, cuneate, or deeply divided, 

with most having a smooth surface (some may have perforations or holes), and smooth, ruffled, or dentate margins. 

The blades are attached to the substratum by a basal holdfast of compacted rhizoidal cells either directly (without 

stipe) or below a stipitate portion, and/or rhizoidal extensions of cells from the lower portions of the blade that 

may run between the two cell layers of the blade. Growth 

is diffuse by cell divisions, primarily along the margins. 

Vegetative cells are uninucleate, with a single laminate or 

cup-shaped chloroplast, and usually one pyrenoid. However, rhizoidal cells are often multinucleate.

Most members of the other group, Ulva sect. enteromorpha, are cylindrical to compressed, hollow, and one 

cell layer (monostromatic) throughout (Figure 13B), while 

a few others may be a flat, distromatic blade with margins and/or a basal portion that are hollow and monostromatic (Figure 13C). These may be unbranched or branched 

(often repeatedly), sometimes with uniseriate or pluriseriate proliferations or branchlets, and are basally attached 

by a system of rhizoidal nonseptate filaments that form a 

holdfast or by rhizoids growing from the lower basal cells. 

These rhizoidal extensions may fill the basal portion of the 

thallus to form a solid stipe. Cells are uninucleate with a 

single (laminate or cup-shaped) chloroplast and one to several pyrenoids; rhizoidal cells are sometimes multinucleate.

Vegetative reproduction can be by fragmentation or 

by developing new upright thalli from basal cells or persistent holdfasts. Life histories are usually an isomorphic 

alternation of generations of a diploid sporophyte and 

haploid gametophytes. Some are capable of parthenogenetic development, and for other species a sexual generation is apparently lacking. All cells of the thallus, except 

some of basal and rhizoidal cells, are capable of becoming 

reproductive. Asexual reproduction is by quadriflagellate 

zoospores or aplanospores produced within the thallus 

cells. Sexual reproduction is by the formation of biflagellate isogametes or anisogametes. In some species the biflagellated swarmers develop parthenogenetically. Spores, 

parthenogenetic gametes, and fertilized gametes initially 

develop into a uniseriate filament that forms a tubular 

monostromatic germling. As the tube develops, it either 

(1) remains one cell layer throughout, or becomes partially 

flattened with or without a hollow basal portion and/or 

hollow margins (Ulva sect. enteromorpha) or (2) becomes 

compressed throughout with the cell walls adhering to 

form a distromatic blade (Ulva sect. ulva).

Remarks. Species of Ulva are found worldwide in marine, estuarine, and freshwater habitats. While 

most are attached, some become unattached and can be 

free-floating or form mats on intertidal mud and sand 

flats. Other unattached species sink and grow in shallow 

to deep subtidal waters (e.g., one species formed extensive 

meadows at 650 m depths in the Hawaiian Islands; Sylvia 

A. Earle, NOAA [currently with Deep Ocean Exploration 

and Research], personal communication).

Species concepts of the “Ulva-Enteromorpha complex” are based on morphology and anatomy and are 

complicated by a high degree of plasticity. Characters 

primarily used to separate the species are (1) morphological, if cylindrical, compressed or flattened, entire or 

branched, and smooth or dentate margins; (2) anatomical, 

one or two layers thick; and (3) cytological, such as cell 

shape and arrangement (regularly or irregularly arranged), 

dimensions and thickness of cell walls (in transections and 

surface views), chloroplast shape, and number of pyrenoids. Many studies have shown these characters to be 

highly variable within species, varying with age, reproductive condition, and influenced by ecological factors such as 

wave exposure, tidal factors, currents, temperature, salinity, and light and biotic factors such as grazing or microorganisms in seawater (e.g., Provasoli and Pintner, 1964, 

1980, 1981; L. Fries, 1975; Bonneau, 1977).
n u m b e r 9 4 • 3 3

Studies of field-collected specimens, combined 

with morphological and reproductive details, cytological developmental patterns in culture, and the apparent 

inability of species to interbreed, have also been used to 

evaluate species concepts (Bliding, 1963, 1968; Koeman 

and van den Hoek, 1981, 1982a, 1982b, 1984; Hoeksema and van den Hoek, 1983; Tanner, 1986; Phillips, 

1988). Originally considered a single genus by Linnaeus 

(1753), it was Link (in Nees, 1820) who divided Ulva and 

described the genus Enteromorpha. Since then, it has long 

been suggested that Enteromorpha and Ulva are closely 

related, perhaps the same genus. Recent molecular analyses support that they are a single genus (Tan et al., 1999; 

Shimada et al., 2003; Hayden et al., 2003; Hayden and 

Waaland, 2004).

Two Asian species, potentially invasive, have been 

recently reported in Pacific Mexico. Ulva fasciata Delile 

(1813), apparently introduced (Mateo-Cid and MendozaGonzález, 2003), is reported from Pacific Baja California 

and Sinaloa to Oaxaca (R. Aguilar-Rosas et al., 2005a). 

A Japanese species, U. pertusa Kjellman (1897), first 

reported in southern California (Hayden and Waaland, 

2004), has now been identified in northwest Baja California (R. Aguilar-Rosas et al., 2008). Hopefully, continued 

monitoring and measures taken, if necessary, will prevent 

their introduction and spread into the Gulf of California.

A broadly expanding species, U. expansa (Setchell) 

Setchell et N. L. Gardner (1920a), has been reported in 

the southern Gulf, from Bahía de La Paz (Howe, 1911, as 

“U. fasciata f. expansa”; Huerta-Múzquiz and MendozaGonzález, 1985; Rocha-Ramírez and Siqueiros-Beltrones, 

1991; Cruz-Ayala et al., 2001; Pedroche et al., 2005).

Twelve species of Ulva, in two separate morphological sections, are present in the northern Gulf of California.

Key to the Species of Ulva in the nort

1a. Thalli flat, linear to rounded blades; 2 cell layers thick throughout [Figure 13A]. . . . . . . . . 

1b. Thalli either: tubular to compressed and hollow and 1 cell layer throughout; or with upper flat or blade-like portion of 2 

cell layers with hollow margins of 1 cell layer, and a tubular hollow basal portion of 1 cell layer [Figure 13B,C] 

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

2a. Blades mostly less than 2 cm tall; in small clumps  . . . . . . . . . . . .

2b. Blades mostly over 5 cm tall; usually not in clumps  

3a. Cells in transection of middle portion of blade, quadrate to only slightly elongate. . . . . .

3b. Cells in transection of middle portion of blade, subquadrate to anticlinally elongate  

4a. Blades digitate, with a lower basal or bicular portion and up to 6 lanceolate segments tapering upward  . . . . U. n

4b. Blades not digitate, either orbicular to wider than tall or linear and much taller than wide, entire or divided, with basal 

portion narrowed toward base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5a. Blades linear, up to 6 times taller than wide, often spirally twisted with ruffled margins. . . . . . . . . U. californica (in part)

5b. Blades not linear; obovate (broadest near upper end) or wider than tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6a. Blades obovate and deeply lobed; margins smooth; transection, thicker in central portion, of cells up to 2 times taller than 

wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. lobata

6b. Blades orbicular (circular and flattened) to wider than tall; margins usually dentate; in transection, more or less same thickness throughout, of elongated cells, up to 3 times taller than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. rigida

7a. Thalli bladelike; flattened upward (2 cells thick), with hollow (1 cell thick) margins and/or tubular basal portions . . . . . .

. . . . 

7b. Thalli tubular to compressed, simple to much branched; hollow and 1 cell layer throughout.

8a. Thalli more or less tubular throughout; axes and branches usually densely covered with numerous short branchlets and 

spinelike projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. acanthophora

8b. Thalli tubular to compressed, simple and unbranched to branched; axes and branches without spiny projections; short 

branchlets, if present, not dense  

9a. Cells in surface view irregularly arranged throughout (not arranged in longitudinal rows in any part of thallus) . . . . . 10

9b. Cells in surface view in distinct to more or less clearly arranged longitudinal rows (often more evident in younger or narrower portions of axes and branches)  .

10a. Thallus tubular throughout (or rarely only slightly compressed); usually not branched (or if branched, only one to few 

simple branches); in transection outer cell wall thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. intestinalis

10b. Thallus compressed, usually widening upward and branched; in transection outer cell wall thick . . . . . . . U. compressa
3 4 • 

Ulva sect. ulva

Members of this section include species of Ulva that 

are flat, bladelike, and distromatic throughout (Figure 

13A). The two cell layers of the blade are developmentally 

independent but closely adherent.

Five of the known species of this section are in the 

northern Gulf of California.

Ulva californica Wille

FIGURE 14A,B

Ulva californica Wille in Collins, Holden and Setchell, 

1899:Exsiccate No. 611; Collins, 1909b:215; Setchell 

and Gardner, 1920b:264; Dawson, 1945c:23; Doty, 

1947:11, pl. 2: figs. 6–10; Dawson, 1960a:31; Dawson, 1961b:374; Abbott and Hollenberg, 1976:78, 

fig. 35; Littler and Littler, 1981:149, 150; Pedroche 

and González-González, 1981:63; González-González, 

1993:443; L. Aguilar-Rosas and Bertsch, 1983:114, 

120; L. Aguilar-Rosas L. et al., 1985:125; HuertaMúzquiz and Mendoza-González, 1985:42; MendozaGonzález and Mateo-Cid, 1985:22; L. Aguilar-Rosas 

and Pacheco-Ruíz, 1986:77; Mendoza-González and 

Mateo-Cid, 1986:419; Tanner, 1986:517, figs. 1–19; 

Sánchez-Rodríguez et al., 1989:39; L. Aguilar-Rosas and 

Pacheco-Ruíz, 1989:81; R. Aguilar-Rosas and Machado-Galindo, 1990:188; Stewart, 1991:27; León-Tejera 

et al., 1993:199; Servière-Zaragoza et al., 1993:482; R. 

Aguilar-Rosas and M. Aguilar-Rosas, 1994:517, 529; 

González-González et al., 1996:139; Núñez-López et 

al., 1998:39; Servière-Zaragoza et al., 1998:169, 177; 

Hayden and Waaland, 2004:378, tbl. 3; Pedroche et al., 

2005:22; Kawai et al., 2007:223–226, figs. 3, 4.

Ulva angusta Setchell et N. L. Gardner, 1920a:283, pls. 

27, 31: fig.1; Setchell and Gardner, 1920b:264, pls. 

22, 26: fig.1; Setchell and Gardner, 1930:137; Dawson, 1944:201; Smith, 1944:45, pl. 4: figs. 1–3; Dawson, 1961b:374; Abbott and Hollenberg, 1976:78, fig. 

34; Silva, 1979:341; L. Aguilar-Rosas and Bertsch, 

1983:114; Stewart and Stewart, 1984:140; L. AguilarRosas et al., 1985:125; Mendoza-González and MateoCid, 1986:419; González-González et al., 1996:139.

Enteromorpha angusta (Setchell et N. L. Gardner) Doty, 

1947:20, pl. 1: figs. 1–6.

Ulva scagelii Chihara, 1968:87, figs. 3–8.

Ulva taeniata sensu Dawson, 1945b:59, 64 [non Ulva taeniata (Setchell) Setchell et N. L. Gardner, 1920a:286].

Algae variable in size and shape in the Gulf: blades 

usually small, up to 2 cm tall and 1.5 cm wide, cuneate 

or more or less triangular-shaped, with undulate margins; 

but blades can also be larger, lanceolate to oblanceolate 

(much taller than wide) up to 15 cm long and to 5 cm 

wide, usually with ruffled margins; attached below by a 

slender stipe and along base of the blade. Two cell layers, 30–40(–60) μm thick in transection; usually thicker 

near base. Cells in surface view polygonal with rounded 

corners, usually irregularly arranged, (5–)10–23 μm in 

diameter by (5–)10–15 μm long; chloroplast with 1–2(–4) 

pyrenoids. Cells in transection more or less quadrate to 

slightly elongate, 5–12 μm wide by 5–18 μm tall.

Quadriflagellate zoospores develop into gametophytes. 

Anisogamous gametes fuse to form a zygote; female gametes capable of developing parthenogenetically; male gametes can also develop without fertilization (Tanner, 1986).

Habitat. On rocks or occasionally epiphytic;

high to low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to La Paz. Pacific coast: Alaska to Baja California 

Sur; Isla Guadalupe; Isla Magdalena; Nayarit to Michoacán; Japan.

Type Locality. Pacific Beach, south of La

Jolla, San Diego County, California.

Remarks. In the northern Gulf, Ulva californica is usually found growing in its smaller, more or less 

triangular-shaped form (cf. Tanner, 1986:514, figs. 1, 4, 

5). On the basis of similarities in morphology, cytology, 

and life histories, the absence of interfertility barriers, and 

preliminary electrophoretic data, two larger Pacific species, 

U. angusta and U. scagelii, were placed in synonymy with 

11a. Thalli cylindrical to slightly compressed; repeatedly and usually densely branched up to several orders; ultimate branchlets multiseriate or partly to entirely uniseriate. . . . . . .

11b. Thalli either: simple and unbranched, or with branches only near base, or a main axis with lateral branches  .

12a. Thallus with lateral branches or proliferations (rarely unbranched); cells in surface view in longitudinal but not transverse rows  . . . . .

12b. Thallus either: unbranched, or with branches only from or near base; cells in surface view in longitudinal and transverse 

rows (although sometimes may be obscure)  . . . . . 
n u m b e r 9 4 • 3 5

U. californica (Tanner, 1986); this has been supported by 

the molecular analyses of Hayden and Waaland (2004). 

The larger thalli of U. californica are not common in the 

northern Gulf but have been reported from Guaymas as 

“U. angusta” (Dawson, 1961b; Mendoza-González and 

Mateo-Cid, 1986).

Ulva lactuca Linnaeu

FIGURE 15

Ulva lactuca Linnaeus, 1753:1163; Setchell and Gardner, 

1920b:265; Collins, 1909b:214; Setchell and Gardner, 

1924a:717; Setchell and Gardner, 1930:138; Dawson, 

1944:202; Smith, 1944:45, pl. 3: figs. 6,7; Dawson, 

1945b:64; Taylor, 1945:42; Dawson, 1959a:5, 8–11; 

Dawson, 1961b:374; Dawson, 1962b:228; Dawson, 

1962c:278; Scagel, 1966:60, pl. 31: figs. I–K; Bliding, 

1968:537, figs. 1–5; Huerta-Múzquiz and Tirado-Lizárraga, 1970:126; Chávez B., 1972:268; Norris, 1973:3, 

17; Abbott and Hollenberg, 1976:83, fig. 39; HuertaMúzquiz , 1978:336, 337, 339; Pedroche and GonzálezGonzález, 1981:63; R. Aguilar-Rosas, 1982:84; L. 

Aguilar-Rosas et al., 1982:52, fig. 8; L. Aguilar-Rosas 

and Bertsch, 1983:115, 120; Huerta-Múzquiz and Mendoza-González, 1985:42; Mendoza-González and Mateo-Cid, 1985:22; Ibarra-Obando and R. Aguilar-Rosas, 

1985:96; L. Aguilar-Rosas and Pacheco-Ruíz, 1986:77; 

Ortega et al., 1987:77, 90, pl. 13: figs. 56, 57; SánchezVargas and Hendrickx, 1987:161; Salcedo-Martínez 

et al., 1988:82; Sánchez-Rodríguez et al., 1989:39; 

Santelices, 1989:243, pl. AA: fig. 4; Dreckmann et 

al., 1990:24, 37; Mendoza-González and Mateo-Cid, 

1992:24; Mateo-Cid and Mendoza-González, 1991:27; 

Rocha-Ramírez and Siqueiros-Beltrones, 1991:30; Martínez-Lozano et al., 1991:23; Mateo-Cid and MendozaGonzález, 1992:24; González-González, 1993:443; Mateo-Cid et al., 1993:51; Stout and Dreckmann, 1993:4; 

León-Tejera et al., 1993:199, 202; Servière-Zaragoza 

et al., 1993:482; Mateo-Cid and Mendoza-González, 

1994a:51, 60; Mateo-Cid and Mendoza-González, 

1994b:34, 44; Mendoza-González et al., 1994:103, 

112; González-González et al., 1996:289; Pacheco-Ruíz 

and Zertuche-González, 1996a:432; Anaya-Reyna and 

Riosmena-Rodríguez, 1996:862; Bucio-Pacheco and 

Dreckmann, 1998:42; Yoshida, 1998:40; MendozaGonzález and Mateo-Cid, 1998:24, 27; RodríguezMorales and Siqueiros-Beltrones, 1999:22; Mateo-Cid 

et al., 2000a:69; Paul-Chávez and Riosmena-Rodríguez, 

2000:146; Cruz-Ayala et al., 2001:190; Pacheco-Ruíz et 

al., 2002:200; Pacheco-Ruíz et al., 2003:447; Hayden 

and Waaland, 2004:373, tbl. 3; Riosmena-Rodríguez 

et al., 2005:101; Hernández-Herrera et al., 2005:146; 

R. Aguilar-Rosas et al., 2005b:36; Pedroche et al., 

2005:28; Mateo-Cid et al., 2006:48; Servière-Zaragoza 

et al., 2007:8; Pacheco-Ruíz et al., 2008:191, 201.

Algae cuneate, ovate to oblanceolate, up to 5 cm tall, 

light green, usually entire, occasionally laciniate, with 

undulate margins; attached by a small discoid holdfast. 

Thickness at margins 40–55 μm; in midportions up to 85 

μm; and in basal region 80–100 μm thick. Cells irregularly arranged in surface view; quadrangular to polygonal, 

12–20 μm by 10–25 μm in diameter, with a single parietal 

chloroplast and 1–2 pyrenoids. Cells in transection, more 

or less quadrate, 15–25(–30) μm tall by 10–25 μm wide.

Habitat. On rocks; high intertidal to shallow

subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Cabo Pulmo. Pacific coast: Alaska to central 

California; northern Baja California to Oaxaca; Costa Rica; 

Ecuador; Galápagos Islands; Peru; Chile; China; Japan.

FIGURE 14. Ulva californica: Habits, small cuneate forms, typically 

found in the northern Gulf. A. EYD-27407. B. JN-5425.
3 6 • 

Type Locality. “In Oceano” (Linnaeus,

1753); probably west coast of Sweden (Womersley, 1984).

Remarks. Ulva lactuca, the type species of the 

genus, has been reported worldwide. In the northwest Gulf 

of California, the effects of an El Niño year (1997–1998) 

on populations of Ulva lactuca were dramatic (PachecoRuíz et al., 2003). Stands of U. lactuca along the Gulf 

coast of Baja California across from Canal de Ballenas 

were significantly reduced in aerial coverage by 70% and 

biomass by 80% during El Niño in comparison to the preEl Niño years (May 1995, 1996) and the post-El Niño 

year (May 2000) recovery.

Ulva lobata (Kützing) Harv

FIGURE 16

Phycoseris lobata Kützing, 1847:54; Kützing, 1849:477; 

Kützing, 1856:10, pl. 27.

Ulva lobata (Kützing) Harvey, 1855:265; Silva et al., 

1996a:748; Silva et al., 1996b;235; Pedroche et al., 

2005:30.

Ulva lobata (Kützing) Setchell et N. L. Gardner, 1920a:284; 

Setchell and Gardner, 1920b:268; Smith, 1944:46, pl. 

4: figs. 4, 5; Taylor, 1945:43; Doty, 1947:10; Dawson, 

1957a:2, 4; Dawson, 1959a:4; Dawson, 1959c:4; Dawson, 1961b:374; Guzmán del Próo et al., 1972:260; Abbott and Hollenberg, 1976:85, fig. 40; Silva, 1979:341; 

Pedroche and González-González, 1981:64; SánchezRodríguez et al., 1989:39; Santelices, 1989:242, pl. AA: 

fig. 6; Rocha-Ramírez and Siqueiros-Beltrones, 1991:30; 

León-Tejera et al., 1993:199; Servière-Zaragoza et al., 

1993:482; González-González et al., 1996:140; Hoffmann and Santelices, 1997:65, fig. 10; León-Alvarez 

et al., 1997:397; Servière-Zaragoza et al., 1998:169; 

Paul-Chávez and Riosmena-Rodríguez, 2000:146; L. 

Aguilar-Rosas et al., 2000:133, 134; Hayden et al., 

2003:279; Hayden and Waaland, 2004:377, tbl. 3.

FIGURE 15. Ulva lactuca: Habit (US Alg. Coll.-36624).
n u m b e r 9 4 • 3 7

Ulva fasciata f. lobata Setchell in Collins, Holden and 

Setchell, 1901b:No. 863; Collins, 1909b:216.

Blades up to 15 cm tall, obovate, mostly deeply lobed 

or divided; margins smooth to ruffled; gradually narrowed 

in lowermost portion, sometimes more or less stipe-like at 

holdfast. Blades in transection 40–50 μm thick at margins 

and noticeably thicker in midportion, up to 90 μm. Cells 

viewed in transection appear vertically elongated, almost 

twice as tall as wide; with a cup-shaped chloroplast.

Habitat. On rocks or mud and sand; mid to

low intertidal.

Distribution. Gulf of California: Santa

Teresa; Bahía de La Paz. Pacific coast: Oregon to Baja 

California; Jalisco to Guerrero; Clipperton Island; Ecuador; Galápagos Islands; Chile; Juan Fernandez Island.

Type Locality. Chile

Remarks. Although Levring (1941, as “Phycoseris lobata”; see also Wynne, 2002) listed U. lobata as 

a synonym of U. nematoidea, molecular analyses have not 

supported this (Hayden and Waaland, 2004), and these 

two are recognized as separate species.

Ulva nematoidea Bory de Saint-Vincen

FIGURE 17A,B

Ulva nematoidea Bory de Saint-Vincent, 1828:190; Levring, 1941:605, figs. 1B–D, pl. 49: fig. 1; Levring, 1960:9; 

Wynne, 2002: figs. 1–3.

Ulva fasciata f. costata M. Howe, 1914:20, pl. 1, pl. 2: figs. 

10–23; Taylor, 1947:60; Dawson et al., 1964:8, pl. 3; 

Acleto O., 1977a:16, fig. 13; Acleto O., 1977b:3, fig. 4.

Ulva costata (M. Howe) Hollenberg, 1971:283, fig. 1, 

nom. illeg. [since it is a later homonym of Ulva costata 

Wollny, 1881:30]; Abbott and Hollenberg, 1976:80, fig. 

36; Santelices and Abbott, 1978:215; Acleto O. 1980:2, 

figs. 1–3; L. Aguilar-Rosas and Bertsch, 1983:114, 120; 

R. Aguilar-Rosas et al., 1984a:151; R. Aguilar-Rosas 

et al., 1984b:159; L. Aguilar-Rosas and Pacheco-Ruíz, 

1985:71; L. Aguilar-Rosas et al., 1985:125; L. AguilarRosas and Pacheco-Ruíz, 1986:77; Acleto O., 1986:38, 

fig. 4; Santelices, 1989:241, pl. AA: fig. 3; R. AguilarRosas and Machado-Galindo, 1990:188; R. Aguilar-Rosas and M. Aguilar-Rosas, 1994:517, 526; 

FIGURE 16. Ulva lobata: Habit (JN-6103a).
3 8 • 

FIGURE 17. Ulva nematoidea: A. Thallus narrowly linear and twisted (EYD-10421). B. Thallus with central planar region and ruffled margins 

(US Alg. Coll.-8799).
n u m b e r 9 4 • 3 9

Mateo-Cid and Mendoza-González, 1994a:51; MateoCid and Mendoza-González, 1994b:34, 44; MendozaGonzález et al., 1994:112; González-González et al., 

1996:139; Hoffman and Santelices, 1997:59, fig. 8; 

Mendoza-González et al., 1999:60; R. Aguilar-Rosas et 

al., 2005b:36.

Ulva dactylifera Setchell et N. L. Gardner, 1920a:285, pl. 

26: fig.1; Setchell and Gardner, 1920b:272, pl. 21: fig.1; 

Setchell and Gardner, 1924a:717; Dawson, 1944:201; 

Dawson, 1959a:6, 11; Dawson, 1961b:374; Abbott 

and Hollenberg, 1976:80, fig. 37; Huerta-Múzquiz, 

1978:339; Pedroche and González-González, 1981:63; 

L. Aguilar-Rosas and Bertsch, 1983:115, 120; GonzálezGonzález, 1993:443; L. Aguilar-Rosas et al., 1985:125; 

Huerta-Múzquiz and Mendoza-González, 1985:42; 

Mendoza-González and Mateo-Cid, 1986:419; Stewart, 1991:28; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:24; Mendoza-González and Mateo-Cid, 1992:23; 

Servière-Zaragoza et al., 1993:482; González-González 

et al., 1996:140; Bucio-Pacheco and Dreckmann, 

1998:42; Rodríguez-Morales and Siqueiros-Beltrones, 

1999:22; L. Aguilar-Rosas et al., 2000:133, 134; PaulChávez and Riosmena-Rodríguez, 2000:146; Pedroche 

et al., 2005:25; Hernández-Herrera et al., 2005:146.

Ulva fasciata f. taeniata Setchell in Collins, Holden and 

Setchell, 1901b: No. 862.

Ulva taeniata (Setchell) Setchell et N. L. Gardner, 

1920a:286, pl. 28; Setchell and Gardner, 1920b:273, pl. 

23; Smith, 1944:48, pl. 3: figs. 1–3; Dawson, 1945b:64; 

Doty, 1947:8, pl. 2: figs. 11, 12, pl. 4: figs. 3, 4; Dawson, 1961b:374; Abbott and Hollenberg, 1976:87, fig. 

43; Santelices and Abbott, 1978:215; Huerta-Múzquiz, 

1978:335; L. Aguilar-Rosas and Pacheco-Ruíz, 1986:77; 

Santelices, 1989:247, pl. AA: figs. 1,2; Rocha-Ramírez 

and Siqueiros-Beltrones, 1991:24; Mateo-Cid and Mendoza-González, 1991:27; Mendoza-González and Mateo-Cid, 1992:24; Servière-Zaragoza et al., 1993:482; 

Mateo-Cid and Mendoza-González, 1994a:51; MateoCid and Mendoza-González, 1994b:34, 44; MendozaGonzález et al., 1994:103, 112; González-González et 

al., 1996:140; Hoffmann and Santelices, 1997:71, fig. 

12; Cruz-Ayala et al., 2001:190; Hayden and Waaland, 

2004:379, tbl. 3; Pedroche et al., 2005:33.

Algae up to 10 cm tall, with upper blade divided into 

4–6 narrow, linear segments, each 5–15 mm wide; often 

twisted, sometimes with a central planar region (midrib), 

and usually with crisped, ruffled margins that may or 

may not be dentate; lower portion of blade orbicular and 

attached along the basal margin or attached by a short 

stipe. Cells in surface view isodiametric, 15–20 μm in 

diameter; in transection (16–)20–40 μm tall by 16–20 μm 

wide. Middle of upper lanceolate segments, up to 190 μm 

thick; with cells in surface view, 12–16 μm in diameter, 

and in transection, elongated, 12–16 wide and 35–60(–80) 

μm tall. Cells of margins in transection, mostly slightly 

elongate, 12–16 μm wide and 12–16 μm tall.

Habitat. On rocks; upper to mid intertidal

Distribution. Gulf of California: Santa

Teresa to Bahía La Paz; Playa Los Cerritos (vicinity Matzalán), Sinaloa. Pacific coast: Oregon to Punta Eugenia, 

Baja California Sur; Sinaloa to Michoacán; Peru; Chile; 

Juan Fernandez Island. (Galápagos Islands and Ecuador 

[Taylor, 1945:43, with a query].)

Type Locality. Concepción, Provincía Concepción, Chile

Remarks. Tanner (1979) and Stewart (1991) 

suggested that U. costata, U. dactylifera, and U. taeniata were conspecific; this has been recently supported 

by molecular evidence (Hayden and Waaland, 2004). 

Although Levring (1941, 1960) considered U. costata

and U. nematoidea to be one species, this has been largely 

ignored. Recently, Wynne (2002) provided morphological 

evidence that these two were the same, with U. nematoidea having nomenclatural priority.

Ulva rigida C. Agar

FIGURE 18

Ulva rigida C. Agardh, 1823:410; Howe, 1914:18; Setchell and Gardner, 1920b:269; Setchell and Gardner, 

1924a:717; Setchell and Gardner, 1930:137; Levring, 1941:606; Dawson, 1944:202; Smith, 1944:47; 

Dawson, 1949:239, 242; Taylor, 1947:60; Papenfuss, 

1960:305, text-fig. 4, pl. 1: fig. 11; Dawson, 1961b:374; 

Dawson et al., 1964:8; Dawson, 1966a:5; Scagel, 

1966:61, pl. 30: figs. A–C, pl. 31: figs. A–H; Bliding, 1968:546, figs. 6–10; Acleto O., 1973:9; Abbott 

and Hollenberg, 1976:87, fig. 41; Huerta-Múzquiz, 

1978:338; Koeman and van den Hoek, 1981:37, figs. 

78–107; Littler and Littler, 1981:151–153; Hoeksema 

and van den Hoek, 1983:74, figs. 20–24; Littler and 

Littler, 1984:15, 24; Mendoza-González and MateoCid, 1985:22; L. Aguilar-Rosas and Pacheco-Ruíz, 

1986:77; Dungan, 1986:295; Phillips, 1988:445, figs. 

24–26; Santelices, 1989:243, pl. AA: fig. 5; R. AguilarRosas and Machado-Galindo, 1990:188; R. AguilarRosas et al., 1990:123; Stewart, 1991:28; Mateo-Cid 

and Mendoza-González, 1991:27; Rocha-Ramírez and 
4 0 • 

Siqueiros-Beltrones, 1991:30; R. Aguilar-Rosas and 

M. Aguilar-Rosas, 1994:517, 526; Mateo-Cid and 

Mendoza-González, 1994b:44; González-González 

et al., 1996:290; Hoffmann and Santelices, 1997:68, 

fig. 11; Mendoza-González and Mateo-Cid, 1998:25; 

Mendoza-González et al., 1999:60; L. Aguilar-Rosas 

et al., 2000:133; Kraft, 2000:532, fig. 11A–E; CruzAyala et al., 2001:190; Abbott and Huisman, 2004:57, 

fig. 12A–D; Hayden and Waaland, 2004:379, tbl. 3; 

Pedroche et al., 2005:32; Mateo-Cid et al., 2006:48; 

Kraft, 2007:46, fig. 19A–E; Pacheco-Ruíz et al., 

2008:191, 201.

Ulva lactuca var. rigida (C. Agardh) Le Jolis, 1863:38; 

Collins, 1903:8, pl. 41: fig.1; Collins, 1909b:215; 

Howe, 1911:490; Huerta-Múzquiz, 1978:335; HuertaMúzquiz and Mendoza-González, 1985:42; PachecoRuíz and Zertuche-González, 1996a:432.

Blades broader than tall, up to 5 cm tall and to 10 

cm wide in the Gulf, dark green; sometimes deeply lobed, 

usually with dentate margins (spines 1–3 cells long); firm 

and comparatively stiff, narrowed abruptly at base to a 

short solid stipe above a small discoid holdfast. Cells in 

surface view irregularly arranged, polygonal to rounded, 

10–15 μm in diameter; with a single chloroplast and 1–3 

pyrenoids. Transection of 2-cell-layer blade, up to 100 μm 

thick (thickest in lowermost portion), with a distinct thick 

hyaline, mucilage layer between cell layers. Cells in transection predominately elongated (most evident in mid to 

lower portions), occasionally squarish, 15–45 μm tall by 

10–15 μm wide.

Habitat. On rocks, usually exposed; high to

low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Bahía La Paz. Pacific coast: Alaska to Colima; 

Peru; Chile; Juan Fernandez Island; Hawaiian Islands.

Type Locality. Cádiz, southern Spain (Papenfuss, 1960)

Remarks. Ulva rigida is distinguished from 

other species of the Gulf by blades that are generally wider 

than tall, darker green in color, and usually with microscopic teeth along the blade margins.

FIGURE 18. Ulva rigida: Habit, showing the wider than tall nature of the thallus (JN-5742).
n u m b e r 9 4 • 4 1

Ulva sect. enteromorpha

(Link in Nees) Endlic

Enteromorpha Link in Nees, 1820:5.

Ulva sect. enteromorpha (Link in Nees) Endlicher, 

1843:19.

Unlike Ulva sect. ulva, thalli of the members of U. sect. 

enteromorpha are completely monostromatic throughout 

or not completely distromatic throughout. This section 

includes species that are either (1) tubular to slightly compressed and only one cell thick (monostromatic) throughout (Figure 13B) or (2) compressed to flattened upward 

to form two-cells-thick (distromatic) blades that have 

hollow, one-cell-thick (monostromatic) margins (Figure 

13C) and/or a one-cell thick (monostromatic) tubular base 

(Figure 13B).

Seven species are known in the northern Gulf of 

California.

Ulva acanthophora (Kützing)

Hayden, Blomster, Maggs, P. C. Sil

Stanhope et Waaland

FIGURE 19A,B

Enteromorpha acanthophora Kützing, 1849:479; Kützing, 1856:pl.34: fig.1; Setchell and Gardner, 1920b:254; 

Setchell and Gardner, 1924a:715, pl. 16: fig. 43, pl. 

38; Dawson, 1944:203; Dawson, 1949:234; Dawson 

et al., 1960b:9, 13; Dawson, 1961b:373; Dawson, 

1962c:278; Huerta-Múzquiz and Tirado-Lizárraga, 

1970:126; Norris, 1973:3; Huerta-Múzquiz, 1978:337, 

338; Huerta-Múzquiz and Mendoza-González, 

1985:42; Mendoza-González and Mateo-Cid, 1985:22; 

Norris, 1985:211; Mendoza-González and Mateo-Cid, 

1986:419; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:30; Martínez-Lozano et al., 1991:23; GonzálezGonzález et al., 1996:136; Pacheco-Ruíz and ZertucheGonzález, 1996a:432; Cruz-Ayala et al., 2001:190.

Ulva acanthophora (Kützing) Hayden, Blomster, Maggs, 

P. C. Silva, Stanhope et Waaland, 2003:288; Pedroche 

et al., 2005:21; Pacheco-Ruíz et al., 2008:201.

Enteromorpha ramulosa var. acanthophora (Kützing) 

Chapman, 1956:419, fig. 67; Paul-Chávez and Riosmena-Rodríguez, 2000:146 [Pedroche et al., 2005:21 

noted this combination was invalid].

Enteromorpha ramulosa sensu Adams, 1994:29, 51, pl. 3 

[non Enteromorpha ramulosa (J. E. Smith) Carmichael 

in W. J. Hooker, 1833:315; basionym: Ulva ramulosa 

J. E. Smith, 1810: pl. 2137, which is a synonym of Ulva 

clathrata (Roth) C. Agardh, 1811:23].

Algae more or less tubular throughout, usually profusely branched, up to 30(–45) cm tall, 1–2 mm in diameter; branches tubular, constricted at base; axes and 

branches beset with numerous short and spinose ultimate 

branchlets. Cells in surface view, mostly 10–15 μm in 

diameter, and not arranged in longitudinal series in main 

axes and branches; but more or less in longitudinal series 

in ultimate branchlets and spinose projections.

Habitat. On rocks and tidal platforms, often

exposed, and in sand in shallow protected waters; mid 

intertidal to shallow subtidal; also dredged 22–36 m 

(Dawson, 1944).

Distribution. Gulf of California: Bahía Tepoca to Bahía La Paz; Isla Espiritu Santo. Pacific coast:

Bahía San Quintín, Baja California; Bahía Tortuga and 

Bahía Almejas (Isla Santa Margarita), Baja California Sur; 

Golfo de Tehuantepec, Oaxaca.

Type Locality. Bay of Islands, northeast

coast of North Island, New Zealand.

Remarks. Within the Gulf of California, Ulva 

acanthophora has been collected in November but appears 

to be more common from February to June. It can be distinguished from other Gulf species by its large size and the 

characteristic, numerous, short branchlets and spine-like 

projections along its axes and branches. The presence of 

the New Zealand species in the Gulf and Pacific Mexico 

shows a very disjunct distribution. It would be interesting to comparatively test Gulf U. acanthophora with the 

New Zealand type and type locality specimens of U. acanthophora (Chapman, 1956; Adams, 1994) using morphological and molecular studies, such as those of Blomster et 

al. (1998, 1999), Hayden et al. (2003), and Hayden and 

Waaland (2004).

Ulva clathrata (Roth) C. Agar

FIGURE 20

Conferva clathrata Roth, 1806:175.

Ulva clathrata (Roth) C. Agardh, 1811:23; Blomster et al., 

1999:579; Pedroche et al., 2005:23; Servière-Zaragoza 

et al., 2007:8; Pacheco-Ruíz et al., 2008:191, 201.

Enteromorpha clathrata (Roth) Greville, 1830:lxvi, 181; 

Collins, 1909b:199; Setchell and Gardner, 1920b:260; 

Dawson, 1959a:7, 10; Dawson, 1961b:373; Dawson, 

1962b:228; Bliding, 1963:107, figs. 64–69; HuertaMúzquiz and Tirado-Lizárraga, 1970:126; Norris, 
4 2 • 

FIGURE 19. Ulva acanthophora: A. Close-up of spiny axes and branches (JN-5455). B. Robust thallus with abundant lateral branches, branchlets, and spines (JN-383).
n u m b e r 9 4 • 4 3

1976a:73, fig. 27; Huerta-Múzquiz, 1978:339; Silva, 

1979:340; R. Aguilar-Rosas, 1982:84; Ibarra-Obando 

and R. Aguilar-Rosas, 1985:96; De la Lanza et al., 

1989:91; Mateo-Cid and Mendoza-González, 1991:26; 

Rocha-Ramírez and Siqueiros-Beltrones, 1991:24; 

Mateo-Cid et al., 1993:51; Servière-Zaragoza et 

al., 1993:482; Mendoza-González et al., 1994:111; 

González-González et al., 1996:283; Riosmena-Rodríguez and Paul-Chávez, 1997:66; Mendoza-González 

and Mateo-Cid, 1998:24; Yoshida, 1998:34; NúñezLópez et al., 1998:38; Rodríguez-Morales and SiqueirosBeltrones, 1999:22; Blomster et al., 1999:575, figs. 2, 

10–16, Mateo-Cid et al., 2000a:69; Paul-Chávez and 

Riosmena-Rodríguez, 2000:146; Cruz-Ayala et al., 

2001:190; Hayden et al., 2003:288, tbl. 4; Abbott and 

Huisman, 2004:46, fig. 5A–C; R. Aguilar-Rosas et al., 

2005b:35.

Enteromorpha crinita Link in Nees, 1820:2 [index]; Dawson, 1945b:22.

Enteromorpha clathrata var. crinita (Link in Nees) 

Hauck, 1884:429; R. Aguilar-Rosas, 1982:85; SánchezRodríguez et al., 1989:39.

Ulva muscoides Clemente y Rubio, 1807:320; Cremades 

and Pérez-Cirera, 1990:490.

Enteromorpha muscoides (Clemente y Rubio) J. Cremades in Cremades and Pérez-Cirera, 1990:489; Silva et 

al., 1996a:737; Núñez-López et al., 1998:38; Blomster 

et al., 1999:575, figs. 1, 3–9, 22–25; Cruz-Ayala et al., 

2001:190.

Enteromorpha plumosa sensu Setchell and Gardner, 

1930:137 [non Enteromorpha plumosa Kützing, 

1843:300; which is now Ulva paradoxa C. Agardh, 

1817:XXII].

Ulva ramulosa J. E. Smith, 1810:pl. 2137.

Enteromorpha ramulosa (J. E. Smith) Carmichael in W. J. 

Hooker, 1833:315; Huerta-Múzquiz and Tirado-Lizárraga, 1970:126; Mendoza-González and Mateo-Cid, 

1998:24.

Algae cylindrical to slightly compressed, up to 10 cm 

tall and 0.5–2.0 mm in diameter, and repeatedly branched, 

with numerous proliferations. Branches and ultimate 

branchlets end in multiseriate or uniseriate rows of cells. 

Cells in surface view, in distinct longitudinal rows in narrower portions, and either similarly ordered or more randomly arranged in broader older portions; cells 8–20 μm 

wide and 8–36 μm long, with a single chloroplast and 1 or 

more pyrenoids.

FIGURE 20. Ulva clathrata: Habit of extensively branched thallus (JN-4994).
4 4 • 

Habitat. On rocks, mudflats and tidal platforms, in crevices, and in tide pools; generally in protected

habitats; high to mid intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Bahía de La Paz; Punta Arena (vicinity of Cabo 

Pulmo). Pacific coast: Alaska to Oaxaca; Isla Clarion (Islas 

Revillagigedo); Isla Clipperton; Hawaiian Islands; Chile; 

China; Japan.

Type Locality. Fehmarn, SW Baltic Sea(Roth,

1806) [neotype locality: Landskoma, Baltic Oresund (neotype illustrated by Bliding, 1963: fig. 69a–d)].

Remarks. Surprisingly, Ulva clathrata has not 

yet been reported from the northern Gulf. Collections in 

the vicinity of Puerto Peñasco (JN-4994; US Alg. Coll.- 

310816) represent a northern extension from Puerto 

Escondido (Dawson, 1959a).

Ulva compressa Linnaeu

Ulva compressa Linnaeus, 1753:1163; Linnaeus, 

1755:433; Blomster et al., 1998:332, figs. 50 (typotype), 55–57; Hayden et al., 2003:289, tbl. 4; MateoCid et al., 2006:48, 58; Pedroche et al., 2005:24; Pacheco-Ruíz et al., 2008:191, 201.

Enteromorpha compressa (Linnaeus) Link ex Nees, 

1820:Index [2]; León-Tejera et al., 1993:199; GonzálezGonzález, 1993:443; Servière-Zaragoza et al., 1993:482; 

Leskinen and Pamilo, 1997:17; Bucio-Pacheco and 

Dreckmann, 1998:42; Yoshida, 1998:34; Paul-Chávez 

and Riosmena-Rodríguez, 2000;144; Abbott and Huisman, 2004:48, fig. 5D.

Enteromorpha compressa (Linnaeus) Greville, 1830:180, 

pl. 18: figs. 1–3; Setchell and Gardner, 1903: 213; Collins, 1909b:201; Setchell and Gardner, 1920b:251, 

pl. 14: figs. 7, 8, pl. 16: fig. 3; Setchell and Gardner, 

1924a:716; Dawson, 1944:203; Dawson, 1949:236; 

Dawson, 1957b:7; Dawson, 1959a:4, 6, 11; Dawson, 1961b:373; Dawson, 1962b:228; Dawson, 

1962c:278; Bliding, 1963:132, figs. 82–84; Dawson, 

1966a:5; Huerta-Múzquiz and Tirado-Lizárraga, 

1970:126; Chávez B., 1972:268; Norris, 1973:3, 17; 

Norris, 1976a:74, fig. 29; Devinny, 1978:358; HuertaMúzquiz, 1978:336; 338; Silva, 1979:340; Pedroche 

and González-González, 1981:63; Koeman and van 

den Hoek, 1982a:288, figs. 2–30; Huerta-Múzquiz and 

Mendoza-González, 1985:42; Mendoza-González and 

Mateo-Cid, 1985:22; Mendoza-González and MateoCid, 1986:419; Dreckmann et al., 1990:24; Mateo-Cid 

and Mendoza-González, 1991:27; Martínez-Lozano 

et al., 1991:23; Mendoza-González and Mateo-Cid, 

1992:23; Mateo-Cid and Mendoza-González, 1992:24; 

Mateo-Cid et al., 1993:51; León-Tejera and GonzálezGonzález, 1993:497; León-Tejera et al., 1993:199; Mateo-Cid and Mendoza-González, 1994a:51; Mateo-Cid 

and Mendoza-González, 1994b:44; Mendoza-González 

et al., 1994:112; González-González et al., 1996:283; 

Mendoza-González and Mateo-Cid, 1996:74, 87, pl. 

23: figs. 102, 103; Blomster et al., 1998:319, figs. 4–6, 

10–15, 30–40, 55–57; L. Aguilar-Rosas et al., 2002:235; 

Dreckmann et al., 2006:153.

Algae mostly compressed, sometimes more or less 

tubular; up to 10 cm tall and mostly to 5 mm wide; upper 

portions usually sparsely branched (rarely unbranched); 

branches more slender near their branching origin and 

usually broadening distally; sometimes with microscopic, 

multiseriate or rarely uniseriate, branchlets; basal portion tapering to holdfast of rhizoidal cells. Cells in surface 

view, more or less quadrate, angular or rounded, 8–15 μm 

in diameter; irregularly arranged throughout; with 1 or 

occasionally 2 pyrenoids.

Habitat. On rocks, usually in exposed habitats; also in mudflats and estuaries, sometimes free-floating, rarely epiphytic; high to mid intertidal (also dredged

4–32 m; Dawson, 1944).

Distribution. Gulf of California: Las Piedras del Burro to Bahía de La Paz. Pacific coast: Alaska to

Costa Rica; Peru; Chile; Hawaiian Islands; China; Japan.

Type Locality. Probably Bognor, Sussex,

England (based on selection of lectotype illustration of 

Dillenius [1742:pl. 9: fig. 8] and typotype [see Stearn, 

1957:129] by Blomster et al. [1998:332, figs. 50, 55–57]).

Ulva flexuosa Wulfen

FIGURE 21A,B

Ulva flexuosa Wulfen, 1803:1; Hayden et al., 2003:289, 

tbl. 4; Pedroche et al., 2005:26; Servière-Zaragoza et 

al., 2007:8.

Conferva flexuosa Roth, 1800:188–190, nom. illeg. [since 

it is a homonym of Conferva flexuosa O. F. Müller, 

1782:5].

Enteromorpha flexuosa (Wulfen) J. Agardh, 1883:126; 

Collins, 1909b:203; Setchell and Gardner, 1920b:255; 

Taylor, 1945:38; Dawson, 1957b:5; Bliding, 1963:73, 

figs. 38–41; Huerta-Múzquiz and Tirado-Lizárraga, 

1970:126; Chávez B., 1972:268; Huerta-Múzquiz 

and Garza Barrientos, 1975:6; Norris, 1976a:76, fig. 

30; Devinny, 1978:358; Silva, 1979:340; Pedroche 
n u m b e r 9 4 • 4 5

and González-González, 1981:63; R. Aguilar-Rosas, 

1982:84; Schnetter and Bula Meyer, 1982:15, pl. 1: figs. 

A–F; L. Aguilar-Rosas et al., 1985:125; Oliva-Martínez 

and Ortega, 1987:127, pl. 3: figs. 4a,b; Ortega et al., 

1987:77, pl. 13: figs. 53–55; L. Aguilar-Rosas and Pacheco-Ruíz, 1989:81; Sánchez-Rodríguez et al., 1989:39; 

De Lara-Isassi et al., 1989:102; Dreckmann et al., 

1990:24; Mateo-Cid and Mendoza-González, 1991:27; 

Mateo-Cid and Mendoza-González, 1992:24; MateoCid et al., 1993:51; González-González, 1993:443; 

Servière-Zaragoza et al., 1993:482; R. Aguilar-Rosas 

and M. Aguilar-Rosas, 1994:517; Mendoza-González 

et al., 1994:112; González-González et al., 1996:284; 

Mendoza-González and Mateo-Cid, 1996:74, 87, 

pl. 24: figs. 104–106; Leskinen and Pamilo, 1997:17; 

Bucio-Pacheco and Dreckmann, 1998:42; Yoshida, 

1998:35; Mendoza-González and Mateo-Cid, 1998:24, 

27; Servière-Zaragoza et al., 1998:169, 180; RodríguezMorales and Siqueiros-Beltrones, 1999:22; López et al., 

2000:340; Cruz-Ayala et al., 2001:190; L. AguilarRosas et al., 2002:235; Abbott and Huisman, 2004:48, 

fig. 6A–C; Wysor, 2004:215; Hernández-Herrera et al., 

2005:146; Dreckmann et al., 2006:153.

Enteromorpha compressa var. flexuosa (Wulfen) Hamel, 

1931a:65, fig. 48 (5,6).

Enteromorpha prolifera var. flexuosa (Wulfen) Doty, 

1947:15; Dawson, 1961b:374.

Enteromorpha lingulata J. Agardh, 1883:143; Setchell and 

Gardner, 1930:137; Taylor, 1945:39; Huerta-Múzquiz 

and Tirado-Lizárraga, 1970:126, 131; Huerta-Múzquiz 

FIGURE 21. Ulva flexuosa (JN-4790): A. Lower portion of small thallus with several lateral branches and rhizoids growing downward toward 

the base. B. Surface view of cell arrangement in regular longitudinal and more or less transverse rows.
4 6 • 

and Garza Barrientos, 1975:6; León-Tejera et al., 

1993:199, 204; Mendoza-González and Mateo-Cid, 

1998:24.

Enteromorpha tubulosa (Kützing) Kützing, 1856:11, 

pl. 32: fig. 2; Dawson, 1945b:59; Taylor, 1945:38, 

“prox.”; Dawson, 1961b:374; Huerta-Múzquiz and 

Tirado-Lizárraga, 1970:126; Chávez B., 1972:268; 

Huerta-Múzquiz, 1978:335, 338; Huerta-Múzquiz and 

Mendoza-González, 1985:42.

Enteromorpha intestinalis var. tubulosa Kützing, 

1845:247.

Algae cylindrical to slightly compressed, up to 10 cm 

tall and 0.5–5.0 mm wide, usually unbranched or sometimes sparsely branched (rarely with secondary branches), 

and without proliferations. Cells in surface view in longitudinal rows and transverse rows (evident in younger, 

narrower portions; not always distinct in older or broader 

portions); angular, quadrate to elongate, 8–15 μm wide by 

10–20 μm long, with a parietal chloroplast and 1–2(–6) 

pyrenoids. Transverse section of 1 cell layer, 20–30 μm 

thick; with cells 12–20(–30) μm tall by 9–20 μm wide.

Habitat. On rocks; high to low intertidal

Distribution. Gulf of California: El Tornillal to La Paz. Pacific coast: British Columbia to Costa

Rica; Panamá; Colombia; Ecuador; Galápagos Islands; 

Chile; Hawaiian Islands; Japan.

Type Locality. Duino (NW of Trieste), Italy,

Adriatic Sea.

Ulva intestinalis Linnae

FIGURE 22

Ulva intestinalis Linnaeus, 1753:1163; Linnaeus, 1755:418; 

Blomster et al., 1998:332, figs. 49 (typotype), 52–54; 

Hayden et al., 2003:289, tbl. 4; Hayden and Waaland, 

2004:377, tbl. 3; Pedroche et al., 2005:27; Mateo-Cid et 

al., 2006:48; Pacheco-Ruíz et al., 2008:191, 201.

Enteromorpha intestinalis (Linnaeus) Link in Nees, 1820:Index [2], 5; Howe, 1911:490; Setchell and Gardner, 

1920b:252; Dawson, 1944:203; Dawson, 1961b:373; 

Dawson, 1962c:278; Bliding, 1963:139, figs. 87–89; 

Dawson, 1966a:5; Huerta-Múzquiz and Tirado-Lizárraga, 1970:126; Brusca and Thomson, 1975:42; Norris, 

1976a:76, fig. 31; Huerta-Múzquiz, 1978:336, 338; Silva, 

1979:340; L. Aguilar-Rosas et al., 1982:61; R. AguilarRosas, 1982:84; Koeman and van den Hoek, 1982a:308, 

figs. 70–94; Littler and Arnold, 1982:309; L. AguilarRosas and Bertsch, 1983:114, 119; L. Aguilar-Rosas et 

al., 1985:125; Huerta-Múzquiz and Mendoza-González, 

1985:42; Ibarra-Obando and R. Aguilar-Rosas, 1985:96; 

L. Aguilar-Rosas and Pacheco-Ruíz, 1986:77; TelloVelazco, 1986:73; L. Aguilar-Rosas and Pacheco-Ruíz, 

1989:81; Sánchez-Rodríguez et al., 1989:39; R. AguilarRosas and Machado-Galindo, 1990:188; Dreckmann et 

al., 1990:24, 37; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:30; González-González, 1993:443; Mateo-Cid et 

al., 1993:51; Stout and Dreckmann, 1993:4; R. AguilarRosas and M. Aguilar-Rosas, 1994:517, 529; MendozaGonzález et al., 1994:112; González-González et al., 

1996:285; Mendoza-González and Mateo-Cid, 1996:74, 

87, pl. 23: figs. 98–101; Anaya-Reyna and RiosmenaRodríguez, 1996:862; Pacheco-Ruíz and ZertucheGonzález, 1996a:432; Leskinen and Pamilo, 1997:17; 

Bucio-Pacheco and Dreckmann, 1998:42; Yoshida, 

1998:35; Mendoza-González and Mateo-Cid, 1998:24, 

27; Blomster et al., 1998:319, figs. 1–3, 16–26, 27–29, 

49, 52–54; Rodríguez-Morales and Siqueiros-Beltrones, 

1999:30; L. Aguilar-Rosas et al., 2000:133; LeónTejera and González-González, 2000:327; Paul-Chávez 

and Riosmena-Rodríguez, 2000:146; Cruz-Ayala et al., 

2001:190; Abbott and Huisman, 2004:49, fig. 7A–C; 

Riosmena-Rodríguez et al., 2005:101; Hernández-Herrera et al., 2005:146; R. Aguilar-Rosas et al. 2005b:35; 

Dreckmann et al., 2006:153.

Enteromorpha marchantiae Setchell et N. L. Gardner, 

1924a:716, pl. 16: figs. 40–42.

Algae usually simple, tubular and smooth throughout 

(sometimes irregularly inflated and constricted), up to 15 

mm in diameter; attenuated toward base, remaining tubular (rarely becoming slightly compressed) and broadening 

upward; rarely branched. Cells in surface view, irregularly 

arranged throughout; 5–10 μm wide and 5–12 (–17) μm 

long; with a single cup-shaped chloroplast and usually 

1(–2) pyrenoid(s).

Habitat. On rocks, occasionally epiphytic on

other algae; high to low intertidal; also dredged 4–50 m 

(Dawson, 1944).

Distribution. Gulf of California: Puerto

Peñasco to Bahía La Paz. Pacific coast: Alaska to Chiapas; 

Peru; Chile; Hawaiian Islands; Japan; China.

Type Locality. “In Mari omni” (Linnaeus,

1753); probably Woolwich, River Thames, London, 

England (based on selection of lectotype illustration of 

Dillenius [1742:pl. 9: fig. 7] and typotype [see Stearn, 

1957:129] by Blomster et al. [1999:332, figs. 49, 52–54]).

Remarks. Although some specimens of Ulva 

intestinalis can sometimes be confused with morphologically 
n u m b e r 9 4 • 4 7

similar specimens of U. compressa, studies have shown them 

to be genetically distinct (Blomster et al., 1998; Hayden and 

Waaland, 2004).

Ulva linza Linnaeu

FIGURE 23

Ulva linza Linnaeus, 1753:1163; Setchell and Gardner, 

1920b:262, pl. 12: figs. 1–4; Levring, 1941:605; Smith, 

1944:44, pl. 3: figs. 4, 5; Hayden et al., 2003:289, tbl. 

4; Hayden and Waaland, 2004:378, tbl. 3; Mateo-Cid 

et al., 2006:48; Pacheco-Ruíz et al., 2008:191, 201.

Enteromorpha linza (Linnaeus) J. Agardh, 1883:134, pl. 

4: figs. 110–112; Collins, 1909b:206; Howe, 1911:490; 

Doty, 1947:18, pl. 1: figs. 7–9; Dawson, 1961b:373; 

Dawson, 1966a:5; Norris, 1976a:76, fig. 32; Silva, 

1979:340; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:30; Mendoza-González and Mateo-Cid, 1992:23; 

Mateo-Cid et al., 1993:51; Servière-Zaragoza et al., 

1993:482; González-González et al., 1996:285, 371; 

Pacheco-Ruíz and Zertuche-González, 1996a:432; Yoshida, 1998:36, pl. 1-3: fig. G; Mendoza-González and 

Mateo-Cid, 1998:24; L. Aguilar-Rosas et al., 2000:133, 

137; L. Aguilar-Rosas et al., 2002:235; Abbott and 

Huisman, 2004:50, fig. 7D; R. Aguilar-Rosas et al., 

2005b:36.

Algae linear to lanceolate, unbranched blades, up to 

40 cm long and up to 10 cm wide; composed of a flattened 

upper portion of 2 cell layers, with margins usually hollow and 1 cell thick; margins smooth or sometimes ruffled; 

and a lower portion that gradually tapers downward to a 

tubular, hollow monostromatic basal portion. Cells with 

a single cup-shaped chloroplast and usually 1 pyrenoid. 

Cells in surface view irregularly arranged in upper flat portions, 10–20 μm in diameter; cells in longitudinal rows in 

basal portions, elongated, up to 35 μm long. Cells in transection rectangular, up to 20 μm tall and to 35 μm wide.

Habitat. On sand/mud covered rocks and

shells in drainage channels of estuaries; occasionally in 

tide pools and shaded habitats; and epiphytic on crustose corallines (L. Aguilar-Rosas et al., 2000); mid to low 

intertidal.

Distribution. Gulf of California: Las Piedras del Burro to La Paz. Pacific coast: Alaska to Baja

California; Jalisco; Oaxaca; Chile; Juan Fernandez Island; 

Hawaiian Islands; Japan.

Type Locality. “In Oceano” (Linnaeus,

1753:1163); Sheerness, Kent, England (Hayden et al., 

2003:289).

Remarks. On the basis of molecular analysis, 

Hayden and Waaland (2004) noted that northeast Pacific 

“Ulva linza” and European U. linza (Tan et al., 1999) may 

be different taxa. It will be interesting to see the results of 

comparative molecular analyses of the Gulf “U. linza,” 

Pacific “U. linza,” and European U. linza (type or type 

locality), to test their taxonomic status and phylogenetic 

relationships.

Ulva radiata (J. Agardh) Hayden, Bloms

Maggs, P. C. Silva, Stanhope et Waala

Enteromorpha radiata J. Agardh, 1883:156; Koeman and 

van den Hoek, 1982b:54, figs. 54–78.

Ulva radiata (J. Agardh) Hayden, Blomster, Maggs, P. C. 

Silva, Stanhope et Waaland, 2003:290, tbl. 4.

FIGURE 22. Ulva intestinalis: Cluster of tubular thalli; most unbranched, a few branched near base (JN-3327).
4 8 • 

Enteromorpha prolifera subsp. radiata (J. Agardh) Bliding, 1963:56, figs. 25–28.

Enteromorpha prolifera sensu Setchell and Gardner, 1920b:254; Setchell and Gardner, 1924a:715; 

Dawson, 1944:204; Doty, 1947:14; Glynn and Stewart, 1973:216; Norris, 1976a:77, fig. 33; Pedroche 

and González-González, 1981:63; Huerta-Múzquiz 

and Mendoza-González, 1985:42; Mendoza-González 

FIGURE 23. Ulva linza: Thalli of three unbranched blades, widening upward; flattened upper portion with tubular margins, above a short 

tubular basal portion (JN-3694).
n u m b e r 9 4 • 4 9

and Mateo-Cid, 1986:419; Martínez-Lozano et al., 

1991:23; Servière-Zaragoza et al., 1993:482; GonzálezGonzález et al., 1996:286; Pacheco-Ruíz and ZertucheGonzález, 1996a:432; Cruz-Ayala et al., 2001:190; Pedroche et al., 2005:32; Pacheco-Ruíz et al., 2008:201 

[non Enteromorpha prolifera (O. F. Müller) J. Agardh, 

1883:129; which is Ulva prolifera O. F. Müller, 1778:7, 

fig. 1].

Ulva prolifera sensu Hayden and Waaland, 2004:377, tbl. 

3 [non Ulva prolifera O. F. Müller, 1778:7].

Enteromorpha tubulosa sensu Setchell and Gardner, 

1924a:716 [non Enteromorpha tubulosa (Kützing) 

Kützing, 1856:11; E. intestinalis var. tubulosa Kützing, 

1845: 247; which is Ulva flexuosa Wulfen, 1803:1].

Algae with a distinct, cylindrical to compressed main 

axis, variable in width from very slender (hairlike) filaments 

to broad, expanded or convoluted masses; usually with 

numerous lateral branches (rarely sparingly branched). 

Cells with a single chloroplast, usually with only 1 pyrenoid; in surface view in more or less longitudinal series 

(usually distinct in some portions); angular, quadrate to 

polygonal, (10–)12–18 μm wide, becoming elongated near 

base, 10–18 μm wide by 12–20(–34) μm long. Cells in 

transection, 10–18 μm wide by 15–30 μm tall.

Habitat. On rocks, forming mats on mudflats

or free-floating; usually in sheltered bays or estuaries; 

intertidal.

Distribution. Gulf of California: Bahía de

Los Ángeles to Bahía La Paz. Pacific coast: Washington to 

southern California; Jalisco; Panamá.

Type Locality. Arctic Norway (Bliding, 1963)

Remarks. Gulf of California specimens are 

tentatively referred to Ulva radiata. This species is probably more widely distributed in the Gulf of California than 

current records indicate. Bliding (1963, as “E. prolifera

subsp. radiata”) proposed that E. prolifera and E. radiata

were different taxa. Hayden and Waaland (2004) found 

the Pacific coast “U. prolifera” (Washington to southern 

California) was only distantly genetically related to European U. prolifera (Blomster et al., 1998) and hypothesized 

that the Pacific Northwest material identified as “U. prolifera” was U. radiata, noting this needed to be tested with 

comparative molecular data from the European U. radiata. Comparisons of Gulf “U. radiata” with U. radiata

from Europe (type locality: Arctic Norway) and the western Pacific are needed to evaluate its taxonomic status.

Cladophorop

Cladophorophyceae C. Hoek, D. G. Mann et Jahns, 

1995:408, nom. invalid. [see McNeill et al., 2006:Art. 

36.2; Pröschold and Leliaert, 2007:130].

Algae are uniseriate unbranched or branched filaments composed of multinucleate cells, with thalli ranging 

from simple filaments to structurally more complex forms. 

Cells of sporophytes and gametophytes are multinucleate 

and with numerous chloroplasts that form a parietal network or a more or less continuous layer. Some of the chloroplasts may also have a single pyrenoid. The principal 

polysaccharide of the cell wall is crystalline cellulose.

Life histories for many of the species are isomorphic. 

Sporophytes produce quadriflagellate zoospores that 

develop into gametophytic thalli. Gametophytes produce 

biflagellate isogametes or anisogametes. After fusion of two 

gametes, the zygote develops into a diploid sporophyte.

Two of the orders are present in the northern Gulf of 

California.

Key to the Orders of Cladophorophyceae in t

1a. Algae unbranched or branched filaments of terete cells throughout; tenacular cells absent; sequential formation of laterals 

at nodes; cell division by centripetal crosswall formation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cladophorales

1b. Algae of diverse forms: large siphonous, tubular cells or inflated, clavate, or globose cells; tenacular cells present; simultaneous development of laterals at nodes; cell division lenticular or segregative  . . . . . . .

Clad

Cladophorales Haeckel, 1894:302.

Cladophorales G. S. West, 1904:56; van den Hoek, 

1984:157–178.

Algae are filamentous and may be (1) simple and 

unbranched or with few to many branches, or (2) branched 

filaments with few to many lateral branches that unite to 

form a network. Most are attached to the substratum by 

rhizoids, although some may become unattached and freefloating. The cells of the filaments have crosswalls and 

are multinucleate, with one or more discoid or reticulate 

chloroplasts and usually one or more pyrenoids. Growth 

is apical or intercalary. Cell division is by nonsynchronous 

centripetal invagination, in which the parent wall divides 

into more or less equal halves (= “Cladophora-type” of 

cell division).
5 0 • 

Vegetative propagation by fragmentation is known 

for many of the species. The life histories, where known, 

are isomorphic (with gametophytes and sporophytes of 

similar morphologies). Sporophytes produce biflagellate 

or quadriflagellate zoospores in sporangia derived from 

vegetative cells. Gametophytes produce biflagellate isogametes or anisogametes.

Remarks. There have been different opinions on 

whether the Cladophorales should also include the Siphonocladales (e.g., van den Hoek, 1984; Bakker et al., 1994; 

Hanyuda et al., 2002; Abbott and Huisman, 2004; Wynne, 

2005), or if the two should be kept as separate orders 

(Womersley, 1984; Leliaert et al., 2003; Kraft, 2007) with 

the type of cell division being a principle taxonomic character. The use of families in the Cladophorales/Siphonocladales complex is also problematic as there seems to be no 

phylogenetic basis that supports the currently accepted families (Leliaert et al., 2007). For now, the families are treated 

herein in these two orders. Members of the Cladophorales 

are mostly marine (although a few, e.g., Cladophora, also 

have freshwater species) and distributed from sub-boreal 

to tropical seas, and members of the Siphonocladales are 

exclusively marine and subtropical to tropical.

There is one family, well represented, in the northern 

Gulf of California.

Clad

Cladophoraceae W. Wille in Warming, 1884:30.

Algae in this family are filamentous, uniseriate, 

branched or unbranched, and vary in color from light to 

dark green. Cells are multinucleate, with parietal chloroplasts containing few to many pyrenoids.

Asexual reproduction for many is by fragmentation. 

Sexual reproduction, where known, is isogamous.

Remarks. In the northern Gulf of California 

the family is represented by three genera, Chaetomorpha, 

Cladophora, and Rhizoclonium. However, the taxonomic 

distinction of the unbranched genera Rhizoclonium and 

Chaetomorpha is questionable (Blair et al., 1982; Blair, 

1983; Silva et al., 1996a), with taxonomy based solely on 

filament diameter, cell sizes, and nature of the holdfasts. 

The generic and species concepts need further study and 

phylogenetic testing.

Key to the Genera of Cladophoraceae in the 

1a. Filaments sparsely to densely branched, to 1 or more orders  . . . . 

1b. Filaments simple and unbranched or with short rhizoidal branches  

2a. Filaments unattached, entangled or free-floating; or if attached, by short rhizoidal branches along their length 

. . . . . . 

2b. Filaments attached by a discoid basal cell or by rhizoids extending only from the basal cell. . . . . . 

Chaetomorpha Kützing

Chaetomorpha Kützing, 1845:203.

Algae are uniseriate, unbranched filaments that 

grow singularly or in clumps and may be free-floating or 

attached by a discoid basal cell or by nonseptate rhizoidal processes from an elongate basal cell. Growth is by 

intercalary cell division only. Cells of the filaments can be 

shorter or up to 10 times longer than wide, and rectangular, isodiametric, or cylindrical to barrel-shaped. Cells are 

multinucleate, with a parietal or reticulate chloroplast and 

many pyrenoids.

Vegetative reproduction is by fragmentation. Life histories are presumed to be an isomorphic alternation of 

stages. Sporangia are formed in undifferentiated cells and 

produce quadriflagellate zoospores (probably meiospores). 

Sexual reproduction is isogamous, with gametangia (also 

formed in an undifferentiated cell) producing biflagellate 

isogametes. Isogametes have also been reported to develop 

parthenogenetically.

Remarks. Five species have been reported to 

occur only in the southern Gulf (Dawson, 1949; Dawson, 1950c; Huerta-Múzquiz and Mendoza-González, 

1985; R. Aguliar-Rosas and Machado-Galindo, 1990; 

Riosmena-Rodríguez and Paul-Chávez, 1997; Pedroche 

et al., 2005): C. linoides Kützing (1847), C. minima Collins et Hervey (1917), C. pachynema (Montagne) Kützing 

(1847), C. spiralis Okamura (1903, 1912), and C. javanica Kützing (1847).

Of the four species that are known in the northern 

Gulf of California, three also occur in the southern Gulf.
n u m b e r 9 4 • 5 1

Chaetomorpha aerea (Dillwyn) Kützing

FIGURE 24B

Conferva aerea Dillwyn, 1806:pl. 80.

Chaetomorpha aerea (Dillwyn) Kützing, 1849:379; Collins, 1909b:324; Hartmann, 1929:485; Dawson, 

1944:208 (in part); Chávez B., 1972:268; HuertaMúzquiz, 1978:338; Silva, 1979:342; Blair, 1983:187, 

figs. 5, 6; Oliva-Martínez and Ortega, 1987:128, pl. 4: 

fig. 2a–d; Ortega et al., 1987:78, pl. 14: figs. 65–67; 

Mateo-Cid and Mendoza-González, 1992:24; Servière-Zaragoza et al., 1993:482; Mendoza-González 

et al., 1994:112; González-González et al., 1996:278; 

Pacheco-Ruíz and Zertuche-González, 1996a:432; Yoshida, 1998:54; L. Aguilar-Rosas et al., 2000:133; Abbott and Huisman, 2004: 65, fig. 17; Pedroche et al., 

2005:34; Pacheco-Ruíz et al., 2008:201.

Chaetomorpha linum sensu Abbott and Hollenberg, 

1976:101 [in part] [non Chaetomorpha linum (O. F. 

Müller) Kützing, 1845:204].

Tufts of erect, straight, uniseriate filaments, up to 4 

(or more) cm tall, attached to substrate by an elongate 

basal cell with a disc-shaped holdfast. Upper to lower cells 

of more or less same diameter throughout, (125–)240–400 

μm in diameter, often more or less squarish to slightly 

elongate, 1.0–2.0 times longer than wide; lowermost cells, 

1.0–2.5 times longer than wide. Basal cell (nonannulate) of 

similar diameter to cells above, but much longer, (4–)5–8 

times longer than wide, up to 1.8 mm long.

Reproduction not observed in Gulf specimens. Fertile 

cells reported from elsewhere, up to 600 μm in diameter 

(Blair, 1983).

Habitat. On rocks; high to mid intertidal

Distribution. Gulf of California: Puertecitos

to Bahía de Los Ángeles. Pacific coast: central California 

to Nayarit; Peru; Chile; Hawaiian Islands; Japan.

Syntype Localities. England and Wales,

UK.

Remarks. Some specimens of Chaetomorpha aerea (e.g., JN-5180a on US Alg. Coll. slide 4144; El 

Desemboque, Sonora) were particularly rich with microscopic endophytes and epiphytes, such as Entocladia viridis, that grew on, sometimes encircling, the cells of the 

basal region.

Some phycologists consider Chaetomorpha aerea and 

C. linum to be growth forms of a single species (e.g., Christensen, 1957; Burrows, 1991). However, herein these two 

are recognized as separate species (e.g., following Kornmann, 1972; Blair et al., 1982; Blair, 1983; Silva et al., 

1996a), until further morphological and molecular systematic studies have been done on types and type-locality specimens of these species, along with needed comparisons to 

elucidate their relationship to Gulf of California “C. aerea.” 

Although crystalline cell inclusions are reported in C. aerea

from other regions (Leliaert and Coppejans, 2004:202, fig. 

47), they have not been observed in Gulf C. aerea.

Key to the Species of Chaetomorpha in the n

1a. Filaments unattached, free-floating or lying on substratum, or entangled with other algae  . . . . . . . . 

1b. Filaments attached by a basal cell or rhizoids from basal cell; forming clumps or tufts  

2a. Upper filament cells 400–900 μm wide, 1–4 times longer than wide; distinctive basal cell very long, up to 9 mm, usually 

with annular constrictions  . . . . . 

2b. Upper filament cells narrower, 175–450 μm wide, only 0.5–2.5 times longer than wide; basal cell shorter, less than 4 mm, 

without annular constrictions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Cells wider than long, 250–280 μm wide by 125–148 μm long (0.5–0.6 times diameter); lowermost 1–2 basal cells 1.25 

times longer than wide, up to 350 μm long. . . . . . .

3b. Cells longer than wide, 625–1100 μm long (0.75–2.5 times diameter), 250–450 μm wide; basal cell 3–5 times longer than 

wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4a. Filaments mostly curly; cells 175–450 μm wide, more or less squarish to slightly longer (0.75–1.5 times diameter); basal 

cell less than 5 times longer than wide. . . . . . . . . 

4b. Filaments straight; 240–400 μm wide; cells longer than wide (mostly 1.5–2.5 times diameter); basal cell up to 5–8 times 

longer than wide  . . . 
5 2 • 

Chaetomorpha antennina

(Bory de Saint-Vincent) Kützin

FIGURES 24A, 25B

Conferva antennina Bory de Saint-Vincent, 1804a:381; 

Bory de Saint-Vincent, 1804b:161.

Chaetomorpha antennina (Bory de Saint-Vincent) Kützing, 

1847:166; Kützing, 1849:379; Collins, 1909b:324; Howe, 

1914:37; Setchell and Gardner, 1920b:203; Setchell and 

Gardner, 1924a:713; Setchell and Gardner, 1930:136; 

Dawson, 1944:207; Dawson, 1945b:59; Taylor, 

1945:52; Dawson, 1949:245, 251; Dawson, 1950c:149, 

figs. 6, 7; Dawson, 1951:52; Dawson, 1954a:3; Dawson, 

1959a:9, 12; Dawson, 1959c:4; Dawson and Beaudette, 

1959:18, 20; Dawson, 1961b:376; Dawson, 1961c:403, 

pl. 1: figs 7, 8; Dawson, 1962b:172, fig. 8 (right); HuertaMúzquiz and Tirado-Lizárraga, 1970:126; Norris, 

1973:3, 19; Abbott and Hollenberg, 1976:101, fig. 58; 

Huerta-Múzquiz , 1978:338–340; Stewart and Stewart, 

1984:140; Huerta-Múzquiz and Mendoza-González, 

1985:42; Mendoza-González and Mateo-Cid, 1985:22; 

Mendoza-González and Mateo-Cid, 1986:419; SalcedoMartínez et al., 1988:82; Sánchez-Rodríguez et al., 

1989:39; Dreckmann et al., 1990:24, 37; Mateo-Cid and 

Mendoza-González, 1991:27; Mendoza-González and 

Mateo-Cid, 1992:24; Mateo-Cid and Mendoza-González, 

1992:24; González-González, 1993:443; Stout and 

Dreckmann, 1993:4; Servière-Zaragoza et al., 1993:482; 

Mateo-Cid and Mendoza-González, 1994a:52; MateoCid and Mendoza-González, 1994b:44; León-Tejera and 

González-González, 1994:493; Mendoza-González et 

al., 1994:112; González-González et al., 1996:278; Silva 

et al., 1996b:229, 234; Mendoza-González and MateoCid, 1996:75, 87, pl. 25, figs. 109, 110; Pacheco-Ruíz 

and Zertuche-González, 1996a:432; León-Alvarez et al., 

FIGURE 24. Species of Chaetomorpha. A. Chaetomorpha antennina: Clump of relatively stiff filaments (EYD-11023). B. Chaetomorpha aerea:

Lower region of filament; long basal cell ending in disc-shaped base with short rhizoids (JN-5180a).
n u m b e r 9 4 • 5 3

1997:397; Galindo-Villegas et al., 1997:4; Mateo-Cid 

and Mendoza-González, 1997:56; Servière-Zaragoza et 

al., 1998:169, 176, 178; Bucio-Pacheco and Dreckmann, 

1998:44; Aguila-Ramírez et al., 1998:511; Yoshida, 

1998:54; Mendoza-González and Mateo-Cid, 1998:25, 

27; Mendoza-González et al., 1999:61; León-Tejera 

and González-González, 2000:237; Mateo-Cid et al., 

2000a:69; Carballo et al., 2002:752; Abbott and Huisman, 2004:66, fig. 18A–D; Pedroche et al., 2005:35–36; 

Hernández-Herrera et al., 2005:146; Dreckmann et al., 

2006:154; Servière-Zaragoza et al., 2007:7; PachecoRuíz et al., 2008:201.

Conferva media C. Agardh, 1824:100.

Cladophora media (C. Agardh) Kützing, 1849:380; 

Huerta-Múzquiz and Tirado-Lizárraga, 1970:120, 131, 

135; Chávez B., 1972:268; Pedroche and GonzálezGonzález, 1981:64.

Cladophora pacifica Kützing, 1849:379 (see Collins 

1909b:324; Howe, 1914:37).

Algae erect, tufted or clumped, of relatively stiff, uniseriate filaments, up to 10 cm tall, attached by an elongated 

basal cell with long slender rhizoids from proximal end. 

Upper cells 400–900 μm in diameter, 1–4 times as long as 

wide; basal cell of similar width but much longer, 5–9 mm 

in length. Basal cell usually with annular constrictions.

Fertile cells swollen and larger than vegetative cells.

Habitat. On rocks, usually in shaded areas;

high to low intertidal.

Distribution. Gulf of California: Bahía de

Los Ángeles to Bahía San Carlos; Isla San Ildefonso; Cabeza 

Ballena to Cabo Pulmo. Pacific coast: southern California 

to Chiapas; Isla San Benedicto (Islas Revillagigedo); Clipperton Island; El Salvador; Panamá; Galápagos Islands; 

Ecuador; Peru; Chile; Hawaiian Islands; China; Japan.

Type Locality. Réunion Island (Bourbon Is.),

Mascarene Islands, Indian Ocean.

Chaetomorpha bangioides E. Y. Daw

FIGURE 25A

Chaetomorpha bangioides E. Y. Dawson, 1950c:149, 

figs. 8–10; Dawson, 1959a:7, 12; Dawson, 1961b:376; 

Espinoza-Ávalos, 1993:333; Servière-Zaragoza et 

al., 1993:479, 485; González-González, 1993:443; 

González-González et al., 1996:128; Servière-Zaragoza 

et al., 1998:169, 180; Pedroche et al., 2005:36.

Algae tufted, up to 1 cm tall; of uniseriate more or 

less contorted filaments and attached by a small, conical 

holdfast of coalesced rhizoids issued from the primary 

basal cell. Cells wider than long (more or less rectangular 

in lateral view), 250–280 μm in diameter, shorter in length 

(mostly 0.5–0.6 times diameter); lowermost 1–2 cells of 

similar diameter as upper cells, but slightly longer (up to 

350 μm in length). Cell walls thick, up to 60 μm.

Reproduction unknown.

Habitat. On rocks; intertidal

Distribution. Gulf of California: Isla Patos;

Puerto Escondido. Pacific coast: Nayarit to Jalisco.

Type Locality. Isla Patos, off Isla Tiburón,

Gulf of California.

Remarks. Chaetomorpha bangioides is named 

for the resemblance of its filament cells to those of Bangia

fuscopurpurea. It differs from other Gulf of California species of Chaetomorpha in the shape of its cells, rectangular 

with the diameter wider than the length, and the absence 

of a very long or otherwise distinctive basal cell (Dawson, 

1950c). This species remains poorly known and in need of 

further collection and study.

Chaetomorpha linum (O. F. Müller) Kützing

Conferva linum O. F. Müller, 1778:7, pl. 771; fig. 2.

Chaetomorpha linum (O. F. Müller) Kützing, 1845:204; 

Dawson, 1949:229; Dawson, 1959a:12; Dawson, 

1961b:377; Devinny, 1978:358; L. Aguilar-Rosas et al., 

1982:54, 61; L. Aguilar-Rosas and Bertsch, 1983:115, 

121; Blair, 1983:178, figs. 1, 2a; Huerta-Múzquiz and 

Mendoza-González, 1985:42; Mendoza-González and 

FIGURE 25. Species of Chaetomorpha. A. Chaetomorpha bangioides: Rectangular (wider than long) cells of different “Bangia-like” 

shapes in upper part of filament (after Dawson, 1950c:fig. 10). B. 

Chaetomorpha antennina: Basal cell, typically very long, with short 

attachment rhizoids, and cell wall with annular constrictions.
5 4 • 

Mateo-Cid, 1985:22; L. Aguilar-Rosas and PachecoRuíz, 1986:77; Mendoza-González and Mateo-Cid, 

1986:419; Ortega et al., 1987:79, 90, pl. 15: fig. 68; 

L. Aguilar-Rosas and Pacheco-Ruíz, 1989:81; R. Aguilar-Rosas and Machado-Galindo, 1990:188; Stewart, 

1991:29; Mateo-Cid and Mendoza-González, 1991:27; 

Rocha-Ramírez and Siqueiros-Beltrones, 1991:24; 

Mendoza-González and Mateo-Cid, 1992:24; MateoCid and Mendoza-González, 1992:24; Mateo-Cid et 

al., 1993:51; León-Tejera and González-González, 

1993:497; Servière-Zaragoza et al., 1993:482; R. Aguilar-Rosas and M. Aguilar-Rosas, 1994:517; MateoCid and Mendoza-González, 1994b:34, 44; MendozaGonzález et al., 1994:112; González-González et al., 

1996:278; Yoshida, 1998:56; Servière-Zaragoza et al., 

1998:169, 177, 180; Rodríguez-Morales and SiqueirosBeltrones, 1999:22; L. Aguilar-Rosas et al., 2000:133, 

134; L. Aguilar-Rosas et al., 2002:235; Wysor, 

2004:216; Pedroche et al., 2005:38–39; RiosmenaRodríguez et al., 2005:101; Hernández-Herrera et al., 

2005:146.

Chaetomorpha aerea f. linum (O. F. Müller) Collins, 

1909b:325.

Chaetomorpha aerea sensu Dawson, 1944:208 [in 

part] [non Chaetomorpha aerea (Dillwyn) Kützing, 

1849:379].

Algae of mostly curved to curled, uniseriate filaments, 

up to 5 (or more) cm long, usually entangled and unattached or occasionally found attached to substrate by 

an elongated, lobed basal cell. Upper cells cylindrical to 

barrel-shaped, (175–)200–450 μm in diameter, and not 

as long as wide or only slightly longer (length 0.75–1.5 

times the width); basal cells, when present, 3–8 times longer than wide.

Reproductive cells barrel-shaped to subglobose, 400–

600 μm in diameter.

Habitat. Estuarine habitats and mangrove

lagoons; mid intertidal to shallow subtidal (also dredged; 

Dawson, 1944).

Distribution. Gulf of California: Puertecitos

to La Paz. Pacific coast: Alaska to Baja California; Sinaloa 

to Colima; Oaxaca; Panamá; Chile; Japan.

Syntype Localities. Nakskov Fjord and

Rødby Fjord, Denmark.

Remarks. The species Chaetomorpha linum

and C. aerea have long been problematic and often confused (Blair, 1983), and Gulf of California specimens 

referred to either of these species should be reexamined to 

determine if only one or both taxa occur in the northern 

Gulf of California. See also comments herein on Chaetomorpha aerea under Remarks for that species.

Cladophora Kützing

Cladophora Kützing, 1843:262

Algae consisting of uniseriate filaments, sparsely to 

profusely branched; usually erect and attached by rhizoids 

from basal cells; or sometimes free-floating, forming small 

to large filamentous masses. Branching is often up to several 

orders, most with variation in pattern of branching and in 

diameter (often decreasing in diameter upward). Cells are 

cylindrical, multinucleate, and with numerous small chloroplasts (usually form a parietal reticulum) and pyrenoids. 

Cell division is apical or intercalary or sometimes both.

Vegetative reproduction by fragmentation occurs in 

many species. Life histories are an alternation of isomorphic generations. The terminal and subterminal cells of 

the filaments become the reproductive cells. Sporophytic 

reproduction is usually by the formation of quadriflagellate or biflagellate zoospores. Gametophytes produce 

biflagellate isogametes.

Remarks. The Gulf species of Cladophora are 

in need of critical study. Their taxonomy, based primarily 

on characters such as habit, cell diameter and length/width 

ratio, and type and degree of branching, is complicated 

by a high degree of variability of their relatively simple 

morphologies. Another character, considered to have systematic value, is the presence or absence of crystalline cell 

inclusions (Leliaert and Coppejans, 2004). Comparative 

morphological studies of Gulf specimens of Cladophora

and type specimens (such as those of van den Hoek, 1963, 

1982; Söderström, 1963; Sakai, 1964; van den Hoek and 

Womersley, 1984; van den Hoek and Chihara, 2000) and 

molecular and phylogenetic analyses are needed to test 

their taxonomic status.

Five taxa are recorded in the southern Gulf: 

Cladophora lehmanniana (Lindenberg) Kützing (1843) 

by Dawson (1959a, as “C. utriculosa Kützing”), C. glomerata (Linnaeus) Kützing (1843) by Huerta-Müzquiz and 

Mendoza-González (1985), C. glomerata var. crassior (C. 

Agardh) van den Hoek (1963) by Huerta-Müzquiz and 

Mendoza-González (1985, as “C. crispata”), C. rivularis

(Linnaeus) van den Hoek (1963) by Huerta-Müzquiz and 

Mendoza-González (1985, as “C. insignis”), and C. stimpsonii Harvey (1860; Dawson, 1959b) by Rocha-Ramírez 

and Siqueiros-Beltrones (1991).

A highly diverse genus, with 11 species of Cladophora

currently recognized in the northern Gulf of California.
n u m b e r 9 4 • 5 5

Cladophora albida (Nees) Kützing

FIGURE 26A

Annulina albida Nees, 1820:Index [1] [see Silva et al., 

1996a:769].

Cladophora albida (Nees) Kützing, 1843:267; León-Tejera 

et al., 1993:199; González-González, 1993:443; Servière-Zaragoza et al., 1993:481; Silva et al., 1996a:769; 

González-González et al., 1996:129, 368; Yoshida, 

1998:60; Nuñez-López et al., 1998:39; van den Hoek 

and Chihara, 2000:122, figs. 55–59, 63B,D; Pedroche 

et al., 2005:40.

Conferva albida Hudson, 1778:595, nom. illeg. [since 

it is a later homonym of Conferva albida Forsskål, 

1775:XII].

Cladophora albida (Hudson) Kützing, 1843:267; Kützing, 

1849:400; Kützing, 1854:pl. 15; Collins, 1909b:336; 

Setchell and Gardner, 1920b:218; Yamada, 1928:fig. 

1; Dawson, 1944:209; Söderström, 1955:275, figs. 

1–6; Dawson, 1959a:12; Dawson, 1960a:31; Dawson, 

1961b:377; Dawson, 1961c:403, pl. 2: figs. 1, 2; Dawson, 1962b:182, fig. 32; van den Hoek, 1963:94, pl. 20: 

figs. 241–243, pl. 21: figs. 244, 246–257, pl. 22: figs. 

258–279, pl. 23: figs. 280–297, pl. 24: figs. 298–316; 

Söderström, 1963:65, figs. 55–61; Sakai, 1964:30, 

Key to the Species of Cladophora in the no

1a. Basal portion procumbent, of densely entangled horizontally low-growing filaments; upper portion of erect filaments 

. . . . . . .

1b. Basal and upper portions erect; of delicate to coarse filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a. Cells of basal region with rhizoids that have cell walls with annular constrictions (occasionally some basal cells also with 

annular constrictions)  . . . . . 

2b. Cells of basal region with rhizoids that lack annular constrictions or completely without rhizoids . . . . . . . . . . . . . . . . . 3

3a. Cell diameter of main branches over 200 μm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3b. Cell diameter of main branches less than 200 μm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

4a. Basal cells without downward growing rhizoids; filaments branched mostly to one side; branchlets usually strongly incurved or curled. . . . . . 

4b. Basal cells with downward growing rhizoids; filaments branched otherwise; branchlets not curled  

5a. Branched sparingly, dichotomous in lower portions; secund to pectinate (occasionally opposite or rarely alternate) in upper 

portions  . . . . . . 

5b. Branched mostly dichotomously to trichotomously throughout  

6a. Stiff tufts; filament cells much longer than wide, lower cells 300–500 μm wide, 20–30 times longer than wide; upper cells 

100–150 μm wide, 4–10 times longer than wide. . . . . .

6b. Lax tufts (not stiff), branching divaricate; cells only 2–4 times longer than wide and smaller diameter throughout; lower 

cells 190–240 μm wide; upper cells 25–40 μm wide. . . . . .

7a. Branching dichotomous or trichotomous, becoming irregularly alternate to pectinate above; cells 120–180(–200) μm in 

diameter in lower portions, narrowing above. . . . . . . . . 

7b. Branching subdichotomous, irregular, alternate or secund; cells of smaller diameter, 20–60(–85) μm  

8a. Upper main axes subdichotomously, irregularly or unilaterally branched; branches formed at distal end of cells; cells of 

similar diameter, 20–30(–40) μm, throughout. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. albid

8b. Upper axes subdichotomously and/or secundly branched; lower cells larger than 30 μm in diameter  

9a. Branched sparingly (widely spaced), subdichotomously; branches of different lengths intermixed; lower cells 50(–85) μm 

wide (5–6 times as long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. serice

9b. Branched more regularly; subdichotomously below to secund above; lower cells less than 60 μm wide (3–5 times as long)

. 

10a. Dense short tufts (<4mm tall) of stiff filaments; cells mostly 4–5 times longer than wide throughout (longest in lower 

cells) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tiburonensi

10b. Delicate hairlike filaments to 2 cm tall; lower cells 3–4 times longer than wide, upper cells 1–2 times longer than wide 

. . . . . . . 
5 6 • 

figs. 10, 11, pl. 17: fig. 1; Söderström, 1965:173, 181; 

Abbott and Hollenberg, 1976:104, fig. 62; Devinny, 

1978:358; Ortega et al., 1987:79, 90, pl. 16: figs. 72, 

73; Schnetter and Bula Meyer, 1982:24, pl. 3: figs. D–F; 

van den Hoek, 1982:100, pl. 15: figs. 133–137, pl. 16: 

fig. 144; L. Aguilar-Rosas and Pacheco-Ruíz, 1986:77; 

Mateo-Cid and Mendoza-González, 1991:27; Stewart, 

1991:30; Mendoza-González and Mateo-Cid, 1992:24; 

León-Tejera et al., 1993:199; Mendoza-González and 

Mateo-Cid, 1996:75, 87, pl. 25: figs. 111–113; Pacheco-Ruíz and Zertuche-González, 1996a:432; Dreckmann et al., 2006:154; Pacheco-Ruíz et al., 2008:201.

Conferva hamosa Kützing, 1843:267.

Cladophora bertolonii var. hamosa (Kützing) Ardissone, 

1887:242; Huerta-Múzquiz and Mendoza-González, 

1985:42.

Cladophora magdalenae Harvey, 1851:pl. 355A; Dawson, 1949:230.

Algae small tufts up to 1 cm tall, of delicate filaments with main axes closely and pseudodichotomously 

branched, mostly 1–2 orders. Branches formed at distal 

end of cells; usually 1 per cell. Filaments thin, of similar 

diameter throughout; mostly 20–30(–40) μm in diameter; 

cells (2–)4–5 times as long as wide. Cells with a reticulate 

chloroplast and numerous pyrenoids.

Habitat. On shaded or exposed rocks, also

epiphytic on Cladophora microcladioides and epizoic on a 

parrotfish beak; intertidal to shallow subtidal.

Distribution. Gulf of California: Bahía de

Los Ángeles to Bahía Kino; Puerto Escondido; Cabo San 

Lucas. Pacific coast: British Columbia to Chiapas; El Salvador; Colombia; Japan.

Type Locality. Isle of Selsey, England

Remarks. Not well known in the Gulf, 

Cladophora albida has been found epiphytic on C. microcladioides in the vicinity of Bahía Kino (Dawson, 1944) 

and Bahía de Los Ángeles (Pacheco-Ruíz and ZertucheGonzález, 1996a) and growing on the beak of a parrotfish 

taken at Puerto Escondido (Dawson, 1959a). A specimen 

from Cabo San Lucas (EYD-3203 [HAFH now UC]) represents a southern record for the Gulf.

Cladophora columbiana Collins

FIGURE 27A

Cladophora columbiana Collins in Setchell et N. L. Gardner, 1903:226; Dawson, 1944:210 [with doubt]; Dawson, 1961b:378; Abbott and Hollenberg, 1976:105, 

fig. 63; Devinny, 1978:358; L. Aguilar-Rosas et 

al., 1985:125; Mendoza-González and Mateo-Cid, 

1985:22; L. Aguilar-Rosas and Pacheco-Ruíz, 1986:77; 

Mendoza-González and Mateo-Cid, 1986:419; R. 

Aguilar-Rosas et al., 1990:123; Rocha-Ramírez and 

Siqueiros-Beltrones, 1991:30; Martínez-Lozano et al., 

1991:22; R. Aguilar-Rosas and M. Aguilar-Rosas, 

1994:517, 526; González-González et al., 1996:129; 

Mendoza-González et al., 1999:61; Paul-Chávez and 

Riosmena-Rodríguez, 2000:146; Pedroche et al., 

2005:42; Riosmena-Rodríguez et al., 2005:101.

Cladophora trichotoma sensu Collins, 1909b:349; Howe, 

1911:492; Setchell and Gardner, 1920b:210, pl. 16: 

fig. 2; Smith, 1944:58, pl. 7: fig. 2; Dawson, 1945b:64; 

Dawson, 1949:219, 222, 228; Huerta-Múzquiz and 

Mendoza-González, 1985:42; González-González et 

al., 1996:131 [non Cladophora trichotoma (C. Agardh) 

Kützing, 1849:414; basionym: Conferva trichotoma

FIGURE 26. Species of Cladophora. A. Cladophora albida: Portion of a sparsely branched axis (after Dawson, 1962b:fig. 32). B. 

Cladophora tiburonensis: Habit of individual frond from short 

dense tuft (after Dawson, 1944: pl. 31, fig. 1).
n u m b e r 9 4 • 5 7

C. Agardh, 1824:121, which is Cladophora pellucida

(Hudson) Kützing, 1843:271; basionym: Conferva pellucida Hudson, 1762:483 (see van den Hoek, 1963:215)].

Algae cushionlike or forming low-growing mats of 

entangled filaments, 3–5 cm tall; attached by rhizoids 

from cells of basal filaments and from lower cells of erect 

filaments; lower basal filaments growing more or less 

horizontal; upper filaments erect, unbranched to sparsely 

branched in lower portion, becoming dichotomously or 

trichotomously to irregularly branched upward; with a 

blunt terminal cell. Cells cylindrical; in main axes 150–

250(–300) μm in diameter, and 400–1500 μm long; apical 

cells narrower (90–)130–170 μm in diameter.

Habitat. On rocks or other hard substratum;

intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas; Bahía del Rincón to La Paz. Pacific 

coast: Alaska to Bahía Tortugas, Baja California Sur.

Type Locality. Port Renfrew, Vancouver

Island, British Columbia, Canada.

Cladophora graminea Collins

FIGURE 28A,B

Cladophora graminea Collins, 1909a:19, pl. 78: fig. 6; 

Setchell and Gardner, 1920b:211; Dawson, 1944:210; 

Smith, 1944:59, pl. 7: fig. 1; Dawson, 1945b:64; Dawson, 1949:222, 245; Dawson, 1951:52; Dawson et al., 

1960a:26, pl. 1: fig. 6; Dawson et al., 1960b:5, 7–9, 

12; Dawson, 1961b:377; Dawson, 1966a:6; Dawson and Neushul, 1966:174; Abbott and Hollenberg, 1976:105, fig. 64; Huerta-Múzquiz, 1978:336; 

Devinny, 1978:358; Littler and Littler, 1981:151, 153; 

Schnetter and Bula Meyer, 1982:25, pl. 3: figs. A–C; L. 

Aguilar-Rosas and Bertsch, 1983:116, 121; L. AguilarRosas and Pacheco-Ruíz, 1986:77; R. Aguilar-Rosas 

et al., 1990:123; Stewart, 1991:31; R. Aguilar-Rosas 

and M. Aguilar-Rosas, 1994:517; Mateo-Cid and 

Mendoza-González, 1994a:57; González-González et 

al., 1996:130; Pacheco-Ruíz and Zertuche-González, 

1996a:432; Núñez-López et al., 1998:39; Pedroche et 

FIGURE 27. Species of Cladophora. A. Cladophora columbiana: Branching habit of a few upper branches (after Setchell and Gardner, 1920b: 

pl. 16, fig. 2). B. Cladophora macdougalii: Portion of two filaments, apical region (left) and upper portion (right) (after M. Howe, 1911: pl. 33, 

fig. 7).
5 8 • 

al., 2005:43; Mateo-Cid et al., 2006:48; Pacheco-Ruíz 

et al., 2008:201.

Algae in tufts, up to 8(–18) cm tall; relatively stiff, 

coarse, dark green to blackish-green; dichotomously or 

trichotomously branched, sometimes becoming alternate 

in upper portions. Branch divisions at more or less same 

angle throughout; branches usually curving in same direction. Cells very long in relation to diameter, in lower portions up to 300 μm in diameter, and 20–30 times as long 

as wide; in upper branches 100–150 μm in diameter, and 

4–10 times as long as wide.

Habitat. On rocks, in shaded areas, crevices

or under other larger algae; mid intertidal to shallow 

subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas. Pacific coast: Oregon to Isla Magdalena (Baja California Sur); Colombia.

Type Locality. Pacific Grove, Monterey

County, California.

Remarks. Dawson (1944, 1966a) referred 

his Gulf specimens to Cladophora graminea with doubt. 

However, the Gulf specimens, with their long cell length 

to width ratio, seem to agree with Pacific coast C. graminea and thus, for now, are retained under this name.

Cladophora hesperia Setchell et N. L. Gardn

FIGURE 29A,B

Cladophora hesperia Setchell et N. L. Gardner, 

1924a:713, pl. 13: fig. 17; Dawson, 1944:211; 

Dawson, 1949:247; Dawson, 1952:431; Dawson, 

1954e:323; Dawson, 1961b:377; Dawson, 1966a:6; 

Stewart, 1982:54; Huerta-Múzquiz and MendozaGonzález, 1985:42; Rocha-Ramírez and Siqueiros-Beltrones, 1991:30; González-González et al., 1996:130; 

Rodríguez-Morales and Siqueiros-Beltrones, 1999:17, 

22; L. Aguilar-Rosas et al., 2000:133; Paul-Chávez and 

Riosmena-Rodríguez, 2000:146; L. Aguilar-Rosas et 

FIGURE 28. Cladophora graminea: A. Habit (JN-3118). B. Axis of long cells, slightly tapering in diameter to the apices (from Collins, 1909a: 

pl. 78, fig. 6).
n u m b e r 9 4 • 5 9

al., 2002:235; Pedroche et al., 2005:44; Mateo-Cid et 

al., 2006:48, 58.

Algae forming dense clumps or sometimes spreading 

thin mats, up to 1.5 cm tall, light green, abundantly and 

densely dichotomously to trichotomously branched, up to 

6 orders, divaricate (branching at a wide angle) from the 

upper end of cells. Main branches of more or less uniform width, 190–240 μm in diameter; at each branch fork 

diameter becomes reduced upward; short branchlets of the 

fourth to sixth order much narrower, mostly 25–40 μm in 

diameter; tapering to blunt apices. Cells mostly 2–4 times 

as long as wide (up to 1 mm in length).

Habitat. On mudflats and rocks, usually

among other turf-forming algae; mid to low intertidal.

Distribution. Gulf of California: Piedras de

La Salina to Bahía Kino; Bahía San Gabriel, Isla Espiritu 

Santo to Cabo San Lucas. Pacific Mexico: Punta Norte, 

Isla Cedros (Baja California) to Velero Point (~10 miles 

[16 km] west of Punta Malarrimo), Bahía Sebastián Vizcaíno (Baja California Sur); Mazatlán (Sinaloa).

Type Locality: “Muddy habitat, at the

southern extremity of Lower California [Baja California 

Sur]” (Setchell and Gardner, 1924a:714); given as “distrito de Cabo San Lucas” by Pedroche et al. (2005:44).

Remarks. Originally described from the southern Gulf of California, Cladophora hesperia has also been 

reported from Pacific Mexico (Dawson, 1954e).

Cladophora macdougalii M. Ho

FIGURE 27B

Cladophora macdougalii M. Howe, 1911:491, pl. 33: fig. 

7; Collins, 1912:96; Setchell and Gardner, 1920b:214; 

Dawson, 1944:211; Dawson, 1961b:378; Hommersand, 1972:70; Espinoza-Ávalos, 1993:333; GonzálezGonzález et al., 1996:130; L. Aguilar-Rosas et al., 

2000:129, 133; Pedroche et al., 2005:45.

Algae in tufts, coarse and rigid, yellow to dark green, 

up to 17 cm tall; branched sparingly dichotomously 

FIGURE 29. Cladophora hesperia: A. Branches widely divaricate and decreasing in diameter with each successive branch order (EYD-832). B. 

Branching habit of upper branches (after Setchell and Gardner, 1924a: pl. 13, fig. 17).
6 0 • 

below, and in upper portions usually secund to pectinate, 

or occasionally opposite or rarely alternate. Cells of main 

filaments 135–310 μm in diameter, 6–15 times as long as 

wide; cells of upper segments 75–110 μm in diameter, 1–4 

times as long as wide; apical cells subacute or blunt.

Habitat. Tidal flats and rocks; probably mid

tidal.

Distribution. Gulf of California: El

Machorro; San Felipe to Puertecitos.

Type Locality. San Felipe, Baja California,

Gulf of California.

Remarks. Thus far, Cladophora macdougalii

is only known from the type and the more recent collections of L. Aguilar-Rosas et al. (2000). Although Collins (1912) noted that C. macdougalii was “quite close” 

to California specimens identified as “C. ovoidea” and 

Setchell and Gardner (1920b) considered Californian 

“C. ovoidea” close to illustrations of Kützing (1853:pl. 

92: fig. 1), Dawson (1944) later preferred to keep these 

two separate. Further, the type material of European C. 

ovoidea Kützing (1843) is now considered to be a synonym of C. sericea (Hudson) Kützing (1843; see van den 

Hoek, 1963, 1982), and the California “C. ovoidea” 

sensu Collins (1909b, 1912) and Smith (1944) was recognized as a separate species, C. sakaii I. A. Abbott (1972). 

(Note that this is a replacement name since the earliest 

name, C. densa W. H. Harvey (1860; Dawson, 1959b), is 

a later homonym of C. densa (Roth) Kützing, 1845.) The 

lowermost basal portions of C. macdougalii have smooth 

cell walls, distinguishing them from those of C. sakaii,

which have annular constrictions (see Abbott and Hollenberg, 1976:fig. 66).

For now, C. macdougalii is still recognized as a species, and will be until more Gulf collections have been 

made and morphological and molecular analyses have 

been completed and compared to other known species.

Cladophora microcladioides Collins

FIGURE 30A,B

Cladophora microcladioides Collins, 1909a:17, pl. 78: 

figs. 2, 3; Setchell and Gardner, 1920b:212, pl. 13: fig. 2; 

Dawson, 1944:209; Smith, 1944:59, pl. 7: fig. 3; Dawson, 1945b:60, 64; Doty, 1947:23; Dawson, 1949:222, 

224, 225; Dawson, 1961b:378; Dawson, 1962c:278; 

Dawson, 1966a:6; Abbott and Hollenberg, 1976:106, 

fig. 65; Devinny, 1978:538; R. Aguilar-Rosas, 1982:84; 

L. Aguilar-Rosas and Bertsch, 1983:116, 121; Stewart and Stewart, 1984:140; Huerta-Múzquiz and 

Mendoza-González, 1985:42; Mendoza-González and 

Mateo-Cid, 1985:22; L. Aguilar-Rosas and PachecoRuíz, 1986:77; Huerta Muzquiz, 1978:336; MendozaGonzález and Mateo-Cid, 1986:419; Sánchez-Rodríguez et al., 1989:39; Dreckmann et al., 1990:24, 37; 

Mateo-Cid and Mendoza-González, 1991:27; Stewart, 

1991:31; González-González, 1993:443; Mateo-Cid et 

al., 1993:51; León-Tejera et al., 1993:199, 204; R. Aguilar-Rosas and M. Aguilar-Rosas, 1994:517; MateoCid and Mendoza-González, 1994a: 52, 61; MendozaGonzález et al., 1994:103, 112; González-González et 

al., 1996:130; Pacheco-Ruíz and Zertuche-González, 

1996a:432; Anaya-Reyna and Riosmena-Rodríguez, 

1996:862; Bucio-Pacheco and Dreckmann, 1998:43; 

Núñez-López et al., 1998:39; Mendoza-González et al., 

1999:61; Rodríguez-Morales and Siqueiros-Beltrones, 

1999:17, 22; L. Aguilar-Rosas et al., 2000:133; LeónTejera and González-González, 2000:327; López et 

al., 2000:339; Paul-Chávez and Riosmena-Rodríguez, 

2000:146; L. Aguilar-Rosas et al., 2002:235; Wysor, 

2004:217; R. Aguilar-Rosas et al., 2005b:36; Hernández-Herrera et al., 2005:146; Pedroche et al., 2005:45; 

Riosmena-Rodríguez et al., 2005:101; Mateo-Cid et al., 

2006:48; Pacheco-Ruíz et al., 2008:201.

Algae loosely tufted, up to 10 cm tall or more. 

Lower filaments 120–170(–200) μm in diameter with 

cells 3–4(–5) times as long as wide; becoming narrower 

above, 60–120 μm in diameter, with cells 2–4 times as 

long as wide; ultimate branchlets 30–50 μm in diameter 

and tapered slightly to rounded apices. Branching dichotomously or trichotomously from main axis, becoming 

irregularly alternate to pectinate above; lateral branches 

progressively shorter toward apex, with shorter secund 

branchlets along adaxial side; ultimate branchlets sometimes recurved.

Habitat. On rocks; mid intertidal to shallow

subtidal to 4(–32) m depths.

Distribution. Gulf of California: Puerto

Peñasco to Cabeza Ballena. Pacific coast: Alaska to Michoacán; Oaxaca; Panama.

Type Locality. San Pedro, Los Angeles

County, California.

Cladophora montagneana Kützing

Cladophora montagneana Kützing, 1849:415; van den 

Hoek, 1982:105, figs. 145–173; van den Hoek and 

Womersley, 1984:204, figs. 66B, 67E–I; Bucio-Pacheco 

and Dreckmann, 1998:43; Pedroche et al., 2005:46.
n u m b e r 9 4 • 6 1

Conferva brachyclados Montagne, 1837:349; Montagne, 

1842a:13, pl. 4 [type of Cladophora montagneana; see 

van den Hoek, 1982:105].

Cladophora brachyclados (Montagne) Harvey, 1858:81.

Cladophora delicatula Montagne, 1850:302; Kützing, 1856: pl. 1: fig. 2; Dawson, 1944:210; Dawson, 

1961b:377; Dawson, 1966a:5 [with a query]; GonzálezGonzález et al., 1996:281; Bucio-Pacheco and Dreckmann, 1998:44; Wysor, 2004:217; Mateo-Cid et al., 

2006:48 [with a query].

Algae delicate hair-like tufts, of narrow filaments up to 

2 cm long; branching often widely spaced, pseudodichotomous to irregular, sometimes in unilateral rows; attached 

by branched basal rhizoids. Cells of main axis and lower 

branches usually 40–50 μm in diameter and 3–4(–5) times 

longer than wide (mostly up to 200 μm long); upper cells 

shorter in length than lower cells, 18–25(–30) μm in diameter, and 1–2 times longer than wide.

Habitat. On rocks and tidal flats; intertidal

Distribution. Gulf of California: Puerto

Peñasco; Rio Mayo. Pacific coast: Michoacán; Panama.

Type Locality. Cuba

Remarks. Collections of Cladophora montagneana in the vicinity of Puerto Peñasco were referred to 

this species with a query by Dawson (1966a, as “C. delicatula”). Although this species has been reported as “C. 

delicatula,” on the Pacific coast from Oregon to southern California (Doty, 1947; Dawson, 1961b), Abbott and 

Hollenberg (1976:105) concluded that Pacific specimens 

identified as “C. delicatula sensu Collins (1909b; Smith, 

1944)” were misidentified and were another species, 

C. albida. Thus, the presence of C. montagneana in the 

northern Gulf needs to be verified. Crystalline cell inclusions were found in C. montagneana from Tanzania (Leliaert and Coppejans, 2004:202, fig. 48), but have not been 

reported in the Gulf “C. montagneana.”

The name C. montagneana given by Kützing (1849) 

was necessary since a new combination of the earliest name 

for this taxon, Conferva brachyclados Montagne (1837), 

FIGURE 30. Cladophora microcladioides: A. Branching pattern of an upper branch (after Setchell and Gardner, 1920b: pl. 13, fig. 2). B. Upper 

lateral branches with pectinate branching pattern (EYD-2928).
6 2 • 

would have created a later homonym of Cladophora 

brachyclados (Kützing) Kützing (1845; basionym: Conferva crispata var. brachyclados Kützing, 1834).

Cladophora prolifera (Roth) Kützing

Confera prolifera Roth, 1797:182, pl. III: fig. 2.

Cladophora prolifera (Roth) Kützing, 1843:271; Kützing, 1853:24, pl. 82: fig.3; Taylor, 1945:57; Dawson, 

1949:230; Dawson, 1957b:8; Dawson, 1961b:378; 

Chávez B., 1972:268; van den Hoek, 1963 [in part, excluding C. rugulosa]:208, pl. 51: figs. 677–682, pl. 52: 

fig. 686; van den Hoek, 1982:166, pl. 32: figs. 318–

327; Mateo-Cid and Mendoza-González, 1991:27; 

Mateo-Cid and Mendoza-González, 1991:24; Mendoza-González and Mateo-Cid, 1992:24; Mateo-Cid 

and Mendoza-González, 1992:24; Mateo-Cid et al., 

1993:51; González-González, 1993:443; Stout and 

Dreckmann, 1993:4; Servière-Zaragoza et al., 1993:482; 

Mateo-Cid and Mendoza-González, 1994b:34; Mendoza-González et al., 1994:112; González-González et 

al., 1996:369; Bucio-Pacheco and Dreckmann, 1998:43; 

Mendoza-González and Mateo-Cid, 1998:25, 27; L. 

Aguilar-Rosas et al., 2000:129, 133, 134; van den Hoek 

and Chihara, 2000 [excluding C. rugulosa]:52, fig. 22; 

Mateo-Cid et al., 2000a:61, 69; L. Aguilar-Rosas et al., 

2002:235; Leliaert and Coppejans, 2003:56, figs. 6A–

C, 7; Leliaert, 2004:32, figs. 6A–C, 7; Pedroche et al., 

2005:47; Mateo-Cid et al., 2006:58.

Algae dark green, forming dense tufts of branched 

filaments, up to 3 cm tall; above a distinctive stipe-like 

region of entangled, simple to branched, rhizoids (with 

annular constrictions) that descend from lower portions of 

basal cells, and form a rhizoidal attachment mat. Branches 

develop on apical end of axis and branch cells. Cells of 

main axes, branches, and basal cells (often with annular 

constrictions), 150–300 μm in diameter and 7–10 times 

as long as wide (1050–3000 μm in length); apical cells, 

95–220 μm in diameter and 2.5–8 times as long as wide 

(320–1760 μm in length), with rounded tip. Rhizoids simple or branched, 40–100 μm in diameter, with characteristic annular constrictions.

Habitat. On rocks and in tide pools, generally

in shady areas; low intertidal to shallow subtidal.

Distribution. Gulf of California: El Tornillal

to Bahía Concepción; Cabo Pulmo. Pacific coast: Laguna 

Ojo de Liebre (Scammon’s Lagoon), Baja California Sur to 

Oaxaca; Costa Rica; Ecuador; Galápagos Islands; China 

(Hainan Island); Taiwan; Japan; Vietnam.

Type Locality. “In mare Corsicam” (Roth,

1797). Lectotype: Miramare, Trieste, Italy (van den Hoek, 

1963).

Remarks. Cladophora prolifera has been 

recently reported in the Gulf of California (Mateo-Cid 

et al., 1993; L. Aguilar-Rosas et al., 2000, 2002). The 

description follows that of van den Hoek and Chihara 

(2000), who noted this species was predominately subtidal and widely distributed in tropical to warm-temperate 

waters of the Atlantic, Pacific, and Indian Oceans. Leliaert 

and Coppejans (2004:202) have reported another taxonomic feature, crystalline cell inclusions, in C. prolifera

from Turkey. The relationship of C. prolifera to a similar 

species, Cladophora rugulosa G. Martens (1868) has been 

uncertain, e.g., the latter was considered to be a synonym 

of C. prolifera by van den Hoek (1963). Recently, Leliaert 

and Coppejans (2003; see also Leliaert, 2004) provided 

morphological evidence that these species are distinct, 

thus supporting others who have kept them as separate 

species (e.g., Papenfuss, 1943; Sakai, 1964; Papenfuss and 

Chihara, 1975; Yoshida, 1998).

Cladophora sericea (Hudson) Kützin

Conferva sericea Hudson, 1762:485.

Cladophora sericea (Hudson) Kützing, 1843:264; van den 

Hoek, 1963 [in part]:77, pl. 17: figs. 184–189, pl. 18: figs. 

190–209, pl. 19: figs. 210–226, 245; Abbott and Hollenberg, 1976:108, fig. 67; van den Hoek, 1982:93, pl. 

15: figs. 138–143; Mendoza-González and Mateo-Cid, 

1985:22; L. Aguilar-Rosas and Pacheco-Ruía, 1986:77; 

Dreckmann et al., 1990:24, pl. 2: fig. 4; Mateo-Cid and 

Mendoza-González, 1991:27; Stewart, 1991:31; MateoCid and Mendoza-González, 1991:27; Mateo-Cid and 

Mendoza-González, 1992:24; Servière-Zaragoza et al., 

1993:482; Mateo-Cid and Mendoza-González, 1994a:52; 

Mendoza-González et al., 1994:112; González-González 

et al., 1996:281, 410; Mendoza-González and MateoCid, 1996:75, 87, pl. 26: figs. 114–118; Galindo-Villegas 

et al., 1997:2, 4; Mateo-Cid and Mendoza-González, 

1997:60, pl. 8: figs. 32, 33; Bucio-Pacheco and Dreckmann, 1998:43; Núñez-López et al., 1998:39; MendozaGonzález and Mateo-Cid, 1998:25; Mateo-Cid et al., 

2000a:59, 69; Abbott and Huisman, 2004:77, fig. 22E; 

Pedroche et al., 2005:48; Dreckmann et al., 2006:154.

Conferva rudolphiana C. Agardh, 1827:636.

Cladophora rudolphiana (C. Agardh) Kützing, 1843:268; 

Collins, 1909b:336; Setchell and Gardner, 1920b:218; 

Dawson, 1944:209; Dawson, 1957b:8; Dawson, 

1961b:378; González-González et al., 1996:131.
n u m b e r 9 4 • 6 3

Cladophora gracilis sensu Doty, 1947:23; Dawson, 

1961b:377; Sakai, 1964:33–38 [non Cladophora gracilis Kützing, 1845:763 (note that Cladophora gracilis (A. 

W. Griffiths ex Mackay) Kützing, 1845:215 is nom. illeg. since the basionym Conferva gracilis A. W. Griffiths 

ex Harvey, 1835:304 is also nom. illeg. as it is a later 

homonym of Conferva gracilis Wulfen, 1803:21; thus, 

the correct new name is Cladophora gracilis Kützing, 

1845:763, which is now a synonym of Cladophora 

flexuosa (O. F. Müller) Kützing, 1843:270; basionym: 

Conferva flexuosa O. F. Müller, 1782: 5, pl. 882 (see 

Söderström, 1963, 1965; Bot et al., 1989; Silva et al., 

1996a; van den Hoek and Chihara, 2000:164)].

Algae up to 5 cm tall, filaments of main axis branched 

sparingly, widely spaced, branches of different lengths 

intermixed; cells of axis 50(–85) μm in diameter and 5–6 

times as long as wide (250–510 μm in length); cells of ultimate branches narrower, 20–30 μm in diameter and 5–7 

times as long as wide (100–210 μm in length).

Habitat. On rocks in tidal creeks and epiphytic on drift Gracilaria; intertidal to shallow subtidal

Distribution. Gulf of California: Empalme,

Guaymas; Punta Arena to Cabo Pulmo. Pacific coast: 

Alaska to Chiapas; Costa Rica; Hawaiian Islands.

Type Locality. Isle of Sheppey, Kent,

England.

Cladophora tiburonensis E. Y. Da

FIGURE 26B

Cladophora tiburonensis E. Y. Dawson, 1944:211, pl. 

31: figs. 1–3; Dawson, 1949:247; Dawson, 1961b:378; 

Espinoza-Ávalos, 1993:333; González-González et al., 

1996:131; Pedroche et al., 2005:48.

Algae forming dense tufts of short stiff filaments, up to 

4 mm tall, branching subdichotomous to secund; branch 

diameter gradually decreasing upward. Basal cells up to 

60 μm in diameter; upper cells about 25 μm in diameter; 

cells mostly 4–5 times as long as wide, occasionally much 

longer below.

Fertile cells 40–70 μm in diameter, conspicuously 

shorter and swollen compared to slender vegetative cells.

Habitat. In tide pools; mid to low intertidal

Distribution. Gulf of California: Isla Turners; Mazatlán, Sinaloa

Type Locality. Isla Turners, off southeast end

of Isla Tiburón, Las Islas de la Cintura, Gulf of California.

Remarks. Cladophora tiburonensis is not well 

known. Dawson (1944), while noting its resemblance to 

C. graminea Collins (1909a), set it apart on the basis of its 

minute size and stiff habit. It is retained here to point out 

that its taxonomic status needs to be investigated.

Cladophora vagabunda (Linnaeus

van den Ho

Conferva vagabunda Linnaeus, 1753:1167; van den Hoek, 

1963: pl. 36: figs. 470, 471 [type illustrations].

Cladophora vagabunda (Linnaeus) van den Hoek, 

1963:144, pl. 33: figs. 434, 436–439; pl. 36: figs. 

470–490, pl. 37: 491–503, pl. 39: 505–514; Schnetter 

and Bula Meyer, 1982:26, pl. 3: figs. H, I; van den 

Hoek, 1982:137, figs. 264–294; Mateo-Cid and Mendoza-Gonzalez, 1991:27; Mateo-Cid and MendozaGonzález, 1992:24; Servière-Zaragoza et al., 1993:482; 

van den Hoek and Chihara, 2000:180, figs. 76–81; 

L. Aguilar-Rosas et al., 2002:235; Leliaert and Coppejans, 2003:70, figs. 14A,B, 15; Abbott and Huisman, 

2004:79, figs. 24A–D; Leliaert, 2004:47, figs. 14A,B, 

15; Pedroche et al., 2005:49; Hernández-Herrera et al., 

2005:307; Mateo-Cid et al., 2006:58.

Conferva expansa Mertens ex Jürgens, 1817:Decade 5: 

[exsiccate] No. 8.

Cladophora expansa (Mertens ex Jürgens) Kützing, 

1843:265; Chávez B., 1972:268.

Conferva fascicularis Mertens ex C. Agardh, 1824:114.

Cladophora fasicularis (Mertens ex C. Agardh) Kützing, 

1843:268; Collins, 1909b:345; Chapman, 1956:447, 

fig. 98; Sakai, 1964:25, fig. 8A–D, pl. 4: fig. 1; Yoshida, 

1998:62.

Cladophora uncinella Harvey, 1860:334; Dawson, 

1959b:28; Sakai, 1964:60, fig. 28, pl. 11: fig. 1; Yoshida, 1998:70; van den Hoek and Chihara, 2000:182.

Algae erect tufts, up to 5 cm tall, main axes pseudodichotomously to pseudotrichotomously branched, up to 

3 orders; branchlets sometimes straight filaments, but usually clusters of short, secund or recurved branchlets, with 

relatively straight to curled apical cell ends; attached by 

branched rhizoids from basal end of lower cells, which may 

partly coalesce. Main filaments 140–260(–300) μm in diameter and (3–)4–8 times longer than wide; apical cells much 

smaller, (20–)30–50 μm in diameter and 2–6(–9) times longer than wide. Cell walls of upper cells mostly 1.0–2.0 μm 

thick; much thicker in lower cells, up to 20 μm thick.

Habitat. On rocks, limestone-clay ledges, and

mudflats and in tide pools, lagoons, and salt marshes; 
6 4 • 

sometimes unattached and floating or may be epiphytic or 

entangled with other algae or aquatic plants; high to low 

intertidal.

Distribution. Gulf of California: Piedras

de La Salina to Puerto Peñasco. Pacific coast: Nayarit to 

Guerrero; Colombia; Hawaiian Islands; Japan; Korea; 

China; Vietnam.

Type Locality. Selsey, Sussex, England (lectotype; van den Hoek, 1963)

Remarks. Cladophora vagabunda and a few 

other taxa sensu van den Hoek (1963) were considered 

to be forms of a single species, C. flexuosa (O. F. Müller) Kützing (1843), by Söderström (1965). Since then, 

however, C. vagabunda has continued to be recognized 

as a distinct taxon (e.g., Silva et al., 1996; Leliaert, 2004). 

DNA-DNA hybridization experiments of Bot et al. (1990) 

and molecular analysis of Bakker et al. (1995) on widely 

geographically separated isolates of C. vagabunda have 

shown considerable divergence, with at least four cryptic 

species. Specimens of Gulf C. vagabunda need to be similarly tested to elucidate their taxonomic status.

Rhizoclonium Kützing

Rhizoclonium Kützing, 1843:261.

Algae are uniseriate, slender, cylindrical filaments, 

up to 60 μm in diameter, that are unbranched and may 

lack or have one- to few-celled rhizoidal laterals. They 

are either attached by a lobed holdfast or lay unattached 

on hard substratum or, more commonly, on mud or sand 

flats. Cells are mostly several times longer than wide and 

contain numerous nuclei and a parietal, reticulate chloroplast with pyrenoids.

Vegetative reproduction is by fragmentation of the 

filaments. It is also by akinetes that are formed within 

some of the cells as they swell and thicken. Life histories, 

where known, are an alternation of isomorphic generations. Asexual reproduction is by biflagellate or quadriflagellate zoospores. Sexual reproduction is by biflagellate 

isogametes.

Remarks. Rhizoclonium occurs worldwide in 

fresh, brackish, and marine waters. The generic distinction 

of Rhizoclonium and Lola A. Hamel et G. Hamel (1929) 

is tenuous and needs to be investigated. For example, the 

only character separating them is isogamous reproduction 

in Rhizoclonium and anisogamous reproduction in Lola. 

Likewise, the generic differences between Rhizoclonium 

and Chaetomorpha, as well as their species separation, 

can be difficult. The morphological characters used, i.e., 

filament diameter, length/width ratio of cells, the absence 

or presence of attachment structures, and the absence or 

presence and number of lateral rhizoidal filaments, have 

been shown to be highly variable (e.g., Koster, 1955; 

Nienhuis, 1975; Nienhuis and de Bree, 1981; Blair, 1983). 

Setchell and Gardner (1920b) concluded that the only useful taxonomic character was filament cell diameter.

One species has been reported in the southern Gulf: 

Rhizoclonium hieroglyphicum (C. Agardh) Kützing (1845) 

by Huerta-Múzquiz and Mendoza-González (1985).

There are two species and two varieties known in the 

northern Gulf of California.

Key to the Species of Rhizoclonium in the nor

1a. Uniseriate filaments mostly straight; cells mostly greater than 30 μm in diameter, up to 75 μm wide; filament cells either 

without or with short rhizoidal proliferations of only a single cell  . . . . . .

1b. Uniseriate filaments mostly twisted or curly; cells mostly less than 30 μm in diameter (occasionally up to 45 μm); filament 

cells either without or with tapering rhizoidal proliferations of 2–5 (or more) cells long . . . . . . . . . . . . . . . . . . . . . . . . . .2

2a. Cells mostly 20–30(–45) μm in diameter and 20–60(–90) μm long; filaments sometimes with few rhizoids . . . . . . . . . . . .

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

2b. Cells smaller in diameter, usually 8–12(–20) μm, 24–50(–70) μm long; filaments rarely with rhizoids . . . . . . . . . . . . . . . .

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

Rhizoclonium riparium (Roth)

Harvey var. ripariu

Conferva riparia Roth, 1806:216; Koster, 1955:336, fig. 1.

Rhizoclonium riparium (Roth) Harvey, 1849:238, pl. 

CCXXXVIII; Setchell and Gardner, 1920b:182; Smith, 

1944:63, pl. 7: fig. 4; Taylor, 1945:55; Doty, 1947:27; 

Hollenberg, 1948:155; Dawson, 1961b:376; Scagel, 

1966:74, fig. 18A–F; Nienhuis, 1975:84–88, figs. 35–

60; Abbott and Hollenberg, 1976:92, fig. 46; Zedler, 

1980:124; Nienhuis and de Bree, 1981:434; R. Aguilar-Rosas, 1982:84; Zedler, 1982:41; Schnetter and 
n u m b e r 9 4 • 6 5

Bula Meyer, 1982:18, pl. 1: figs. Q–S, pl. 2: figs. A, 

B; Blair, 1983:203; Huerta-Múzquiz and MendozaGonzález, 1985:42; Mendoza-González and MateoCid, 1986:419; Mateo-Cid and Mendoza-González, 

1991:27; Stewart, 1991:32; Mateo-Cid and MendozaGonzález, 1992:25; León-Tejera et al., 1993:199; 

Servière-Zaragoza et al., 1993:482; Mateo-Cid and 

Mendoza-González, 1994b:44; Mendoza-González et 

al., 1994:112; Pedroche et al., 1995:112; GonzálezGonzález et al., 1996:289; Mendoza-González and Mateo-Cid, 1996:76, 88, pl. 27: fig. 122; Sánchez-Lizaso 

and Riosmena-Rodríguez, 1997:56; Núñez-López et 

al., 1998:39; L. Aguilar-Rosas et al., 2000:133; PaulChávez and Riosmena-Rodríguez, 2000:146; CruzAyala et al., 2001:190; Abbott and Huisman, 2004:83, 

fig. 25D; Wysor, 2004:218; Pedroche et al., 2005:50; 

Hernández-Herrera et al., 2005:146; Dreckmann et al., 

2006:154.

Algae unbranched, uniseriate filaments of cylindrical 

cells, mostly 1–2 times longer than wide (sometimes up to 

6 times); (10–)15–30(–45) μm in diameter by 15–90(–270) 

μm long; without or with a few short rhizoidal proliferations of 2–5 (or more) cells; often unattached. Chloroplast 

reticulate, with 6 or more pyrenoids.

Vegetative reproduction by fragmentation. Reproduction not reported in Gulf specimens. Reproduction has 

been reported in Europe. Asexual reproduction is by biflagellate and quadriflagellate zoospores, and sexual reproduction by isogamous gametes (Nienhuis, 1975).

Habitat. Entangled with other algae or aquatic

plants, in tide pools, or on sand flats and mudflats of salt 

marshes, lagoons, and estuaries; or free-floating and unattached (also reported as epiphytic on Gelidium johnstonii

and Spyridia filamentosa; L. Aguilar-Rosas et al., 2000); 

high to low intertidal.

Distribution. Gulf of California: Puertecitos to Punta Arena (north of Cabo Pulmo). Pacific coast:

Alaska to Chiapas; Panama; Colombia; Galápagos Islands; 

Chile; Hawaiian Islands; China; Japan.

Type Locality. Norderney, East Frisian

Islands, Germany.

Rhizoclonium riparium var. implexum 

(Dillwyn) Rosenvinge

Conferva implexa Dillwyn, 1809:46, (Suppl.) pl. B; 

Koster, 1955:336, fig. 2.

Rhizoclonium riparium var. implexum (Dillwyn) 

Rosenvinge, 1893:915, fig. 34; Taylor, 1945:55; Silva 

et al., 1996a:786; Mateo-Cid et al., 2000a:69; LeónTejera and González-González, 2000:327; Pedroche et 

al., 2005:50; Servière-Zaragoza et al., 2007:8.

Rhizoclonium implexum (Dillwyn) Kützing, 1845:206; 

Setchell and Gardner, 1920b:183; Smith, 1944:62, 

pl. 8: fig. 3; Taylor, 1945:55; Koster, 1955: 344, 348; 

Dawson, 1961b:375; Scagel, 1966:73, pl. 33A,B; Abbott and Hollenberg, 1976:92, fig. 45; Huerta-Múzquiz 

and Mendoza-González, 1985:42; Mendoza-González 

and Mateo-Cid, 1985:22; Mendoza-González and Mateo-Cid, 1986:419; Mateo-Cid and Mendoza-González, 

1994a:57; González-González et al., 1996:138; 

Galindo-Villegas et al., 1997:2, 4; Yoshida, 1998:72; 

Abbott and Huisman, 2004:82, fig. 25C.

Lola implexa (Dillwyn) G. Hamel, 1931a:120, fig. 37 (6–

7); Ortega et al., 1987:78, pl. 14: figs. 60–62; Perrot, 

1965:506; González-González et al., 1996:137.

Rhizoclonium kerneri Stockmayer, 1890:582; Collins, 

1909b:329; Setchell and Gardner, 1920b:185; Taylor, 

1945:55; Dawson, 1961b:375; González-González, 

1993:443; Ortega et al., 1987:77, pl. 13: figs. 58, 59; 

Servière-Zaragoza et al., 1993:485; González-González 

et al., 1996:289; Servière-Zaragoza et al., 1998:169.

Rhizoclonium kochianum Kützing, 1845:206; Dawson, 

1959a:12; Dawson, 1960a:31; Dawson, 1961b:376; 

Earle, 1972:85; González-González et al., 1996:139.

Algae unbranched, uniseriate filaments of cylindrical 

cells, mostly 3–5 times longer than wide, (8–)10–12(–20) 

μm in diameter by (24–)30–50(–70) μm long; mostly lacking or rarely with short, rhizoidal proliferations. Chloroplasts reticulate, with numerous pyrenoids.

Habitat. Epiphytic or entangled with other

algae or plants, on rocks, in tide pools, or on sand and 

mudflats of salt marshes, lagoons, and estuaries; or unattached and free-floating; high to low intertidal.

Distribution. Gulf of California: Guaymas

to Punta Arena (north of Cabo Pulmo). Pacific coast: 

Alaska to Oaxaca; Isla Clarión (Islas Revillagigedo); Panama; China; Japan; Hawaiian Islands; Galápagos Islands.

Type Locality. Bantry, Ireland

Remarks. The taxonomy of Rhizoclonium 

implexum has been diversely interpreted. Koster (1955) 

placed R. kerneri and R. kochianum in synonymy with R. 

implexum and also questioned whether R. implexum was 

distinct from R. riparium or if the two were one highly 

variable species. Since then, these latter two have been variously treated: as separate species (e.g., Abbott and Hollenberg, 1976; Womersley, 1984; Yoshida, 1998; Abbott 

and Huisman, 2004); as a single species with a variety, 
6 6 • 

R. riparium var. implexum (Dillwyn) Rosenvinge, 1893 

(e.g., Silva et al., 1996a; Pedroche et al., 2005); and as 

an unnamed form of R. riparium (Blair, 1983). Herein, 

while treated as a single species following Nienhuis 

(1975), Nienhuis and de Bree (1981), Blair (1983), Burrows (1991), and John et al. (2004), the two varieties are 

recognized until further studies can be done.

Rhizoclonium tortuosum (Dillwyn) Kützing

FIGURE 31

Conferva tortuosa Dillwyn, 1805:pl. 46.

Rhizoclonium tortuosum (Dillwyn) Kützing, 1845:205; 

Setchell and Gardner, 1920b:185; Chapman, 1939:19; 

Doty, 1947:25; Dawson, 1961b:376; Blair, 1983:198, 

figs. 12, 13; Hoffman and Santelices, 1997:91, fig. 19; 

Bucio-Pacheco and Dreckmann, 1998:44; Pedroche et 

al., 2005:51.

Chaetomorpha tortuosa (Dillwyn) Kleen, 1874:45; Scagel 

et al., 1989:62.

Lola tortuosa (Dillwyn) Chapman, 1953:65; Chapman, 

1956:463.

Rhizoclonium lubricum Setchell et N. L. Gardner in N. 

L. Gardner, 1919:492, pl. 42: fig. 5A,B; Setchell and 

Gardner, 1920b:185, pl. 9: fig. 5a,b; Taylor, 1945:56; 

Doty, 1947:26.

Lola lubrica (Setchell et N. L. Gardner in N. L. Gardner) 

A. Hamel et G. Hamel, 1929:1094; Hamel, 1931a:118, 

fig. 37 (1–5); Dawson, 1944:208; Dawson, 1961b:376; 

Abbott and Hollenberg, 1976:92, fig. 47; Ortega et al., 

1987:78, pl. 14: figs. 63, 64; Scagel et al., 1989:74; 

Stewart, 1991:32; González-González et al., 1996:137; 

Paul-Chávez and Riosmena-Rodríguez, 2000:146; Pedroche et al., 2005:49.

Rhizoclonium riparium f. validum Foslie in Wittrock et 

Nordstedt, 1884:No. 624; Foslie, 1890:139.

Algae of slippery filaments, up to 40 cm long, often 

entangled or forming free-floating mats; composed of 

unbranched filaments, (25–)30–75 μm in diameter; newer 

cells (25–)35–100 μm long; mature cells 120–300 μm long; 

attached by simple short rhizoids. Cells with a reticulate 

chloroplast, and numerous pyrenoids; cell wall 0.2 μm 

thick.

Vegetative reproduction by fragmentation. Reproduction not observed in Gulf material. A life history involving an isomorphic alternation of generations has been 

reported, with asexual reproduction by quadriflagellate 

zoospores, and sexual reproduction that is anisogamous, 

with biflagellate gametes (Hamel and Hamel, 1929; Blair, 

1983).

Habitat. On rocks or other hard substrata,

often entangled with other algae; on mud-sand flats in 

lagoons and shallow bays; or sometimes unattached and 

free-floating; mid to low intertidal.

Distribution. Gulf of California: Guaymas;

Bahía San Gabriel; Isla Espíritu Santo; Laguna de Agiabampo (Sonora). Pacific coast: Alaska to Michoacán.

Type Locality. Lectotype: Swansea, Glamorgan, Wales, UK (Blair, 1983:198)

Remarks. Rhizoclonium tortuosum has been 

variously treated; some recognize it as a distinct species, 

others recognize it as part of a species complex. For example, Burrows (1991) considered R. riparium to be a single 

species conspecific with R. tortuosum and R. implexum. 

Herein, R. tortuosum is tentatively recognized as a separate species until detailed comparative morphological 

studies and molecular analyses have been conducted on 

Gulf material for comparisons to the European types of 

these taxa (see Silva et al., 1996a:936 for discussion of 

nomenclature and taxonomic problems of R. tortuosum).

FIGURE 31. Rhizoclonium tortuosum: Terminal portion of young 

filament with short segments (left) and terminal portion of filament 

with long segments (right) (after Gardner, 1919: pl. 42, fig. 5A,B).
n u m b e r 9 4 • 6 7

Siphonocladal

Siphonocladales (Blackman et Tansley) Oltmanns, 1904: 

134.

Algae are of various forms, composed of large tubular, siphonous, or vesiculate cells that are formed by either 

lenticular cells, or by internal segregative cell division (not 

always obvious). In segregative division the protoplast 

cleaves into protoplasmic spheres, which, in turn, may 

either form new cell walls, remain in the parental cell, or 

be released as vegetative propagules. Endogenous expansion of these spheres within the parental vesicle results 

in the formation of pseudoparenchymatous tissue (e.g., 

Dictyosphaeria). Cells divide in some members by centripetal invagination of cell walls, but with new branches 

that remain in open connection (i.e., lack septa) at the 

base of the parent cell (e.g., Cladophoropsis). New lateral 

branches may also arise in some genera by expansion of 

lenticular cells; i.e., septa are initiated prior to elongation 

of the vesicular branch on the surface of the parent vesicle 

(e.g., Valonia).

Cells are cylindrical or inflated, globose, clavate, or 

vesiculate. There is simultaneous formation of laterals at 

nodes (rather than sequential), and branching can be in 

three directions. Tenacular cells that are formed are one of 

four different kinds, and secondary anastomoses are usually formed.

Remarks. The taxonomy of the families, as 

well as their generic composition, of the CladophoralesSiphonocladales complex is unresolved (see Leliaert 

et al., 2003, 2007, and Remarks under Cladophorales 

herein). Although segregative cell division is a primary 

characteristic of the families Siphonocladaceae and the 

Boodleaceae, it has not been reported or described for all 

the genera generally considered to be members of these 

families.

Three of the currently accepted families in the 

Siphonocladales are represented in the northern Gulf of 

California.

Key to Families of Siphonocladales in the

1a Thalli erect, of relatively large, elongated cylindrical cells that bear irregularly to regularly, radially arranged lateral or 

whorled branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphonocladacea

1b. Thallus a large individual vesicle; or cushionlike, of simple to branched, cylindrical to clavate, siphon cells  

2a. Thalli variable, filamentous to netlike; of branched, cylindrical or siphon cells secondarily attached to each other; cells 

formed either by segregative division or by centripetal invagination of cell walls . . . . . . . . . . . . . . . . . . . . . . Boodleaceae

2b. Thallus a single globose vesicle, or composed of elongate siphon cells, some with branches, usually entangled,; main axis 

not distinct; growth by lenticular cells  . . . . .

Boodl

Boodleaceae Børgesen, 1925:19, 24.

Algae are of various morphologies; mostly, they are 

tufts, clumps, or mats of clavate, cylindrical, or filiform 

cells with vesicular branches that often become constricted 

near their base, and are usually attached to the substratum 

by rhizoids. Development of new cells, forming laterals 

or branches, is primarily by segregative cell division, i.e., 

cytoplasm divides into portions that expand into protrusions before a cell wall is formed, or, in some, is by centripetal invagination of the cell walls. Some may also initiate 

new growth by the formation of lenticular cells. Cells are 

multinucleate with reticulate chloroplasts that lack pyrenoids. Crystalline cell inclusions may be present or absent.

Reproduction is poorly known. Life histories, in those 

studied, are an alternation of isomorphic generations. 

Biflagellate and quadriflagellate zoospores have been 

observed in only a few of the genera. Sexual reproduction, 

where known, is by biflagellate isogametes.

Remarks. Members of the Boodleaceae found 

in the Gulf of California have usually been referred to the 

Siphonocladaceae (e.g., Pedroche et al., 2005). However, 

the type genus, Siphonocladus F. Schmitz (1879), does 

not group with the other genera generally assigned to 

the family in molecular phylogenetic analyses (Leliaert et 

al., 2003; Leliaert and Coppejans, 2006). Although some 

genera such as Phyllodictyon have been included in the 

Anadyomenaceae Kützing (1843), there is no phylogenetic 

support for this, and that family is restricted to the genera 

Anadyomena J. V. Lamouroux (1812) and Microdictyon

Decaisne (1841) as originially proposed by Kützing (Leliaert et al., 2007). Therefore, as noted by Kraft (2007) and 

Leliaert (University of Ghent, personal communication, 

2008), the family Boodleaceae is available for the genera 
6 8 • 

other than Siphonocladus, Anadyomena, and Microdictyon. Herein the Boodleaceae is interpreted to include 

Boodlea, Struvea, Cladophoropsis, and Phyllodictyon.

Two genera of this family have been recorded for the 

southern Gulf: Struvea Sonder (1845) by Huerta-Múzquiz 

and Mendoza-Gonzalez (1985, as “Struvea sp.”) and 

Boodlea (Dickie) G. Murray et DeToni (in Murray, 1889) 

with the report of B. composita (Harvey) Brand (1904) 

by Mateo-Cid et al. (1993) and Mendoza-González et al. 

(1994).

Two of the genera are known in the northern Gulf of 

California.

Key to the Genera of Boodleaceae in the n

1a. Algae of terete, branched filaments (not netlike); branches formed singly on distal end of filament and elongate without 

forming a basal crosswall (septae)  . . . . . . 

1b. Algae a simple or branched stalk(s) with an upper netlike portion (formed by attachment of tenacular cells of tips of apical 

cells of lateral branches to adjacent filaments); or the upper portion only partially netlike (reduced or incomplete or sometimes lacking) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllodictyo

Cladophoropsis Børgesen

Cladophoropsis Børgesen, 1905:288.

Algae generally form green, cushionlike turfs or mats 

of interweaving filaments, usually lack a distinct axis, and 

attach to substratum by branched septate rhizoids, adherent cell walls, or hapteroid cells. The filaments, more or 

less uniform in diameter, have irregularly spaced, usually sparse, lateral branches. Laterals are produced from 

nonseptate protrusions at the distal end of intercalary and 

apical cells that elongate to various lengths and gradually 

taper toward the apex. Septa (crosswalls), although initially 

lacking, can develop secondarily at intercalary and branch 

points. Cells are multinucleate, with numerous discoid 

chloroplasts, each has a single pyrenoid; crystalline inclusions present. Cells divide by centripetal invagination of 

cell walls. Growth is by apical and intercalary cell division. 

Annular constrictions and lenticular cells are absent. Rhizoids, produced above the septum, can become well developed and produce new axes. Adventitious rhizoids develop 

where branches come in contact with the substratum.

Propagation is by aplanospores, probably produced 

by segregative division. Life histories are presumed to be a 

biphasic alternation of generations.

Remarks. Generic limits of Cladophoropsis are unclear, and only a few of the described species 

are apparently distinctive. The taxonomy is complicated 

by having few morphological characters, two of which, 

growth form and branching pattern, are environmentally 

influenced (e.g., Leliaert et al., 2003). There has also been 

confusion in the interpretation of septate and nonseptate 

lateral branches. Further problems were found in the 

species sampling in phylogenetic studies (Kooistra et al., 

1993). Leliaert and Coppejans (2004) noted the absence 

or presence of different morphological types of crystalline cell inclusions to be of systematic value at the species 

level.

One species, Cladophoropsis membranacea (Hofman Bang ex C. Agardh) Børgesen (1905; Leliaert and 

Coppejans, 2006), has been reported in the southern Gulf 

(Huerta-Múzquiz and Mendoza-González, 1985), and 

Pacific Mexico (Dawson, 1960a, as “Cladophora membranacea”; see also Dawson, 1961b; Huerta-Múzquiz and 

Garza-Barrientos, 1975; Pedroche et al., 2005).

Two species are known in the northern Gulf of 

California.

Key to the Species of Cladophoropsis in the 

1a. Filaments relatively slender, 65–100(–125) μm in diameter; cells very long, up to 15 mm, 30–100 times longer than wide

. . . . . . .

1a. Filaments much wider, usually over 350 μm in diameter; cells of main filaments not as long, 2.3–18 times longer than 

wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. macromere
n u m b e r 9 4 • 6 9

Cladophoropsis gracillima E. Y. Daw

FIGURE 32A–C

Cladophoropsis gracillima E. Y. Dawson, 1950c:149, 

151, figs. 12, 13; Dawson, 1957b:7; Dawson, 1960a:32; 

Dawson, 1961b:379; Dawson, 1962b:228; Dawson, 

1966a:6; González-González et al., 1996:131; Rodríguez-Morales and Siqueiros-Beltrones, 1999:17; Leliaert, 2004:237, figs. 77 (isotype), 78; Pedroche et al., 

2005:52; Mateo-Cid et al., 2006:48, 58.

Algae loosely filamentous tufts, of sparsely, irregularly branched, cylindrical filaments, 2–4 cm long; entangled by creeping or entwining branches to other algae and 

attached by rhizoids to substratum. Chloroplast a slightly 

open to dense, parietal reticulatum with 1–4 pyrenoids. 

Cells very long, (0.3–)1.0–15 mm in length (30–100 times 

longer than wide), with diameter in cells of main filaments 

(50–)65–120(–130) μm; in basal cells 50–160 μm; and 

in apical cells (60–)80–120(–135) μm. Branches arising 

beneath a septum of the filament cell and usually remain 

in open connection between the branch cell and filament. 

Cell walls 2 μm thick.

Reproduction unknown for this species.

Habitat. On rocks and in tide pools, usually

entangled with other small algae; mid to low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Puerto Refugio, Isla Ángel de la Guarda; Calerita (vicinity of La Paz) to Punta Palmilla. Pacific coast: 

Costa Rica.

Type Locality. Punta Palmilla, Baja California Sur

Remarks. Dawson (1950c) separated 

Cladophoropsis gracillima from the morphologically similar species on the basis of its entangled habit, slender filaments, and much longer cells (30–100 times longer than 

diameter). Crystalline cell inclusions were briefly noted 

in a list as present in C. gracillima (Leliaert and Coppejans, 2004:202). This apparently was an error since Leliaert (2004:239) and Leliaert and Coppejans (2006:670) 

considered C. gracillima, because of its lack of crystalline 

inclusions and absence of tenacular cells, to be of uncertain taxonomic affinity. When more Gulf specimens are 

collected, studies are needed to evaluate the distinctiveness 

and phylogenetics of this taxon.

Cladophoropsis macromeres W. R. Taylor

Cladophoropsis macromeres W. R. Taylor, 1928:64, pl. 

4: figs. 15, 16; Mendoza-González and Mateo-Cid, 

1986:419; González-González et al., 1996:281; Leliaert, 2004:116, figs. 3C,D, 5A,B, 8A–E; Pedroche et al., 

2005:52; Leliaert and Coppejans, 2006:662, figs. 3, 4, 

18–22.

Algae of cylindrical, loosely branched entangled filaments, forming mats up to 15 cm wide and 2–15 cm thick; 

usually found free floating or lying unattached on substratum. Main filaments (280–)350–510 μm in diameter; cells 

2.3–18 times longer than wide, with cells walls up to 10 

μm thick; upper filaments narrower, (140–)280–360(–400) 

FIGURE 32. Cladophoropsis gracillima: A–C. Drawings of type 

specimen (EYD-3233; after Dawson, 1950c: figs. 12–13). A. Upper 

portion of a branched filament showing the manner of septation 

of branches. B, C. Small portions of a skein-like alga showing extremely long cells and infrequent branching.
7 0 • 

μm in diameter with cells up to 60 times longer than wide, 

and cell walls 2–4 μm thick. Branching mostly irregular, sometimes unilateral in upper portions. Chloroplasts 

polygonal to rounded in parietal reticulum, usually with 

a single large pyrenoid. Prismatic calcium oxalate crystals 

present in most cells.

Habitat. Free-floating or cast ashore on sand

flats or mudflats; sometimes entangled with other algae or 

aquatic plants; in protected tide pools or shallow waters; 

intertidal.

Distribution. Gulf of California: Guaymas

Type Locality. Fort Jefferson on Garden Key,

Dry Tortugas National Park, Monroe County, Florida.

Remarks. Cladophoropsis macromeres, a 

western Atlantic species, is known in warm, calm, shallow waters (Taylor, 1960; Wynne, 2005); the description 

is based on Leliaert and Coppejans (2006). It has been 

recorded once in the northern Gulf from drift material 

(Mendoza-González and Mateo-Cid, 1986), and more 

collections would be helpful to verify its presence in the 

Gulf of California.

Phyllodictyon J. E. Gr

Phyllodictyon J. E. Gray, 1866:70.

Algae are erect, of one or more reticulate (netlike) 

blades, above a monosiphonous (rarely branched) stalk 

that is attached by basal rhizoids. Blades are complanate, 

with an apical, uniaxial orientation that is maintained 

throughout development with primary and secondary laterals arising from opposite, astipitate protrusions along 

distal sides of previously cut-off segments. The distinctive 

network of the blade is formed as the initial laterals and 

subsequent higher-order distichously arranged laterals 

develop by nonsynchronous centripetal wall invaginations 

(Cladophora-type cell division). Intercalary cell divisions 

are common. Tenacular cells on the apices of second- and 

third-order (rarely more) laterals may contact and fuse 

with adjacent laterals to form the filament network of the 

blade. Cells are multinucleate and have numerous discoid 

chloroplasts, each with a single pyrenoid.

Life histories are not known, but zoospores have been 

reported.

Remarks. Phyllodictyon and the similarlooking genus Struvea Sonder (1845) are differentiated 

from each other primarily by their mode of cell division: 

“Cladophora-type” in Phyllodictyon versus the segregative cell division in Struvea (Kraft and Wynne, 1996), a 

generic distinction initially supported by molecular evidence (e.g., Kooistra et al., 1993; Bakker et al., 1994). 

However, Leliaert and Coppejans (2007) noted that while 

Phyllodictyon, Struvea, Cladophoropsis, Chamaedoris 

Montagne (1842b), Boodlea G. Murray et G. B. De Toni 

(in Murray, 1889), and Struveopsis Rhyne et Robinson 

(1969) were all nonmonophyletic genera and their generic 

separation was problematic, that for now and for practical 

utility, they should be distinguished by their considerable 

and consistent habit (morphological) differences.

One species, Phyllodictyon anastomosans (Harvey) 

Kraft et M. J. Wynne (1996) has been reported in the 

southern Gulf (Rocha-Ramírez and Siqueiros-Beltrones, 

1991; Mateo-Cid et al., 2000a; both as “Struvea anastomosans”), and Pacific Mexico (Dawson, 1949, 1950d, as 

“S. delicatula”; Chávez B., 1972, Huerta-Múzquiz, 1978, 

Mateo-Cid and Mendoza-González, 1991, Mateo-Cid 

and Mendoza-González, 1992, León-Tejera et al., 1993, 

and González-González, 1993, all as “S. anastomosans”; 

and Mendoza-González and Mateo-Cid, 1998).

Two of the species are known in the Gulf of California.

Key to the Species of Phyllodictyon In the nort

1a. Algae with narrow main axes, 200–380 μm diameter, and upper netlike blades; tenacular cells at tips of apical cells and at 

base of main axial cells; lenticular cells absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pulcherrimu

1b. Algae with broader main axes, 950–1120 μm in diameter, upper bladelike portion either lacking or reduced (partial); tenacular cells only at tips of apical cells; lenticular cells present. . . . . .

Phyllodictyon pulcherrimum J. E. Gr

FIGURE 33

Phyllodictyon pulcherrimum J. E. Gray, 1866:70; Kraft 

and Wynne, 1996:131–135, 139, figs. 26–28; John et 

al., 2004:23; Leliaert, 2004:fig. 52 (holotype); Pedroche 

et al., 2005:54; Leliaert et al., 2008:8, 10, tbl. 3.

Struvea pulcherrima (J. E. Gray) Murray et Boodle, 

1888:281; Schneider and Searles, 1991:80, fig. 70.

Cladophoropsis pulcherrima (J. E. Gray) Leliaert et 

Coppejans in Leliaert, 2004:192, comb. nov. prov., figs. 

5C,D, 52–55.

Struvea ramosa Dickie, 1874b:316; Dawson, 1966b:55, 

fig. 1B; González-González et al., 1996:139.
n u m b e r 9 4 • 7 1

Algae erect, up to 20 cm tall, with reticulate blade(s) 

above a monosiphonous stalk, up to 1.5 mm in diameter, 

and attached below by branching, basal rhizoids. Lower 

portion of stalk usually unbranched; upper portion with a 

series of opposite branches, up to 3 orders, that form netlike blades, up to 12 cm long and up to 10 cm wide. Main 

axes 200–380 μm in diameter. Lateral branches, developed below a septum at distal end of axis cell, with up to 6 

branches per cell. Tenacular cells abundant at tips of apical 

cells, and base of cells on main axes. Lenticular cells absent.

Habitat. On rock; subtidal, 17 m depth

Distribution. Gulf of California: Punta

Concepción.

Type Locality. Gulf of Mexico

Remarks. The phylogenetic relationship of 

the western Atlantic Phyllodictyon pulcherrimum (type 

locality Gulf of Mexico) to Gulf of California P. pulcherrimum should be tested. Calcium oxalate crystalline cell 

inclusions, thought to be of systematic value, have been 

described in P. pulcherrimum from other regions (Leliaert 

and Coppejans, 2004; Leliaert et al., 2008) but are not 

reported in Gulf of California P. pulcherrimum.

FIGURE 33. Phyllodictyon pulcherrimum. A. Axis with several well-developed open reticulate fronds above a monosiphonous stalk (EYD25872). B. Portion of net-like blade.
7 2 • 

Phyllodictyon robustum

(Setchell et N. L. Gardner) Leliaert et W

FIGURE 34A–E

Cladophoropsis robusta Setchell et N. L. Gardner, 

1924a:714, pl. 13: fig. 16 [holotype specimen only; see 

Leliaert et al., 2008:3–4, 9, figs. 2, 3, 15, 16, 23 (illustrations of holotype)]; Dawson, 1944:206; Dawson, 

1957b:8; Dawson, 1961c:404, pl. 4: fig.3, pl. 5: fig. 6; 

Dawson, 1962b:228; Chávez B., 1972:268; MendozaGonzález and Mateo-Cid, 1986:419; Rocha-Ramírez 

and Siqueiros-Beltrones, 1991:24; Mateo-Cid and Mendoza-González, 1992:24; Mateo-Cid et al., 1993:51; 

Servière-Zaragoza et al., 1993:482; González-González 

et al., 1996:131; Servière-Zaragoza et al., 1998:168.

Cladophoropsis? robusta sensu Dawson, 1959a:9–11, fig. 

3A; Dawson, 1961b:379.

Phyllodictyon robustum Leliaert et Wysor in Leliaert et 

al., 2008:230, figs. 1–46.

Struveopsis robusta (Setchell et N. L. Gardner) Rhyne et 

H. Robinson, 1969:470; Stout and Dreckmann, 1993:6, 

17; Kraft and Wynne, 1996:140; Pacheco-Ruíz and Zertuche-González, 1996a:432; Bucio-Pacheco and Dreckmann, 1998:42; L. Aguilar-Rosas et al., 2000:129, 133;

Mateo-Cid et al., 2000a:69; L. Aguilar-Rosas et al., 

2002:235; Wysor, 2004:219, fig. 7; Pedroche et al., 

2005:54; Mateo-Cid et al., 2006:58.

Pseudostruvea robusta (Setchell et N. L. Gardner) Egerod, 

1975:47; Espinoza-Ávalos, 1993:333; GonzálezGonzález et al., 1996:138; Pacheco-Ruíz and Zertuche-González, 1996a:432; Rodríguez-Morales 

and Siqueiros-Beltrones, 1999:22; Cruz-Ayala et al., 

2001:190; Pacheco-Ruíz et al., 2008:202.

Willeella mexicana E. Y. Dawson, 1950c:151, fig. 11 

(holotype); Papenfuss and Egerod, 1957:83; Dawson, 

1957b:8; Dawson, 1959a:11–12; González-González et 

al., 1996:141; Leliaert, 2004:49; Leliaert and Coppejans, 2006:673; Leliaert et al., 2008: figs. 4 (holotype), 

5–7 (isotype).

Cladophoropsis mexicana (E. Y. Dawson) Leliaert et Coppejans in Leliaert, comb. nov. prov., 2004:184, fig. 47.

Algae erect, 2–4(–5) cm tall, highly variable in morphology; usually of few to several, monosiphonous main 

axes, unbranched to laterally branched up to 3 orders; 

attached below by entangled basal rhizoids. Main axes 

diameter 370–900(–1300) μm, increasing upward; branch 

diameter mostly 600–900 μm; apical cells with rounded 

tips, 950–1120 μm in diameter, 1.5–3.2 mm in length. 

Lower portion of main axis (stipitate), usually unbranched 

in first 1.0–2.5 cm; upper portion either remaining 

unbranched or becoming bladelike, with few, paired lateral branches, developed below a septum at distal end of 

axis cell, 2–4 mm in length, of (1–)3–6 cells, with up to 

4 branches per cell. Septa either absent or present after 

growth of a few millimeters in length. Tenacular cells, 

apparently rare, at tips of apical cells; abundant on rhizoidal cells in culture (Leliaert et al., 2008). Lenticular cells 

present. Calcium oxacalate crystals present in most cells; 

usually rectangular, polygonal or irregular, 8–40(–60) μm 

wide, with dentate margins; rarely octahedral.

Reproductive cells, with numerous raised pores, 

develop from rows of 2–4 slightly swollen apical cells 

(after Leliaert et al., 2008).

Habitat. Growing on rocks or sometimes

sponges; in tidepools and low intertidal to shallow 

subtidal.

Distribution. Gulf of California: Punta

Gorda to Bahía La Paz; Cabo Pulmo. Pacific coast: Nayarit 

to Oaxaca; El Salvador; Costa Rica; Panamá.

Type Locality. Punta Colorado (adjacent to

Bahía Bocochibampo), vicinity of Guaymas, Sonora, Gulf 

of California.

Remarks. Since its original description from 

the Gulf of California, Cladophoropsis robusta has a 

notable history of taxonomic changes in generic placement. Papenfuss and Egerod (1957) suggested that C.

robusta could be a Willeella Børgesen (1930). Dawson 

(1959a:12, as “C.? robusta”) disagreed and noted in the 

type of C. robusta, “lens-shaped incipient branchlets” 

before branch formation that were “somewhat suggestive 

of Struvea” in the similar way its branches were cut off 

by a “lens-like cell” (lenticular cells). Dawson questioned 

its generic placement and concluded that C.? robusta and 

Willella mexicana Dawson (1950c) were the same species, 

more closely allied with Valoniopsis than to Willeella. 

Subsequently, C. robusta was transferred to other genera, 

Struveopsis by Rhyne and Robinson (1969) and then later 

to Pseudostruvea by Egerod (1975).

Leliaert (2004) and Leliaert and Coppejans (2006) initially observed the holotype of C. robusta (CAS now UC) 

was not in agreement with the original description or illustration of Setchell and Gardner (1924a:714, pl. 13: fig. 

16) and noted the type to be “Valoniopsis pachynema.” 

They further suggested that most records published as 

“C. robusta” were Willeella mexicana. Most recently, Leliaert et al. (2008) reported re-examination of the holotype 

of C. robusta showed it to have tenacular cells at tips of 

some branches and that the branches developed exclusively 
n u m b e r 9 4 • 7 3

FIGURE 34. Phyllodictyon robustum: A. Clump with some axes separated to show branching pattern (EYD-18555). B–D. Holotype of 

Cladophoropsis robusta (I. M. Johnston-135): B. Two specimens, axes with a few short, opposite branches near apices (after Setchell and Gardner 1924a: pl. 13, fig. 16). C. Prostrate and erect axes of another specimen with branches developing from lenticular cells. D. Close-up of lenticular cell buds on lower portion of an axis (C, D, after Leliaert et al., 2008:figs. 2, 3). E. Branching pattern of frond (after Dawson, 1950c:fig. 

11; holotype of Willeella mexicana, EYD-1789).
7 4 • 

by lenticular cells, which together with its lack of “blades,” 

gave it a “Valoniopsis-like” appearance. They also found 

an isotype specimen of W. mexicana to have some branches 

initiated by lenticular cells and tenacular cells at tips of some 

apical cells. Results from their critical examination of type 

material, morphological and culture studies, and molecular analyses resolved the generic status of C. robusta as 

belonging to Phyllodictyon, and supported P. robustum and 

W. mexicana as being a single species (Leliaert et al., 2008).

Sipho

Siphonocladaceae F. Schmitz, 1879:20, as “Siphonocladiaceae.”

Algae are erect and either are turfs or clumps of axes 

with irregular to regular, radially arranged branches that 

are composed of relatively large, branched, elongated 

cylindrical to clavate, siphon cells (e.g., Siphonocladus). 

Other members are initially or may remain solid or hollow, more or less spherical thalli that often develop into 

irregular or lobed shapes or can become open and sheetlike and are composed of large, globose cells that produce 

a pseudoparenchymatous structure (e.g., Dictyosphaeria). 

Tenacular cells occur in some and may function as structural support or as adventitious rhizoids. Cell division is 

exclusively segregative in main axes and branches of turfbranched forms and in the spherical to lobed forms. There 

can also be cell division by centripetal invagination in the 

rhizoids. Cells are multinucleate, with a reticulate chloroplast or numerous chloroplasts that either lack pyrenoids 

or have one or more pyrenoids.

Vegetative reproduction in some is by fragmentation. 

Life histories, where known, are an isomorphic alternation of generations, involving quadriflagellate zoospores, 

and isogamous biflagellate gametes. (However, at least 

one, Siphonocladus pusillus, has been suggested to have a 

macrodiploid phase, with the gametes being the only haploid phase [Jónsson and Puiseaux-Dao, 1959].)

Remarks. Which genera comprise the Siphonocladaceae, as well as the family’s relationship to other 

families of Cladophorales-Siphonocladales complex, is in 

a state of flux (Leliaert et al., 2003; Kraft, 2007). Herein 

the taxonomy of the family and order is based on the genus 

Siphonocladus F. Schmitz (1879; Børgesen, 1905; generitype: Siphonocladus wilbergii F. Schmitz (1879); now = 

S. pusillus (C. Agardh ex Kützing) Hauck, 1884; basionym: 

Valonia pusilla C. Agardh ex Kützing, 1849). The family 

also includes the genera Dictyosphaeria and Ernodesmis.

Although not yet reported in the northern Gulf, one 

genus, Dictyosphaeria Decaisne (1842a; Decaisne ex 

Endlicher, 1843), is represented in the southern Gulf by 

reports of three species: D. versluysii Weber-van Bosse 

(1905) by Dawson (1944), Rodríguez-Morales and 

Siqueiros-Beltrones (1999), and Paul-Chávez and Riosmena-Rodríguez (2000); D. australis Setchell (1926) by 

Dawson (1949); and D. cavernosa (Forsskål) Børgesen 

(1932) by Anaya-Reyna and Riosmena-Rodríguez (1996) 

and Riosmena-Rodríguez et al. (2005).

One of the genera is found in the northern Gulf of 

California.

Ernodesmis Børgesen

Ernodesmis Børgesen, 1912b:259.

Algae forming bushy clumps composed of elongate cells, with a primary upright clavate cell (or basal 

stalk) that is attached below by a holdfast of rhizoids 

and bears a distal whorl of branches. Whorl clusters are 

composed of 3–10 siphon branches. Each siphon branch 

of the whorl can initiate new branches at its apex, and 

in turn, each of these individual whorl branches can produce another whorl of siphon branches; this process may 

be repeated to produce several orders of whorled clusters. Secondary adventitious holdfasts may develop at the 

base of older branches, where they come in contact with 

the substratum. New uprights are produced from welldeveloped rhizoids. Tenacular cells are sparse, occurring 

only at junction of branch cells. Annular constrictions 

present on the basal stalk may also be present on the 

lower part of the branches. Cells are multinucleate, with 

numerous plate-like chloroplasts that form a reticulum, 

and contain 1 pyrenoid. Segments developed by lenticular cell division and sometimes also by segregative division in main stalks and by centripetal invagination in the 

rhizoids.

Vegetative reproduction is by fragmentation. Reproduction and life histories are poorly known.

Remarks. The familial placement of Ernodesmis within the Cladophorales-Siphonocladales complex is 

uncertain. Solely on a morphological basis it would seem 

that Ernodesmis could be related to Valonia and Valoniopsis and therefore would be in the family Valoniaceae. 

However, the molecular analysis (Leliaert et al., 2003) 

supported a phylogenetic relationship of Ernodesmis to 

Siphonocladus, as earlier observed by Børgesen (1913). 

Thus, for now, Ernodesmis is tentatively considered a 

member of the Siphonocladacaeae.

A monotypic genus of the Caribbean and known 

throughout the western Atlantic, the one species has also 

been reported in the Gulf of California.
n u m b e r 9 4 • 7 5

Ernodesmis verticillata (Kützing) Børgesen

FIGURE 35

Valonia verticillata Kützing, 1847:165.

Ernodesmis verticillata (Kützing) Børgesen, 1912b:259, 

figs. 10–12; Børgesen, 1913:66, figs. 52–54; Dawson, 

1949:244; Dawson, 1959a:4, 8, 11; Dawson, 1961b:379; 

Dawson, 1966a:6; Dawson, 1966b:55; Papenfuss and 

Chihara, 1975:309, figs. 1–3; Huerta-Múzquiz and 

Garza-Barrientos, 1975:4; Huerta-Múzquiz and Mendoza-González, 1985:44; Mateo-Cid et al., 1993:51; 

González-González et al., 1996:286; La Claire et al., 

1997:831; Rodríguez-Morales and Siqueiros-Beltrones, 

1999:22; Paul-Chávez and Riosmena-Rodríguez, 

2000:146; Mateo-Cid et al., 2000a:61, 70; Pedroche et 

al., 2005:55; Mateo-Cid et al., 2006:49, 58; ServièreZaragoza et al., 2007:7.

Algae light green tufts, up to 2.5 cm (sometimes more) 

tall; thallus a single elongate, cylindrical stalk cell, 10–20 

mm in length and up to 2.5 mm in diameter, that bears a 

whorl of cylindrical branches at its apex. Each branch of 

the initial whorl, in turn, bears another whorl of similar 

but smaller cells; and this whorl pattern may repeat itself 

up to 6 times.

Habitat. On rocks, usually in shaded areas, in

crevices, tidepools, and lagoons; mid to low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Cabeza Ballena. Pacific coast: Isla Socorro 

(Islas Revillagigedo).

Type Locality. St. Croix, U.S. Virgin Islands

Remarks. Crystalline cell inclusions have been 

reported in specimens of Caribbean Ernodesmis verticillata from the type locality and in Gulf of California specimens from Punta Palmilla, Baja California Sur (Leliaert 

and Coppejans, 2004).

Valo

Valoniaceae Kützing, 1849:507.

Members of this family lack a distinct central axis 

and are either relatively simple forms ranging from single 

giant vesicular cells to aggregations of vesicles, siphons, 

or cylindrical cells, or are more complex forms that produce clusters of elongated vesicles or cells from the primary vesicle. Growth is by lenticular cell division in which 

lenticular segments are formed inside the vesicles. These 

segments may either remain lenticular, or expand outward 

to develop branches (i.e., septa are initiated prior to elongation of the vesicular branch on the surface of the parent 

vesicle, e.g., Valonia) or may develop hapteroid cells that 

attach the alga to the substratum or adjacent vesicles.

Although biflagellate and quadriflagellate swarmers 

have been observed, for most species the life histories are 

not completely known. Sexual reproduction is only known 

in Valonia macrophysa Kützing (1843) and involves biflagellate isogametes with an isomorphic alternation of generations (Chihara, 1953, 1959).

One genus in the family, Valonia C. Agardh (1823), is 

recorded in the southern Gulf on the basis of the reports 

of V. macrophysa Kützing (1843) by Huerta-Múzquiz and 

Mendoza-González (1985) and Rodríguez-Morales and 

Siqueiros-Beltrones (1999).

One genus is known in the northern Gulf of California.

Valoniopsis Børgesen

Valoniopsis Børgesen, 1934:10.

Algae forming cushion-like mats, composed of interwoven club-shaped, filiform, or elongated cylindrical 

siphons (cells) that initially grow more or less perpendicular to the substratum, then become repent and entangled, 

and attach by adventitious rhizoids that develop from the 

tips of downward arching branches that come in contact 

with the substratum. Branching is unilateral, pseudodichotomous or irregular, and obscures apical orientation. Well-developed attachment rhizoids can give rise to 

new erect segments. Siphons with one to several (1–)2–3 

branches that are developed from cell division by lenticular cells (not from protrusions) distally and laterally on 

FIGURE 35. Ernodesmis verticillata: Upper portion showing successive series of whorled branches (after Børgesen, 1913:fig. 52).
7 6 • 

the siphons; septa are formed only at branch base. Annular constrictions and tenacular cells are absent. Cells with 

numerous discoid chloroplasts, each with single pyrenoid 

covered with a starch sheath and bisected by traversing 

thylakoids. Cell division in the rhizoids is by centripetal 

invagination.

Reproduction is apparently unknown. Life histories are presumed to be similar to other algae of the 

Siphonocladales.

Remarks. Valoniopsis can sometimes be confused with Cladophoropsis, but Valoniopsis has lenticular 

cells and septa at branches, whereas Cladophoropsis has no 

lenticular cells and mostly nonseptate (occasionally septate) 

branches. The distinction between Valoniopsis and some 

species of Valonia is less certain, and the validity of their 

generic separation is debatable (Abbott, 1986; Kraft, 2000; 

Abbott and Huisman, 2004). Biochemically, both genera 

produce similar amino and fatty acids (Sivalingam, 1989).

There are only two known species of Valoniopsis, and 

both occur in the northern Gulf of California. One is an 

endemic species in the Gulf of California, and the other is 

a widespread tropical-subtropical species.

Key to the Species of Valoniopsis of the n

1a. Lateral branches of the siphons, mostly 300–400 μm in diameter, and constricted at branch junctions  . . . .  V

1b. Lateral branches of the siphons, mostly 450–1000 μm in diameter, and not constricted at branch junctions . . . . . . . . . . .

. . . . . . 

Valoniopsis hancockii E. Y. Da

FIGURE 36A

Valoniopsis hancockii E. Y. Dawson, 1944:207, pl. 31: fig. 

9; Dawson, 1961b:379; Abbott, 1986:164; EspinozaÁvalos, 1993:333; González-González et al., 1996:140; 

Pedroche et al., 2005:56; Pacheco-Ruíz et al., 2008:202.

Algae green, usually cushionlike, of small entangled 

masses of cylindrical, coenocytic siphons (filaments), 

with only occasional septa, mostly at branches. Prostrate 

siphons very crooked, 125–250 μm in diameter; ascending 

siphons straighter, 300–400 μm in diameter, with infrequent short lateral branches of about the same diameter 

that are sharply constricted at their base; apices broadly 

rounded.

Habitat. On sand and rocky shores, low

intertidal.

Distribution. Gulf of California: Puerto

Refugio, Isla Ángel de la Guarda.

Type Locality. Puerto Refugio, Isla Ángel de

la Guarda, Islas de la Cintura, Gulf of California.

Remarks. A northern Gulf endemic, Valoniopsis hancockii (currently known only from the type 

collection) is distinguished by siphons that only occasionally have septa and has basal constrictions of the branch 

siphons (filaments) (Dawson, 1944; Abbott, 1986). Morphological and molecular studies on further collections of 

V. hancockii, particularly from the type locality, are necessary before this taxon can be properly understood.

Valoniopsis pachynema (G. Martens) Børgesen

FIGURE 36B,C

Bryopsis pachynema G. Martens, 1868:24, pl. 4: fig. 2.

Valoniopsis pachynema (G. Martens) Børgesen, 1934:10, 

figs. 1, 2; Dawson, 1944:206; Dawson, 1949:234, 238; 

Dawson, 1951:54; Dawson, 1954e:324; Issac, 1957:85, 

pl. 28: figs. 6, 7; Papenfuss and Egerod, 1957:84; 

Dawson, 1959a:6, 12; Dawson, 1961b:379; Dawson, 

1966a:6; Littler and Littler, 1981:151: tbl. 3, 152: fig. 

4; Stewart, 1982:54; Littler and Littler, 1984:22; Abbott, 1986:163; Servière-Zaragoza et al., 1993:482; 

González-González et al., 1996:291; La Claire et al., 

1997:831; Yoshida, 1998:51; Rodríguez-Morales and 

Siqueiros-Beltrones, 1999:22; L. Aguilar-Rosas et al., 

2000:133; Kraft, 2000:580, fig. 27C,D; Paul-Chávez 

and Riosmena-Rodríguez, 2000:146; Abbott and Huisman, 2004:93, fig. 32A; Mateo-Cid et al., 2006:48, 58; 

Pacheco-Ruíz et al., 2008:202.

Valonia pachynema (G. Martens) Weber-van Bosse, 

1913:61.

Cladophoropsis robusta sensu Taylor, 1945:51 [see Dawson, 1959a:12] [non Cladophoropsis robusta Setchell 

et N. L. Gardner, 1924b:714, which is Phyllodictyon 

robustum (Setchell et N. L. Gardner) Leliaert et Wysor 

in Leliaert et al., 2008:10].

Algae dark green, forming dense cushionlike 

clumps, up to 3 cm high and to 7 cm in diameter, irregularly branched near segment apices with 1 or more, 
n u m b e r 9 4 • 7 7

usually curved, branchlets, and attached below by rhizoids. Siphons coenocytic, cylindrical, (300–)450–1000 

μm in diameter, interlaced and ascending. Branches with 

basal septa (lenticular crosswall); younger branches erect, 

older branches repent.

Habitat. On rocks, occasionally entwined with

other algae of turf communities; high to low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Bahía Agua Verde; Isla Espíritu Santo to Bahía 

de La Paz. Pacific coast: Velero Point (south Bahía Sebastián Vizcaíno) and Punta Malarrimo, Baja California Sur; 

Isla Isabel, Nayarit; Hawaiian Islands; China; Japan.

Syntype Localities. Benkulen and Pulau

Tikus, near Bengkulu, Sumatra, Indonesia (vide Silva et 

al., 1996a).

Remarks. Crystalline cell inclusions, considered to be of systematic value, have been reported in 

Valoniopsis pachynema from several locales in the Indian 

Ocean (Leliaert and Coppejans, 2004).

Bryopsido

Bryopsidophyceae Bessey, 1907:287.

Algae of this class are almost all exclusively marine and 

vary from relatively simple forms to structurally complex 

forms that are siphonous or tubular and coenocytic (i.e., 

lack cell walls and are multinucleate), with cytoplasm also 

containing numerous fusiform or ellipsoidal chloroplasts 

and a large central vacuole. Many are homoplastidic, with 

only chloroplasts that may contain or lack pyrenoids; others are heteroplastidic, with both chloroplasts and amyloplasts (colorless starch storage plastids). Cytoplasmic 

streaming occurs in the siphons, with organelles moving 

with the cytoplasm. Accessory pigments are siphonoxanthin and siphonein. Cell wall constituents are manna, 

xylan, and glucan (cellulose) in varying proportions; glucan is not in the crystalline form as found in members of 

the class Chlorophyceae Wille ex Warming (1884).

Reproduction is by biflagellate, quadriflagellate, or 

multiflagellate zoospores. In species that produce biflagella or quadriflagella, the flagella are in eleven and five 

o’clock positions, with overlapping basal bodies. Other 

members have stephanokont zoospores, i.e., multiple flagella arranged in whorls at the cell’s anterior end. Sexual 

reproduction is anisogamous.

The class is usually considered to have a single order, 

Bryopsidales (e.g., Silva et al., 1996a; Abbott and Huisman, 2004; herein). However, some phycologists consider 

the class to contain two orders and include the Halimedales C. van den Hoek, D. G. Mann et H. M. Jahns (1995).

Bryops

Bryopsidales J. H. Schaffner, 1922:133.

FIGURE 36. Species of Valoniopsis. A. Valoniopsis hancockii: 

Portion of type showing short branches with constrictions at 

their base (drawing of type specimen, EYD-[1940]-215, after 

Dawson, 1944: pl. 31, fig. 9). B. Valoniopsis pachynema: Portion showing branching and a branch bending downward with 

tip attaching to substratum by means of haptera (after Børgesen, 1934: pl. 13, fig. 1f ). C. Valoniopsis pachynema: Clump of 

branched, cylindrical, siphonlike segments (photo of live specimen from Puerto Peñasco, Sonora, Mexico).
7 8 • 

Codiales Setchell, 1929:584.

Caulerpales J. Feldmann, 1946:753.

Siphonales Warming, 1884:33, as “Siphoneae.”

Algae are variable in shape and size, ranging from 

simple (unbranched) to branched and to complex systems 

of branching tubular siphons or filaments. Some form 

globose vesicles; others are of filaments or siphons that 

are freely branched or formed into thalli with distinctive 

morphologies. Most are attached by prostrate rhizoids. 

Transverse septa are generally lacking in vegetative portions but may be present at the base of reproductive sporangia or gametangia. Cells are coenocytic with numerous 

nuclei and chloroplasts that lack or contain pyrenoids. In 

some, the vegetative portion of the thallus is traversed by a 

system of internal strands of cell wall material (trabeculae) 

that form an elaborate network, thought to strengthen the 

outer siphon walls. All members possess two distinctive 

xanthophylls, siphonein and siphonoxanthein, which are 

not found in other members of the Chlorophyta.

Some reproduce vegetatively by rhizome-like structures, fragmentation, or propagules. Sporophytic reproduction is by multiflagellate zoospores or aplanospores, 

usually developed within morphologically differentiated 

sporangia. Sexual reproduction is isogamous, anisogamous, or oogamous, with gametangia producing biflagellate gametes.

Remarks. Although the Bryopsidales is the 

more widely accepted name for this order (e.g., Silva et 

al., 1996a; Abbott and Huisman, 2004), the name Caulerpales has also been recently used (Graham and Wilcox, 

1999; Lee, 1999). Members of this very large order are 

widespread in distribution and are particularly abundant 

in tropical to subtropical regions.

Phylogenetic analyses of the Bryopsidales suggested 

that the suborders Bryopsidineae Hillis-Colinvaux (1984) 

and Halimedineae Hillis-Colinvaux (1984) are distinct 

monophyletic lineages (Lam and Zechman, 2006). Family-level monophyly was supported for the Bryopsidaceae, 

Derbesiaceae and Codiaceae (Bryopsidineae), and the 

Caulerpaceae and Halimedaceae (Halimedineae), but not 

for the Udoteaceae (Lam and Zechman, 2006).

Five families are represented in the northern Gulf of 

California.

Key to the Families of Bryopsidales in the 

1a. Algae differentiated into three distinct portions: an erect, cylindrical to frondose upper portion; arising from prostrate, 

stolonlike runners; and attached by rhizoids at various points; not calcified . . . . . . . . . . . . . . . . . . . . . . . . . Caulerpaceae

1b. Algae of various simple to structurally complex forms: filamentous or siphonous or vesiculate; not calcified or calcified.

2a. Algae of coenocytic filaments, tubular cells or siphons or vesicles; not structurally complex; not calcified .

2b. Algae multiaxial, of coenocytic filaments formed into complex thalli that are firm or spongy; calcified or not calcified. 

3a. Macrothalli filamentous, of one or more percurrent or non-percurrent axes with lateral branches; gametophytes and sporophytes isomorphic in some; others heteromorphic with filamentous macro-gametophytes, and micro-sporophytes of 

creeping prostrate filaments (protonema) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bryopsidaceae

3b. Heteromorphic macrothalli; sporophytes and gametophtyes very different looking: sporophytes filamentous, without a 

percurrent axis, subdichotomously to irregularly branched; gametophytes a single vesicle  . . . . . .

4a. Thalli not calcified; spongy; prostrate and cushionlike or erect, terete to flattened axes and branches; without segments

. . . . . 

4b. Thalli of calcified cylindrical to flattened segments that alternate with narrow, flexible noncalcified joints 

. . . . . . 

Bryopsid

Bryopsidineae Hillis-Colvinaux, 1984:287.

The suborder Bryopsidineae is characterized by homoplasty, the absence of a concentric lamellar system, and 

cell walls that are composed primarily of the polysaccharides, either mannan, or xylan and cellulose. Allelochemicals (secondary natural products) are generally absent. 

Reproduction is non-holocarpic. Reproductive structures 

usually have a septum (plug) at their base.

Remarks. Species of the Bryopsidineae are 

tropical to subboreal in distribution. The suborder is represented by three families, Bryopsidaceae, Derbesiaceae, 

and Codiaceae, in the northern Gulf of California.

Bryops

Bryopsidaceae Bory de Saint-Vincent, 1829:203

Algae are coenocytic, siphonous filaments that are 

simple or sparsely to densely branched and are attached to 
n u m b e r 9 4 • 7 9

the substratum by a rhizoidal holdfast. Filamentous thalli 

lack internal trabeculae, may be with or without a distinct 

percurrent axis, and are branched pinnately, radially, or 

irregularly from a few to several orders. The main component of siphon walls is glucan (cellulose) and xylan in 

the gametophytes, while in the sporophytes it is mannan.

Reproduction can be by fragmentation, regeneration 

from extruded protoplasts, or parthenogenesis. Life histories, where known, generally involve filamentous phases: 

(1) an alternation of gametophytic Bryopsis and a sporophytic microthallus of creeping prostrate filaments (protonema), (2) an alternation of gametophytic “Bryopsis-like” 

phase with a filamentous Derbesia phase (i.e., Bryopsidella 

Feldmann ex H. Rietema, 1975), or (3) zygotes produced 

by Bryopsis that develop directly into another gametophyte. Stephanokont zoospores (with a ring or whorl of 

several flagella) are produced in differentiated sporangia 

with a basal septum in filamentous sporophytes. A few of 

the species are known to produce aplanospores.

Sexual reproduction is non-holocarpic. Gametophytic 

thalli can be either monoecious or dioecious. Biflagellate 

anisogametes are produced in the ultimate branchlets segregated from the rest of the filamentous thallus by a basal 

membrane.

One genus of the family, Bryopsis, is represented in 

the northern Gulf of California.

Bryopsis J. V. Lamou

Bryopsis J. V. Lamouroux, 1809d:333.

Algae are erect, composed of a cylindrical, coenocytic, percurrent axial filament that is oppositely or more 

or less radially or irregularly branched, often appearing 

featherlike, and attached below by rhizoids. Growth is 

indeterminate. Filaments are without septa and contain 

numerous disc-shaped chloroplasts, each containing 1 

pyrenoid. Siphon walls of presumed sporophytes contain 

mannan, while those of gametophytes reportedly contain 

cellulose and xylan.

Asexual reproduction is by fragmentation, aplanospores, and regeneration from extruded protoplasts. Life 

histories generally involve an alternation of filamentous 

phases, with a feather-like or plumose gametophyte and 

either a sporophytic microthallus of creeping filaments 

(protonema) or an erect filamentous Derbesia-like phase. 

In some of the species cultured, filamentous or protonema 

sporophytes produced stephanokont zoospores that either 

developed into gametophytes or into a microthallus that 

produced anisogametes (Rietema, 1975; van den Hoek, 

1981; Tanner, 1981). Sexual reproduction, with thalli 

being monoecious or dioecious, involves the tips of the 

ultimate branchlets that separate from the branch by formation of a basal septum, or plug, at maturity and develop 

into gametangia and produce biflagellate anisogametes.

Remarks. The morphology of species of Bryopsis is so variable and sometimes overlapping (e.g., 

Womersley, 1984) that understanding the species limits 

will probably best be facilitated by molecular systematics. 

The Gulf of California species of Bryopsis, as well as those 

of the eastern Pacific, are in need of taxonomic, culture life 

history, molecular, and phylogenetic studies, such as the 

studies of Krellwitz et al. (2001).

One species, B. muscosa J. V. Lamouroux (1809b), 

has been reported in the southern Gulf by Dawson (1959a, 

1961b).

Five other taxa, four species and one variety, are currently known in the northern Gulf of California.

Key to the Species of Bryopsis in the n

1a. Main axes radially branched (sometimes irregular)  . . . . . .

1b. Main axes pinnately or oppositely branched in one plane or more or less in one plane (featherlike in appearance)  

2a. Main axes with both determinate and longer indeterminate branchlets. . . . . . . . . . . . . . . . . . . . . . . . . . . B. salvadoreana

2b. Main axes with only determinate branchlets  

3a. Main axes over 400 μm in diameter (up to 1000 μm); lower portions without branches and upper portions with abundant, 

pinnately arranged branchlets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. corticulans

3b. Main axes smaller in diameter, 200–400 μm; with frequent, occasional, or sparse branchlets limited to upper or terminal 

portions  

4a. Main axes over 250 μm in diameter, up to 400 μm in diameter; ultimate branchlets opposite along sides of branches in one 

plane  . . . . . . . . . . . 

4b. Main axes 210–250 μm in diameter; branchlets absent in lower portions; branchlets in upper portion usually sparse and 

more or less in one plane or partly radial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pennata var. minor
8 0 • 

Bryopsis corticulans Setchell

Bryopsis corticulans Setchell in Collins, Holden, and 

Setchell, 1899: Exsiccate No. 626; Collins, 1909b:404; 

Setchell and Gardner, 1903:230; Setchell and Gardner, 

1920b:160, pl. 15: figs. 4, 5, pl. 27; Smith, 1944:73, 

pl. 9: fig.3; Dawson, 1951:52; Dawson, 1961b:381; 

Abbott and Hollenberg, 1976:111, fig. 70; Devinny, 

1978:358; Pedroche and González-González, 1981:64; 

L. Aguilar-Rosas and Bertsch, 1983:116; MendozaGonzález and Mateo-Cid, 1986:419; Stewart, 1991:33; 

Servière-Zaragoza et al., 1993:481; González-González 

et al., 1996:126; Aguilar-Ramírez et al., 1998:511; Pedroche et al., 2005:57.

Algae tufted, up to 5–10(–20) cm tall; main axes 400–

1000 μm in diameter; lower portions without branches, 

upper portions abundantly pinnately branched. Branches 

with downward growing rhizoids from their base; ultimate branchlets, 150–300 μm in diameter, abruptly constricted at base.

Habitat. On rocks; mid to low intertidal

Distribution. Gulf of California: Bahía

Kino. Pacific coast: southern British Columbia to Islas San 

Benito, Baja California; Jalisco.

Type Locality. Pacific Grove, Monterey

County, California (Smith, 1944).

Remarks. Bryopsis corticulans is only known 

in the Gulf of California from the report of MendozaGonzález and Mateo-Cid (1986). The description is based 

on Abbott and Hollenberg (1976). The species complex, 

comprising three morphologically similar taxa, B. corticulans, B. hypnoides, and B. plumosa (Hudson) C. Agardh 

(1822), needs to be studied to clarify the taxonomic status 

and relationship of these species to each other and to the 

Gulf B. corticulans.

Bryopsis hypnoides J. V. Lamou

Bryopsis hypnoides J. V. Lamouroux, 1809d:333; Lamouroux, 1809b:135, pl. 1; fig. 2a,b; Setchell and Gardner, 

1920b:159; Smith, 1944:73, pl. 9: fig. 2; Taylor, 1945:60 

[with doubt]; Dawson, 1949:244; Dawson, 1961b:381; 

Abbott and Hollenberg, 1976:113, fig. 71; HuertaMüzquiz, 1978:339; Pedroche and González-González, 

1981:64; L. Aguilar-Rosas and Bertsch, 1983:116, 121; 

Huerta-Muzquiz and Mendoza-González, 1985:44; L. 

Aguilar-Rosas and Pacheco-Ruíz, 1986:77; MendozaGonzález and Mateo-Cid, 1986:420; Sánchez-Rodríguez et al., 1989:39; Dreckmann et al., 1990:24, pl. 2: 

fig. 2; Mateo-Cid and Mendoza-González, 1991:27; 

Rocha-Ramírez and Siqueiros-Beltrones, 1991:31; 

Mendoza-González and Mateo-Cid, 1992:24; Mateo-Cid and Mendoza-González, 1992:25; MateoCid et al., 1993:52; González-González, 1993:443; 

Servière-Zaragoza et al., 1993:481; R. Aguilar-Rosas 

and M. Aguilar-Rosas, 1994:517; Mateo-Cid and 

Mendoza-González, 1994a:52; Mateo-Cid and Mendoza-González, 1994b:45; Mendoza-González et al., 

1994:103, 112; González-González et al., 1996:275, 

365; Mendoza-González and Mateo-Cid, 1996:64, 67, 

76, 88, pl. 28: figs. 126–128; Bucio-Pacheco and Dreckmann, 1998:43; Mendoza-González and Mateo-Cid, 

1998:25; Servière-Zaragoza et al., 1998:168; Yoshida, 

1998:139; L. Aguilar-Rosas et al., 2000:133, 134, 136, 

137; Mateo-Cid et al., 2000a:70; Cruz-Ayala et al., 

2001:190; Abbott and Huisman, 2004:96, fig. 33A; 

Wysor, 2004:219; Pedroche et al., 2005:58; Mateo-Cid 

et al., 2006:59; Dreckmann et al., 2006:154.

Algae tufted, up to 2–4 cm tall; main axes 200–250(–

400) μm in diameter, branching often profuse, usually 

radial or sometimes irregular; branches become progressively narrower and smaller distally; ultimate branchlets 

up to 50 μm in diameter and up to 5 mm long, abruptly 

constricted at the base, and gradually tapered at upper end. 

Rhizoids grow downward from base of primary branches.

Habitat. On rocks or shells; epiphytic on

Gelidium pusillum (L. Aguilar-Rosas et al., 2000); mid to 

low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas; Bahía Concepción to Cabo Pulmo. 

Pacific coast: British Columbia to Chiapas; Panama; Chile; 

Hawaiian Islands; Japan.

Type Locality. Near Cette, Mediterranean

coast of France.

Remarks. Studies of Bryopsis hypnoides from 

California (Burr and West, 1970) and Europe (Rietema, 

1975) showed them to be monoecious, whereas those of 

the northeastern United States were dioecious (Urban, 

1969). Bryopsis hypnoides is part of a species complex 

that includes B. corticulans and B. plumosa; Gulf specimens need to be cultured and morphologically and molecularly analyzed in comparions with type locality material 

of all three of these species to elucidate their taxonomy 

(see also Remarks under B. corticulans).

Bryopsis pennata J. V. Lamouroux var. pen

FIGURE 37

Bryopsis pennata J. V. Lamouroux, 1809d:333, pl. 3, 

fig. 1 a,b; Egerod, 1952:370, fig.7; Dawson, 1957b:9; 
n u m b e r 9 4 • 8 1

Dawson, 1959a:6, 8, 12, 14; Dawson, 1961b:381; Dawson, 1961c:404, pl. 4: fig.4; Dawson, 1966a:7; HuertaMüzquiz, 1978:339;González-González, 1993:443; 

Servière-Zaragoza et al., 1993:481; González-González 

et al., 1996:275; Pacheco-Ruíz and Zertuche-González, 

1996a:432; Mendoza-González and Mateo-Cid, 

1998:25, 27; Mendoza-González et al., 1999:61; LeónTejera and González-González, 2000:327; Abbott and 

Huisman, 2004:98, fig. 33B; López et al., 2004:10; 

Wysor, 2004:219; Pedroche et al., 2005:58; Mateo-Cid 

et al., 2006:48, 59; Dreckmann et al., 2006:154.

Bryopsis plumosa var. pennata (J. V. Lamouroux) Børgesen, 1911:147; Dawson, 1944:212; Rocha-Ramírez 

and Siqueiros-Beltrones, 1991:31; León-Tejera and 

González-González, 1993:497; González-González et 

al., 1996:127.

Algae of 1 or more erect filaments, up to 5 cm tall; main 

axes and branches with more or less opposite branchlets 

of similar length in upper portions and in one plane (giving a lanceolate to oblong featherlike appearance); axes 

sometimes only sparsely branched; arising from a rhizoidal holdfast. Main axis 250–400 μm in diameter; branchlets 40–75(–150) μm in diameter and up to 2.0 mm long.

Habitat. On rocks, usually in shaded areas,

crevices, and tide pools; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Cabeza Ballena. Pacific coast: Nayarit; 

Guerrero to Oaxaca; Costa Rica; El Salvador; Panama; 

Chile; Hawaiian Islands.

Type Locality. Antilles, West Indies

Bryopsis pennata var. minor J. Agar

Bryopsis pennata var. minor J. Agardh, 1887:23; Silva 

et al., 1996a: 807; Bucio-Pacheco and Dreckmann, 

1998:43; López et al., 2000:338; Mateo-Cid et al., 

2000a:70; Pedroche et al., 2005:59; Pacheco-Ruíz et 

al., 2008:202.

Bryopsis pennatula J. Agardh, 1847:6; Kützing, 1856:27, 

pl. 76: fig. 2; Setchell and Gardner, 1920b:158; Taylor, 1945:61; Dawson, 1949:250, 251; Dawson, 

1959a:12; Dawson, 1961b:381; Smith, 1969:624; 

Huerta-Múzquiz and Tirado-Lizárraga, 1970:126; Abbott and Hollenberg, 1976:113, fig. 72; L. Aguilar-Rosas and Bertsch, 1983:116, 121; Stewart and Stewart, 

1984:140; L. Aguilar-Rosas et al., 1985:125; HuertaMúzquiz and Mendoza-González, 1985:44; L. AguilarRosas and Pacheco-Ruíz, 1986:77; Mendoza-González 

and Mateo-Cid, 1986:420; Sánchez-Rodríguez et al., 

1989:38; Dreckmann et al., 1990:24, 37; Mateo-Cid 

and Mendoza-González, 1991:27; Rocha-Ramírez and 

Siqueiros-Beltrones, 1991:31; Mendoza-González and 

Mateo-Cid, 1992:24; León-Tejera et al., 1993:199, 

204; González-González, 1993:443; Stout and Dreckmann, 1993:5; Servière-Zaragoza et al., 1993:443; 

R. Aguilar-Rosas and M. Aguilar-Rosas, 1994:517; 

González-González et al., 1996:127; Mateo-Cid and 

Mendoza-González, 1994b:45; Mendoza-González 

et al., 1994:112; Mendoza-González and Mateo-Cid, 

1996:77, 88, pl. 2: figs. 129–131; Anaya-Reyna and Riosmena-Rodríguez, 1996:862; Silva et al., 1996a:807; 

Galindo-Villegas et al., 1997:4; Aguila-Ramírez et al., 

1998:511; Rodríguez-Morales and Siqueiros-Beltrones, 

1999:22; Paul-Chávez and Riosmena-Rodríguez, 

2000:146; López et al., 2000:340; Carballo et al., 

2002:753; Hernández-Herrera et al., 2005:146; Riosmena-Rodríguez et al., 2005:101; Lam and Zechman, 

2006:671; Servière-Zaragoza et al., 2007:3, 7.

Algae of erect, cylindrical main axes, 210–250 μm 

in diameter, up to 3 cm tall; branchlets absent in lower 

portions; branchlets in upper portion sparse (often with 

only a few uppermost branchlets near the terminal portion) and more or less distichously or partially radially 

arranged, short, less than 1 mm long, 20–30 μm in diameter, strongly constricted at base, with rounded apices.

Habitat. On rocks and in tide pools; high to

low intertidal.

FIGURE 37. Bryopsis pennata: Frond with distichous branching 

pattern giving a featherlike appearance (EYD-27451).
8 2 • 

Distribution. Gulf of California: Punta

Cheuca, Sonora; Bahía de La Paz to Cabo Pulmo, Baja 

California Sur; Mazatlán, Sinaloa. Pacific coast: central 

California to Baja California; Nayarit to Chiapas; Galápagos Islands.

Type Locality. San Agustín, Oaxaca, Pacific

Mexico (Setchell and Gardner [1920b:159] noted “St. 

Augustin” as cited by Agardh (1847) was probably in the 

vicinity of Pochutla and Punta de Huatulco, Oaxaca [see 

also Liebmann, 1869:viii]).

Remarks. Agardh (1887), in describing Bryopsis pennata var. minor, cited B. pennatula J. Agardh (1847) 

in the protologue, thus apparently reducing B. pennatula to 

a variety of B. pennata. Therefore, as noted by Silva et al. 

(1996a), these two taxon names have the same type, with 

B. pennatula being a synonym of B. pennata var. minor.

Specimens identified as “B. pennatula” from the Gulf 

of California and Pacific Mexico should be reexamined to 

verify they belong here.

Bryopsis salvadoreana E. Y. Da

Bryopsis salvadoreana E. Y. Dawson, 1961c:405, pl. 5: 

fig. 2, pl. 6; Dawson, 1962b:228; Wysor, 2004:221, 

figs. 11, 12.

Algae of erect dense tufts, (2–)3–5 cm tall; cylindrical 

main axes, 200–400 μm in diameter; with both determinate and indeterminate branchlets more or less distichously 

arranged; determinate branchlets constricted at base; 

indeterminate branchlets irregularly distichous, becoming long, 3–12 mm, and may further divide, upper portion 

sometimes with one or more divisions (some appearing 

lobed), apices rounded; attached by a dense basal mass of 

entangled rhizoids.

Habitat. On boulders; intertidal

Distribution. Gulf of California: Isla Raza

(Las Islas de la Cintura). Pacific coast: Punta Chiriquín 

and Isla Meanguera, El Salvador; Panama.

Type Locality. Arrecife de Sacate, off Punta

Chiriquín, Golfo de Fonseca, El Salvador.

Remarks. Gulf of California specimens from 

Isla Raza (EYD-26103; US Alg. Coll.-37352) are tentatively refered to Bryopsis salvadoreana. Although they are 

similar to the type specimen of B. salvadoreana (EYD21758; US Alg. Coll-5162), the indeterminate branchlets 

of the Gulf material are not as abundant or apparently 

as well developed, and most are not as long and lack the 

fully developed upper divisions of the type. More Gulf collections and type locality material are needed for further 

study, along with molecular analyses, to test their taxonomic and phylogenetic status.

Derbes

Derbesiaceae Hauck, 1884:421.

Halicystaceae G. M. Smith, 1930:227, as “Halicystidaceae”; Smith, 1944:69; Papenfuss, 1955:129.

Algae of this family have two morphologically very 

different life history phases: a filamentous, siphonous 

sporophyte and vesiculate gametophytes. Sporangia with 

a basal septum are borne laterally on the sporophyte 

filaments and produce zoospores with numerous flagella 

(stephanokontic). Gametangia are formed in isolated portions of the cytoplasm inside the gametophytic vesicles 

and produce biflagellate anisogametes.

Remarks. Although life histories for many 

species, including the generitype, involve an alternation of 

heteromorphic forms, variations in life histories of some 

species of Derbesia have also been reported: (1) macrogametes that develop new sporophytes by parthenogenesis, 

(2) development of vesicular phase directly from sporangia without the production of zoospores, (3) sporophytes 

that grow from zoospores, and (4) a haploid gametophyte 

that alternates with heterokaryotic sporophyte (the latter 

characterized by the presence of haploid nuclei from the 

two parents, e.g., D. tenuissima (Moris et De Notaris) P. 

Crouan et H. Crouan, 1867). Under these circumstances, 

nuclear fusion occurs in young sporangia shortly before 

meiosis and cleavage of zoospores.

One genus is known in the northern Gulf of California. A vesiculate “Halicystis-phase” is also reported in the 

northern Gulf. Its life history is unknown, but it may be 

involved with a Derbesia or possibly another genus (see 

below for various reported life histories).

Derbesia Solier

Derbesia Solier, 1846:452; Solier, 1847:157.

Halicystis Areschoug, 1850:446.

Thalli are of two morphologically different forms. The 

sporophyte is composed of siphonous cylindrical filaments 

that are usually erect, and loosely unilaterally, irregularly 

or occasionally dichotomously branched, and attached by 

a prostrate system of irregularly branched rhizoidal filaments or lobed haptera. Although usually absent, simple 

or compound transverse septa sometimes develop in the 

siphonous filaments, mostly at base of branches. The 
n u m b e r 9 4 • 8 3

gametophytes are spherical to pyriform single-cell vesicles, “Halicystis-phase.” Both the filaments and vesicles 

are multinucleate and contain numerous disc-shaped or 

spindle-shaped small chloroplasts, which in some species 

have one to three pyrenoids, while other species are without pyrenoids.

Asexual reproduction can occur by fragmentation and 

proliferation of extruded protoplasts. The diploid, filamentous sporophyte, Derbesia-phase, produces stephanokont 

zoospores in modified lateral sporangia separated from 

the vegetative filament by a septum (or plug) or a double 

septum near their bases. A haploid vesicular gametophyte, 

Halicystis-phase, has been shown in the life histories for 

some species of Derbesia. A gametangium appearing as 

a transverse band within the vesicle develops biflagellate 

anisogametes that are released forcibly through papillae 

on the vesicle wall.

Remarks. Filamentous species of Derbesia

occur in the intertidal and down to 23 m depths of tropical to boreal seas worldwide. There are at least three types 

of life histories known to involve species of Derbesia. Culture studies have shown life histories with a sporophytic 

filamentous Derbesia-phase and a gametophytic vesicular 

Halicystis-phase (Hollenberg, 1935, 1936; Kornmann, 

1938; Drew, 1955; Page, 1970; Rietema, 1975). Sporophytic “Derbesia-like” phases have also been linked 

to the gametophytes of some species of Bryopsis (Hüstede, 1964; Rietema, 1975), and apparent sporangia of a 

New England species of Derbesia developed directly into 

vesicular gametophytes without producing any zoospores 

(Sears and Wilce, 1970).

Two species and one unidentified species are reported 

in the southern Gulf: Derbesia turbinata M. Howe et Hoyt 

(1916) by Dawson (1966b), D. hollenbergii W. R. Taylor 

(1945) by Dawson (1959a), and “Derbesia sp.” by RochaRamírez and Siqueiros-Beltrones (1991). The Gulf species 

of Derbesia need to be investigated morphologically, with 

culture studies and DNA analyses.

One species and one unidentified species of filamentous 

Derbesia are reported in the northern Gulf of California.

Key to the dfferent life history phases of Derbe

1a. Sporophyte (Derbesia): cylindrical, coenocytic filaments without a distinct percurrent axis; branched sparsely to moderately, predominately unilateral or sometimes irregular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1b. Gametophyte: a spherical vesicle (a single large cell), up to 15 mm in diameter, above a very short stalk . . . . . . . . . . . . . .

. . . . . . . . 

2a. Filaments branched mostly unilaterally; 50–65 μm in diameter  . . . . . . .

2b. Filaments sparsely and irregularly branched; 40–70 μm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Derbesia sp.

Derbesia marina (Lyngbye) Solie

FIGURE 38

Vaucheria marina Lyngbye, 1819:79, pl. 22A (sporophyte).

Derbesia marina (Lyngbye) Solier, 1846:453; Solier 

1847:157; Collins, 1909b:407, pl. 17: fig. 152; Setchell and Gardner, 1920:164, pl. 15: fig. 3; Smith, 

1944:71, pl. 8: fig. 4; Dawson, 1956:34, fig. 15; 

Dawson, 1961b:380; Dawson, 1962c:278; Abbott 

and Hollenberg, 1976:115, fig. 73; Huerta-Múzquiz, 

1978:336; Devinny, 1978:538; Kobara and Chihara, 

1980:214, figs. 8–12; Kobara and Chihara, 1981:2, 

figs. 1–3, 7–20; Pedroche and González-González, 

1981:64; Lewbel et al., 1981:165; Huerta-Múzquiz 

and Mendoza-González, 1985:44; Mendoza-González 

and Mateo-Cid, 1985:22; L. Aguilar-Rosas and Pacheco-Ruíz, 1986:77; Dreckmann et al., 1990:26, 37; 

Mateo-Cid and Mendoza-González, 1991:24, 27; 

Mateo-Cid and Mendoza-González, 1992:25; MateoCid and Mendoza-González, 1993:9–11; Mateo-Cid 

et al., 1993:52; Stout and Dreckmann, 1993:5; MateoCid and Mendoza-González, 1994a:57; Mateo-Cid 

and Mendoza-González, 1994b:45; León-Tejera and 

González-González, 1994:492; Mendoza-González et 

al., 1994:113; González-González et al., 1996:293; Pacheco-Ruíz and Zertuche-González, 1996a:432; Mendoza-González and Mateo-Cid, 1996:76, 88, pl. 28: 

figs. 123–125; Bucio-Pacheco and Dreckmann, 1998:43; 

Mendoza-González and Mateo-Cid, 1998:25; Yoshida, 

1998:144, fig. 1-13: E–G; Servière-Zaragoza et al., 

1998:169; Rodríguez-Morales and Siqueiros-Beltrones, 

1999:22; López et al., 2000:339; Wysor, 2004:223; 

Riosmena-Rodríguez et al., 2005:101; Pedroche et al., 

2005:60; Mateo-Cid et al., 2006:59; Dreckmann et al., 

2006:154.
8 4 • 

Derbesia vaucheriaeformis sensu Chávez B., 1972:268 

[non Derbesia vaucheriaeformis (Harvey) J. Agardh, 

1887:34].

Gastridium ovale Lyngbye, 1819:72, pl. 18: fig. B 

(gametophyte).

Halicystis ovalis (Lyngbye) Areschoug, 1850:447; Setchell and Gardner, 1920b:155, pl. 14: fig. 3; Hollenberg, 

1935:782, pls.1–4; Smith, 1944:70, pl. 9: fig. 1; Dawson et al., 1960a:10, 14, pl. 1: fig. 5; Dawson et al., 

1960b:5, 6, 8, 11, 13; Dawson, 1961b:380; Scagel, 

1966:116, pl. 2: figs. A–F; Abbott and Hollenberg, 

1976:115, fig. 75; Kobara and Chihara, 1980:214, 

figs. 1–7, 13; Lewbel et al., 1981:165; L. Aguilar-Rosas and Bertsch, 1983:116, 122; R. Aguilar-Rosas et 

al., 1990:123; Stewart, 1991:115; Mendoza-González 

and Mateo-Cid, 1992:24; Mateo-Cid et al., 1993:52; 

González-González et al., 1996:137; Pacheco-Ruíz and 

Zertuche-González, 1996a:432; L. Aguilar-Rosas et al., 

2000:133; Pacheco-Ruíz et al., 2008:202.

Sporophytes small filamentous tufts, up to 15 mm tall; 

filaments of long, cylindrical coenocytic cells, 50–65 μm 

in diameter; branching mostly unilateral, with subacute 

apices; attached by irregularly branched prostrate filaments. 

Chloroplasts without pyrenoids. Sporangia formed on ultimate branchlets, borne on a short pedicel with a crosswall 

at apex of pedicel; ovoid to pyriform, 150–200 μm long 

and 100–160 μm in diameter; producing zoospores.

Gametophytes (Halicystis-phase of Derbesia marina) 

erect, spherical vesicles, up to 15 mm in diameter; above a 

very short stalk attached to calcareous substratum by basal 

rhizoids that often penetrate it. Vesicle with a large vacuole, and peripheral cytoplasm containing multiple nuclei 

and numerous chloroplasts without pyrenoids. Vegetative 

reproduction by regeneration of new vesicles from apparently perennial attachment rhizoids. Sexual reproduction 

by biflagellate anisogametes, produced in separate gametophytic vesicles. Male vesicles develop slender gametes in 

brownish-green fertile areas of the cytoplasm. Female vesicles develop larger and rounder gametes in greenish-black 

fertile areas. Gametes forcibly released through 1 or more 

papillae (appearing as whitish dots) on the vesicle wall.

Remarks. Culture life history studies have 

linked gametangial Halicystis to sporophytic Derbesia 

from the Pacific coast of North America (Hollenberg, 

1935, 1936; Scagel, 1966), Japan (Kobara and Chihara, 

1980, 1981), Europe (Kornmann, 1938; Feldmann, 1950; 

Eckhardt et al., 1986), and Bermuda (Page, 1970). Thus, 

species of Halicystis are usually referred to as the “Halicystis-phase of Derbesia.”

Habitat. On rocks, epiphytic on crustose corallines; intertidal

Distribution. Gulf of California: Puerto

Peñasco to Bahía Concepción; Bahía del Rincón to San 

Jose del Cabo; Matzatlán, Sinaloa. Pacific coast: Alaska to 

Baja California; Nayarit to Chiapas; Panama; Japan.

Type Locality. Kvivig, Strømø, Faeroe

Islands, Denmark.

Derbesia sp.

Derbesia sp. of Dawson, 1966a:7; Mateo-Cid et al., 

2006:48.

Algae forming small, filamentous tufts, up to 10 mm 

long, of filaments sparsely and irregularly branched, without a percurrent axis, and 40–70 μm in diameter. Siphonous cells with numerous chloroplasts that lack pyrenoids.

Habitat. Growing on rocks or sometimes

entangled on gorgonians; mid to low intertidal.

Distribution. Gulf of California: Punta Pelícano, vicinity of Puerto Peñasco

Remarks. Although lacking a distinct main 

axis and reproductive structures, the material reported 

FIGURE 38. Derbesia marina: Sporophyte: Filaments bearing lateral 

sporangia, each with a single basal septum (after Collins, 1909b: pl. 

17, fig. 152).
n u m b e r 9 4 • 8 5

as “Derbesia sp.” (Dawson, 1966a) somewhat resembles 

sparsely and irregularly branched forms of Bryopsis pennata. More collections, particularly fertile specimens, are 

needed to determine which species of filamentous Derbesia are present in the northern Gulf.

Halicystis-phase

FIGURE 39A,B

“Halicystis ovalis” sensu Norris and Bucher, 1976:3, fig. 2.

Northern Gulf specimens: spherical vesicles, up to 10 

mm in diameter, and dark emerald green. Vesicle above a 

very short stalk; lowermost portion of basal stalk attached 

to Amphiroa by rhizoidal growths.

Habitat. Epiphytic on the articulated coralline, Amphiroa; low intertidal to shallow subtidal

Distribution. Gulf of California: Isla Willard (Bahía San Luis Gonzaga) to Bahía Concepción

Remarks. Although similar in morphology to 

the Pacific coast Halicystis ovalis-phase, the northern Gulf 

material referred to as “Halicystis-phase” were epiphytic 

on the articulated coralline, Amphiroa vanbosseae (Norris and Bucher, 1976), whereas the Pacific coast Halicystis 

ovalis-phase of Derbesia marina (Abbott and Hollenberg, 

1976; Scagel et al., 1989) and the Japanese H. ovalisphase of D. marina and H. parvula-phase of D. tenuissima

(Kobara and Chihara, 1980) are only reported on crustose 

corallines.

The life history of Gulf of California Halicystis, or even 

if the vesicles are gametangial or sporophytic, is unknown. 

Thus far, known life histories that involve a “Halicystis” 

either have an alternate heteromorphic phase (which is 

either a Derbesia or an alternate that is another genus, 

i.e., not a Derbesia) or have a repeating life history of only 

Halicystis-forms (i.e., with no alternate phases). Therefore, Gulf of California Halicystis, including those of the 

southern Gulf (reported epiphytic on mangroves; MateoCid et al., 1993, as “vesicular gametangium [of] Derbesia 

marina”), need to be tested for host specificity and need culture studies to elucidate their life history and reproductive 

condition (whether gametangial, as generally presumed, or 

sporophytic), and molecular analyses to determine their 

taxonomic status and phylogenetic relationships.

Codia

Codiaceae Kützing, 1843:302, 308.

Thalli may be growing prostrate or erect, lack calcification, and are multiaxial, spongy, unbranched or 

branched, variable forms, that can be globose, cushionlike, cylindrical, compressed, flattened, or blade-like, and 

are attached by compacted masses of rhizoids. Internally 

the thalli are composed of branching, tubular, mostly 

FIGURE 39. Halicystis-phase: A, B. Gametophytes: Globose, coenocytic thalli (JN-5732); epiphytic on the articulated coralline Amphiroa.
8 6 • 

nonseptate, multinucleate filaments (siphons). Branching of the filaments (siphons) is mostly sympodial, with 

the outer branched filaments forming a cortical layer of 

utricles at the surface. Cells contain numerous small nuclei 

and discoid chloroplasts (without pyrenoids), and no amyloplasts, with cell walls containing mostly mannan. Accessory pigments include siphonoxanthin and siphonein.

Asexual reproduction is by fragmentation or by parthenogenesis in some. Sexual reproduction is by biflagellate anisogametes. Gametangia are developed laterally on 

the utricles and cut off by a basal septum.

The family consists of five genera, with the speciesrich genus Codium represented in the northern Gulf of 

California.

Codium Stackhouse

Codium Stackhouse, 1797:xvi, xxiv.

Algae are spongy, non-calcareous macrothalli of definite shapes, with species that are either prostrate or repent 

globose or pulvinate forms, or erect branching cylindrical, 

compressed, or complanate forms. Internally, the thalli are 

composed of branched filaments that are organized into 

a medulla of longitudinal, densely intertwined, colorless 

coenocytic filaments (medullary siphons) and an outer 

surface layer of inflated, cylindrical or clavate, photosynthetic utricles with an apical wall that is usually thickened 

and often ornamented in a characteristic pattern. Utricles 

are developed primarily by the enlargement of sympodial 

branches of medullary siphons and secondarily by budding 

or by production of additional utricle-forming medullary 

siphons from basal portion of existing utricles. Rhizoidal 

filaments (siphons), which grow into the medulla, are also 

produced from basal portion of the utricles. Colorless 

hairs are produced along the sides of the utricles, with a 

basal plug which remains as a prominent scar after the 

hair falls off.

Asexual reproduction is by parthenogenesis, fragmentation, or the cutting off of modified aborted gametangia. 

Species may be monoecious or dioecious. Gametangia are 

cylindrical or fusiform to ovate above a short pedicel with 

a basal plug and are borne laterally on the utricles. Biflagellate anisogametes are extruded in gelatinous masses upon 

rupture of the apices of the gametangia. Male gametes are 

very small, with one or two chloroplasts, and female gametes are much larger and have numerous chloroplasts. Life 

histories are not fully known. The zygote develops into an 

amorphous, prostrate vesicle that produces erect elongate 

vesicles; these, in turn, initiate primary utricle-producing 

siphons that eventually consolidate into a multiaxial thallus. Some species are known to have a filamentous derbesioid or vaucherioid (Vaucheria-like) stage in their life 

history (Fletcher et al., 1989; Yang et al., 1997).

Remarks. Codium is a widespread genus with 

species occurring from the intertidal down to at least 70 

m depths (Norris and Olsen, 1991). Although species are 

found in almost every sea (except the Arctic and Southern 

Ocean), the largest number of species of Codium are in 

subtropical to temperate regions.

Several species that were originally described from the 

Gulf of California (Setchell and Gardner, 1924a; Dawson, 

1944) have been placed in synonymy with other species on 

the basis of comparative morphological studies (Pedroche et 

al., 2002). These species concepts need to be tested by molecular phylogenetic and morphological analyses. One species, 

Codium picturatum Pedroche et P. C. Silva (1996), described 

from Colima (Pacific Mexico), has also been reported from 

the southern Gulf, and includes records of “C. setchellii”

(Pedroche et al., 2002, 2005; Riosmena-Rodríguez et al., 

2005) (note “C. setchellii” of Dawson [1949] and Brusca 

and Thomson [1975] may also belong here).

Currently, there are three species known in the northern Gulf of California. Two of these are also known from 

the Pacific coast of Mexico, and one, C. brandegeei, is 

apparently a Gulf-endemic species.

Key to the Species of Codium in the no

1a. Thalli usually very long, up to 85 cm tall; axes and branches mostly cylindrical, branches often flattened at dichotomies; 

very large utricles, most greater than 900 μm in length  . . . . . . . . .

1b. Thalli shorter in overall length; axes and branches variable, of three forms: cylindrical throughout or cylindrical but 

compressed at the dichotomies or compressed but with cylindrical ultimate branches; utricles less than 900 μm (mostly 

550–850 μm long)  

2a. Utricles claviform, constricted below; utricle apices conical, hyaline, and very thick (up to 85 μm thick) . . C. brandegee

2b. Utricles mostly cylindrical (if constricted, only slightly); utricle apices truncate or rounded (not conical), hyaline, and thinner (up to 46 μm thick)  . . . . .
n u m b e r 9 4 • 8 7

Codium amplivesiculatum

Setchell et N. L. Gardn

FIGURES 40, 41A,B

Codium amplivesiculatum Setchell et N. L. Gardner, 1924a:709, pl. 15: figs. 28, 29, pl. 35; Dawson, 

1944:217; Dawson, 1959a:7, 16; Dawson, 1961b:381; 

Norris, 1973:4, 17; Norris, 1985:208; EspinozaÁvalos, 1993:333; González-González et al., 1996:132; 

Pacheco-Ruíz and Zertuche-González, 1996a:432; 

Núñez-López et al., 1998:39; Pedroche et al., 2002:40, 

figs. 42–53; R. Aguilar-Rosas et al., 2005b:36; Pedroche et al., 2005:69; Servière-Zaragoza et al., 2007:7; 

Pacheco-Ruíz et al., 2008:202; Riosmena-Rodríguez 

and Holguín-Acosta, 2008:1, figs. 1–12.

Codium decorticatum sensu Howe, 1911:494; Setchell 

and Gardner, 1930:130; Taylor, 1939:9; Huerta-M, 

1978:339; Mateo-Cid and Mendoza-González, 1992:25 

[in part; see Pedroche et al., 2005:72]; Mateo-Cid et 

al., 1993:52 [with a query]; González-González et al., 

1996:369 [Gulf specimens only]; Silva et al., 1996b:235 

[non Codium decorticatum (Woodward) M. Howe, 

1911:494].

Codium dichotomum sensu Taylor, 1945:68; Dawson, 

1949:229 [in part; see Pedroche et al., 2002:40] [non 

Codium dichotomum (Hudson) S. F. Gray, 1821:293].

Codium fernandezianum sensu Taylor, 1945:71 [in part; 

see Pedroche et al., 2002:40] [non Codium fernandezianum Setchell, 1937a:592].

Codium longiramosum Setchell et N. L. Gardner, 

1924a:710, pl. 15: fig. 27, pl. 37; Taylor, 1945:70 [in 

part]; Dawson, 1961b:382; Dawson, 1966b:55; Stewart and Stewart, 1984:140; González-González et al., 

1996:134.

Codium magnum E. Y. Dawson, 1950f:298, fig. 1; Dawson, 1961b:382; Dawson, 1962c:278, 280; HuertaMúzquiz, 1978:336; Huerta-Múzquiz and MendozaGonzález, 1985:44; Ibarra-Obando and R. AguilarRosas, 1985:96; Mendoza-González and Mateo-Cid, 

1985:22; Sánchez-Rodríguez et al., 1989:36, 38–39; 

Rocha-Ramírez and Siqueiros-Beltrones, 1991:30; Mateo-Cid and Mendoza-González, 1994a:52; GonzálezGonzález et al., 1996:134; Rodríguez-Morales and 

Siqueiros-Beltrones, 1999:22; Riosmena-Rodríguez et 

al., 2005:101.

Algae erect, mostly cylindrical, up to 85 cm tall 

(largest to 10 m in length; Pedroche et al., 2002); simple or branched dichotomously to subdichotomously (or 

sometimes trichotomously); branches usually flattened at 

dichotomies; attached below by a spongy mass of rhizoidal filaments. Lower branches (6–)13–20 mm in diameter, 

often with long interdichotomies, up to 40 cm in length; 

dichotomies flattened, up to 10 cm wide. Upward branches 

gradually narrowing to apices, (1–)3–10 mm in diameter. 

Medullary filaments, 25–110 μm in diameter, with lateral 

cell wall thickenings (‘cellulose plugs”). Utricles cylindrical to clavate, (840–)1180–1930(–3025) μm in length, and 

(140–)230–410(–655) μm in diameter in upper portions, 

and near the base (40–)110–235(–385) μm in diameter; 

utricle apices truncate or rounded, 4.5–8.0 μm thick; utricle wall sides 1.5–2.3 μm thick; hairs and scars present, 

never abundant, about 220 μm below the apices.

Gametangia elongated, fusiform to ovate, wider at 

their base, (160–)255–340(–430) μm long by (46–)90–

140(–170) μm in diameter; borne about (370–)450–600

(–750) μm below the utricle apex on short pedicel, up to 

15 μm long.

Habitat. Attached or free-floating in esteros or

lagoons; on rocks; low intertidal to shallow subtidal, usually 0.5 to 17 m depths.

Distribution. Gulf of California: Puerto

Refugio, Isla Ángel de la Guarda to Punta Arena (north of 

Cabo Pulmo). Pacific coast: Bahía de San Quintín to Bahía 

Magdalena; Isla Cedros; Isla Guadalupe; Isla Clarión and 

Isla Socorro (Islas Revillegigdo); Nayarit to Jalisco.

Type Locality. Drift, most likely Isla

Estanque (also known as Pond Island and Isla La Bibera); 

however, Setchell and Gardner (1924a:709) only stated 

“near south end of [Isla] Ángel de la Guarda, Gulf of 

California.”

Remarks. Codium amplivesiculatum is the 

largest species of Codium, and may be the largest green 

alga in the Gulf of California. Two other species, the Gulf 

of California C. longiramosa Setchell et N. L. Gardner 

(1924a) and the Pacific Baja California C. magum E. Y. 

Dawson (1950f), were recently treated as synonyms of C. 

amplivesiculatum by Pedroche et al. (2002). Hommersand 

(1972) and Pedroche et al. (2005:70) noted that the problematic separation of the Gulf C. amplivesiculatum and the 

Japanese C. cylindricum Holmes (1896; Yoshida, 1998) 

and their phylogenetic relationship needs to be clarified.

Codium brandegeei Setchell et N. L. Gardn

FIGURES 41C–F, 42

Codium brandegeei Setchell et N. L. Gardner, 1924a:712, 

pl. 14: figs. 25, 26, pl. 30; Dawson, 1944:215; 
8 8 • 

FIGURE 40. Codium amplivesiculatum: Habit of a large specimen (JN-5700).
n u m b e r 9 4 • 8 9

FIGURE 41. Species of Codium: Detail of utricles, some with gametangia and/or hairs, showing variation in shape and size. A, B. Codium amplivesiculatum: Large, wide utricles; (drawn from the type specimen, I. M. Johnston-74, after Setchell and Gardner, 1924a: pl. 15, figs. 28, 29). 

C–F. Codium brandegeei: Variation in utricle size: C, D. Utricles drawn from the type of the synonym C. cervicorne (Marchant-8, after Setchell 

and Gardner, 1924a: pl. 14, figs. 19, 20). E, F. Utricles of C. brandegeei (drawn from the type specimen, T. S. Brandegee-28, after Setchell and 

Gardner, 1924a: pl. 14, figs. 25, 26). G, H. Codium simulans: Variation in utricles (drawn from the type specimen, I. M. Johnston-8, after 

Setchell and Gardner, 1924a: pl. 14, figs. 21, 22).
9 0 • 

González-González et al., 1996:132; Pedroche et al., 

2002:54–55, figs. 19, 66–80; Pedroche et al., 2005:70; 

Mateo-Cid et al., 2006:49, 59; Pacheco-Ruíz et al., 

2008:202.

Codium anastomosans Setchell et N. L. Gardner, 

1924a:711, pl. 16: figs. 36, 37; Dawson, 1944:218; 

Dawson, 1961b:381; Martínez-Lozano et al., 1991:22; 

Espinoza-Ávalos, 1993:333; González-González et al., 

1996:132.

Codium cervicorne Setchell et N. L. Gardner, 1924a:712, 

pl. 14: figs. 19, 20, pl. 32b; Dawson, 1961b:381; Martínez-Lozano et al., 1991:22; González-González et al., 

1996:132.

Codium cuneatum sensu Dawson, 1944:218 [non Codium 

cuneatum Setchell et N. L. Gardner, 1924a:708].

Codium macdougalii E. Y. Dawson, 1944:218, pl. 35: fig. 

1; Dawson, 1961b:382; Dawson, 1966a:7; Dawson, 

1966b:55; Norris, 1973:4, 17, 19; Espinoza-Ávalos, 

1993:333; González-González et al., 1996:134; L. Aguilar-Rosas et al., 2000:133.

Codium reductum Setchell et N. L. Gardner, 1924a:707, 

pl. 14: figs. 23, 24, pl. 33; González-González et al., 

1996:134.

Codium simulans sensu Dawson, 1944:216; Dawson, 

1945a:22 [non Codium simulans Setchell et N. L. Gardner, 1924a:706].

Codium tomentosum sensu Mendoza-González and Mateo-Cid, 1986:420 [non Codium tomentosum Stackhouse, 1797:xxiv].

Algae erect, more or less cylindrical throughout, up to 

15 cm tall; branching irregularly dichotomous; attached by 

discoid base of rhizoidal filaments. Branches cylindrical; in 

lower portions sometimes narrow and constricted at dichotomies; in midportions broadening; and in terminal portions 

sometimes swollen to club-shaped, up to 1.6 cm in diameter. Medulla filaments 23–45(–50) μm in diameter. Utricles slender, cylindrical to clavate, constricted below, giving 

upper portion spoon-shaped appearance, (460–)540–820(–

1150) μm long by (70–)95–200(–290) μm in diameter, and 

in basal portion (30–)40–70(–112) μm in diameter; sometimes with secondary utricles; utricle apices blunt to dome 

shaped, outer end with hyaline, laminated, conical cell wall, 

(30–)50–85 μm thick; hairs and scars present.

Gametangia fusiform or cylindrical, (130–)200–270(–

330) μm long by (40–)60–90(–125) μm in diameter; usually one per utricle, borne about (230–)280–390(–500) 

μm below the apices.

Habitat. On rocks and tidal platforms, often

exposed; mid intertidal to shallow subtidal, down to 10 

m depths.

Distribution. Gulf of California: Punta Pelícano (vicinity of Puerto Peñasco) to Guaymas; Bahía Concepción to Bahía de La Paz

Type Locality. “Probably at La Paz,” Baja

California Sur, Gulf of California (Setchell and Gardner, 

1924a).

Remarks. Codium brandegeei is apparently 

endemic to the Gulf of California, as the Pacific Mexico 

reports of “C. brandegeei” from Nayarit and Jalisco (Servière-Zaragoza et al., 1993, as “C. cervicorne”) and Guerrero (Salcedo-Martínez et al., 1988, as “C. cervicorne”)

were noted to be C. isabelae W. R. Taylor (1945) by 

Pedroche et al. (2005:71).

Although the Gulf C. macdougalii is now considered to be conspecific with C. brandegeei (Pedroche et 

al., 2002), in the field the “C. macdougalii-morph” can 

be distinguished from the other Gulf species of Codium

by the broad, swollen, club-shaped upper portions of its 

branches (Figure 42).

Codium simulans Setchell et N. L. Gardn

FIGURES 5B, 41G,H, 43

Codium simulans Setchell et N. L. Gardner, 1924a:706, pl. 

14: figs. 21, 22, pl. 31; Setchell and Gardner, 1930:133; 

Dawson, 1944:216; Dawson, 1949:233–234, 237, 

242; Dawson, 1959a:18; Dawson, 1961b:382; Norris, 

FIGURE 42. Codium brandegeei: Habit of “C. macdougalii-form” 

with swollen, club-shaped branches (JN-5020).
n u m b e r 9 4 • 9 1

1973:4, 17, 19; Brusca and Thomson, 1975:42; HuertaMúzquiz, 1978:338; Littler and Littler, 1981:151, 

153; Stewart and Stewart, 1984:140; Littler and Littler, 1984:15, 23, 25; Norris, 1985:209, fig. 16.2; 

Salcedo-Martínez et al., 1988:82; Rocha-Ramírez and 

Siqueiros-Beltrones, 1991:30; Martínez-Lozano et al., 

1991:22; González-González et al., 1996:135; AnayaReyna and Riosmena-Rodríguez, 1996:862; PachecoRuíz and Zertuche-González, 1996a:432; L. AguilarRosas et al., 2000:133; Pedroche et al., 2002:61, figs. 

65, 81–89; R. Aguilar-Rosas et al., 2005b:36; Pedroche 

et al., 2005:77; Hernández-Herrera et al., 2005:146; 

Riosmena-Rodríguez et al., 2005:101; Mateo-Cid et al., 

2006:49, 59; Servière-Zaragoza et al., 2007:7; PachecoRuíz et al., 2008:191, 202.

Codium anastomosans sensu Martínez-Lozano et al., 

1991:22 [non Codium anastomosans Setchell et N. L. 

Gardner, 1924a:711; which is now C. brandegeei Setchell et N. L. Gardner, 1924a:712 (see Pedroche et al., 

2005:780)].

Codium cervicorne sensu Martínez-Lozano et al., 1991:22; 

Servière-Zaragoza et al., 1998:168 [non Codium cervicorne Setchell et N. L. Gardner, 1924a:712 (see Pedroche et al., 2005:780)].

Codium conjunctum Setchell et N. L. Gardner, 1924a:706, 

pl. 15: figs. 32, 33, pl. 32a; Dawson, 1949:236, 238; 

Dawson, 1944:217; Sánchez-Vargas and Hendrickx, 

1987:161; Salcedo-Martínez et al., 1988:82; GonzálezGonzález et al., 1996:132.

Codium cuneatum Setchell et N. L. Gardner, 1924a:708, 

pl. 16: figs. 34–35, pl. 34; Setchell and Gardner, 

1930:133; Dawson, 1944:218; Dawson, 1945a:22; 

Dawson, 1949:227; Silva, 1951:99, pls. 3–6, figs. 23–

32; Dawson et al., 1960a:10, 26, pl. 5: fig. 2; Dawson et al., 1960b:6–14; Dawson, 1961b:382; Dawson, 

1966a:7; Dawson and Neushul, 1966:174; Chávez B., 

1972:268; Guzmán del Próo et al., 1972:260; Norris, 

1973:4, 18; Abbott and Hollenberg, 1976:116, fig. 76; 

Lewbel et al., 1981:165; L. Aguilar-Rosas and Bertsch, 

1983:116, 122; Stewart and Stewart, 1984:140; Mendoza-González and Mateo-Cid, 1985:22; HuertaMúzquiz and Mendoza-González, 1985:44; Mendoza-González and Mateo-Cid, 1986:420; SánchezRodríguez et al., 1989:38, 39; R. Aguilar-Rosas et 

al., 1990:123; Mateo-Cid and Mendoza-González, 

1991:27–28; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:18, 30; Martínez-Lozano et al., 1991:22; Mendoza-González and Mateo-Cid, 1992:24, 25; MateoCid and Mendoza-González, 1992:25; Mateo-Cid et al., 

1993:52; Servière-Zaragoza et al., 1993:482; MateoCid and Mendoza-González, 1994a:52; Mateo-Cid and 

Mendoza-González, 1994b:34; Mendoza-González et 

al., 1994:113; González-González et al., 1996:132; Pacheco-Ruíz and Zertuche-González, 1996a:432; NúñezLópez et al., 1998:39; Cruz-Ayala et al., 1998:194; 

Rodríguez-Morales and Siqueiros-Beltrones, 1999:22; 

Sánchez-Rodríguez et al., 2001:148; Pacheco-Ruíz et 

al., 2008:191.

Codium dichotomum sensu Taylor, 1945:68 [in part; see 

Pedroche et al., 2002:78] [non Codium dichotomum 

(Hudson) S. F. Gray, 1821:293].

Codium latum subsp. palmeri sensu Pedroche et al., 

2002:34, figs. 33, 34, 37 [non Codium latum subsp. 

palmeri (E. Y. Dawson) P. C. Silva, 1962:208; 

basionym:Codium palmeri E. Y. Dawson, 1945a:23 

(see Pedroche et al., 2005:76, 77)].

Codium tomentosum sensu Howe, 1911:493 [non Codium tomentosum (Hudson) Stackhouse, 1797:xxivi].

Codium unilaterale Setchell et N. L. Gardner, 1924a:710, 

pl. 15: figs. 30–31, pl. 36; Martínez-Lozano et al., 

1991:22; González-González et al., 1996:135.

Algae erect, up to 35 cm tall; branching dichotomous 

to subdichotomous; branches 4–15 mm in diameter and 

variable in shape, either cylindrical throughout; cylindrical, and only slightly flattened to cuneate at branch dichotomies (2.0–25 mm broad); or compressed, with only stipe 

and branch tips cylindrical; and attached by a disc of rhizoidal filaments. Medullary filaments 19–77 μm in diameter. Utricles cylindrical, subcylindrical to slightly claviform, 

(330–)450–800(–1300) μm long by (70–)200–250(–440) 

μm in diameter in upper portions and (30–)40–120(–385) 

μm in diameter near the base; apices of utricle smooth, 

mostly rounded, with hyaline, apical cell wall, 20–46 μm 

thick. Hairs or scars present, single to numerous, about 

120 μm below the thickened distal end of the utricle.

Gametangia, 1 or 2 per utricle, mostly subfusiform 

to fusiform, (130–)180–290(–330) μm long by (50–)60–

110(–162) μm in diameter and usually narrowed at the 

base; borne on a short pedicel (up to 10 μm in length), 

(90–)230–350(–670) μm below utricle tip.

Habitat. On rocks and tidal platforms, usually

exposed; mid intertidal to shallow subtidal, down to 20 m 

depths.

Distribution. Gulf of California: Puerto

Peñasco to Cabeza Ballena. Pacific coast: Santa Cruz 

Island (California Channel Islands) to Guerrero; Isla Guadalupe; Isla Clarión (Islas Revillagigedo).

Type Locality. Isla San Marcos, off the Baja

California Sur coast between Caleta de San Lucas and 

Punta Chivato (separated by Canal de San Marcos [Craig 

Channel]), Gulf of California.
9 2 • 

Remarks. The most common species of 

Codium in the northern Gulf, C. simulans is found from 

February to September. Silva (1951:101, as “C. cuneatum”) noted it “to be a unique species whose young stages 

are more or less repent, intricate, and basally anastomosing, but whose adult thalli are erect and openly branched...

[and] degree and incidence of flattening are variable.” As 

currently understood (Pedroche et al., 2002), C. simulans comprises a wide variety of different morphologies 

(many of which were previously separate species) that 

may be cylindrical throughout or mostly cylindrical but 

compressed at the dichotomies or mostly compressed with 

only the stipe and ultimate branches cylindrical.

Subtidal specimens of Codium sp. A (Figure 44; 

JN-5286) from Puerto Refugio (Isla Ángel de la Guarda) are 

somewhat similar to “C. cuneatum-form” of C. simulans

but are much larger, wider and flattened throughout. While 

they are tentatively referred to this species, they should be 

further studied to see if they represent a new taxon.

Uncertain Record:

Codium fragile (Suringar) Hari

Acanthocodium fragile Suringar, 1868:258.

Codium fragile (Suringar) Hariot, 1889:32.

Remarks. Although Codium fragile is widely 

reported on the Pacific coast from Alaska to Baja California (e.g., Abbott and Hollenberg, 1976; GonzálezGonzález et al., 1996; Scagel et al., 1989; Pedroche et al., 

2005), its reported presence in the northern Gulf (Martínez-Lozano et al., 1991) needs to be verified.

Halimedin

Halimedineae Hillis-Colvinaux, 1984:287.

The suborder Halimedineae is characterized by heteroplasty, presence of a concentric lamellar system, and cell 

walls that are composed primarily of xylan, with cellulose 

absent (or only barely present). Allelochemicals (secondary natural products) are usually present. Reproduction is 

mostly holocarpic (i.e., entire cytoplasm of the coenocytic 

thallus is transformed into gametes). Reproductive structures are usually without septa.

Remarks. Species of the suborder Halimedineae are mostly tropical to subtropical, with a few temperate, in distribution. Many members are known to produce 

unique natural products, diterpenoid metabolites (Paul 

and Fenical, 1985, 1986).

FIGURE 43. Codium simulans: Habit, with terete to slightly compressed axis and branches, flattened at dichotomies (JN-5713).
n u m b e r 9 4 • 9 3

FIGURE 44. Codium sp. A: Specimen somewhat similar to “Codium cuneatum-form” of C. simulans: Flat, with exceptionally broad axes, 

possibly a new species (JN-5286).
9 4 • 

One of its families, the Udoteaceae J. Agardh (1887), 

is represented in the southern Gulf by two species Chlorodesmis hildebrandtii A. et E. S. Gepp (1911; see Dawson, 

1959a, 1961b; González-González et al., 1996; MateoCid et al., 2000a) and Geppella decussata E. Y. Dawson 

(1959a) described from Isla San Francisco (off south end 

of Isla San José, Baja California Sur). There is a problem with the generic placement of G. decussata, as the 

generitype of Geppella Børgesen (1940), G. mortensenii 

Børgesen, is now conspecific with Rhipiliopsis mortensenii (Børgesen) Fraghaly et Denizot (1979). Therefore, the 

generic status of the Gulf G.? decussata is uncertain (see 

also Tanaka and Itono, 1977; Dong and Tseng, 1983) and 

in need of reinvestigation.

Two families of the suborder Halimedineae, the Caulerpaceae and Halimedaceae, are represented in the northern as well as the southern Gulf of California.

Caul

Caulerpaceae Kützing, 1843:302, 307.

Algae are differentiated into distinct erect portions 

and prostrate, creeping stolon-like runners, that issue rhizoids that attach it to sand or hard substratum. The erect 

portions exhibit a great variety of forms and sizes, from 

tubular to foliose, and can be simple to much branched. 

The vegetative portions of these algae are coenocytic, multinucleate, and with two types of plastids, chloroplasts 

and amyloplasts. Structurally, there is an internal web 

or network of trabeculae that originate from the interior 

walls of the thallus and are unique to this family.

Asexual reproduction is by fragmentation and also 

by vegetative propagation of the stolons. Zooidangia are 

apparently known only in the non-holocarpic Caulerpella. 

Its reproductive structures are in external stalked-whorls, 

developed from the basal portion of a fertile branch that is 

separated from the lateral branch by a septum (crosswall). 

Spores are released via a discharge tube at apex of fertile 

branch. Sexual reproduction has been reported in a few 

species of holocarpic Caulerpa (e.g., Clifton and Clifton, 

1999). The species of Caulerpa thus far studied are diploid, forming biflagellate anisogametes within the contents 

of the branches by the separation of the cytoplasm without development of septa (crosswalls). Anisogametes are 

released through superficial papillae.

Remarks. The Caulerpaceae is composed of 

two genera. Caulerpa is found in tropical to subtropical 

seas worldwide and in temperate waters of the Southern 

Hemisphere, and Caulerpella Prud’homme van Reine et 

Lokhorst (1992) is found in tropical and subtropical seas.

One genus, Caulerpa, is known in the Gulf of 

California.

Caulerpa J. V. Lamou

Caulerpa J. V. Lamouroux, 1809d:332.

Algae included in Caulerpa are easily recognized by 

their differentiated fronds: erect upper branches of diverse 

morphologies that arise from prostrate, creeping stolons 

(rhizome-like runners), which are attached to the substratum at various points by groups of colorless rhizoids. The 

erect axes and branches are cylindrical to blade-like and 

simple or pinnately, radially, or bilaterally branched; their 

shape and pattern are species specific and may be elaborate and variable. The thallus is coenocytic throughout, 

i.e., multinucleate, with numerous chloroplasts and amyloplasts (specialized for starch accumulation), a cell wall 

principally of xylan, and an elaborate internal network of 

trabeculae (slender internal strands of cell wall material) 

that traverses the cytoplasm throughout the thallus, presumably structurally bracing the cell wall. Growth is apical and indeterminate.

Asexual reproduction is by fragmentation. Sexual 

reproduction, where known, is holocarpic (i.e., the entire 

cytoplasm is transformed into gametes), and the thallus 

remains unmodified, and no crosswalls are formed. Both 

dioecious and monoecious species are reported, with 

anisogametes extruded in gelatinous strands through 

papillae on the siphon wall.

Remarks. Members of Caulpera grow from 

the intertidal down to depths of at least 81 m (Littler 

et al., 1985, 1986; Norris and Olsen 1991). Exhibiting 

extreme morphological plasticity, their habit is influenced 

by both biotic and physical factors of their environment, 

e.g., light and water motion, which has resulted in numerous described subspecies, varieties, forms, or ecads. When 

wounded or grazed by herbivorous fish or invertebrates 

(e.g., sacoglossan molluscs) the thallus responds by extruding a yellowish gelatinous mass that hardens in a few minutes to form a wound plug (Dreher et al., 1978; Dawes 

and Goddard, 1978; Menzel, 1980, 1988). Several species produce secondary metabolites, caulerpin (Santos and 

Doty, 1968; Maiti et al., 1978) and caulerpicin (Doty and 

Aguilar-Santos, 1966; Mahendran et al., 1979; Nielsen 

et al., 1982; Vest et al., 1983) or caulerpenyne (Amico 

et al., 1978; Faulkner, 1984; Amade and Lemée, 1998), 

that serve as a chemical defense, making them unpalatable or even toxic to certain herbivores (e.g., Norris and 

Fenical, 1982; Paul and Fenical, 1986, 1987). Some species of sacoglossan mollusks, specialist herbivores, ingest, 
n u m b e r 9 4 • 9 5

transfer and store the natural products, along with the 

chloroplasts, of Caulerpa, in their tissues (e.g., Doty and 

Aguilar-Santos, 1970; Trench, 1973).

Three species and one variety are reported in the southern Gulf: C. antoensis Yamada (1940; = C. arenicola W. R. 

Taylor, 1950) by Huerta-Múzquiz and Mendoza-González 

(1985) and Rocha-Ramírez and Siqueiros-Beltrones (1991); 

C. peltata J. V. Lamouroux (1809c) by Dawson (1959a), 

Huerta-Múzquiz and Mendoza-González (1985), RochaRamírez and Siqueiros-Beltrones (1991), and Mateo-Cid 

et al. (2000a); and C. racemosa (Forsskål) J. Agardh 

(1873) by Dawson (1944), Huerta-Muzquiz (1978), 

Mendoza-González and Mateo-Cid (1986), Anaya-Reyna 

and Riosmena-Rodríguez (1996), Rodríguez-Morales and 

Siqueiros-Beltrones (1999, as “C. racemosa var. uvifera”), 

Paul-Chávez and Riosmena-Rodríguez (2000), and Riosmena-Rodríguez et al. (2005); and C. racemosa var. turbinata (J. Agardh) Eubank (1946) by Dawson (1944, as 

“C. racemosa var. chemnitzia”), Dawson (1949, 1959a), 

Huerta-Múzquiz and Mendoza-González (1985, as “C. 

racemosa var. chemnitzia”), Rocha-Ramírez and SiqueirosBeltrones (1991), and Mateo-Cid et al. (2000a).

Three species are present in the northern Gulf of 

California.

Key to the Species of Caulerpa in the 

1a. Axes and branches flattened throughout, with primary branches and numerous pinnate (oppositely branched) branchlets

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

1b. Axes, primary branches and branchlets cylindrical throughout  

2a. Cylindrical erect axes with oppositely branched branchlets (featherlike appearance); branchlet apices acute, with pointed 

tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sertularioide

2b. Cylindrical erect axes simple and unbranched or sparsely and irregularly branched; 500–1000 μm in diameter; apices 

rounded or more or less clavate  . . . . . .

Caulerpa mexicana f. pectinata (Kützing) W. R.

Taylor

FIGURE 45

Caulerpa pectinata Kützing, 1849:495.

Caulerpa mexicana Sonder ex Kützing f. pectinata (Kützing) W. R. Taylor, 1960:141; Apartado et al., 2002:99, 

figs. 2, 5; Pedroche et al., 2005:64.

Caulerpa pinnata C. Agardh f. pectinata (Kützing) 

Weber-van Bosse, 1898:291, pl. 24: fig. 3; Dawson, 

1944:213; Dawson, 1961b:380; Dawson, 1966a:7; 

Dawson, 1966b:55, fig. 1A; González-González et al., 

1996:127; Mateo-Cid et al., 2006:48; Pacheco-Ruíz et 

al., 2008:202.

Caulerpa crassifolia (C. Agardh) J. Agardh f. pectinata 

(Kützing) Collins, 1909b:414.

Algae erect, with flat, fronds of pinnate branches, up 

to 5 cm tall, arising from a short stalk above the spreading stolon; attached below by rhizoids issued from stolons. 

Fronds simple or forked; branches lanceolate to oblong in 

shape, 2–5 cm long and narrow, up to 5 mm wide; composed of a flattened main axes and closely set, opposite flat 

short branchlets in one plane; branchlets not contracted 

at base, not closely adjacent, and curved slightly upward 

with acuminate tips.

FIGURE 45. Caulerpa mexicana f. pectinata: Habit, narrow pinnate 

fronds; axes and branchlets are flattened (JN-5861).
9 6 • 

Habitat. On sand covered rock, tidal platforms and in tide pools; often under larger algae such

as Sargassum; mid intertidal to shallow subtidal (also 

dredged in 12–22 m depths; Dawson, 1944).

Distribution. Gulf of California: Puerto

Peñasco; Puerto Refugio, in shallows of channel between 

Isla Mejía and Isla División (northwest end of Isla Ángel 

de la Guarda).

Type Locality. La Guaira (La Guayra),

seaport town for Caracas, Federal District, northern 

Venezuela.

Remarks. Generally considered a tropical 

western Atlantic taxon (Taylor, 1960; Apartado et al., 

2002), Caulerpa mexicana f. pectinata has also been 

reported in the northern Gulf by Dawson (1944, 1966a, 

1966b, as “C. pinnata f. pectinata”). Gulf specimens need 

to be morphologically and molecularly compared with 

those of the western Atlantic, particularly with those from 

the type locality, to test their phylogenetic relationship.

Caulerpa sertularioides (S. G. Gmelin) M. Ho

FIGURE 46A,B

Fucus sertularioides S. G. Gmelin, 1768:151, pl. 15: fig. 4.

Caulerpa sertularioides (S. G. Gmelin) M. Howe, 

1905:576; Dawson, 1944:213; Taylor, 1945:63; 

Dawson, 1949:244–245; Dawson, 1959a:4, 6, 7, 14; 

Dawson, 1961b: 381; Huerta-Múzquiz and TiradoLizárraga, 1970:126; Huerta-Múzquiz and TiradoLizárraga, 1970:121–122, 127, 131, 134, 136; Chávez 

B., 1972:268; Earle, 1972:85; Huerta-Múzquiz, 

1978:338, 340; Huerta-Múzquiz and MendozaGonzález, 1985:44; Mendoza-González and MateoCid, 1986:420; Ortega et al., 1987:80; Sánchez-Vargas 

and Hendrickx, 1987:161; Salcedo-Martínez et al., 

1988:82; Sánchez-Rodriguez et al., 1989:39; RochaRamírez and Siqueiros-Beltrones, 1991:24; Mateo-Cid 

and Mendoza-González, 1991:28; Martínez-Lozano 

et al., 1991:23; Mateo-Cid and Mendoza-González, 

1992:25; León-Tejera and González-González, 

1993:497; León-Tejera et al., 1993:199, 204; GonzálezGonzález, 1993:443; Mateo-Cid et al., 1993:52; Mendoza-González et al., 1994:113; González-González et 

al., 1996:277, 366–367; Anaya-Reyna and RiosmenaRodríguez, 1996:862, 863; Galindo-Villegas et al., 

1997:4; Mendoza-González and Mateo-Cid, 1998:25, 

27–28; Yoshida, 1998:103; Cruz-Ayala et al., 1998:192; 

Rodríguez-Morales and Siqueiros-Beltrones, 1999:22; 

Paul-Chávez and Riosmena-Rodríguez, 2000:146; 

López et al., 2000:339; Mateo-Cid et al., 2000a:60, 70; 

Cruz-Ayala et al., 2001:190; Scrosati, 2001:722; Beltrán-Magallanes, 2002:44; Carballo et al., 2002:753; 

Abbott and Huisman, 2004:124, fig. 45B,C; Hodgson 

et al., 2004:32; López et al., 2004:10; Wysor, 2004:224; 

Riosmena-Rodríguez et al., 2005:101; Pedroche et al., 

2005:67; Servière-Zaragoza et al., 2007:3, 7.

Algae with erect (feather-like) fronds, in the northern 

Gulf mostly less than 7.0 cm tall, arising from a short 

cylindrical stalk above the horizontal stolon, attached 

below by rhizoids. Fronds 2–7(–11) cm long and up to 

1 cm wide, most simple and unbranched or occasionally branched; with a terete main axes bearing opposite cylindrical branchlets; branchlets not constricted at 

base, usually closely adjacent, and curved upward, with 

pointed tips.

Habitat. In sandy habitats and in crevices and

tide pools; attached to sand, on rocks or broken shells; 

low intertidal to shallow subtidal.

Distribution. Gulf of California: Bahía San

Carlos; Bahía Concepción to Punta Los Frailes; Mazatlán (Bahía de Altata to Estero del Urías), Sinaloa. Pacific 

coast: Bahía Magdalena, Baja California Sur; Islas Revillagigedo; Nayarit to Chiapas; Panama; Hawaiian Islands; 

China; Japan.

Type Locality. “. . . nonnisi [only] in coralliis americanis” (Gmelin, 1768:151); “no definite locality

is given” (Howe, 1905:576); “possibly tropical Atlantic 

America” (Hodgson et al., 2004:32).

Remarks. Collections (US Alg. Coll.) from 

Bahía San Carlos (north of Guaymas, Sonora) establish 

Caulerpa sertularioides in the northern Gulf. Two forms 

of this species have been previously recorded in the southern Gulf Caulerpa sertularioides f. brevipes (J. Agardh) 

Svedelius (1906) has been reported in Laguna de Agiabampo, southern Sonora (Ortega et al. 1987),and C. sertularioides f. longiseta (Bory de Saint-Vincent) Svedelius 

(1906) from La Paz by Huerta-Múzquiz and MendozaGonzález (1985) and Cruz-Ayala et al. (2001).

Ecological monitoring of Caulerpa sertularioides

over 2 years in Bahía Balandra (vicinity of La Paz) by 

Scrosati (2001) showed a high population abundance 

during the relatively high temperatures of El Niño conditions and, in contrast, an absence during the relatively 

low temperatures of La Niña. Scrosati (2001) suggested 

that quantitative monitoring of Gulf of California seaweeds could be useful in predicting long-term effects of El 

Niño and La Niña on changes of marine algal distribution 

and abundance.
n u m b e r 9 4 • 9 7

Caulerpa vanbosseae Setchell et N. L. Gardn

FIGURE 47A,B

Caulerpa vanbosseae Setchell et N. L. Gardner, 1924a:704, 

pl. 13: figs. 13–15; Dawson, 1944:212; Dawson, 

1949:234; Dawson, 1959a:8–10, 14; Dawson et al., 

1960b:13; Dawson, 1961b:381; Dawson, 1966a:7; 

Littler and Littler, 1984:15, 23; Sánchez-Rodríguez et 

al., 1989:38, 39; Mateo-Cid et al., 1993:52; GonzálezGonzález et al., 1996:127; Paul-Chávez and RiosmenaRodríguez, 2000:146; Cruz-Ayala et al., 2001:190; Pedroche et al., 2005:69; Mateo-Cid et al., 2006:48, 59; 

Pacheco-Ruíz et al., 2008:202.

Algae cylindrical throughout, erect portions simple 

and unbranched or irregularly branched (sometimes more 

or less alternately or oppositely branched); forming loosely 

entangled, sometimes spreading, clumps up to 6 cm tall; 

attached below by rhizoids issued from creeping stolons. 

Erect axes and branches, and horizontal stolons of more 

or less similar diameter, (400–)500–1000 μm in diameter. 

Terminal portions of axes and branches often more or less 

clavate or rounded. Reproduction not known.

FIGURE 46. Caulerpa sertularioides: A. Habit (JN-4471). B. Prostrate stolons bearing erect axes with distichously arranged branchlets; both 

axes and branchlets are cylindrical (JN-4277).
9 8 • 

Habitat. On rocks or broken shells, in crevices

and tide pools; low intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Bahía La Paz. Pacific coast: Punta Entrada 

(Bahía Magdalena) and Isla Santa Margarita (Bahía Las 

Almejas), Baja California Sur.

Type Locality. Vicinity of La Paz, Baja California Sur, Gulf of California

Remarks. Caulerpa vanbosseae is apparently 

known only in the Gulf of California and Pacific Baja 

California Sur. A few specimens of Gulf C. vanbosseae

bear a morphological resemblance to some of the cylindrical, sparsely, irregularly branched forms of deep-water C. 

ashmeadii Harvey (1858) from the eastern Gulf of Mexico 

(US Alg. Coll.).

Halimed

Halimedaceae Link, 1832:115.

Most members of this family have a multiaxial construction, and most are calcified (aragonite mineral form 

FIGURE 47. Caulerpa vanbosseae: Erect upper portions of simple to irregularly branched cylindrical axes, above horizontal, cylindrical, stolonlike rhizomes. A. Habit (JN-6562). B. Habit (JN-4981).
n u m b e r 9 4 • 9 9

of calcium carbonate). Development is initially uniaxial; 

as more siphonous filaments are formed, they become 

consolidated into a multiaxial thallus, with a filamentous 

medulla and a cortex of utricles. Cells have numerous 

chloroplasts and leucoplasts, with xylan as a principal cell 

wall constituent.

Sexual reproduction is holocarpic. Biflagellate anisogametes are formed in differentiated gametangia that 

develop above the surface layer of the segments. Anisogametes are discharged through papillae.

A monotypic family; the single genus Halimeda is represented in the northern Gulf of California.

Halimeda J. V. Lamou

Halimeda J. V. Lamouroux, 1812:186, as “Halimedea.”

Algae are erect or spreading and variously branched 

and are composed of articulated sequences of calcified 

segments that alternate with narrow, flexible noncalcified 

joints. Attachment to the substratum is by either a large, 

dense fibrous holdfast; a small holdfast of rhizoidal filaments; or by several rhizoidal masses developed from the 

segments or nodes. Calcified (aragonite) segments may 

be cylindrical, compressed or flat, or spherical, simple or 

lobed. Segments are internally composed of a multiaxial 

(rarely uniaxial), filamentous medulla that branches outward to form 2–6 layers of subsurface utricles and a surface layer of utricles. Outer surface utricles may be loosely 

to closely adherent. Growth is indeterminate. The flexible 

joints are composed of coenocytic medullary filaments 

that are closely parallel and remain separate or are fused 

in pairs, in small groups, or into a single group. Filaments 

contain numerous nuclei, chloroplasts and amyloplasts, 

and have cell walls primarily composed of xylan. Accessory pigments are siphonoxanthin and siphonein.

Asexual reproduction is by fragmentation. New thalli 

can also develop at ends of filaments that grow out of segments, as well as from filaments of basal stolons. Sexual 

reproduction, where known, is holocarpic. Biflagellate 

anisogametes are formed in gametangia on branched stalks 

grouped together on the surface of the calcified segments.

Remarks. Halimeda is widely distributed in 

tropical to subtropical seas, from intertidal to depths of at 

least 150 m (Littler et al. 1985, 1986). Halimeda has long 

been known to play a critical role in the formation of tropical beach sands (van Overbeek and Crist, 1947). Growth 

patterns, calcification, and defense chemicals are synchronized on a daily cycle to maximize growth by avoiding 

some daytime herbivores (Hay, 1988, 1997b). Holocarpic 

sexual reproduction of Halimeda also follows a diel pattern (Clifton, 1997; Hay, 1997a; Clifton and Clifton, 

1999). Initiated after sunset, it is a brief phenomenon, with 

almost all of the alga’s protoplasma put into gamete production. Overnight, the green calcified segments become 

white, with stalked green gametangia on their surface and 

edges, and right before dawn, the gametes are released, 

leaving behind the dead calcified structure.

Currently there is one species, H. discoidea, known in 

the northern Gulf of California.

One additional species, Halimeda opuntia (Linnaeus) 

J. V. Lamouroux (1816), is recorded in the southern Gulf 

(Rocha-Ramírez and Siqueiros-Beltrones, 1991).

Halimeda discoidea Decaisne

FIGURE 48A–C

Halimeda discoidea Decaisne, 1842b:102; Collins, 

1909b:400; Howe, 1911:492; Setchell and Gardner, 1920b:177, pl. 13: fig. 3; Setchell and Gardner, 

1924a:704; Dawson, 1944:214; Taylor, 1945:73; Dawson, 1949:244, 245; Taylor, 1950:85, pl. 45: fig. 1; 

Dawson, 1957b:9; Dawson, 1959a:4, 6, 8, 18; Hillis, 

1959:352, pl. 2: fig. 5, pl. 5: fig. 11, pl. 6: fig. 11, pl. 7: 

figs. 9, 10, pl. 8: figs. 5–8, pl. 11; Dawson, 1960c:97; 

Dawson et al., 1960b:13, 14; Dawson, 1961b:382; 

Dawson, 1961c:406, pl. 7: fig. 1; Dawson, 1962b:177, 

229, fig. 22; Huerta-Múzquiz, 1978:338; HillisColinvaux, 1980:136, fig. 41; Pedroche and GonzálezGonzález, 1981:64; González-González, 1993:443; 

Huerta-Múzquiz and Mendoza-González, 1985:44; 

Salcedo-Martínez et al., 1988:82; Sánchez-Rodríguez 

et al., 1989:39; Mateo-Cid and Mendoza-González, 

1991:25; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:24; Martínez-Lozano et al., 1991:22; Tsuda and 

Kamura, 1991:69, pl. 4: fig. 1; Mateo-Cid and Mendoza-González, 1992:25; León-Tejera et al., 1993:199, 

202; Servière-Zaragoza et al., 1993:482; Mateo-Cid 

and Mendoza-González, 1994b:45; León-Tejera and 

González-González, 1994:26; González-González et al., 

1996:287; Bucio-Pacheco and Dreckmann, 1998:43, 

45; Yoshida, 1998:114; Hillis et al., 1998:671; 

Mendoza-González and Mateo-Cid, 1998:26; ServièreZaragoza et al., 1998:169, 179; Rodríguez-Morales and 

Siqueiros-Beltrones, 1999:22; López et al., 2000:339; 

Paul-Chávez and Riosmena-Rodríguez, 2000:146; Abbott and Huisman, 2004:131, fig. 49B–E; López et 

al., 2004:10; Wysor, 2004:225; Hernández-Herrera 

et al., 2005:146; Pedroche et al., 2005:80; Riosmena-
1 0 0 • 

FIGURE 48. Halimeda discoidea: A. Habit (after Setchell and Gardner, 1920b: pl. 13, fig. 3). B. Surface view of decalcified segment with hexagonal pattern formed by abutting surface utricles; occasionally, two adjacent utricles merge across one of their facets, consequently forming 

polygons (EYD-25921). C. Section of segment showing large secondary utricle bearing smaller cup-shaped peripheral utricles (EYD-25921).
n u m b e r 9 4 • 1 0 1

Rodríguez et al., 2005:101; Servière-Zaragoza et al., 

2007:7; Pacheco-Ruíz et al., 2008:202.

Algae forming small clumps of subdichotomously 

to trichotomously branched fronds (branching in several 

planes), up to 10 cm high; of lightly to moderately calcified flat segments that alternate with very short noncalcified joints of entangled medullary filaments fused in pairs 

(rarely 3); attached below by a single, relatively small, mass 

of rhizoidal filaments. Flat segments variable in shape, 

mostly obovate to reniform, and 1.0–2.5 cm wide; upper 

segments usually flexible, leathery. Surface utricles forming 

hexagonal pattern in surface view; cup-shaped in transection, 25–45 μm in diameter; laterally adherent to adjacent 

surface utricles (even after decalcification) along their distal margins for 15–20 μm; borne on notably larger, inflated 

secondary internal utricles, 60–175 μm in diameter.

Habitat. On rocks, and in crevices and tide

pools; low intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Refugio, Isla Ángel de la Guarda; Isla Partida; Guaymas 

to San Jose del Cabo. Pacific coast: Isla Socorro and Isla 

Clarion (Islas Revillagigedo); Isla Magdalena (southwest 

of Punta Entrada, Bahía Magdalena, Baja California) to 

Costa Rica; Panama; Hawaiian Islands; Bikini Island; 

China; Japan.

Type Locality. “Not certain” (Silva et al.,

1996a). Although the herbarium label (PC) indicates it 

was collected “Kamtschatka” (Russia) during the voyage of the exploring ship Venus (Decaisne, 1842b:102), 

Hillis (1959:352; see also Hillis-Colvinaux, 1980) noted 

that “at least the locality [is] incorrect,” as waters of the 

Kamchatka Peninsula are far too cold for this tropical to 

subtropical species.

Remarks. In the Gulf of California, Halimeda 

discoidea is usually found in the warmer waters of the 

southern Gulf, but it is also known from a few collections 

in the northern Gulf.

Dasycladoph

Dasycladophyceae C. van den Hoek, D. G. Mann et Jahns, 

1995:436, nom. invalid. [see McNeill et al., 2006:Art. 

36.2; Pröschold and Leliaert, 2007:130].

Algae of this class are exclusively marine, siphonous, 

and composed of a primary main axis or stalk that bears 

whorls of simple or branched laterals that are either free 

or joined distally to form a cortex. Each thallus is a single 

coenocytic cell with a parietal layer of cytoplasm and a 

large central vacuole. Many are slightly to moderately 

calcified. Siphons contain numerous nuclei and fusiform 

to ellipsoid chloroplasts that do not have pyrenoids but, 

instead, often have storage grains of fructan and starch. 

These grains may also be found in the cytoplasm. Cytoplasmic streaming of the organelles, including chloroplasts, 

grains, and nuclei, occurs along numerous “lanes” (striae) 

that run parallel to the longitudinal axis of the cell. The cell 

wall of the vegetative thallus is composed of mannan, and 

the wall of the gametangial cyst is predominantly cellulose.

Life histories, where known, involve a haploid gametophytic stage, the macrothallus, that alternates with a 

zygote stage containing a giant diploid nucleus (the only 

diploid nucleus in the life history). Sexual reproduction is 

by fusion of biflagellated gametes. Gametes may be formed 

in gametangial cysts of lateral branches or in disc rays.

There is one order represented in the northern Gulf of 

California.

Dasy

Dasycladales Pascher, 1931:328.

Dasycladales Bessey, 1907: 287.

Structurally composed of coenocytic cells, these algae 

have radial symmetry and are often calcified and attached 

below by rhizoids. The siphonous main axis or stalk bears 

whorls of simple or divided laterals that can be free or 

united and are sometimes branched to several orders.

Reproduction is by operculate cysts that function as 

gametangia and are developed from a nucleus that has 

divided into numerous small secondary nuclei and presumably undergoes meiosis to produce gametes. These cysts 

release biflagellate isogametes, which upon fusion, form a 

quadriflagellate zygote. This zygote settles and develops into 

a siphonous system from which the primary axis develops.

Remarks. The extensive fossil record of the 

Dasycladales is unique among the Chlorophyta. Most of 

the 150 genera of Dasycladales are fossils, extending back 

to the Ordovician (e.g., Pia, 1927; Nitecki, 1970). Currently, there are only 11 known living genera, of which 

three extend from the Cretaceous.

In the southern Gulf, one of its two families, the 

Dasycladaceae Kützing (1843), is represented by two 

genera, Batophora J. Agardh (1854) and Neomeris

Lamouroux (1816), with records of Batophora oerstedii

J. Agardh (1854; Ortega et al., 1987) and Neomeris annulata Dickie (1874a; Dawson, 1959a; Huerta-Múzquiz, 

1978; Mateo-Cid et al., 2000a:70; Hinojosa-Arango and 
1 0 2 • 

Riosmena-Rodríguez, 2001; Riosmena-Rodríguez et al., 

2005).

The Polyphysaceae of the Dasycladales is represented 

throughout the Gulf of California.

Poly

Polyphysaceae Kützing, 1843:302, 311.

Acetabulariaceae Nägeli, 1847:158, 252.

Algae a unicell, with a stalk bearing two kinds of 

whorled laterals that can be arranged in two different ways: 

(1) numerous whorls of hyaline branched laterals (may be 

pale green when immature) and a subterminal or terminal 

disc or cap of a whorl of gametangial rays (which may be 

lost after reproductive maturity and later replaced) or (2) 

alternating whorls of hyaline (or pale green) laterals and 

green discs or caps of gametangial rays. Gametangial rays 

with a circle of short protuberances (superior corona) on the 

upper side that bear hairs and/or scars after hairs are shed; 

rays may or may not have a ring of additional segments 

beneath (inferior corona), which if present, lacks hairs.

Reproduction is by gametes that develop in cysts 

within the gametangial rays of the cap or disc.

Remarks. The Polyphysaceae was shown to be 

a monophyletic group distinct from the family Dasycladaceae (Olsen et al., 1994).

One of the genera, Parvocaulis S. Berger, Fettweiss, 

Gleissberg, Liddle, U. Richter, Sawitsky et Zuccarello 

(2003), has been reported in the southern Gulf with records 

of two species Parvocaulis pusilla (M. Howe) S. Berger et 

al. (2003) by Huerta-Múzquiz and Mendoza-González 

(1985), Rocha-Ramírez and Siqueiros-Beltrones (1991), 

and Baynes (1999) (all as “Acetabularia pusilla (M. Howe) 

Collins, 1909b”) and Parvocaulis parvulus (Solms-Laubach) S. Berger et al. (2003) by Mateo-Cid et al. (2000a, as 

“Acetabularia parvula Solms-Laubach, 1895”).

One genus, Acetabularia, is known in the northern 

Gulf of California.

Acetabularia J. V. Lamou

Acetabularia J. V. Lamouroux, 1812:185.

Algae are light to moderately calcified and are composed of an erect, siphonous stalk (up to 1 mm in diameter) above a lobed, rhizoidal base attached to solid 

substratum. The cylindrical stalk bears successive whorls 

of deciduous, branched sterile laterals (hairs) in its upper 

portions that can leave scars, and at its upper end it bears, 

terminally or subterminally, one or more caps or discs of 

laterally joined rays. Near the base of each ray are borne 

small projections on the upper and lower surfaces: the 

projections form a whorl, the superior (apical) corona 

with colorless hairs and the inferior corona without hairs. 

Cytoplasm is peripheral and contains numerous discoid 

plastids and lacks pyrenoids and surrounds a large central 

vacuole. Chloroplast ultrastructure is variable and with or 

without stacked grana. Starch grains are in both the chloroplast and cytoplasm, but they are not membrane-bound 

in the latter. Vegetative cells have a single, diploid, primary nucleus in a branch of the rhizoid. Primary nuclear 

division is meiotic, followed by a series of mitotic divisions 

that form secondary nuclei that migrate up the stalk and 

establish themselves in the rays of the cap.

Asexual reproduction is by fragmentation or growth 

of new thalli from basal rhizoids or regeneration after 

wounding from grazing herbivores or physical force, 

such as wave action. Gametangial rays develop thickwalled, operculate cysts by uninucleate aggregations of 

cytoplasm, and subsequent division of the nuclei produces biflagellate isogametes. The thick-walled cysts are 

released from the rays and can survive for some time on 

their own. When conditions are suitable, an ostiole opens 

in the cyst, and the gametes swim out. Two gametes fuse 

to form the diploid zygote that settles and develops into 

a new thallus.

Remarks. Species of Acetabularia are quite 

distinctive with a narrow stalk that bears a beautiful, delicate, parasol-like, discoid, or cup-shaped cap. They are 

found in tropical to subtropical seas, often in brackish to 

hypersaline shallow waters. On the basis of the pioneering 

studies of Woronine (1862) and De Bary and Strasburger 

(1877) the life history of Acetabularia has long been 

reported to be three years before the reproductive caps 

with cysts were formed, but more recently, Berger and 

Liddle (2003) showed the life history can be completed in 

one growing season.

Three western Atlantic species have also been reported 

in the southern Gulf of California: Acetabularia farlowii

Solms-Laubach (1895) and A. crenulata J. V. Lamouroux 

(1816) were reported by Ortega et al. (1987) and RochaRamírez and Siqueiros-Beltrones (1991). More recently, 

L. Aguilar-Rosas et al. (2006) added another species, Acetabularia schenckii K. Möbius (1889), to Bahía de La Paz 

and Sinaloa and also added new distribution records of A. 

farlowii to Sinaloa and Pacific Baja California.

One species is recorded in the upper Gulf of California.
n u m b e r 9 4 • 1 0 3

Acetabularia caliculus J. V. Lamouroux in 

et Gaimard

FIGURE 49A,B

Acetabularia caliculus J. V. Lamouroux in Quoy et Gaimard, 

1824:621, pl. 90: figs. 6, 7; Yoshida, 1998:154; Berger 

et al., 2003:527, figs. 8, 31. As Acetabularia “calyculus”: Dawson, 1954d:396, fig. 13h; Dawson, 1966a:8; 

Dawson, 1966b:55; Valet, 1969:617, pl. 44: figs. 7–9; 

Huerta-Múzquiz and Tirado-Lizárraga, 1970:127, 132, 

136; Womersley, 1971:119, fig. 15; Huerta-Múzquiz, 

1978:340; Womersley, 1984:295, figs. 101B, 102B–D; 

Huerta-Múzquiz and Mendoza-González, 1985:42; 

González-González et al., 1996:126; Anaya-Reyna and 

Riosmena-Rodríguez, 1996:862; Núñez-López et al., 

1998:39; Rodríguez-Morales and Siqueiros-Beltrones, 

1999:22; Paul-Chávez and Riosmena-Rodríguez, 

FIGURE 49. Acetabularia caliculus: A. Cluster of several mature thalli, with calcified stalk and cup-shaped disc of laterally adhering gametangial 

rays; some discs with central tuft of deciduous hairs (issued from the corona superior) still intact (liquid preserved; EYD-27569). B. Numerous 

spherical reproductive cysts enclosed in gametangial rays of mature disc; note smooth, truncate outer margin of each ray and faint seam of 

calcium carbonate along ray lateral margins (US Alg. Coll. slide 5533).
1 0 4 • 

2000:146; Cruz-Ayala et al., 2001:190; RiosmenaRodríguez et al., 2005:101; Pedroche et al., 2005:84; 

Mateo-Cid et al., 2006:48; Dreckmann et al., 2006:154.

Alga erect, usually growing in clusters, up to 3 cm tall; 

a rigid, thin cylindrical calcified stalk with several whorls 

of ephemeral branched laterals (green when young, later 

colorless) in upper portion and a subterminal, cup-shaped 

disc, 3–5(–7) mm in diameter; attached below by a rhizoidal 

base. Disc solitary of (20–)22–30 laterally adjoined rays; 

outer margin of rays truncate in most discs or knobbed in 

some discs. Corona superior with short blunt lobes that 

each bear 2–3(–4) branched colorless hairs or hair scars; 

inferior corona inconspicuous, each segment a single blunt 

lobe beneath the ray at the place of attachment.

When reproductive, each gametangial ray filled with 

45–75 spherical cysts containing gametes; each cyst 100–

150(–175) μm in diameter (US Alg. Coll.-95320, slide 

5533).

Habitat. Growing on shells in shell/sand and

sand/mud bottoms of rapid flowing tidal channels; mid to 

low intertidal.

Distribution. Gulf of California: Bahía

Cholla (Choya), vicinity of Puerto Peñasco; Bahía La Paz. 

Pacific coast: Baja California Sur; Oaxaca to Chiapas; 

China; Japan.

Type Locality. Baie des Chiens-Marins

(Shark Bay), Western Australia.

Remarks. Acetabularia caliculus is apparently 

known only from shallow bays in the Gulf of California. 

The outer margins of the disc rays in the northern Gulf 

specimens of A. caliculus (Figure 49B) are both truncate 

(in agreement with those reported by Berger and Kaever, 

1992) and sometimes knobbed (as observed in A. caliculus

of Lord Howe Island, Australia; Kraft, 2008: fig. 104C,D). 

Morphological and molecular studies, such as those of 

Olsen et al. (1994) and Berger et al. (2003), are needed to 

test the taxonomy and phylogenetics of Gulf A. caliculus in 

relation to those from the Western Australian type locality.

The original spelling of the species name, caliculus 

(Lamouroux in Quoy and Gaimard, 1824), agrees with 

the Latin usage (calix (L.), a small cup [Brown, 1956]); 

thus, it is not necessary to change the original spelling to 

Greek usage, i.e., calyculus (calyx, kalyx [Gr.]).
STRAME

Stramenopiles D. J. Patterson, 1989:372.

Stramenopiles are a very large, extremely diverse, group of eukaryotes that 

range from the microscopic to the largest of all marine plants, and include autotrophs, heterotrophs, and mixotrophs. Most are planktonic or benthic and are 

found in terrestrial, freshwater, and marine environments, although a few are 

aerial and some are endosymbionts. The discovery that heterokont algae are 

related to some nonheterokont algae and to various nonphotosynthetic organisms (including some previously thought to be fungi or protozoa, such as protozoan-like unicells or loosely associated colonies, hypochytrids, water molds, 

and downy mildews) has led to their being grouped together. The ancestor of 

the basal Stramenopiles may have been an alga, and then some groups could 

have arisen through plastid loss or, possibly, branched off before the appearance of plastids within the group. Plastid reduction as well as reacquisition has 

also occurred in some lineages. Stramenopiles are grouped together on the basis 

of their cytology and ultrastructure, supported by biochemical and molecular 

studies (e.g., Moestrup and Andersen, 1991; Williams, 1991a, 1991b; Leipe et 

al., 1994; Reith, 1995; Medlin et al., 1995, 1997; Saunders et al., 1997; Daugbjerg and Andersen, 1997; Delwiche, 1999; Patterson, 1989, 1999; Goertzen and 

Theriot, 2003; Andersen, 2004a, 2004b; Adl et al., 2005).

Members include both unicellular motile flagellates and multicellular organisms; typically, all members have cells or produce cells with heterokont flagella 

(two dissimilar flagella) at some point in their life history, e.g., zoospores or 

gametes. One of the flagella is an anterior tinsel flagellum with mastigonemes 

(unique three-parted, tubular hair-like structures) in two opposite rows. This 

feature, together with the flagella typically being supported by four microtubule 

roots in a distinctive pattern, and mitochondria with tubular cristae are defining characteristics of the Stramenopiles. Most members either have this tinsel 

flagellum or were derived from organisms that had this type of flagellum. The 

other flagellum is reverse-directed, smooth, and usually shorter or is reduced to 

a basal body in some. A few Stramenopiles do not produce cells with the typical 

Part I

BROWN ALGAE
1 0 6 • 

heterokont flagella, and the flagella have been altogether 

lost in a few lines, most notably the diatoms.

Photosynthetic Stramenopiles include microscopic 

biflagellated or nonflagellated unicellular, colonial, and 

loosely filamentous algae (e.g., chrysophytes, diatoms) 

and multicellular algae that range from siphonous filaments (e.g., Vaucheria) to the structurally complex kelps 

(e.g., Macrocystis, Laminaria). Some have siliceous scales 

and spines or silicate skeletons, and their species may also 

be found in the fossil record; others are without cell walls 

or scales, and a few may be amoeboid. Cells of stramenopile algae have chloroplasts containing chlorophyll a and 

chlorophylls c1 and c2 (some may also have c3) and, usually, the accessory pigments fucoxanthin, heteroxanthin, 

vaucheriaxanthin, or violaxanthin. Plastids have three 

thylakoids per lamellae, lack nucleomorphs, and are surrounded by four membranes; the outermost is continuous 

with the endoplasmic reticulum. Reserve polysaccharides 

include chrysolaminarin or laminarin, formed in vesicles 

outside of the plastid.

Remarks. Classification systems have not 

always grouped the currently accepted members of the 

Stramenopiles together (for a review, see Hausmann et al., 

2003), and they have been variously placed in the Chromista Cavalier-Smith (1981), Heterokonta Cavalier-Smith 

(1986), Heterokontophyta Ø. Moestrup (1992), and 

Protista or Protoctista (e.g., Whittaker, 1969; Margulis 

et al., 1990; Corliss, 1994; Cavalier-Smith, 1986, 1998; 

Margulis and Schwartz, 1998). Recently, Adl et al. (2005) 

recognized the Stramenopiles (Patterson, 1989, 1999) to 

comprise 20 groups within the Chromalveolata Adl et al. 

(2005), one of seven clusters of eukaryotes that represent 

the basic groups (or traditional kingdoms) in their nameless ranked higher-level classification.

One very large multicellular group of Stramenopiles, the Phaeophyceae, is well represented in the Gulf of 

California.

Phaeophyc

Phaeophyceae Kjellman, 1891:176.

Melanophyceae Rabenhorst, 1863:274.

Fucophyceae Warming, 1884:38, as “Fucoideae.”

Phaeophyceae Hansgirg, 1886:17, 27.

The Phaeophyceae (brown algae) are multicellular 

and exceptionally variable in form and size, ranging from 

microscopic to macroscopic, and include filamentous, 

crustose, tubular, globose, and foliose forms and very 

large, structurally complex species. The 266 genera comprising the brown algae are almost all marine; only four to 

five genera occur in freshwater (Wehr and Sheath, 2003). 

The Phaeophyceae vary in color from light yellow-brown 

to dark blackish-brown, with a few that are chalky or 

whitish due to their surface calcification (e.g., some species of Padina and Newhousia [Kraft et al., 2004]). The 

class contains the largest of all the algae, with some species of the “giant kelp,” Macrocystis C. Agardh (1820), up 

to 70(–100) m long (Foster and Schiel, 1985; Womersley, 

1987). Most of the marine brown algae are benthic, often 

dominating intertidal and subtidal communities, and are 

found in depths down to 88 m (Littler et al., 1985, 1986); 

a few apparently live as free-floating forms, e.g., species of 

Sargassum in the Sargasso Sea.

Growth may be apical, intercalary, trichothallic, or 

diffuse. Phaeophycean hairs are often present. Cells are 

uninucleate, with the photosynthetic pigments in plastids 

containing thylakoids in groups of three and with or without a pyrenoid. Plastids are singular or present in various numbers and may be parietal, ribbonlike, or discoid; 

physodes (vesicles produced within the plastids) are usually present. The photosynthetic pigments, chlorophylls 

a, c1, and c2, are masked by the accessory pigments ß

carotene and the xanthophylls (particularly, fucoxanthin, 

violaxanthin, and diathoxanthin), thus giving the characteristic brown color to the group. Photosynthetic reserves 

include the soluble carbohydrates laminarin (a beta-linked 

polymer of glucose) and mannitol. Cell walls are firm and 

composed of two layers, the inner of cellulose microfibrils 

and a gelatinous outer layer that contains alginic acid and 

fucinic acid.

Life histories are either isomorphic, heteromorphic 

(usually with a larger sporophyte), or diplontic (meiosis to 

form gametes, e.g., Fucus). Sporophytes develop unilocular 

or plurilocular sporangia; the latter produce more than one 

spore. Sexual reproduction is isogamous, anisogamous, or 

oogamous, with parthenogenesis known in some species. 

Gametangia are typically plurilocular, with the exception being oogamous females. Motile cells (i.e., zoospores 

and gametes) have two laterally inserted, unequal-length 

flagella: the larger is anterior tinsel type, and the smaller 

is posterior whiplash type. However, in the Fucales, the 

flagella are more or less of equal length, and at least one 

group, the Dictyotales, produce nonmotile aplanospores 

and uniflagellate sperm that lack a posterior flagellum.

Remarks. In the northern Gulf of California, 

the Phaeophyceae have about the same numbers of species 

as the green algae, but both have fewer species than the red 

algae. Ecologically, the brown algae generally comprise 
n u m b e r 9 4 • 1 0 7

the bulk of the northern Gulf’s algal biomass, and seasonally, there is greater abundance and more species of brown 

algae during the winter and spring (see also McCourt, 

1984a, 1984b; Huerta-Múzquiz and Mendoza-González, 

1985; Pacheco-Ruíz and Zertuche-González, 1996b). 

Three northern Gulf regions that have high brown algal 

species richness are Puerto Peñasco (Norris, 1975), Bahía 

de Los Ángeles (Pacheco-Ruíz et al., 2008), and Bahía de 

La Paz (L. Aguilar-Rosas and R. Aguilar-Rosas, 1993).

In recent years, the presumed relationships among 

the brown algae have been somewhat uncertain (e.g., 

Saunders et al., 1992). The traditional concept, based on 

morphology and life histories, that the Ectocarpales are 

basal to the other brown algal orders (e.g., Kylin, 1933; 

Papenfuss, 1951, 1955; van den Hoek et al., 1995) has 

not been supported by molecular analyses, and new systematic relationships have been proposed (e.g., de Reviers and Rousseau, 1999; Rousseau et al., 2001; Draisma 

et al., 2001, 2002, 2003). The Phaeophyceae, as presented herein, has several subdivisions to reflect the most 

recent contributions by Cho et al. (2006) and Phillips et 

al. (2008), which elucidate the evolutionary relationships 

among the brown algae. These subdivisions are grouped 

as orders, primarily on the basis of life histories and gene 

sequence data (see synopsis of ordinal characters given by 

Adl et al., 2005).

Members of nine of the orders of Phaeophyceae are 

represented in the northern Gulf of California.

Key to the Orders of the Phaeo

in the northern Gulf

1a. Algae crustose (in at least one of its life history phases); plastids without pyrenoids. 

1b. Algae not crustose; erect, filamentous, or cylindrical, compressed or flattened; plastids with or without pyrenoids  

2a. Crusts discoid to spreading or initial crustose stage that develops into low-growing clumps of mostly horizontal, short 

branches; filamentous structure of 2–3 distinct layers; growth by terminal cells of horizontal and erect filaments or intercalary from basal meristems of cortical filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ralfsiales

2b. Crusts prostrate, encrusting, multilobed; thin; parenchymatous structure; growth by marginal row of apical cells (sporophyte)  . . . . . . . . . .

3a. Algae usually small tufts of uniseriate or multiseriate, unbranched to branched free filaments . . . . . . . . . . . . . . . . . . . . 4

3b. Algae not tufted; usually moderate to large size and various forms: cylindrical, compressed or flattened; simple to branched; 

or saccate, subglobose, or foliose; or frondose; some parenchymatous, others of loosely to densely compacted filaments 

.

4a. Uniseriate filaments, simple to branched; growth apical or intercalary, localized or diffuse; ribbon-shaped or discoid plastids with pyrenoid  . . . . . . . . . . 

4b. Multiseriate branched filaments; growth by single conspicuous apical cell, initially divides transversely, followed by transverse division of subapical cells, then longitudinal divisions, and, in some, secondary transverse divisions; plastids discoid 

and no pyrenoids. . . . . . . 

5a. Growth trichothallic (apical meristem surmounted by filament or tuft of filaments). 

5b. Growth not trichothallic; either intercalary, or by a single apical cell, or group or marginal row of apical cells  

6a. Algae erect, flat, fan-shaped; growing margin fringed with trichothallic filaments (gametophytes) . . . . Cutleriales, in part

6b. Algae not fan-shaped; axes and branches either compressed to flat, narrow to broad, apical and lateral meristems with 

single trichothallic filament or terete (wiry) with apical tufts of trichothallic filaments and determinate branchlets with 

terminal tufts of filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7a. Algae wide, flat blades, simple to sparsely branched blades, or narrow, compressed axes and branches (>1.5 mm wide); 

growth from apical and lateral meristems with single terminal trichothallic filament  . . . . . . .

7b. Algae thin, wiry, terete to subcylindrical axes and branches (<1 mm wide) with terminal tufts of trichothallic filaments and 

distinctive determinate branchlets with sporangial sori surrounding portion of branchlet below terminal tuft of hairs 

. . . . . . 

8a. Growth intercalary; ribbon-shaped plastids with pyrenoid; internally centrally solid or hollow; medulla filamentous, surrounding a core of uniaxial or multiaxial filaments or of large parenchymatous cells  . . . . . . . . . . 

8b. Growth by conspicuous or inconspicuous apical cell or a group or row of apical cells; discoid plastids without pyrenoids; 

internally solid either, parenchymatous with medulla, cortex, and surface meristoderm, or medulla of filament-like hyphael 

cells and cortical assimilatory filaments, or large squarish to rectangular medullary cells and small cuboidal cortical cells 

.
1 0 8 • 

Ishige

Ishigeales G. Y. Cho et S. M. Boo in G. Y. Cho et al., 

2004:934.

Algae are terete, compressed, or foliose branched 

fronds that arise from a small holdfast or expanded crustose base. Growth is from a small inconspicuous meristematic apical cell. Anatomically, there is a distinct medulla 

of hyphael cells that connect to adjacent hyphael cells or 

to the inner cortical cells. The cortex is pseudoparenchymatous, composed of cells of assimilatory filaments. Cortical cells have several discoid plastids but lack pyrenoids 

(note that although rare, a “rudimentary pyrenoid” has 

been reported in I. okamurae [Cho et al., 2004:921, fig. 

5d]). Phaeophycean colorless hairs, originating from the 

outer medullary cells, are clustered within pits (cryptostomata) on the thallus surface and project above it.

Members have an isomorphic life history. Terminal 

unilocular sporangia are developed by transformation of 

cells of the outermost cortical layer and produce spores. 

Uniseriate plurilocular sporangia (gametangia) that lack 

sterile terminal cells originate from the assimilatory filaments and produce isogametes.

Remarks. Species of Ishige may grow attached 

to hard substrate or be epiphytic on other algae. The life 

history is presumed to be isomorphic (Tanaka, 1993; 

Hori, 1993; Cho et al., 2004; Kawai et al., 2005). In culture studies of I. okamurae from Japan, Ajisaka (1989) 

reported biflagellated spores, produced from plurilocular structures on the macrothalli, that developed into 

discs. The discs grew profusely branched filaments that 

later developed into the erect, dichotomously branched 

macrothalli. In contrast, Arasaki (1943, as “I. foliacea”) 

reported a heteromorphic life history in the culture study 

of Japanese I. sinicola; however, the culture may have been 

contaminated, making the results doubtful (see Kawai et 

al., 2005).

Currently, the Ishigeales contains one family. Recent 

molecular phylogenetic studies of Lim et al. (2007) showed 

another genus, Diplura Hollenberg (1969; Tanaka and 

Chihara, 1981b), was also close to the Ishigeales and 

suggested that the genus be removed from the Ralfsiaceae (Ralfsiales) and placed in a new family within the 

Ishigeales.

The single family is represented in the northern Gulf 

of California.

Ishige

Ishigeaceae Okamura in Segawa, 1935:65.

Ishigeaceae is a monotypic family that has the characteristics of its only genus, Ishige.

Remarks. Although the Ishigeaceae has been 

usually placed in the Chordariales (e.g., Yoshida, 1998), 

the family was shown to have a distant phylogenetic relationship with the Chordariales (or Ectocarpales sensu 

lato) and to have diverged early in the phaeophycean lineage (Kawai et al., 2005), basal to the other brown algal 

orders (Cho et al., 2004, 2006; Phillips et al., 2008). On 

the basis of its unique morphological, molecular, and 

ultrastructural features, Cho et al. (2004) established an 

order for the family.

The family is represented in the northern Gulf of California by one species of Ishige.

Ishige Yend

Ishige Yendo, 1907:154.

Algae are erect, filiform, or compressed to complanate, dichotomously to subdichotomously branched 

fronds above a short stipe that is attached below by a discoid holdfast. Small hair pits (cryptostomata) are present 

on thallus surfaces, with phaeophycean hairs developing 

from the cells of the medulla, or lowermost cortical cells . 

Internally, the medulla is composed of intertwined, usually 

dense, colorless, hyphael cells (appearing filament-like), a 

subcortical layer of pseudoparenchymatous cells, and an 

9a. Algae (Gulf members) of terete axes and branches, leaflike blades, and buoyant vesicles; internally with medulla, cortex, 

and surface meristoderm; apical cell sunken in terminal pit; gametangia in pitlike conceptacles within receptacles (reproductive branchlets); oogamous reproduction (sporangia lacking) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fucales

9b. Algae not differentiated as above; thallus flat, strap-shaped to bladelike; reproductive structures, including sporangia, on 

thallus surface (not in pitlike conceptacles)  .

10a. Algae with a single apical cell or group or marginal row of apical cells; medulla 1 or more layers of large cells, cortex of 1 

or more layers of small cells; oogamous reproduction  . . . . . 

10b. Algae with an inconspicuous apical cell; medulla of hyphael cells (filament-like), cortex of anticlinal rows of cuboidal cells; 

isogamous reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ishigeales
n u m b e r 9 4 • 1 0 9

outer cortex of assimilatory filaments (rows of anticlinally 

arranged small pigmented cells). Pyrenoids are absent in I. 

sinicola, and in I. okamurae they are “rudimentary” (Cho 

et al., 2004) and very rarely observed (Hori, 1971).

Unilocular sporangia and uniseriate plurilocular sporangia are either on the same thallus or on separate thalli. 

Both structures are found in the upper portions of branch 

segments. The unilocular sporangia are terminal, developing 

from the outermost cells of assimilatory filaments. Plurilocular sporangia originate from the assimilatory filaments, lack 

any sterile terminal cells, and are in uniseriate rows within 

sori (reported from Japanese I. foliacea [Lee et al., 2003])

Remarks. Although the external habit of Ishige

is somewhat similar to some members of the Dictyotaceae, 

they are easily separated by their internal vegetative anatomy. Species of Ishige have a medulla of entangled hyphael 

cells (appearing filamentous) and a cortex of assimilatory 

filaments, whereas the medulla of members of the Dictyotaceae is composed of one or more layers of large cells and 

a cortical layer of one to several small cells.

Interestingly, of the three known species, the northern Gulf I. sinicola is found on hard substratum, whereas 

I. foliacea of eastern Asia (Japan, China, Taiwan and 

Korea), may be either epilithic or epiphytic on I. okamurae (Cho et al., 2004). 

There is one species, apparently endemic, described 

from the northern Gulf of California.

Ishige sinicola (Setchell et N. L. Gardne

Chihara

FIGURE 50A,B

Polyopes sinicola Setchell et N. L. Gardner, 1924a:784, 

pl. 28: fig. 61, pl. 42b; Dawson, 1954c:267.

Ishige sinicola (Setchell et N. L. Gardner) Chihara, 

1969:3, fig. 2; Hommersand, 1972:70; Littler and Littler, 1981:151, tbl. 3, figs. 4–6; González-González et 

al., 1996:153; Pacheco-Ruíz and Zertuche-González, 

1996b:171; L. Aguilar-Rosas et al., 2000:132; MateoCid et al., 2006:49, 58; Pacheco-Ruíz et al., 2008:203; 

Pedroche et al. 2008:59; K. M. Lee et al. 2009: [in 

press], fig. 3 A–E.

Carpopeltis sinicola (Setchell et N. L. Gardner) Kylin, 

1956:221.

Ishige foliacea sensu Dawson, 1944:234, pl. 53: fig. 2; 

Dawson, 1954c:267; Dawson, 1960c:98; Dawson, 

1961b:394; Dawson, 1966a:10; Norris, 1973:6 [non 

Ishige foliacea Okamura in Segawa, 1935:65; Okamura, 1936:239, figs. 130: 1, 2, 5, fig. 131; which is

Ishige okamurae sensu Yendo, 1907 (in part; only the 

foliose type):155, pl. II: figs. 1–3, 5, 6, 8; and non Ishige 

okamurae Yendo, 1907, as “I. okamurai” (only the filiform type):154].

Algae erect, dark brown to blackish-brown, with flat, 

straplike, regularly to irregularly dichotomously branched 

fronds, up to 10 cm tall and up to 6 mm wide; arising from 

a crustose holdfast. Branch segments, after the dichotomy 

either more or less same length or one longer than the 

other; branches becoming narrower in width upwards. 

Medulla dense; of colorless, entangled hyphael cells. Cortex of filaments of up to 10–13 small cuboidal cells in 

anticlinal rows; in transection, cortical filaments shorter 

near thallus apices, up to 50 μm thick, becoming thicker 

and of more cells below, 70–140 μm in middle portion 

and 120–165 μm thick near stipe. Cryptostomata are rare; 

they contain only a few phaeophycean hairs (produced 

from lowermost cortical cells), a small number of which 

may project above thallus surface. Pyrenoids absent.

Unilocular sporangia terminal on cortical filament 

(Dawson, 1944). Plurilocular sporangia unknown in 

northern Gulf material.

Habitat. On rocks; midtidal to lowermost

intertidal; occasionally in the shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Ensenada Bocochibampo (northeast of 

Guaymas).

Type Locality. Isla Partida, Las Islas de la

Cintura, Gulf of California.

Remarks. In Mexico, the distribution of Ishige 

sinicola is apparently restricted to the northern Gulf. 

According to Pedroche et al. (2008) the southern Gulf 

record from La Paz was erroneously attributed to Dawson 

(1960c) by Riosmena-Rodriguez and Paul-Chávez (1997). 

A perennial species, it is most abundant, with the broadest fronds, during the winter and least abundant, with the 

narrowest fronds, in summer (Dawson, 1941, 1944).

Dawson (1944, as “I. foliacea”) noted that northern Gulf and Japanese specimens were similar in habit, 

and later that another species, Polyopes sinicola Setchell 

et N.L. Gardner (1924a), was incorrectly described as a 

red alga and was actually the same as the Gulf I. foliacea

(Dawson 1954c). Later Chihara (1969) made the combination I. sinicola (Setchell et N. L. Garnder) Chihara and 

considered it to be conspecific with I. foliacea.

Ishigea sinicola was one of the Gulf species thought 

to have a disjunct distribution, known only in the Gulf 

of California and Japan/eastern Asia (Dawson, 1960c; 

Yoshida, 1998; Kitayama, 2006; Pedroche et al., 2008). 
1 1 0 • 

FIGURE 50. Ishige sinicola: A. Specimens illustrating variation in morphology (JN-2980, JN-4621, and JN-4166). B. Portion of a transection 

from lower part of frond; note medulla of entangled hyphael cells and anticlinal rows of cortical filaments (JN-5824).
n u m b e r 9 4 • 1 1 1

Collections from Japan, Korea and the northern Gulf were 

recently studied by K. M. Lee et al. (2009). Their molecular analyses showed the northern Gulf I. sinicola to be distinct from I.okamurae, and those reported as “I. sinicola” 

from Japan (e.g., Yoshida, 1998) and Korea (E.-Y. Lee et 

al., 2003; Kawai et al. 2005) for which they reinstated I. 

foliacea (K. M. Lee et al. 2009). Other east Asian (western Pacific) records of “I. sinicola,” such as those reported 

from China (Tseng, 1983) and Taiwan (Lewis and Norris, 

1987) need to be investigated, and are probably I foliacea

as well.

Sphacelarial

Sphacelariales Migula, 1908:173, 237.

Sphacelariales Oltmanns, 1922:83.

Algae are erect, terete, unbranched or branched filaments that are attached by a mono- or polystromatic 

discoid holdfast or creeping basal filaments (rhizoids or 

stolons) or aggregated cells endophytic in a host. Filaments 

are uncorticated in some species or corticated by rhizoidal 

growth in others. In the species that branch, there are few 

to many determinate or indeterminate branches. Growth 

is by a prominent apical cell that, at first, divides transversely to form a short apical region; below this region the 

cells longitudinally divide (except in Sphacella) to develop 

the characteristic pseudoparenchymatous filament. In 

some taxa these longitudinal segments may also undergo 

secondary transverse divisions and either do not enlarge 

or may continue to grow in length and width. Hairs are 

known in some. Cells contain several discoid chloroplasts 

without pyrenoids.

Vegetative reproduction can be by fragmentation 

or by propagules (specialized branchlets). Life histories, 

where known, are isomorphic or slightly heteromorphic. 

Sporophytic and gametophytic thalli are similar in appearance. Unilocular sporangia (zoidangia) produce zoospores, 

and neutral plurilocular sporangia produce neutral spores. 

Sexual reproduction is isogamous, anisogamous, or oogamous. Plurilocular gametangia produce isogametes or 

anisogametes. Unilocular oogonia develop an oogonium.

Remarks. Most of the species are marine, with 

only one known from freshwater (Prud’homme van Reine, 

1993). Three families are usually recognized in the Sphacelariales. On the basis of molecular and morphological 

analyses of members from these three families, Draisma et 

al. (2002) found problems in resolving their phylogenetic 

relationships and discussed alternative circumscriptions.

There is one family represented in the Gulf of California.

Spha

Sphacelariaceae Decaisne, 1842a:329, 341.

Algae are small to medium sized, usually erect, and filamentous, and may either be solitary individuals or form 

tufts or mats. Thalli arise from a rhizoidal or stoloniferous 

base or a mono- or polystromatic discoid holdfast. The filament-like main axes and branches are similar and either 

uncorticated or corticated. The cortication may be formed 

by rhizoids or by short branchlets growing from the axis. 

Growth of the erect filaments is by transverse division 

of the single apical cell, and in most genera, subsequent 

segments are formed by longitudinal divisions below the 

apical region in a regular and characteristic fashion. This 

produces few to many longitudinal cell walls in regular 

transverse tiers. Secondary transverse divisions occur in 

the longitudinal segments of some species. There is no secondary growth in the secondary segments.

Life histories, where known, are isomorphic or slightly 

heteromorphic. Reproduction is by unilocular and plurilocular sporangia, which are apparently rare. There is also 

vegetative reproduction by propagules (e.g., Sphacelaria 

subgen. Propagulifera) or fragmentation, and parthenogenesis has been reported from “neutral” plurilocular and 

unilocular gametangia. Species are dioecious (some probably are monoecious), with plurilocular gametangia that 

produce isogametes or anisogametes.

Remarks. In contrast to Sphacelaria, the only 

other genus in the Sphacelariaceae, Sphacella Reinke 

(1890), is composed of only monosiphonous filaments 

(lacking longitudinal divided segments or secondary transverse divisions), but this genus has not been found in the 

Gulf of California.

One genus is represented in the northern Gulf of 

California.

Sphacelaria Lyngbye in Horne

Sphacelaria Lyngbye in Hornemann, 1818:pl. 1600.

Sphacelaria Lyngbye, 1819:xxxi, 103.

Algae are erect, terete filaments that form small, sometimes bushy, tufts that grow on rocks attached by small 

discoid holdfasts or are epiphytic and attached by spreading or penetrating rhizoids. Filaments may be unbranched 

or sparsely to densely branched with determinate or indeterminate branches and with or without corticating rhizoids. Phaeophycean hairs are commonly found on some 

species but are absent on others. Growth is by a usually 
1 1 2 • 

conspicuous meristematic apical cell on the axes and lateral branches that at first divides transversely and later 

divides below the initial apical segments by longitudinal 

divisions that produce more and smaller cells in the tiers 

without increasing the filament diameter. Additionally, 

secondary transverse divisions may occur in the longitudinal segment cells of some species. Cells have numerous 

discoid plastids and are without pyrenoids.

Vegetative reproduction by propagules is common in 

some of the species (i.e., S. subgen. Propagulifera). The 

features of the propagules (e.g., various sizes, shapes, and 

numbers of the lateral apical cells and central apical cell) 

are important characters for discriminating the taxa and are 

generally species specific. Sporophytic and gametophytic 

algae are nearly identical in vegetative structure. Unilocular sporangia (zoidangia), neutral plurilocular sporangia, 

and plurilocular gametangia are on separate individuals, or 

several kinds may occur together on the same alga.

Remarks. Three species are reported in the 

southern Gulf. Sphacelaria brevicornis Setchell et N. 

L. Gardner (1924a, as “Sphacelaria brevicorne”) was 

described from La Paz and has again been reported from 

there (Huerta-Múzquiz and Mendoza-González, 1985). 

Dawson (1944), in agreement with Setchell and Gardner (1924a), noted S. brevicornis from Isla Espiritu Santo 

was similar to S. cornuta Sauvageau (1901). However, 

Prud’homme van Reine (1982) thought S. brevicornis 

was probably the same as S. tribuloides but later considered S. brevicornis to be of uncertain taxonomic position (Prud’homme van Reine, 1993). The others were S. 

divaricata Montagne (1849; Riosmena-Rodriguez and 

Paul-Chávez, 1997, as “S. didichotoma”), and an unidentified species, reported by Rocha-Ramírez and SiqueirosBeltrones (1991, as “Sphacelaria sp.”).

There are four species known in the northern Gulf of 

California.

Key to the Species of Sphacelaria in the no

1a. Propagules slender (Y-shaped), the upper end with 2(–3) projecting, long, thin, cylindrical unbranched arms (S. subgen. 

Propagulifera sect. Furcigerae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rigidula

1b. Propagules without projecting arms; stout, tribuliform (more or less triangular), broader at upper end with 1 central and 

2–3 lateral rounded to pointed apical cells (S. subgen. Propagulifera sect. Tribuloides)  

2a. Axes unbranched or sparsely branched; secondary transverse walls absent within longitudinal segment cells; mature propagules with protruding, relatively pointed lateral apical cells (conical horns)  . . . . . . .

2b. Axes branched, often densely, in either the upper or lower portions; secondary transverse walls absent or present within 

longitudinal segment cells; propagules with nonprotruding to slightly protruding lateral apical cells  

3a. Main axes usually without branches in lower portions, often dense with laterals above; secondary transverse cell walls 

occasional within longitudinal segments; propagules with round, conical, slightly protruding lateral apical cells [Figure 

52A] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. californica

3b. Main axes densely branched from base, only sparsely branched above; secondary transverse cell walls absent; propagules 

with 2 rectangular, nonprotruding (to barely) protruding lateral apical cells [Figure 52B]  . . . . . .

Sphacelaria subgen. Propagulifera

sect. Furcigerae Prud’homme van Rei

Sphacelaria subgen. Propagulifera sect. Furcigerae

Prud’homme van Reine, 1982:203.

Erect filaments are irregularly branched and arise 

above a monostromatic discoid base or stolons or rhizoids. 

Lateral branches are of similar morphology to the axes 

and have indeterminate growth. Propagules are cylindrical with long arms. The subapical cell is divided unequally 

and diagonally.

Sphacelaria rigidula Kützing

FIGURE 51A–C

Sphacelaria rigidula Kützing, 1843:292; Kützing, 1855:25, 

pl. 86: fig. 1; De Toni, 1895a:50; Prud’homme van Reine, 1982:203, figs. 508–554; R. Aguilar-Rosas and 

Machado Galindo, 1990:188; Yoshida et al., 1990:281; 

Stewart, 1991:51; González-González, 1993:443; 

León-Tejera and González-González, 1993:498; LeónTejera et al., 1993:200; Mateo-Cid et al., 1993:50; 

Prud’homme van Reine, 1993:149; Servière-Zaragoza 
n u m b e r 9 4 • 1 1 3

et al., 1993:482; Kitayama, 1994:72, figs. 22–28; 

González-González et al., 1996:300; Yoshida, 1998:200, 

fig. 2-3 D; L. Aguilar-Rosas et al., 2000:132; MendozaGonzález et al., 2000:27, figs. 34–37; L. Aguilar-Rosas 

et al., 2002:235; Lipkin and Silva, 2002:41; Abbott and 

Huisman, 2004:190, fig. 72C; López et al., 2004:10; R. 

Aguilar-Rosas et al., 2005b:35; Keum et al., 2005:4, 

figs. 1–12; Riosmena-Rodríguez et al., 2005:101; Mateo-Cid et al., 2006:49, 57; Servière-Zaragoza et al., 

2007:8; Pedroche et al., 2008:33.

Sphacelaria furcigera Kützing, 1855:27, pl. 90: fig. 2; 

Setchell and Gardner, 1924a:724, pl. 19: fig. 58; Setchell and Gardner, 1925:396, pl. 37: fig. 29; Setchell 

and Gardner, 1930:138; Dawson, 1944:224; Dawson, 1951:52; Dawson, 1954d:400, fig. 14h; Dawson, 

1957b:9; Dawson, 1959a:18; Dawson, 1959c:4; Dawson et al., 1960a:10; Dawson, 1966a:10; Dawson and 

Neushul, 1966:174; van den Hoek and Flinterman, 

1968:193; Abbott and Hollenberg, 1976:218, fig. 181; 

Pacheco-Ruíz, 1982:70; Stewart, 1982:54; Schnetter 

and Bula Meyer, 1982:59, pl. 8: fig. I; Stewart and 

Stewart, 1984:141; Huerta-Múzquiz and MendozaGonzález, 1985:44; Sánchez-Rodríguez et al., 1989:41; 

González-González et al., 1996:299; Hoffmann and 

Santelices, 1997:138, fig. 34.

Algae small filamentous tufts, up to 1 cm tall; axes 

and branches uniseriate near tips, each with a conspicuous 

apical cell; cells below multiseriate with 1–7 longitudinal 

walls per segment in side view; attached by rhizoidal filaments or discoid base. Axes sparsely, irregularly branched; 

axes and branches of similar diameter, (16–)25–45 μm in 

diameter; segments as long as or longer than wide. Secondary transverse cell walls occasional within longitudinal 

sections of segments. Phaeophycean hairs present.

Propagules Y-shaped; stalk basally narrowed, terminally with a slightly protruding central apical cell, and 

2(–3) lateral, long, slender, cylindrical arms of more or 

less the same length (170–400 μm) and diameter (20–25 

μm); each arm with a conspicuous apical cell. Unilocular sporangia on 1-cell pedicel, mostly spherical, 55–80 

μm in diameter. Plurilocular sporangia on 1-cell pedicel, 

FIGURE 51. Sphacelaria rigidula: A. Upper part of filament with prominent apical cell, below the secondary segments with longitudinal walls, 

and two phaeophycean hairs (JN-4041). B. Y-shaped mature propagule (JN-3271). C. Erect filament with young propagule and a phaeophycean hair (JN-3715).
1 1 4 • 

elongate-ovoid, 55–63 μm long and 40–50 μm in diameter; two kinds, one with numerous small locules, another 

with fewer and larger locules (“male and female gametangia” of Setchell and Gardner, 1924a:pl. 19, fig. 58).

Habitat. Partially endophytic within host cells

and epiphytic above host surface; on various algae, notably on Sargassum and Padina, and epizoic on a parrotfish “beak” (Dawson, 1959a); mid intertidal to shallow 

subtidal.

Distribution. Gulf of California: El Coloradito to Ensenada de San Francisco (vicinity of Guaymas);

La Paz to Punta Palmilla (vicinity of San José del Cabo). 

Pacific coast: Alaska to southern California; Isla Guadalupe; Punta de la Asunción, Baja California Sur to Oaxaca; 

Isla San Benedicto (Islas Revillagigedo); Clipperton Island; 

Costa Rica; Colombia; Chile; Hawaiian Islands; Korea; 

Japan.

Type Locality. On Cystoseira triquetra,

Nuweiba, Sinai, on the Gulf of Aqaba, Red Sea, Egypt.

Remarks. Sphacelaria rigidula is a relatively 

common epiphyte during the spring in the Gulf. It has 

been widely reported in temperate to tropical waters as “S. 

furcigera” (Prud’homme van Reine, 1982). A Pacific coast 

species, S. subfusca Setchell et N. L. Gardner (1924b) from 

southern California, is morphologically close to S. rigidula. Although S. subfusca was considered to be a synonym 

of S. rigidula (Abbott and Hollenberg, 1976; Kitayama, 

1994), it was kept as a separate species by Prud’homme 

van Reine (1982).

Sphacelaria subgen. Propagulifera

sect. Tribuloides Prud’homme van Rei

Sphacelaria subgen. Propagulifera sect. Tribuloides

Prud’homme van Reine, 1982:178.

Algae erect filaments, composed of axes that are more 

or less pinnately to irregularly branched, arising above a 

monostromatic or polystromatic discoid base, or stolons 

or rhizoids. Propagules are tribuliform (i.e., more or less 

triangular-shaped), broader at upper end, with 1 central 

and 2 lateral apical cells.

Remarks. Keum et al. (1999) found two types 

of division sequences of the upper lateral apical cells of 

the propagule that produced their shape and number of 

cell segments below the propagule waist: (1) round, produced by oblique divisions to form round, conical protruding (horned) apical cells (i.e., Asian S. californica; see 

Keum et al., 1999: figs. 33–36; Figure 52A herein), and (2) 

rectangular, developed by an initial longitudinal and then 

a transverse division to form rectangular, nonprotruding 

(flat) apical cells (for examples, see Keum et al., 1999: fig. 

37 of Asian S. californica and also figs. 51–53 of Asian 

S. novae-hollandica; see also S. hancockii in Figure 52B 

herein). The number of segments under the waist of propagule (cf. Keum et al., 1999:fig. 33 versus Figure 52 herein) 

also affects the shape of the propagule.

Specimens of species recorded from the Gulf of California, along with type and type locality specimens of 

European S. tribuloides, southern California S. californica, and Gulf S. hancockii, need critical comparative 

morphological, molecular, and phylogenetic analyses to 

resolve their taxonomic status.

Three species are currently known in the northern 

Gulf of California.

Sphacelaria californica (Sauvageau)

Setchell et N. L. Gardn

FIGURE 52A

Sphacelaria plumula var. californica Sauvageau, 1901:108, 

fig. 21.

Sphacelaria californica (Sauvageau) Setchell et N. L. Gardner, 1925:395, pl. 37: figs. 23–27; Abbott and Hollenberg, 1976:216, fig. 179; Boo and Chou, 1986:99, 

figs. 16–22; Sánchez-Rodríguez et al., 1989:41; Yoshida et al., 1990:281; Kitayama et al., 1991:189–

193, figs. 1–22; Stewart, 1991:51; Prud’homme van 

Reine, 1993:150; Kitayama, 1994:52, figs. 7–11, 

56G; González-González et al., 1996:164; Yoshida, 

1998:197, fig. 2-3 G; Draisma et al., 1998:186–187, 

tbl. 2; Keum et al., 1999:24, figs. 1–41; MendozaGonzález et al., 2000:25, figs. 12–15; L. Aguilar-Rosas 

et al., 2002:132; Riosmena-Rodríguez et al., 2005:101; 

Servière-Zaragoza et al., 2007:8; Pedroche et al. 2008 

(in part; S. californica only):31.

Sphacelaria tribuloides sensu De A. Saunders, 1898:158, 

pl. 26: figs. 1–8 [non Sphacelaria tribuloides Meneghini, 

1840:[2])].

Algae filamentous tufts, relatively stiff, up to 2.0 cm 

tall; main axes 25–50 μm in diameter in lower portion, 

widening to 60–95 μm in diameter in midportions; lower 

portion without laterals or only very few; attached by small 

polystromatic disc or, if epiphytic, by rhizoids. Upper portions of axes irregularly branched (sometimes more or less 

pinnate) with primary and secondary lateral branches; laterals narrowed at base. Apical cell prominent, 40–60 μm 

in diameter, 90–200 μm long; uppermost segments of uniseriate cells; segments below multiseriate, longitudinally 

divided by 1–5 walls; occasionally, longitudinal divided 
n u m b e r 9 4 • 1 1 5

cells with a few secondary transverse walls. Phaeophycean 

hairs may be absent or sparse to abundant.

Propagules ellipsoidal when young, tribuliform when 

mature, 140–170 μm long, 80–105 μm wide; with 1 central and 2 lateral, slightly protruding, round, conical apical cells (Figure 52A). Unilocular sporangia, spherical to 

oval, 40–50 μm long, 30–50 μm in diameter. Plurilocular 

sporangia obovate, borne on a 1-celled pedicel, 30–40 μm 

by 50–70 μm.

Habitat. On rocks and in tide pools; high to

mid intertidal.

Distribution. Gulf of California: Puertecitos

to San José del Cabo. Pacific coast: Isla Guadalupe; southern California to southernmost Baja California Sur; Islas 

Revillagigedo; Jalisco to Colima; China; Korea; Japan.

Type Locality. San Diego, San Diego

County, California.

Remarks. Gulf records of Sphacelaria californica (L. Aguilar-Rosas et al., 2002; Riosmena-Rodríguez 

et al., 2005) were not seen by the author; the description is 

based on Setchell and Gardner (1925), and the sporangia 

are based on Kitayama (1994). Although Kitayama et 

al. (1991) referred to this taxon as “S. californica Sauvageau ex Setchell et Gardner,” Sauvageau (1901:108) 

validly published this taxon as a variety, S. plumula var. 

californica. However, in the same publication, Sauvageau 

(1901:109), as “Sphacelaria californica?,” also suggested 

it was possibly a species by referring to it with a taxonomic query, but the first valid combination of the taxon 

as a species was made by Setchell and Gardner (1925).

Sphacelaria hancockii E. Y. Daw

FIGURE 52B

Sphacelaria hancockii E. Y. Dawson, 1944:225, pl. 31: fig. 

8; Dawson, 1954e:325; Dawson, 1959a:8, 18; Dawson 

et al., 1960a:12; Dawson, 1961b:387; Huerta-Múzquiz 

and Tirado-Lizárraga, 1970:135; Huerta-Múzquiz 

and GarzaBarrientos, 1975:7; Stewart and Stewart, 

1984:141; González-González et al., 1996:164; L. Aguilar-Rosas et al., 2000:132.

FIGURE 52. Two species of Sphacelaria. A. Sphacelaria californica: Propagule with lateral, rounded, slightly protruding apical cells and filament with both longitudinal divisions and two secondary transverse divisions in surface view (after Setchell and Gardner, 1925: pl. 37, fig. 7). B. 

Sphacelaria hancockii: Propagule with two lateral, rectangular, nonprotruding apical cells and filament without secondary transverse divisions 

(drawn from type specimen, EYD-[1940]-643, after Dawson, 1944: pl. 31, fig. 8).
1 1 6 • 

Sphacelaria mexicana W. R. Taylor, 1945:86, pl. 3: figs. 

1–8; Dawson, 1954e:325; Servière-Zaragoza et al., 

2007:8.

Sphacelaria californica sensu [in part, with reference 

Gulf of California S. hancockii specimens only] Abbott and Hollenberg, 1976:216; Mendoza-González 

et al., 2000:25; Pedroche et al., 2008:31 [non Sphacelaria californica (Sauvageau) Setchell et N. L. Gardner, 

1925:395].

Small filamentous tufts, up to 12 mm tall, densely 

branched from base and sparsely, irregularly branched 

above; attached to rock by dense mat of entangled rhizoids. Erect filaments (30–)40–50(–60) μm in diameter; 

segments about as long as wide, with 1–3(–4) longitudinal 

divisions in side view, secondary transverse cell walls not 

seen. Phaeophycean hairs abundant, up to 10 cells long; 

0.5–1.0 mm long and to 15 μm in diameter.

Propagules bibuliform (as seen from below or the top) 

and tribuliform (triangular-shaped, as seen in side view), 

borne on pedicels of 2–3 cells, 100–110 μm long by 75–90 

μm wide; upper end with 1 central and 2 lateral rectangular, flat apical cells, either not protruding (Figure 52B) or 

only barely protruding.

Habitat. On rocks, sometimes entangled with

other turf algae, such as Cladophora, Ectocarpus, or Ceramium; mid to low intertidal.

Distribution. Gulf of California: Puertecitos

to Isla Turners (off southern end of Isla Tiburón); Isla Carmén; San José del Cabo; Mazatlán, Sinaloa. Pacific coast: 

Isla Guadalupe; Bahía Sebastián Vizcaíno to Punta Abreojos, Baja California; Isla San Benedicto and Isla Socorro 

(Islas Revillagigedo); Colima to Oaxaca.

Type Locality. San José del Cabo, Baja California Sur

Remarks. Dawson (1954e), in reporting 

S. hancockii from Baja California, noted that it and S. 

mexicana W. R. Taylor from Pacific Mexico both had 

bibuliform propagules (cf. Taylor, 1945:pl. 3: fig. 7) and 

considered the latter to be a synonym.

Sphacelaria hancockii may be related to S. californica

and has been considered to be a synonym (Abbott and Hollenberg, 1976; Mendoza-González et al., 2000). However, 

the two differ morphologically: S. hancockii is densely 

branched basally and sparsely above; has narrower filaments, 30–60 μm in diameter; and has smaller propagules, 

100–110 μm long by 75–90 μm wide. S. californica usually has no branches below and is densely branched above; 

has mostly larger diameter filaments, 40–95 μm; and has 

longer propagules, 140–170 μm by 80–105 μm. Detailed 

studies of propagule development, along with vegetative 

differences of Gulf S. hancockii, Gulf S. californica, and 

the type of S. californica, are required to determine if the 

characteristics differ consistently enough to be considered 

separate species (see Remarks under Sphacelaria subgen. 

Propagulifera sect. Tribuloides).

For now, it seems best to recognize them as different species, until more collections are made, especially of 

reproductive material, to facilitate detailed studies, including molecular and phylogenetic analyses to test the status 

of the Gulf taxon.

Sphacelaria tribuloides Meneghini

FIGURE 53A–C

Sphacelaria tribuloides Meneghini, 1840:[2]; Dawson, 

1959a:5, 18; Dawson, 1961b:387; Huerta-Muzquiz 

and Tirado-Lizárraga, 1970:127, 130; Prud’homme 

van Reine, 1982:179, figs. 422–454; Huerta-Múzquiz 

and Mendoza-González, 1985:44; Mateo-Cid et al., 

1993:50; Yoshida et al., 1990:28; Prud’homme van 

Reine, 1993:149; González-González, 1993:443; LeónTejera et al., 1993:204; León-Tejera and GonzálezGonzález, 1993:498; Servière-Zaragoza et al., 

1993:482; Kitayama, 1994:59, figs. 12–15; GonzálezGonzález et al., 1996:300, 381; Yoshida, 1998:202, 

fig. 2-3 E; L. Aguilar-Rosas et al., 2000:132; MendozaGonzález et al., 2000:28, figs. 38–41; Abbott and Huisman, 2004:190, fig. 72D,E; Mateo-Cid et al., 2006:49, 

57; Pedroche et al., 2008:34.

Algae forming tufts of relatively stiff filaments; axes 

mostly unbranched or only sparsely, irregularly branched; 

axes and branches 25–50(–70) μm in diameter; attached 

below by discoid or stoloniferous base. Segments of axes 

and branches 25–60 μm in length; segments below uppermost uniseriate segments with 1–3(–5) longitudinal divisions in side view, each section 5–12 μm wide. Secondary 

transverse cell walls within longitudinal divided sections 

not present. Phaeophycean hairs often present.

Propagules tribuliform, borne on pedicels of 1–3 cells, 

90–130 μm long by 80–110 μm wide, upper end with 1 

central apical cell and 2 large lateral apical cells, slightly 

protruding to somewhat apparent conical horns.

Habitat. On rocks or epiphytic; intertidal

Distribution. Gulf of California: Playa

Santa Teresa to Puerto Peñasco; Bahía Concepción to Isla 

San Diego. Pacific coast: Guerrero to Oaxaca; Isla Socorro 

(Islas Revillagigedo); Hawaiian Islands; Japan.

Type Locality. La Spezia (Spèsa), on the

Ligurian Sea, northern Italy.
n u m b e r 9 4 • 1 1 7

Remarks. The description of Sphacelaria tribuloides is based on specimens from Puerto Peñasco and 

on Prud’homme van Reine (1982) and Kitayama (1994).

Dic

Dictyotales Bory de Saint-Vincent, 1828:142.

Dictyotales Kjellman, 1896:291.

Algae are erect, flattened and membranous, branched 

in one plane, and variously attached by a holdfast of rhizoidal filaments or by rhizoids along lower blade margin 

or, in a few species, by stoloniferous attachments. Growth 

is by an apical meristem, either a single cell, a group of apical cells, or an apical row of marginal cells. Sporophytes 

and gametophytes are parenchymatous in structure, with 

a medulla of one or more layers of large colorless cells and 

a single- to multiple-layer cortex of small cells with discoid 

plastids without pyrenoids. Phaeophycean hairs are either 

absent or present in bands or scattered tufts.

The life history phases, where known, are isomorphic. Sporophytes produce unilocular sporangia that are 

scattered or in rows or groups and with or without an 

indusium or paraphyses. Each sporangium develops one, 

four, or eight nonmotile spores. Sexual reproduction is 

oogamous. Gametophytes are usually dioecious, with 

oogonia and antheridia usually in sori. There is one egg 

produced per oogonium. Antheridia are plurilocular, and 

each of the motile antherozoids have a single flagellum.

Remarks. Members of this order are of widespread occurrence in tropical and subtropical seas. However, some are found in temperate seas and are particularly 

well developed in temperate South Australia (Womersley, 

1987). On the basis of nrDNA sequences, the phylogeny 

of Rousseau et al. (2001) concluded the Dictyotales were 

the earliest lineage to diverge during the evolution of the 

Phaeophyceae.

The one family of the order is well represented 

throughout the Gulf of California.

Dic

Dictyotaceae J. V. Lamouroux ex Dumortier, 1822:72, 101.

Algae are simple or branched in one plane. They can 

be straplike (e.g., Dictyota) or fanlike or bladelike (e.g., 

FIGURE 53. Sphacelaria tribuloides: A. Prominent apical cell of erect filament and secondary segments with longitudinal walls. B. Propagule 

with two lateral rounded protruding apical cells. C. Mature propagule with lateral apical cells elongated to conical horns. A–C. JN-4038.
1 1 8 • 

Padina). Structurally, they are composed of a medulla 

of large colorless cells in one or more layers and a cortical layer of one to several rows of small pigmented cells. 

Growth is by a single apical cell, group of apical cells, or 

marginal row of apical cells. Phaeophycean hairs are present in tufts or zones and are simple, delicate, colorless, and 

mostly deciduous.

Reproductive structures are superficial on the thallus. 

Sporophytic reproduction is by nonmotile aplanospores, 

formed by sporangia that are isolated or in sori. Sporangia 

are mostly divided into a group of four (tetraspores) or 

eight (octospores) aplanospores. Oogonia are solitary or 

grouped in sori. Each oogonium forms a nonmotile single 

egg (ova). Plurilocular antheridia are grouped in sori and 

produce motile antherozoids.

Remarks. Genera of the Dictyotaceae have traditionally been grouped into two taxonomic tribes, following, but not named by, Kjellman (1896): (1) the Dictyoteae 

Lamouroux (1809a) for genera characterized by growth 

from a single apical cell and (2) the Zonarieae Lamouroux 

(1809a) for genera with a group or marginal row of apical cells. However not all the species fit into these tribes. 

For example, some species of Padina exhibit both kinds 

of growth, with a “Vaughaniella-form” that has growth 

by a single apical cell and can undergo a morphogenetic 

change to growth by a marginal row of apical cells (Lewis 

et al., 1987). Sulfuric acid accumulation has been reported 

in some species of Dictyopteris, Spatoglossum, and Zonaria C. Agardh (1817) (Sasaki et al., 1999), and its presence 

or absence has been suggested to be an informative taxonomic character (Sasaki et al., 2004, 2005).

The genera Dictyota, Dilophus and Rugulopteryx

are morphologically similar, separated primarily on the 

basis of their vegetative anatomy. Dictyota sensu stricto 

is distinguished by both a single-layered medulla and cortex throughout (Figure 54B). Dilophus sensu stricto has 

a medulla of two or more layers and a cortex of one cell 

layer. Rugulopteryx has a one cell layer medulla centrally 

and two or more layers in the margins, and a cortex of 

one-cell layer (Figure 54D).

More recent phylogenetic analysis has shown Glossophora, Glossophorella and Pachydictyon to be congeneric 

with Dictyota, and revealed two genera, Canistrocarpus

and Rugulopteryx, that could be distinguished from Dictyota (De Clerck et al., 2006; Hwang et al., 2009). However, the taxonomic status of Dilophus J. Agardh (1882) 

remains problematic. Setchell and Gardner (1925) and 

Hörnig et al. (1992a, 1992b, 1993) concluded that the 

medulla character alone was not enough to separate Dilophus and included it in Dictyota. Others have retained 

Dilophus (e.g., Dawson, 1950a; Womersley, 1987; Phillips, 1992; Huisman, 2000). Phillips (1992) recognized 

that the number of cells in the stalk that bears a sporangium (one or two-cells) in Australian Dilophus was of taxonomic importance, and some molecular phylogenies have 

supported their generic separation (e.g., Hoshina et al., 

2004; Kraft et al., 2004). De Clerck et al. (2006) kept the 

two genera separate, noting this cannot be resolved until 

the generitype, Dilophus gunnianus J. Agardh (1882), has 

been studied.

Next to Sargassum, species of the Dictyotaceae, notably Dictyota and Padina are among the most abundant 

brown algae in the Gulf. In addition to the genera known 

in the northern Gulf, there are three others known in the 

southern Gulf: Lobophora J. Agardh (1894), with the 

report of L. variegata (J. V. Lamouroux) Womersley ex E. 

C. Oliveira (1977) by Dawson (1959a, as “Pocockiella variegata”) and Mateo-Cid et al. (2000b); Taonia J. Agardh 

(1848) with the record of T. lennebackerae Farlow ex J. 

Agardh (1894) by Mateo-Cid et al. (2002b); and Dilophus J. Agardh (1882) with the report Dilophus pinnatus 

E. Y. Dawson (1950a; = Dictyota pinnata (E. Y. Dawson) 

Hörnig, Schnetter et Prud’homme von Reine, 1993) (Espinoza-Ávalos, 1993; Mateo-Cid et al., 2000a, 2000b), and 

elsewhere in Pacific Mexico from Nayarit (León-Tejera 

et al., 1993; González-González, 1993), Jalisco (Pedroche and González-González, 1981; Servière-Zaragoza et 

al., 1993) and Michoacán (Stout and Dreckmann, 1993), 

and in South America from Pacific Colombia (Schnetter 

and Bula Meyer, 1982). In West Africa, material identified 

as Dilophus pinnatus was noted to be a growth form of 

Padina durvillei (Lawson and John, 1987).

There are five genera of Dictyotaceae represented in 

the northern Gulf of California.

Key to the Genera of Dictyotaceae in the no

FIGURE 54A–I

1a. Thallus of flat, repeatedly branched straplike segments; growth from a single apical cell [Figure 54A] . . . . . . . . . . . . . . 2

1b. Thallus of flat, broad unbranched to branched fronds or blades; growth from a single clustered group of apical cells [Figure 

54E] or separate groups of clustered apical cells [Figure 54G] or a continuous marginal row of apical cells [Figure 54I] . . . . 3
n u m b e r 9 4 • 1 1 9

Dictyopteris J. V. Lamou

FIGURE 54E,F

Dictyopteris J. V. Lamouroux, 1809d:332.

Algae are erect or prostrate and are generally clumped 

and composed of flat fronds in one plane or twisting and 

attached by a thickened, discoid to conical holdfast of 

compacted rhizoids. Branching pattern is subdichotomous 

to irregular. Fronds and branches are generally straplike, 

up to 25 mm wide, and have a pronounced midrib, up to 

30 cells thick. Lateral veins are present in some but are 

lacking in the northern Gulf species. Growth is by an apical group of up to 15 cells. There is a single layer of small 

cortical cells, each with several chloroplasts, and a multilayered medulla of larger cells. The midrib is conspicuous 

and much thicker, of 8–30 cells. The thallus wings (on 

each side of the midrib) are multilayered in most species 

(Figure 54F) and only 2 cell layers in a few of the species. 

Phaeophycean hairs are mostly scattered in groups over 

the surfaces.

Reproductive structures are solitary, in groups, or in 

sori and either embedded or projecting above both sides 

of the thallus surface. Sporophytes have subspherical to 

ovoid sporangia that are solitary or borne in sori that are 

scattered over the branches or near the midrib, are without a stalk or on one- to two-celled stalks, and produce 

a tetrad of spores. Gametophytes are dioecious. Oogonia 

are solitary or in small groups. Antheridia are composed 

of many tiered locules in large, whitish sori and are surrounded by one to three layers of sterile cells.

Remarks. Among the Gulf’s brown algae, 

Dictyopteris is most easily recognized by its straplike, 

subdichotomously branched fronds with conspicuous 

midribs. Sasaki et al. (1999) used presence or absence of 

sulfuric acid to separate morphologically similar species 

of Dictyopteris and suggested it was a useful taxonomic 

character (Sasaki et al., 2004, 2005).

Two of the smaller, distromatic species have been 

reported in the southern Gulf: Dictyopteris delicatula J. V. 

Lamouroux (1809d), (Huerta-Múzquiz and MendozaGonzález, 1985; Rocha-Ramírez and Siqueiros-Beltrones, 

1991; Mateo-Cid et al., 2000b; Riosmena-Rodríguez et 

al., 2005; Mendoza-González and Mateo-Cid, 2005); 

and D. repens (Okamura) Børgesen (1924) by Dawson (1959b). Another, D. polypodioides (de Candolle) 

J. V. Lamouroux (1809b; see Silva et al. 1987), has been 

reported from Isla Guadeloupe and Pacific Baja California (Dawson, 1950d; Stewart and Stewart, 1984, both as

D. membranacea (Stackhouse) Batters 1902).

One species is known in the northern Gulf of 

California.

Dictyopteris undulata Holm

FIGURE 55

Dictyopteris undulata Holmes, 1896:251, pl. 8: fig. 1; Abbott, 1972:260, fig. 1; Abbott and Hollenberg, 1976:212, 

fig. 176; Devinny, 1978:358; Huerta-Múzquiz, 

1978:336; Lewbel et al., 1981:165; L. Aguilar-Rosas, 

1982:31; Stewart and Stewart, 1984:141; MendozaGonzález and Mateo-Cid, 1985:24; Benson, 1986:448; 

Sánchez-Rodríguez et al., 1989:40; R. Aguilar-Rosas et 

al., 1990:123; Stewart, 1991:48; Rocha-Ramírez and 

Siqueiros-Beltrones, 1991:32; Martínez-Lozano et al., 

1991:23; Lluch-Cota et al., 1993:336; Mateo-Cid and 

Mendoza-González, 1994a:50; Kajimura, 1995:89; 

González-González et al., 1996:145; Pacheco-Ruíz and 

Zertuche-González, 1996b:171; Tanaka, 1998:75, figs. 

1–15; Yoshida, 1998:211; Mateo-Cid et al., 2000b:196, 

figs. 13, 14, 64, 65, 94; Mendoza-González and MateoCid, 2005:57, figs. 60–69; Mateo-Cid et al., 2006:49, 

2a. Medulla usually a single cell layer with cortex of 1–2 cell layers [Figure 54B] or a medulla of 2–4 cell layers with cortex of 

2–5 cell layers [Figure 54C]  . . . 

2b. Medulla 1 cell layer in central portion and 2–8 cell layers at the branch margins; cortex of 1 cell layer throughout [Figure 

54D] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rugulopteryx

3a. Fronds with conspicuous midrib [Figure 54E]; internally of 1 or more cell layers at margins, increasing number of cell layers toward center of blade (midrib) [Figure 54F]. . . . . . 

3b. Fronds without midribs; internally, cell layers not increasing in number toward center of blade  

4a. Fan-shaped or cup-shaped blades, sometimes lobed or torn; with distinctive inrolled margins [Figure 54I]; noncalcified or 

slightly to heavily calcified  . . 

4b. Fronds irregularly divided or branched, narrow to broadly lanceolate; without inrolled margins [Figure 54G]; not 

calcified  . . . . . .
FIGURE 54. Northern Gulf of California genera of Dictyotaceae: Comparative diagrammatic illustrations of the apical growth region (meristematic cells stippled) and transections of internal anatomy. A. Dictyota and Rugulopteryx: Growth from single apical cell. B. Dictyota transection: Most species have single medullary layer surrounded by single layer of pigmented cortical cells. C. Dictyota transection: A few species (e.g., 

D. coriacea) may have more than one medullary cell layer at margins and more than one cortical cell layer. D. Rugulopteryx transection: One 

medullary cell layer centrally and more than one medullary cell layer at the margins, and a single cortical cell layer. E. Dictyopteris: Growth 

from a group of apical cells at center of distal blade margin. F. Dictyopteris transection: With thicker medulla at center midrib and progressively 

fewer cell layers outward toward the margins. G. Spatoglossum: Growth from separate groups of apical cells along distal margin of blade. H. 

Spatoglossum transection: Medullary layers of irregularly arranged cells and single cortical cell layer. I. Padina: Longitudinal section of inrolled 

margin: Growth from an apical row of meristematic cells along entire distal edge of blade, which is enclosed within the inrolled margin. Number of cell layers below single apical row is initially one and then becomes two (as shown); in Gulf species, the medullary cell layers increase in 

number toward the central portions and basal region of the blade (drawn by Alice R. Tangerini).
n u m b e r 9 4 • 1 2 1

57; Pacheco-Ruíz et al., 2008:203; Pedroche et al., 

2008:36.

Haliseris undulata (Holmes) Okamura, 1899b: Exsiccate 

No. 41; Okamura, 1902a:110; Okamura, 1907:51, pl. 

XI: figs. 1–10.

Neurocarpus undulata (Holmes) Okamura, 1928:190 

[Corrigenda]; Okamura, 1936:172, fig. 88 (1–5).

Dictyopteris polypodioides sensu Hariot, 1891:217 [non 

Dictyopteris polypodioides (de Candolle) J. V. Lamouroux, 1809d:332].

Haliseris polypodioides sensu McClatchie, 1897:354 

[non Haliseris polypodioides (de Candolle) C. Agardh, 

1820:142; = Dictyopteris polypodioides (de Candolle) 

J. V. Lamouroux, 1809d:332].

Dictyopteris zonarioides Farlow, 1899:73; Dawson, 

1951:52; Dawson, 1952:431; Dawson, 1959a:18; 

Dawson et al., 1960a:34, pl. 3: fig. 4, pl. 4: fig. 2; Dawson et al., 1960b:11; Dawson, 1961b:388; Dawson, 

1966a:11; Guzmán del Próo et al., 1972:260; Norris, 

1973:4; Fenical et al., 1973:2383; Espinoza-Ávalos, 

1993:332; González-González et al., 1996:145.

Neurocarpus zonarioides (Farlow) M. Howe, 1914:69; 

Setchell and Gardner, 1924a:728; Setchell and Gardner, 

1925:656, pl. 34: fig. 4, pl. 35: fig. 11, pl. 36: fig. 21, pl. 

38: fig. 39, pl. 95; Dawson, 1944:229.

Algae erect, flat, irregularly dichotomously branched 

fronds, up to 35 cm tall, (3–)10–20 mm wide, with a 

prominent percurrent midrib and lacking any veins, often 

with undulate margins; ultimate branches shorter than 

lower segments; with rounded apices; arising from a more 

or less conical holdfast. In lower portions the wider blade 

may be worn away, leaving the midribs. Midribs may be 

tomentose. Thallus in transverse section up to 12 layers 

of cells and up to 400 μm thick. Growth from an apical 

group of meristematic cells.

FIGURE 55. Dictyopteris undulata: Habit; note the prominent midrib (JN-5801).
1 2 2 • 

Sporangia grouped in sori, 750–1000 μm long, along 

both sides of the midrib. Oogonial and antheridial sori 

scattered over frond. Oogonium 95–150 μm tall by 

95–125 μm wide.

Habitat. On rocks, usually in shaded areas

and tide pools; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to San José del Cabo. Pacific coast: southern California to Baja California Sur; Japan; China; Taiwan.

Type Locality. Misaki, Kanagawa Prefecture, central Japan

Remarks. A seasonal species, Dictyopteris 

undulata occurs in the northern Gulf from February to 

August and appears to reach its peak in size as well as in 

numbers of individuals in late spring. Although previously 

collected during 1972–1975 at Punta la Gringa (vicinity 

of Bahía de los Ángeles), D. undulata was not encounted during the 1985–1986 surveys of Pacheco-Ruíz and 

Zertuche-González (1996b).

Some of the Gulf specimens with narrow fronds resemble Dictyopteris johnstonei N. L. Gardner (1940), a species described from Santa Cruz Island (California Channel 

Islands) and reported southward to Bahía de Todos Santos, Pacific Baja California (Silva, 1957; Mateo-Cid et al., 

2000b; Mendoza-González and Mateo-Cid, 2005; Pedroche et al., 2008), and Isla San Benito (Dawson, 1961b; 

Abbott and Hollenberg, 1976). Silva (1957) has suggested 

the taxonomic status of California D. johnstonei Gardner (1940) needs to be resolved. Reinvestigations should 

include Gulf specimens morphologically similar to Pacific 

coast D. johnstonei (including its type locality) in comparative morphological studies and molecular analyses, 

to each other, and to the distinctive D. undulata of the

Gulf, Japan (type locality), and Pacific coast (including 

type locality specimens of Neurocarpus zonarioides, considered a synonym of D. undulata).

Dictyota J. V. Lamou

FIGURE 54A–C

Dictyota J. V. Lamouroux, 1809a:38.

Dictyota J. V. Lamouroux, 1809d:331.

Glossophora J. Agardh, 1882:108.

Glossophorella Nizamuddin et Campbell, 1995:258.

Pachydictyon J. Agardh, 1894:81.

Algae are erect or repent (prostrate), composed 

of flat ribbonlike fronds that are clumped or bushy in 

appearance, and yellow- to dark brown, and some are iridescent. Branching may be regularly or irregularly dichotomous or alternate throughout; branches are up to 20 

mm wide, with smooth, dentate, crenulate, or ciliate margins and usually obtuse apices. Most species are attached 

by a single, or several rhizoidal holdfasts, although a few 

species are attached by stoloniferous holdfasts. Secondary attachments may occur in some when thallus or its 

branches come in contact with the substratum. Phaeophycean hairs and surface proliferations may be present 

in tufts scattered over the thallus. Growth is by a single 

apical cell at the tip of each rounded or acute branch 

apex. The medulla of one or more layers of large colorless cuboidal cells is easily differentiated from the cortex 

of one or more layers of small pigmented cells, each with 

numerous discoid chloroplasts.

Reproductive structures are mostly in sori (some with 

solitary sporangia), scattered and protruding from both 

sides of the thallus surface. Sporangia, borne on a single 

stalk cell, are divided into four aplanospores and may be 

solitary, grouped in sori, or around hair tufts, over the 

upper portions of the frond. Gametangia are grouped in 

sori. Oogonia are in rows within sori, with each oogonium 

on a single stalk cell. Antheridia are produced in whitish, antheridial sori. Each antheridium, also on a single 

stalk cell, is composed of many tiers of locules and is surrounded by hyaline, elongated sterile cells.

Remarks. Some members of Dictyota exhibit 

wide morphological variation, and this phenotypic plasticity and few taxonomic characters have resulted in confusion regarding discrete differences among many of the 

species. The taxonomic limits of Gulf species are poorly 

known, but “typical” forms of each can be recognized in 

the field. Detailed observations, temporal and spatial, on 

field populations will be helpful in understanding the morphological variation. Culture and life history studies and 

molecular analyses will aid in understanding species and 

phylogenetic relationships. Monographic studies of the 

genus in the Gulf and Pacific Mexico, such as those of De 

Clerck (1999, 2003) for the Indian Ocean and De Clerck 

et al. (2006), are critically needed.

Three species, all originally described from the western Atlantic, are recorded in the southern Gulf: D. bartayresiana J. V. Lamouroux (1809a) reported by Mateo-Cid 

et al. (2000b) and L. Aguilar-Rosas et al. (2000); “D. cervicornis” reported by Mateo-Cid et al. (1993, 2000b) and 

Cruz-Ayala et al. (2001) (note that D. cervicornis Kützing 

(1859) is now Canistrocarpus crispatus (J. V. Lamouroux) 

De Paula et De Clerck in De Clerck et al., 2006); and “D. 
n u m b e r 9 4 • 1 2 3

volubilis” reported by Huerta-Múzquiz and MendozaGonzález (1985), Rocha-Ramírez and Siqueiros-Beltrones 

(1991), Cruz-Ayala et al. (2001), and Riosmena-Rodríguez et al. (2005) (note that D. volubilis Kützing (1849) 

is now D. dichotoma (Hudson) J. V. Lamouroux, 1809a; 

see De Clerck, 2003).

Currently seven species, including one variety, are recognized in the northern Gulf.

Key to the Species of Dictyota in the nor

1a. Medulla of 2–4 (or more) cell layers; cortex of (1–)2–5 cell layers [Figure 54C]. . . . . .

1b. Medulla usually 1 cell layer; cortex of only 1(–2) cell layer(s) [Figure 54B]  

2a. Thalli decumbent overlapping fronds; of short, relatively broad segments. . . . . . . . . . . . . . . . . . . . . . . . . . D. concrescen

2b. Thalli erect, fronds of strap-shaped segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Frond margins with minute denticulations  . . . . . 

3b. Frond margins smooth or sometimes with marginal proliferations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4a. Attached by spreading, simple to divided, terete, stoloniferous growths, up to 500 μm in diameter . . . . . . D. stolonifer

4b. Attached by filamentous rhizoids, sometimes compacted into a holdfast  

5a. Fronds irregularly dichotomously branched (resulting from some branch segments growing longer than others) . . . . . . . .

. . . . . . .

5b. Fronds dichotomously branched throughout (more or less even in lengths)  

6a. Fronds usually more than 5 mm wide; branched at angles usually less than 45o; margins entire or sometimes with marginal 

proliferations  . . . . 

6b. Fronds narrower, less than 4 mm wide; branched at wide angles (mostly 90o–120o); margins entire 

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

Dictyota concrescens W. R. Taylor

FIGURE 56A,B

Dictyota concrescens W. R. Taylor, 1945:89, pl. 10: fig. 

2; Dawson, 1950a:92; Dawson, 1961b:388; SalcedoMartínez et al., 1988:82; Rocha-Ramírez and SiqueirosBeltrones, 1991:24; Cruz-Ayala et al., 2001:190; Riosmena-Rodríguez et al., 2005:101; Pedroche et al., 

2008:38.

Dictyota friabilis sensu Dawson, 1954d:401, fig. 16a,b; 

Dawson et al., 1960b:6, 14; Dawson, 1961b:388; Dawson, 1962b:190, fig. 50; Dawson, 1966a:12; Dawson, 

1966b:56; Stout and Dreckmann, 1993:7; GonzálezGonzález et al., 1996:146; Silva et al., 1996b:234; Mateo-Cid et al., 2000a:68; Mateo-Cid et al., 2000b:203, 

figs. 31–34, 100; Mateo-Cid et al., 2006:49; Pedroche 

et al., 2008 (in part):43 [non Dictyota friabilis Setchell, 

1926:91 (see De Clerck and Coppejans, 1999:tbl. 1; De 

Clerck, 2003:89)].

Algae decumbent, often forming low-growing mats 

of overlapping, delicate, short, and broad blades, up to 

5 cm long and 6 mm wide; subdichotomously branched, 

with one segment of the dichotomy usually larger and 

longer than the other; blades occasionally attached to 

each other by, usually marginal, rhizoids; base attached to 

substratum by masses of rhizoids. Transection 90–110 μm 

thick. Cortex a single layer of cortical cells, 5–18 μm tall 

by 10–14 μm in diameter. Medulla a single layer of large 

cells, 48–50 μm tall by 60–80 μm in diameter.

Sporangia globose, up to 160 μm in diameter, scattered over thallus. Oogonia scattered on suface in upper 

portions of branches; 90–110 μm tall by 80–85 μm in 

diameter.

Habitat. Creeping, low growing, and loosely

attached; often overgrowing other turf algae; mid to low 

intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Bahía de La Paz; Punta Arena to Cabo Pulmo. 

Pacific coast: Islas San Benito (off Baja California) to 

Bahía Magdalena (Baja California Sur); Rocas Alijos (west 

of Baja California Sur); Michoacán to Oaxaca.

Type Locality. Punta Hughes (vicinity of

Bahía Magdalena), about 5.6 km (3.5 miles) southeast of 

Cabo San Lázaro, Baja California Sur, Mexico.

Remarks. Gulf specimens are in general agreement with Pacific Mexico Dictyota concrescens (Taylor, 

1945); thus, for now, they are referred to this taxon. 

However, it has been noted that the type specimen of 
1 2 4 • 

D. concrescens may be inadequate for clear interpretation (Dawson, 1950a). Although Cribb (1954) observed 

D. concrescens to be similar to another low-growing species, D. friabilis Setchell (1926), the northern Gulf D. 

concrescens differs in the field from D. friabilis, primarily 

in lacking its pale-bluish iridescent color (often with noniridescent longitudinal stripes and/or noniridescent broad 

transverse bands; see De Clerck, 2003:89; De Clerck et al., 

2005:104, fig. 75). Comparative morphological studies 

and molecular and phylogenetic analyses that include type 

locality D. concrescens and Gulf of California specimens 

are needed to clarify its taxonomic status.

Dictyota coriacea (Holmes

Hwang, H. S. Kim et W. J

FIGURE 57

Glossophora coriacea Holmes, 1896:251.

Dictyota coriacea (Holmes) Hwang, H. S. Kim et W. J. 

Lee, 2004:189; De Clerck et al., 2006:1285.

Pachydictyon coriaceum (Holmes) Okamura, 1899a:39, 

pl.1: figs. 31–34; Okamura, 1907:105, pl. 23: figs. 1–6, 

pl. 24: figs. 1–18; Okamura, 1936:165, fig. 84 (1–5); 

Dawson, 1950a:85–86; Dawson, 1950b:67; Dawson, 

1950e:268; Dawson, 1951:52; Dawson, 1959a:18; 

Dawson et al., 1960a:38, pl. 3: figs. 1–2; Dawson et 

al., 1960b:8–11; Kumagae, 1969:91, text figs. 1–4, 

pl. [1]: figs. 1–15; Norris, 1973:5; Abbott and Hollenberg, 1976:209, fig. 173; Norris and Bucher, 1976:4; 

Devinny, 1978:358; Littler and Littler, 1981:149, fig. 

2; L. Aguilar-Rosas, 1982:31; Lewbel et al., 1981:162; 

Pacheco-Ruíz, 1982:70; Mendoza-González and Mateo-Cid, 1985:24; L. Aguilar-Rosas and Pacheco-Ruíz, 

1989:81; Sánchez-Rodríguez et al., 1989:40; R. Aguilar-Rosas and Machado-Galindo, 1990:188; R. Aguilar-Rosas et al., 1990:123; Stewart, 1991:49; R. Aguilar-Rosas and M. Aguilar-Rosas, 1994:518; Mateo-Cid 

and Mendoza-González, 1994a:50; González-González 

et al., 1996:155; Pacheco-Ruíz and Zertuche-González, 

1996b:171; Yoshida, 1998:223, pl. 2–4: fig. D, pl. 2–9: 

fig. C; Mendoza-González et al., 1999:59; Mateo-Cid 

et al., 2000b:205, figs. 41–44, 78, 101; Pacheco-Ruíz et 

al., 2008:203; Pedroche et al., 2008:46.

Glossophorella coriacea (Holmes) Nizamuddin in Nizamuddin and Campbell, 1995:259, fig. 3.

Dictyota johnstonii Setchell et N. L. Gardner, 1924a:730, 

pl. 18: figs. 54–56, pl. 39; Setchell and Gardner, 

FIGURE 56. Dictyota concrescens: Low-growing, relatively broad, subdichotomously branched to subpalmate thalli. A. Isotype specimen 

(W. R. Taylor 34-598). B. Habit (JN-4695).
n u m b e r 9 4 • 1 2 5

1925:653; Dawson, 1944:228 [in part]; Norris, 1973:5; 

Pacheco-Ruíz et al., 2008:203.

Pachydictyon binghamiae (J. Agardh) E. Y. Dawson, 

1950a:84; León-Tejera and González-González, 

1993:497; González-González et al., 1996:155 [non 

Dictyota binghamiae J. Agardh, 1894:72 (see Dawson, 

1950e:267–268; Hollenberg and Abbott, 1966:19[629]; 

Abbott and Hollenberg, 1976:209, fig. 173; De Clerck, 

2003:164, 182)].

Algae erect, olive-brown to dark brown, coriaceous, 

of flat, straplike, subdichotomously branched fronds, up 

to 25 cm tall and to 13 mm wide; multilayered medulla 

and cortex, 400–500 μm thick (in transection); attached 

FIGURE 57. Dictyota coriacea: Habit (JN-5522).
1 2 6 • 

by a fibrous holdfast of rhizoidal cells developed from 

lower portion of the main axes. Branches mostly with one 

of the branch segments longer than the other above the 

dichotomous division (occasionally of more or less uniform length); margins smooth; branch apices rounded, 

with a conspicuous single large apical cell. Cortical cells 

15–24 μm tall by 30–45 μm wide; 1–2 cortical cell layers toward the center of blade; 2–4 (or more) cell layers 

in blade margins of mid to upper portions of the thallus, 

often with more cortical cell layers in blade margins of the 

lower portion. Medulla of large cells, 75–145 μm tall by 

140–195 μm wide: 1–2(–3) cell layers toward the center 

and 2–4 (or more) cell layers near blade margins.

Reproductive structures scattered on both surfaces 

of the upper branches. Sporangia solitary or in groups; 

62–70 μm tall by 45–60 μm in diameter. Dioecious (where 

known). Oogonia aggregated in elliptical to elongate, 

dark-brown to “blackish” sori; 85–117 μm tall by 77–94 

μm in diameter. Antheridia unknown in Gulf material (as 

reported in Japan: in numerous oval, whitish sori).

Habitat. On rocks; mid intertidal to 10 m

depths.

Distribution. Gulf of California: Bahía de

Los Ángeles to Isla San Ildefonso; Cabo San Lucas. Pacific 

coast: Oregon; central California to Bahía Magdalena, 

Baja California Sur; Oaxaca; Panama; China; Taiwan; 

Korea; Japan.

Type Locality. Enoura, “neighborhood of

Tokio [Tokyo]” (Holmes, 1896:149), Kanagawa, Honshu ̄, Japan.

Remarks. Gulf Dictyota coriacea may be 

separated in the field from other Gulf Dictyota species 

by its larger size, thicker fronds, darker color, and relatively coarser texture. Its Gulf distribution appears to be 

restricted to the central Gulf, Las Islas de la Cintura (Midriff Islands), and a few of the southern islands.

There has been some taxonomic confusion regarding 

the name of the Gulf specimens. Initially Dawson (1950a) 

concluded that the Japanese Pachydictyon coriaceum (Holmes) Okamura was a synonym of Californian P. binghamiae

(J. Agardh) Dawson, but he later (Dawson, 1950e) discovered that the type of Dictyota binghamiae J. Agardh was 

a different taxon and those he previously identified as “P. 

binghamiae,” including Gulf material, were P. coriaceum. 

Since then, this interpretation has been followed (e.g., Hollenberg and Abbott, 1966; Abbott and Hollenberg, 1976).

De Clerck (2003) noted that additional research on 

taxa of Dictyotaceae with a multilayered cortex, such as 

Pachydictyon J. Agardh (1894), was needed to clarify their 

relationship with Dictyota. Hwang et al. (2004) studied 

both Japanese and Korean P. coriaceum and, on the basis 

of molecular analyses, concluded them to be a species of 

Dictyota and restricted the distribution of D. coriacea

to the western Pacific. Therefore the Gulf of California 

D. coriacea and the eastern Pacific D. coriacea (southern California; Pacific Mexico) need to be comparatively 

tested with the western Pacific (eastern Asia) D. coriacea

(type locality: Japan) to see if they are the same species as 

currently understood. These studies should include type 

material of Dictyota johnstonii Setchell et N. L. Gardner 

(1924a), currently considered a synonym of D. coriacea. If 

Gulf specimens are found to be a species separate from the 

Japanese type locality D. coriacea, D. johnstonii would be 

the earliest valid name for the Gulf taxon.

Dictyota crenulata J. Agar

FIGURE 58A,B

Dictyota crenulata J. Agardh, 1847:7; Setchell and Gardner, 1924a:730, pl. 18: figs. 50, 51; Setchell and Gardner, 1925:655; Taylor, 1939:8; Dawson, 1944:228; 

Taylor, 1945:90, pl. 10: fig. 1; Dawson, 1950a:88; 

Dawson, 1954a:3; Dawson, 1959a:5, 18; Dawson, 

1961b:388; Brusca and Thomson, 1975:42; McEnroe et al., 1977:184; Huerta-Múzquiz and MendozaGonzález, 1985:46; Salcedo-Martínez et al., 1988:82; 

Santelices, 1989:289, pl. GG: fig. 7; Rocha-Ramírez 

and Siqueiros-Beltrones, 1991:31; Lluch-Cota et 

al., 1993:366; León-Tejera and González-González, 

1993:497; Servière-Zaragoza et al., 1993:482; MateoCid and Mendoza-González, 1994b:43; GonzálezGonzález et al., 1996:294, fig. 16; Anaya-Reyna and 

Riosmena-Rodríguez, 1996:383; Mateo-Cid et al., 

2000a:68; Mateo-Cid et al., 2000b:199, figs. 20, 21, 71, 

96; Cruz-Ayala et al., 2001:190; De Clerck, 2003:145, 

166; Riosmena-Rodríguez et al., 2005:101; Altamirano-Cerecedo and Riosmena-Rodríguez, 2007:575, 

figs. 1–6; Servière-Zaragoza et al., 2007:8; Pedroche et 

al., 2008:39.

Dilophus crenulatus (J. Agardh) Nizamuddin et Gerloff, 

1979:867, pl. 1: figs. 1, 2, pl. 2: figs. 1, 2, pl. 7: figs. 1, 

2 [note: lectotype (LD, Herb. Agardh-49042A) is illustrated in pls. 1, 2, 7].

Algae up to 15(–25) cm tall, of flat, dichotomously 

branched straplike fronds, (1–)2–4(–5.5) mm wide; margins with characteristic, conspicuous minute dentation; 

attached by multicellular rhizoids. Branching usually 

at wide angles; sometimes becoming subdichotomous; 
n u m b e r 9 4 • 1 2 7

branches narrower below and widening upward; surface 

sometimes with numerous proliferations. Transection of 

3–6 cell layers (1–4 medulla: 2 cortical), 90–150 μm thick, 

up to 500 μm thick near base. Cortex a single layer; cortical cells rounded, 20–50 μm in diameter. Medulla single 

layer to multilayered (latter evident in basal portion); 

medullary cells 55–100 μm tall by 35–50 μm wide.

Reproductive structures mostly in sori, scattered over 

branch surface, mostly in upper portions. Sporangia may 

be singular or a few in sorus, rounded, 75–90 μm in diameter. Oogonia in groups of 30–45 (in surface view), 4–6 

(in transection view) in sori; 75–80 μm tall by 66–70 μm 

in diameter.

Habitat. On rocks; low intertidal to shallow

subtidal.

Distribution. Gulf of California: Bahía Concepción to Cabo San Lucas. Pacific coast: Bahía Vizcaíno

to Todos Santos, Baja California Sur; Islas Revillagigedo; 

Jalisco to Oaxaca; Costa Rica; Chile.

Type Locality. San Agustín, Oaxaca, Mexico (Agardh, 1847, “St. Augustin, Stilla Oceanen [Pacific

Ocean]”).

Remarks. Described from southern Pacific 

Mexico, Dictyota crenulata may reach its northern distribution limit in the central Gulf of California (Dawson, 

1950a, 1961b). Altamirano-Cerecedo and RiosmenaRodríguez (2007) found that thalli of Gulf D. crenulata

near its northern limit (central Gulf) were shorter and 

wider than those of the southern Gulf, which were longer 

and narrower. Setchell and Gardner (1924a:730, pl. 18: fig. 

50) and Dawson (1944) observed numerous small surface 

proliferations on specimens of Gulf D. crenulata that were 

developing in situ sporelings, apparently from germination of their spores. More recently, Altamirano-Cerecedo 

FIGURE 58. Dictyota crenulata: A. Habit (JN-4091a). B. Close-up of blades with characteristic dentate margins (JN-4114).
1 2 8 • 

and Riosmena-Rodríguez (2007) quantified surface and 

marginal proliferations.

Dictyota dichotoma var. intricata 

(C. Agardh) Grevil

FIGURE 59A–C

Zonaria dichotoma var. intricata C. Agardh, 1820:134.

Dictyota dichotoma var. intricata (C. Agardh) Greville, 

1830:58; Schnetter et al., 1987:193, figs. 2, 3; De 

Clerck, 1999:176, figs. 23A–C, 24A–E, ApI-12; De 

Clerck, 2003:76, figs. 23A–C [23B, holotype of D. divaricata], 24A–E; Servière-Zaragoza et al., 2007:8.

Dictyota divaricata J. V. Lamouroux, 1809b:43; Taylor, 

1945:90 (as “D. divaricata, prox.”); Dawson, 1950a:91; 

Dawson, 1959a:18; Dawson, 1961b:388; Dawson, 

1961c:407, pl. 11: fig. 2; Dawson, 1966a:12; HuertaMúzquiz and Tirado-Lizárraga, 1970:127; Chávez B., 

1972:268; Pedroche and González-González, 1981:67; 

Schnetter and Bula Meyer, 1982:65, pl. 9: figs. E–G; 

Stewart and Stewart, 1984:141; Huerta-Muzquiz and 

Mendoza-González, 1985:46; Salcedo-Martínez et al., 

1988:82; Sánchez-Rodríguez et al., 1989:40; RochaRamírez and Siqueiros-Beltrones, 1991:31; MateoCid et al., 1993:50; González-González, 1993:443; 

León-Tejera and González-González, 1993:497; 

Servière-Zaragoza et al., 1993:482; Mateo-Cid and 

Mendoza-González, 1994b:34; González-González et 

al., 1996:294; Pacheco-Ruíz and Zertuche-González, 

1996b:171; León-Tejera and González-González, 

2000:328; Mateo-Cid et al., 2000a:68; Mateo-Cid 

et al., 2000b:201, figs. 25–27, 98; Cruz-Ayala et al.,

2001:190; L. Aguilar-Rosas et al., 2002:235; Hernández-Herrera et al., 2005:146; Dreckmann et al., 

2006:154; Mateo-Cid et al., 2006:49; Pacheco-Ruíz et 

al., 2008:203; Pedroche et al., 2008:41.

Algae of erect, flat, dichotomously branched fronds, 

up to 4(–8) cm tall and 1.5–3(–4) mm wide; attached by 

rhizoids. Fronds branched with long and narrow dichotomies, often at wide angles; more or less uniform width 

throughout or narrower at each dichotomy upward 

and narrowest at apices. Transection 90–100 μm thick. 

Medulla and cortex, each a single layer of cells. Cortical 

cells 15–18 μm tall by 12–16 μm wide; medullary cells 

60–75 μm tall by 54–56 μm wide.

Sporangia single on a stalk cell, scattered on both surfaces of frond. Oogonia 90–95 μm tall by 82–86 μm in 

diameter; in sori without an indusium, scattered on upper 

portions of both surfaces. Antheridia unknown in Gulf 

specimens.

Habitat. On rocks or occasionally entangled

with other algae; low intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Cabo San Lucas. Pacific coast: Isla Guadalupe; 

Punta Pequeña, Baja California Sur to Islas Revillagigedo; 

Jalisco to Chipas; Colombia.

Type Locality. Cádiz, Spain

Remarks. Northern Gulf specimens previously called “Dictyota divaricata” are for now referred 

to D. dichotoma var. intricata on the basis of their morphological similarity, until needed comparative culture 

and molecular studies can be done to elucidate their taxonomic status. There still is much taxonomic and nomenclatural confusion regarding the “Dictyota dichotoma

complex,” including D. divaricata. Hörnig and Schnetter 

(1988) concluded western Atlantic “D. divaricata” differed from those of the European Atlantic Dictyota dichotoma-complex and described them as a separate species, 

D. pulchella Hörnig et Schnetter. Further, there is some 

evidence that D. dichotoma (Hudson) J. V. Lamouroux 

(1809a) is restricted to warm-temperate and cold-temperate waters of the eastern Atlantic Ocean and Mediterranean Sea (Schnetter et al., 1987; De Clerck, 2003) and 

subtropical waters of the Canary Islands (Tronholm et al., 

2008), and D. dichotoma var. intricata could be restricted 

to warm-temperate and subtropical regions of the Indian 

Ocean (De Clerck, 2003: 81).

Although some Gulf of California specimens are 

somewhat morphologically similar to D. ceylanica Kützing (1859), that species is apparently restricted to the tropical to subtropical Pacific and Indian oceans (De Clerck, 

2003). Further, De Clerck (2003) has shown D. dichotoma var. intricata can be separated from D. ceylanica by 

its surface proliferations, size of thallus, and size of medullary and cortical cells; this separation is confirmed by 

molecular studies of De Clerck et al. (2001).

Dictyota flabellata (Collins)

Setchell et N. L. Gardn

FIGURE 60

Dilophus flabellatus Collins in Collins, Holden and Setchell, 1901a:No. 834.

Dictyota flabellata (Collins) Setchell et N. L. Gardner, 

1924b:12 [in part, excluding Monterey, California, 
n u m b e r 9 4 • 1 2 9

FIGURE 59. Dictyota dichotoma var. intricata: Specimens representing variation in morphology. A, C. Blade width distinctly narrows at dichotomies upward toward apices (A, JN-4087; C, JN-4094). B. Blade width with more gradual narrowing toward apices (JN-5630).
1 3 0 • 

specimens]; Setchell and Gardner, 1925:652 [in part, 

excluding Monterey, California, specimens], pl. 34: fig. 

3, pl. 35: fig. 7, pl. 36: figs. 13–17; Dawson, 1950a:89; 

Dawson, 1952:431; Dawson, 1959a:10, 18; Dawson et 

al., 1960a:34, pl. 3: fig.3; Dawson, 1961b:388; Dawson, 1966a:12; Huerta-Múzquiz and Tirado-Lizárraga, 

1970:127; Earle, 1972:85; Norris, 1973:5; Brusca and 

Thomson, 1975:42; Abbott and Hollenberg, 1976:207, 

fig. 172; Robertson and Fenical, 1977:1071; McEnroe 

et al., 1977:182; Devinny, 1978:385; Littler and Littler, 1981:151, tbl. 3; figs. 4, 7; Lewbel et al., 1981:165; 

L. Aguilar-Rosas, 1982:31; L. Aguilar-Rosas et al., 

1982:54; R. Aguilar-Rosas, 1982:84; Littler and Arnold, 

1982:309; Pacheco-Ruíz, 1982:70; Stewart and Stewart, 

1984:141; Huerta-Múzquiz and Mendoza-González, 

1985:46; Mendoza-González and Mateo-Cid, 1985:24; 

Mendoza-González and Mateo-Cid, 1986:421; SánchezRodríguez et al., 1989:40; Rocha-Ramírez and SiqueirosBeltrones, 1991:31; Stewart, 1991:48; Martínez-Lozano 

et al., 1991:23; León-Tejera and González-González, 

1993:497; Mateo-Cid et al., 1993:50; Lluch-Cota et 

al., 1993:366; Mateo-Cid and Mendoza-González, 

1994a:50; González-González et al., 1996:146; PachecoRuíz and Zertuche-González, 1996b:171; Anaya-Reyna 

and Riosmena-Rodríguez, 1996:862; Riosmena-Rodríguez et al., 1998:25; L. Aguilar-Rosas et al., 2000:132; 

Mateo-Cid et al., 2000b:202, figs. 28–30, 74, 75; L. Aguilar-Rosas et al., 2002:235; De Clerck, 2003:150; Abbott 

and Huisman, 2003b:182, figs. 21, 22; Abbott and Huisman, 2004:204, fig. 77D; Riosmena-Rodríguez et al., 

2005:101; Mateo-Cid et al., 2006:49, 57; Pacheco-Ruíz 

et al., 2008:203; Pedroche et al., 2008:42.

Dictyota hesperia Setchell et N. L. Gardner, 1924a:731, 

pl. 18: figs. 52, 53; Dawson, 1944:228; Dawson, 

1950a:85, 89.

Dictyota johnstonii sensu Dawson, 1944:228 [in part]; 

Dawson, 1950e:268 [non Dictyota johnstonii Setchell 

et N. L. Gardner, 1924a:730, which is Dictyota coriacea (Holmes) Hwang, H. S. Kim et W. J. Lee, 2004].

Algae of one to several erect, straplike fronds, dichotomously branched up to 7 orders; up to 15 cm tall and 

4–10(–15) mm wide; 120–135 μm thick; with smooth surfaces, entire margins, and rounded branch apices; attached 

by discoid holdfast of densely compacted rhizoids. Cortex 

and medulla each a single cell layer. Cortical cells 12–15 

μm tall by 18–20 μm in diameter; medulla cells 60–90 μm 

tall by 75–76 μm wide.

Reproductive structures in sori, scattered on both 

surfaces of mid to upper portions of fronds. Dioecious. 

Oogonia 90–96 μm tall by 85–90 μm in diameter; in 

groups of up to 30 in sori with indusium. Antheridial sori, 

63–65 μm tall by 250–360 μm wide; antheridia 57–60 μm 

tall by 18–20 μm wide.

Habitat. On rocks or rarely epiphytic on other

algae; mid intertidal to shallow subtidal.

Distribution. Gulf of California: El Tornillal to Cabo Pulmo. Pacific coast: central California to

southern Baja California Sur; Jalisco; Oaxaca; Panama; 

Hawaiian Islands.

Type Locality. La Jolla, San Diego County,

California.

Remarks. The most common species of Dictyota in the northern Gulf of California, D. flabellata has 

been found growing from January to August. Individual 

fronds apparently do not persist throughout the entire season, but rather, successive generations continue to replace 

one another. Dawson (1950a) noted Taylor’s (1945) specimens of “D. flabellata” from Costa Rica were probably 

“D. dichotoma” and “D. flabellata” from Pacific Panama 

were D. coriacea.

Dictyota stolonifera E. Y. Da

FIGURE 61A–C

Dictyota stolonifera E. Y. Dawson, 1962d:392, pl. 5: 

fig. A; González-González et al., 1996:147; De Clerck, 

FIGURE 60. Dictyota flabellata: Habit (JN-3474).
n u m b e r 9 4 • 1 3 1

2003:136, figs. 51A–E, 52A–E; Abbott and Huisman, 

2003b:182; Abbott and Huisman, 2004:206, fig. 78C.

Dilophus stoloniferus (E. Y. Dawson) Schnetter et Bula 

Meyer, 1982:62, pl. 9: figs. H–O.

Algae of 1 to several, subdichotomosly branched, flat 

straplike fronds, up to 6 cm tall and 3–4 mm wide, above 

a short compressed stipe, and attached to substratum by 

distinctive multilayered, stoloniferous growths, up to 500 

μm in diameter. Branches lack proliferations, with smooth 

surface and hair tufts; margins entire; terminal portions 

of similar width as rest of frond (not reduced); rounded 

apices with concentric growth bands (not always evident), 

600–1200 μm wide. Medulla a single cell layer, and 2 or 

more cell layers at margins; some medullary cell walls with 

secondary thickenings. Cortex a single cell layer of small 

pigmented cells.

Reproductive structures apparently scattered. Sporangia 70–100 μm in diameter, solitary or in small groups; 

each borne on a single stalk cell. Oogonia in scattered sori, 

up to 200 μm in diameter. Antheridia not seen.

Habitat. On rocks and tidal platforms and in

tide pools; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco. Pacific coast: Costa Rica; Nicaragua; Isla Gorgona Parque Nacional, Colombia; Hawaiian Islands.

Type Locality. Intertidal; volcanic reef,

Masachapa, Pacific Nicaragua.

Remarks. Dictyota stolonifera can be distinguished in the field from other species of Gulf Dictyota, 

primarily by its simple to divided stoloniferous attachment growths. An eastern Pacific species, it has also been 

reported in the Indian Ocean (Coppejans et al., 2000; De 

Clerck, 2003). Species limits of D. stolonifera need to 

FIGURE 61. Dictyota stolonifera: Several thalli, each with basal stolons. A. JN-3865a. B. JN-4113a. C. EYD-27409.
1 3 2 • 

be tested, as it may include more than one taxon. Also, 

the generic status of D. stolonifera needs to be reinvestigated as the species had also been placed in Dilophus.

Although it is very similar to Rugulopteryx, its sporangia 

are on two-celled stalks, whereas those of D. stolonifera

are reported to be on one-celled stalks.

Dictyota vivesii M. Ho

FIGURE 62

Dictyota vivesii M. Howe, 1911:497, pl. 27; Setchell and 

Gardner, 1925:654; Dawson, 1944:229; Taylor, 1945:89; 

Salcedo-Martínez et al., 1988:82; Stout and Dreckmann, 

1993:7; González-González et al., 1996:147.

Dictyota dichotoma sensu Dawson, 1950a:90; Dawson, 1954d:401, fig. 15; Dawson, 1959a:18; Dawson, 

1961b:388 [in part]; Martínez-Lozano et al., 1991:23; 

Mateo-Cid et al., 2000b:200, figs. 22–24, 72; Mateo-Cid 

et al., 2006:49; Pedroche et al., 2008 (in part):39 [non Dictyota dichotoma (Hudson) J. V. Lamouroux, 1809d:331; 

basionym: Ulva dichotoma Hudson, 1762:476].

Algae of 1 or more erect, straplike to oblong fronds, 

up to 15 cm tall and 5–8 mm wide; branching up to 10 

orders, mostly dichotomously branched in lower portions, 

becoming subdichotomous upward; attached by a fibrous 

holdfast. Branches in mid to upper portions 135–160 μm 

thick; basal portion up to 240 μm thick; mostly with entire 

margins; apices mostly 1–2 mm wide. Medulla cells thin 

walled, 95–250 μm tall by 55–115 μm wide. Cortical cells 

elongated, 19–65 μm tall by 10–30 μm wide.

Sporangia in sori, scattered over frond surfaces. 

Gametangial structures not described by Howe (1911) 

FIGURE 62. Dictyota vivesii: Habit (EYD-10938).
n u m b e r 9 4 • 1 3 3

(Oogonia without indusium, in small groups on surface 

among hairs; oogonium pyriform, 80–93 μm tall and 

60–75 μm in diameter. Antheridial sori 340–360 μm wide, 

antheridia 36–55 μm tall and 18–25 μm in diameter [after 

Mateo-Cid et al., 2000b, as “D. dichotoma”]).

Habitat. On rocks and occasionally in tide

pools; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Bahía de La Paz; San José del Cabo. Pacific 

coast: Baja California; Islas Revillagigedo; Michoacán.

Type Locality. La Paz, Baja California Sur,

Gulf of California.

Remarks. Described from the southern Gulf,

Dictyota vivesii has been confused with “D. dichotoma” 

(e.g., Dawson, 1944, 1950a). Although D. dichotoma

has been extensively reported worldwide in tropical and 

subtropical areas, D. dichotoma sensu stricto is probably 

restricted to the northeast Atlantic and Mediterranean Sea 

(De Clerck, 1999, 2003).

While large specimens of Dictyota vivesii are readily 

recognized, smaller forms can be difficult to separate from 

some specimens of D. flabellata (Dawson, 1950a, as “D.

dichotoma”). In the northern Gulf, D. vivesii is less frequent in occurrence than D. flabellata, and as with other 

Dictyota taxa of the Gulf, it is in need of critical study, 

including molecular and phylogenetic analyses, to elucidate 

its taxonomic status. Some “Dictyota dichotoma” specimens of Rocha-Ramírez and Siqueiros-Beltrones (1991), 

Servière-Zaragoza et al. (1993), Pacheco-Ruíz and Zertuche-González (1996b), L. Aguilar-Rosas et al. (2000), 

and Pacheco-Ruíz et al. (2008) may also belong here.

Padina Adans

FIGURE 54I

Padina Adanson, 1763:13, 586; Silva, 1952:298–299; Papenfuss, 1977:275.

Algae are composed of one to several flat, fanlike or 

cup-shaped blades that sometimes are clumped together, 

basally attached by branched rhizoids compacted into a 

conical holdfast or rhizoids along lower blade margins. 

Blades may be entire, split, or eroded, with conspicuous 

(rarely faint) concentric lines or bands of phaeophycean 

hairs on one or both sides of the blade. Growth is by a continuous row of apical cells within an inrolled margin along 

the terminal edge of blade (herein the upper [dorsal] side 

of the blade is defined as the side that the inrolled margin 

is curled toward, and the back of the roll [opposite side] is 

referred to as the lower [ventral] side). Depending on the 

species, blades are two to many cell layers thick, and some 

vary from few near the inrolled margin to more numerous 

cell layers toward their base. Species may be uncalcified or 

lightly to moderately calcified on one or both sides of the 

blade. Internally, there is a medulla of one to several layers of large colorless cells and a cortex of a single cortical 

layer of pigmented cells on each side of the blade.

Sporangia are scattered in patches or in sori in concentric zones between the concentric bands of hairs. 

Gametophytes are usually dioecious. Oogonia are in sori 

between the hair lines. Antheridia are in sori, across from 

the oogonial sori in monoecious species, and in between 

the concentric zones of hairs in dioecious species.

Remarks. Lewis et al. (1987) found two distinct 

morphologies in a species of Padina. In experimental field 

studies, small, turf-forming, narrow Vaughaniella Børgesen (1950; see Cribb, 1951) (= Dictyerpa Collins, 1901; 

= “Dilophus-like”) thalli were found in the presence of 

fish herbivory. However, in the absence of herbivory the 

Vaughaniella-form of the same individual thallus, through 

a morphogenetic change, rapidly developed directly into the 

fan-shaped Padina. Allender and Kraft (1983) noted creeping “Vaughaniella stages” in some of the Australian species 

of Padina. Culture studies of vegetative propagation in a 

Japanese Padina similarily showed a growth form that initially developed from a single apical cell and then developed 

meristems along its margin (Kajimura, 1993). Thus, some 

species of Padina, but apparently not all, have a morphologically different “Vaughaniella” form in their development, and the presence or absence of this morphogenetic 

growth form may prove helpful in the taxonomy of Padina.

Three species are recorded in the southern Gulf (CruzAyala et al., 2001; Riosmena-Rodríguez et al., 2005; 

Ortuño-Aguirre and Riosmena-Rodríguez, 2007); Padina

concrescens Thivy (in Taylor, 1945), P. sanctae-crucis

Børgesen (1914), and P. ramonribae Ávila-Ortiz et Pedroche (2005).

Six taxa, five species and one variety, are known in the 

northern Gulf of California.

Key to the Species of Padina in the nor

1a. Blades forming low-growing clustered rosettes; calcified, often overlapping and attached to each other and to substratum 

by rhizoids (without a stipe or holdfast). . . . . . . . . . . . .

1b. Blades erect, calcified or noncalcified, with a stipe and holdfast  
1 3 4 • 

Padina caulescens Thivy

FIGURE 63A,B

Padina caulescens Thivy in W. R. Taylor, 1945:99; Dawson, 1957b:11, fig. 3 (type specimen); Dawson, 1959a:5, 

18; Dawson, 1961b:389; Dawson, 1961c:407, pl. 7: fig. 

2; Dawson, 1962b:200, fig. 82; Chávez B., 1980:48; 

Mendoza-González and Mateo-Cid, 1986:421; Mateo-Cid et al., 1993:50; León-Tejera et al., 1993:200; 

González-González, 1993:443; Servière-Zaragoza et 

al., 1993:482; Mendoza-González et al., 1994:110; 

González-González et al., 1996:155, fig. 18; L. Aguilar-Rosas et al., 2000:132; Ávila-Ortiz and Pedroche, 

2005:143, figs. 1 (isotype), 2–5; Hernández-Herrera et 

al., 2005:146; Pedroche et al., 2008:48.

Padina durvillei sensu Mateo-Cid et al., 2000b:208 [in 

part] [non Padina durvillei Bory de Saint-Vincent, 

1827a:591].

Algae of 1 to several, entire to much divided, noncalcified to lightly calcified flabellate blades; up to 20 cm tall 

and to 15 cm wide; arising from a conspicuous flattened 

stipe, up to 2 cm long and to 1.5 mm thick (at lowermost 

portion), above a stupose rhizoidal holdfast. Blades often 

divided by repeated splitting; subdivisions up to 8 cm long 

and 0.5–1.5 cm wide; with zones of hairs about 1 mm 

apart (when present) on lower surface and opposed to hair 

lines on upper surface. Near inrolled margins, blades of 2 

cell layers, up to 45 μm thick; midportions of 6–8 cell layers; lower portion of blade (7–)9–12 cell layers; 220–250 

μm thick at base. Cortical cells 20–25 μm tall by 20–40 

μm wide. Medullary cells in midportion up to 20–30 μm 

tall by 40–64 μm wide; in lowermost portion (16–)20–30 

μm tall by 50–75(–100) μm wide.

Gametophytes dioecious. Oogonia single or paired in 

sori, up to 250 μm wide; in more or less continuous or 

broken bands between hair lines on blade surface. Antheridia in sori; patchy, more or less scattered on blade surface.

Habitat. Low intertidal to shallow subtidal,

down to 7(–9) m depths.

Distribution. Gulf of California: Playa El

Coloradito to Bahía Agua Verde. Pacific coast: Isla María 

Magdalena (Islas Tres Marías), Nayarit; El Salvador; 

Costa Rica.

Type Locality. Isla María Magdalena, Islas

Tres Marías, Nayarit, Pacific Mexico.

Remarks. Mateo-Cid et al. (2000b), without 

comment, listed Padina caulescens as a synonym of P. 

durvillei. Ávila-Ortiz and Pedroche (2005) recognized P. 

caulescens as a distinct species but restricted its distribution to Isla María Magdalena, noting other Pacific coast 

herbarium specimens, previously identified as “P. caulescens,” belonged to other species.

While in general agreement with tropical Pacific 

Mexico P. caulescens sensu stricto, the northern Gulf 

specimens herein referred to P. caulescens have shorter 

stipe-like regions, up to 2 cm in length versus 5–6 cm 

lengths reported by Ávila-Ortiz and Pedroche (2005). Gulf 

of California P. caulescens needs to be further studied and 

molecularly and phylogenetically compared to type locality P. caulescens to test their taxonomic status.

Padina crispata Thivy

FIGURE 64A–C

Padina crispata Thivy in W. R. Taylor, 1945:100; Dawson, 

1957b:10, fig. 2 (type specimens); Dawson, 1959a:4–

6; Dawson, 1961b:389; Dawson, 1962b:200, fig. 82; 

2a. Blades 4–8 cell layers thick in mid to lower portions (2–6 medullary; 2 cortical)  

2b. Blades 8 or more cell layers thick in lower portions (6–16 medullary; 2 cortical).

3a. Blades up to 15 cm tall, usually as wide as or wider than tall; slightly calcified; lower portion of 6–8 cell layers (4–6 medullary; 2 cortical) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. gymnospor

3b. Blades up to 5 cm tall, torn or with divisions narrower than tall; noncalcified to lightly calcified (mostly on upper side); mid 

to lower portion of 4–6 cell layers (2–4 medullary; 2 cortical); short stipe up to 5 mm long  . . . . P. mexicana v

4a. Blades thick (300–450 μm thick), noncalcified, dark-brown; somewhat leathery; lower portion of 8–18 cell layers (6–16 

medullary; 2 cortical)  . . . . . 

4b. Blades thinner (130–250 μm thick), noncalcified to lightly calcified; light-brown to olive-green; lower portion of 7–12 cell 

layers (5–10 medullary; 2 cortical) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5a. Blades calcified on lower side near margins (noncalcified on upper side); with a stipe up to 5 mm long; lower portion of 

blade of 8–10 cell layers (6–8 medullary; 2 cortical); 130–200 μm thick  . . . . .

5b. Blades noncalcified to lightly calcified on both sides; with a long, conspicuous stipe, up to 2 cm long; lower portion of blade 

9–12 cell layers (7–10 medullary; 2 cortical); 220–250 μm thick  . . . . . .
n u m b e r 9 4 • 1 3 5

Huerta-Múzquiz and Tirado-Lizárraga, 1970:127; 

Chávez B., 1972:268; Huerta-Múzquiz and GarzaBarrientos, 1975:7; Chávez B., 1980:47; Schnetter and 

Bula Meyer, 1982:64, pl. 10: fig. H; Huerta-Múzquiz 

and Mendoza-González, 1985:46; L. Aguilar-Rosas 

and R. Aguilar-Rosas, 1993:202; González-González, 

1993:443; León-Tejera and González-González, 

1993:497; Mateo-Cid et al., 1993:50; Stout and Dreckmann, 1993:7; Ávila-Ortiz and Pedroche, 1999:357, 

figs. 1–5; L. Aguilar-Rosas et al., 2000:132; Mateo-Cid 

et al., 2000b:207, figs. 45, 46, 79, 103; López et al., 

2004:10; Ávila-Ortiz and Pedroche, 2005:146, figs. 6 

(isotype), 7–23; Servière-Zaragoza et al., 2007:8; Pedroche et al., 2008:48.

Padina tetrastromatica sensu Setchell and Gardner, 

1930:149 [with a query], pl. 8: fig. 41; Chávez B., 1980:49; 

Dawson, 1961b:389 [with a query]; Servière-Zaragoza 

et al., 1993:482; González-González et al., 1996:156; 

Ávila-Ortiz and Pedroche, 1999:355 [non Padina tetrastromatica Hauck, 1887:43, which is Padina antillarum 

(Kützing) Piccone, 1886:36 (see Wynne, 1998b:271; 

Wynne and De Clerck, 1999:286, figs. 1–10)].

Algae flat, erect flabellate blades, up to 6.0 cm tall and 

to 6 cm wide; uncalcified on upper side of blade, calcified 

on upper portion of lower side of blade; blade margins 

entire to divided or torn, into narrow crispate lobed blades, 

1–4 cm wide; with a distinct stipe 0.5(–1.0) cm long and 

up to 1.0 mm thick; attached by stupose rhizoidal holdfast. 

Hair lines in zones 0.75 to 3.0 mm apart, either opposite 

on both sides of blade or only on upper surface. Outer 

margin near inrolled margins of 2 cell layers, 30–48 μm 

thick, shortly becoming 2–4 cell layers, 65–120 μm thick; 

midportion of 4–6 cell layers, 120–130 μm thick; lower 

portion of 6–8(–9) layers, 160–220 μm thick. Cortical 

cells: upper side 15–25 μm tall by 10–20 μm wide; lower 

side (12.5–)17–25 μm tall by 20–30 μm wide. Medullary 

cells 20–50(–65) μm tall by 45–75(–105) μm wide.

Sporangia 116–160 μm tall by 63–75 in diameter; in 

sori without an indusium; on lower surface of blade; in 

irregular rows or scattered between hair zones. Oogonial 

sori, linear to round, up to 250 μm in diameter, with an 

complete indusium when immature, later indusium only 

partially covering or absent; oogonia up to 69 μm tall by 

33–58 μm in diameter. Antheridial sori up to 690 μm, 

with an indusium; on lower surface in lines or patches; 

antheridia 50–67 μm long by 19–30 μm wide (after ÁvilaOrtiz and Pedroche, 1999, 2005).

Habitat. Intertidal

Distribution. Gulf of California: Playa El

Coloradito; Bahía Concepción to Cabeza Ballena. Pacific 

coast: Punta Banda (Baja California) to Laguna San Ignacio (Baja California Sur); Isla Socorro (Islas Revillagigedo); 

Isla María Madre (Islas Tres Marias); Nayarit to Oaxaca; 

El Salvador; Costa Rica; Panama; Colombia.

Type Locality. Gulfo Dulce, Pacific Costa

Rica.

Remarks. Ávila-Ortiz and Pedroche (1999) 

found Pacific Mexico specimens that had been identified as “Padina tetrastromatica” were actually P. crispata. Their Pacific Mexico specimens were calcified along 

the inner side of the inrolled margins, whereas Dawson 

(1957b) noted that the blades of Costa Rican Padina crispata had noncalcified upper surfaces and chalky, relatively 

FIGURE 63. Padina caulescens: A. Habit (JN-4748). B. Clump of 

four blades; note the flattened, relatively long stipe of each blade 

(JN-4748).
1 3 6 • 

heavy calcified lower (inner side) surfaces. The taxonomic 

or ecological significance of this difference in calcification 

needs to be investigated. Although Thivy (in Taylor, 1945) 

originally described oogonia with an indusium, MateoCid et al. (2000b) noted their specimens were without an 

indusium, and illustrations of Ávila-Ortiz and Pedroche 

(2005:figs. 20, 21) showed oogonial sori without and with 

partial and complete indusia. A Gulf specimen, noted as 

not being in good condition (Chávez B., 1980, as “P. tetrastromatica”), may also belong here.

Padina durvillei Bory de Saint-Vincen

FIGURE 65

Padina durvillei Bory de Saint-Vincent, 1827a:591, as “P. 

durvillaei”; Bory de Saint-Vincent, 1827b: Atlas, pl. 

21: fig. 1A–C; Bory de Saint-Vincent, 1828:147; Farlow, 1902:91; Howe, 1911:497; Setchell and Gardner, 

1924a:729; Setchell and Gardner, 1925:661, pl. 93; 

Setchell and Gardner, 1930:150; Dawson, 1944:230; 

Taylor, 1945:101; Dawson, 1951:52; Dawson, 1952:431; 

Dawson, 1957b:11; Dawson, 1959a:19; Dawson et 

al., 1960a:38, pl. 5: fig. 3; Dawson, 1961b:389; Dawson et al., 1964:22, pl. 18: fig. A; Dawson, 1966a:11; 

Chávez B., 1972:268; Hommersand, 1972:70; Norris, 1973:5; Brusca and Thomson, 1975:42; HuertaMúzquiz, 1978:337; Chávez-B., 1980:47; Pedroche 

and González-González, 1981:65; Littler and Littler, 

1981:151; Schnetter and Bula Meyer, 1982:65, pl. 10: 

fig. I; Stewart and Stewart, 1984:141; Huerta-Múzquiz 

and Mendoza-González, 1985:46; Mendoza-González 

and Mateo-Cid, 1985:24; Norris, 1985:213, fig. 16.6; 

Mendoza-González and Mateo-Cid, 1986:421; SalcedoMartínez et al., 1988:82; Sánchez-Rodríguez et al., 

1989:41; Dreckmann et al., 1990:27; Rocha-Ramírez 

and Siqueiros-Beltrones, 1991:31; Martínez-Lozano 

et al., 1991:23; Mateo-Cid and Mendoza-González, 

1991:24; Mateo-Cid et al., 1993:50; León-Tejera et 

al., 1993:200; González-González, 1993:443; Stout 

FIGURE 64. Padina crispata (EYD-20957): A. Habit, showing typically divided, narrow crispate lobes. B. Upper portion of blade calcified on 

the lower side. C. Noncalcified upper side of blade.
n u m b e r 9 4 • 1 3 7

and Dreckmann, 1993:8; León-Tejera and GonzálezGonzález, 1993:497; Servière-Zaragoza et al., 1993:482; 

Mateo-Cid and Mendoza-González, 1994a:50; Mendoza-González et al., 1994:110; González-González et 

al., 1996:156; Pacheco-Ruíz and Zertuche-González, 

1996b:171; Anaya-Reyna and Riosmena-Rodríguez, 

1996:862; Riosmena-Rodríguez et al., 1998:45; L. Aguilar-Rosas et al., 2000:132; Mateo-Cid et al., 2000a:68; 

Mateo-Cid et al., 2000b:208 [in part, not including P. 

caulescens], figs. 48–50, 80, 81, 104; L. Aguilar-Rosas 

et al., 2002:235; López et al., 2004:10; Ávila-Ortiz and 

Pedroche, 2005:152, figs. 24 [watercolored illustration 

of type], 25–36; Hernández-Herrera et al., 2005:146; 

Riosmena-Rodríguez et al., 2005:102; Mateo-Cid et al., 

2006:49, 58; Servière-Zaragoza et al., 2007:8; PachecoRuíz et al., 2008:203; Pedroche et al., 2008:50.

Algae of 1 to several, flabellate (fanlike) blades, up to 

25(–40) cm tall and to 20 cm wide, dark-brown, coriaceous; above a short stipe, up to 1.0 cm long and 0.1–0.8 

mm wide, arising from stupose rhizoidal holdfast. Blades 

not calcified; margins entire or divided, with divisions 

becoming laciniate, reniform, or flabellate; surface with 

conspicuous concentric hair zones. Inrolled margins of 

FIGURE 65. Padina durvillei: Leathery, flabellate blades with alternating concentric bands of hairs and reproductive structures (US Alg. Coll.-

163415).
1 3 8 • 

blades of 2 cell layers, below becoming 6–7 cell layers and 

up to 100–180(–230) μm thick; midportion of 8(–10) cell 

layers, (160–)230–290 μm thick; lower portion of 10–18 

layers, 300–450 μm thick. Cortical cells (12–)15–30 μm 

tall by 15–37 μm wide. Medullary cells 21–40 μm tall by 

100–180 μm wide.

Sporangia ovoid, 100–135(–150) μm long and (45–)

70–90 μm in diameter; sori with indusium (when immature) in irregular lines near margins, becoming scattered 

and in patches in mid to lower portions of blade on both 

sides. Oogonia 140 μm tall by 90 μm in diameter, with 

wall to 12 μm thick, in sori with or without indusium. 

Antheridia 50 μm tall by 17.5 μm in diameter; in sori in 

irregular lines and patches on both surfaces (Ávila-Ortiz 

and Pedroche, 2005).

Habitat. On rocks; mid intertidal to shallow

subtidal, to 7.5 m depths.

Distribution. Gulf of California: Puerto

Peñasco to Cabo San Lucas; Mazatlán, Sinaloa. Pacific 

coast: Isla Guadalupe; Isla Cedros; Isla Socorro (Islas 

Revillagigedo); Punta Rosalía, Baja California to Oaxaca; 

Costa Rica; Panama; Colombia; Galápagos Islands; Ecuador; Peru.

Type Locality. Concepcíon, Chile (Bory de

Saint-Vincent, 1827a) (note that Dawson et al. [1964:22] 

suspected this locality may be incorrect, stating that “from 

the present known distribution of the species it is questionable whether the alga was taken from such cool southern waters”).

Remarks. Padina durvillei is generally considered a eastern Pacific species, but it has also been 

reported in Africa (Price et al., 1978; Lawson and John, 

1987; Stegenga et al., 1997). Although the species name 

has been traditionally spelled “durvillaei,” the spelling 

was corrected to “durvillei” by Price et al. (1978:139) in 

accordance with the Botanical Code (see McNeill et al., 

2006:Art. 60.11, Rec. 60C.1). Oogonial sori in Padina 

durvillei have been reported with an indusium (e.g., 

Mateo-Cid et al., 2000b) and without an indusium (ÁvilaOrtiz and Pedroche, 2005).

In terms of algal biomass, Padina durvillei is second 

only to species of Sargassum as the most abundant brown 

alga in the northern Gulf. In winter, as observed in Puerto 

Peñasco, there is new settlement and rapid development of 

new recruits of P. durvillei. It also appears there is annual 

formation of new blades from a perennial base. Development of new blades begins in late fall and is most apparent in December, and maximum size is reached in January 

and February. During the spring and into the summer, the 

blades become split and divided, and epiphytes begin colonizing them. From July to October the blades show signs 

of senescence, becoming deeply split, often eroded, and 

with the densest growth of epiphytes.

Padina gymnospora (Kützing) Sonder

FIGURE 66

Zonaria gymnospora Kützing, 1859:29, pl. 71: fig. 2 [see 

Allender and Kraft, 1983:86, fig. 7E (holotype picture)].

Padina gymnospora (Kützing) Sonder, 1871:47; Allender 

and Kraft, 1983:89; Mendoza-González and MateoCid, 1986:421; Mateo-Cid et al., 1993:50; GonzálezGonzález, 1993:443; León-Tejera and GonzálezGonzález, 1993:498; León-Tejera et al., 1993:200; 

Servière-Zaragoza et al., 1993:482; González-González 

et al., 1996:297, 340: fig. 19, 378; L. Aguilar-Rosas 

et al., 2000:132; Mateo-Cid et al., 2000a:68; MateoCid et al., 2000b:210, figs. 51–53, 82, 107; Abbott and 

Huisman, 2004:215, fig. 82A–D; Hernández-Herrera 

et al., 2005:146 [non Padina gymnospora sensu Vickers, 1905:58; Børgesen, 1914:46, fig. 155; Taylor, 

1960:237; Earle, 1969:165; which is Padina boergesenii

Allender et Kraft, 1983:87].

Padina vickersiae Hoyt in Howe, 1920:595; Taylor, 1960:236, pl. 34: fig. 1 [see Allender and Kraft, 

1983:89].

Algae fanlike blades, up to 15 cm tall and 5–15 cm wide, 

usually as wide as or wider than long, yellowish-brown, 

simple with entire margins, ruffled; sometimes divided or 

torn into several lobes; calcified on both surfaces; above a 

short, stupose holdfast. Blades near inrolled margins of 2 

cell layers, 30–50 μm thick; midportion of 4–6 cell layers, 

75–80 μm thick; lower portion of 6–8 cell layers, 180–200 

μm thick. Cortical cells 18–30 μm tall by 18–30 μm wide. 

Medullary cells 21–39 μm tall by (75–)100–180 μm wide. 

Hairs in concentric lines on both sides of blades.

Sporangia in sori with an indusium; sporangium up to 

90 μm in diameter; solitary or grouped in noncontinuous 

bands between hair lines on both blade surfaces. Oogonia 

in sori; oogonium 80–100 μm tall by 65–90 μm in diameter. Antheridia in sori up 240 μm wide.

Habitat. Mid intertidal to shallow subtidal

Distribution. Gulf of California: Playa El

Coloradito to Estero Tastiota; Bahía Concepción to Cabo 

Pulmo. Pacific coast: Nayarit to Jalisco; Oaxaca; Hawaiian Islands; China; Japan.

Type Locality. St. Thomas, West Indies (U.S.

Virgin Islands).

Remarks. The Gulf of California specimens are tentatively referred to Padina gymnospora. 
n u m b e r 9 4 • 1 3 9

Ávila-Ortiz and Pedroche (2005) found that herbarium 

specimens of tropical Pacific Mexico “P. gymnospora” 

they examined were either P. crispata or P. mexicana

and noted that P. gymnospora should be applied to only 

material from the Atlantic. Thus, the Gulf of California “P. gymnospora” is in need of critical comparative 

studies, including molecular analyses with Caribbean P. 

gymnospora (type locality), to test their taxonomic and 

phylogenetic relationships.

Padina mexicana E. Y. Dawson var. mexi

FIGURE 67

Padina mexicana E. Y. Dawson, 1944:231, pl. 52: fig. 

2; Dawson, 1959a:19; Dawson, 1961b:389; Dawson, 

1966a:11; Chávez B., 1980:48; Huerta-Múzquiz and 

Mendoza-González, 1985:46; Mateo-Cid and Mendoza-González, 1991:24; Rocha-Ramírez and SiqueirosBeltrones, 1991:31; Mateo-Cid et al., 1993:50; Servière-Zaragoza et al., 1993:482; Mendoza-González 

et al., 1994:110; Mateo-Cid and Mendoza-González, 

1994b:43; González-González et al., 1996:156; Pacheco-Ruíz and Zertuche-González, 1996b:171; AnayaReyna and Riosmena-Rodríguez, 1996:862; RiosmenaRodríguez et al., 1998:45; Ávila-Ortiz and Pedroche, 

1999:357; L. Aguilar-Rosas et al., 2000:132; MateoCid et al., 2000a:68; Mateo-Cid et al., 2000b:211, 

figs. 54–56, 83–86, 106; Cruz-Ayala et al., 2001:190; 

Ávila-Ortiz, 2003:70, figs. 1–9: López et al., 2004:10; 

Ávila-Ortiz and Pedroche, 2005:158, figs. 37 (isotype), 

38–41; Riosmena-Rodríguez et al., 2005:102; MateoCid et al., 2006:49; Pacheco-Ruíz et al., 2008:203; Pedroche et al., 2008:54.

Algae low-growing, more or less prostrate, fan-shaped 

blades, light chalky whitish to grey-brown; blades often 

overlapping, forming clustered rosettes, attached to each 

other and to substratum by multicellular rhizoids along the 

lower portions of blades (without a stipe). Blades mostly 

entire, 1.5–3(–5) cm tall and 1.5–5.0 cm wide; moderately 

calcified (mostly on lower surface); concentric bands of 

hairs alternate with calcification. Blades near inrolled margin of 2 cell layers, 35–80(–100) μm thick; midportion of 

4 cell layers, 90–170 μm thick; lower portions of 5–6 cell 

layers, 150–200(–280) μm thick. Cortical cells 17–33 μm 

FIGURE 66. Padina gymnospora: Blades with ruffled margins and concentric bands of hairs between light calcification (JN-3425b).
1 4 0 • 

tall by 22–36 μm wide; medullary cells 19–37 μm by 

24–48(–56) μm wide.

Sporangia in sori covered with a thin indusium; 

90–120 μm long by 50–90 μm in diameter; patchy or 

discontinuous in irregular lines between the concentric 

bands of calcium carbonate only on the upper side. Oogonia 92–96 μm tall (Mateo-Cid et al., 2000b) by 73 μm in 

diameter; in sori with indusium (when immature), in concentric lines (Ávila-Ortiz and Pedroche, 2005). Antheridia 

not known.

Habitat. On sand-covered rocks and tidal

platforms; mid intertidal to shallow subtidal, to 3.3 m 

depth.

Distribution. Gulf of California: Puerto

Peñasco to Cabo Pulmo. Pacific coast: Todos Santos, Baja 

California Sur; Sinaloa to Oaxaca.

Type Locality. Isla Turners, off southeast

Isla Tiburón, Las Islas de la Cintura, Gulf of California.

Remarks. Padina mexicana, apparently an 

annual species, appears in spring (beginning of the warmwater season) and is found through summer, usually until 

August. It is distinguished from other Gulf species of 

Padina by its smaller size, more or less prostrate, openrosette clusters of chalky calcified fronds that are attached 

by rhizoids along the base of blade (i.e., lacks a stipe or 

holdfast). Although generally considered a Gulf of California to tropical Pacific Mexico species, it has also been 

reported from West Africa (Price et al., 1978; John et al.,

2003, 2004).

Padina mexicana var. erecta Ávila-Ortiz

Padina mexicana var. erecta Ávila-Ortiz, 2003:70, figs. 

10–16; Ávila-Ortiz and Pedroche, 2005:161, figs. 42–

45; Pedroche et al., 2008:55.

Algae erect, up 5 cm tall, with flat blades usually 

divided, uncalcified on both surfaces or calcified on lower 

surface; with a distinct stipe about 5 mm in length. Blade 

near inrolled margin of 2 cell layers, 40–80 μm thick; mid 

to low portions of blade of 4–6 cell layers, 80–120 μm 

thick in middle, and 130–220 μm thick in lower to basal 

FIGURE 67. Padina mexicana var. mexicana: Blades adhering to each other to form rosettelike clusters (photo of living specimen viewed from 

above, Puerto Peñasco, Sonora, Mexico).
n u m b e r 9 4 • 1 4 1

portions. Cortical cells 16–25 tall by 24–36 μm wide on 

upper surface and 24–30 μm tall by 24–36 μm wide on 

lower surface.

Sporangial sori with indusium; mostly on lower surface in continuous bands, occasionally on upper surface 

in irregular lines, alternating with zones of calcium carbonate, and occasionally in patches or discontinuous in 

irregular lines on lower surface; sporangia 80–130 μm tall 

by 50–80 μm in diameter. Gametangial thalli dioecious. 

Oogonial sori with an indusium; in regular, discontinuous, or irregular bands on both surfaces; oogonia 120 μm 

tall by 50 μm in diameter. Antheridial sori with indusium; 

on lower surface in discontinuous lines; antheridia 52 μm 

tall by 27 μm in diameter.

Habitiat. On rocks; intertidal

Distribution. Gulf of California: Puerto

Peñasco; Bahía Concepción; Bahía de La Paz to Cabo San 

Lucas. Pacific coast: Todos Santos, Baja California Sur; 

Sinaloa to Oaxaca.

Type Locality. Puerto Vicente Guerrero,

Guerrero, Pacific Mexico.

Remarks. In the field, Padina mexicana var. 

erecta is distinguished from P. mexicana var. mexicana by 

its erect habit, evident stipe, sporangia on both sides of 

blade, and blades that are with or without calcification 

(Ávila-Ortiz, 2003).

Rugulopteryx De Clerck et Coppejans

FIGURE 54A, D

Rugulopteryx De Clerck et Coppejans in De Clerck, Leliaert, Verbruggen, Lane, De Paula, Payo, and Coppejans 

2006:1286.

Algae are erect, of strap-like flattened fronds that are 

dichotomously to irregularly branched, and attached by 

rhizoids restricted to basal portions of thallus, some with 

basal stoloniferous holdfasts. Fronds have entire margins 

and rounded apices, with surfaces that may be undulate 

or rugose, or flat and smooth; surface proliferations are 

found in some species. Medulla is composed of large cubical to rectangular, colorless cells. Central portions of the 

medulla are one cell layer thick, and two to eight cell layers 

at the branch margins and stoloniferous holdfasts. Cortex 

is composed of a single cell layer (occasionally two-cell 

layers in basal portion) of small cells. Each cortical cell 

with numerous discoid chloroplasts. Growth is by a single, 

protruding or emarginate, apical cell. Phaeophycean hairs 

are scattered in tufts.

Reproductive structures are scattered, and either 

within concavities of the frond surfaces, or protruding 

from the frond surface. Sporangia are ovoid to pyriform 

and may be solitary, in groups, or in sori. Borne on twocelled basal stalks, the sporangia are surrounded by an 

inconspicuous involucrum, and produce four aplanospores (tetraspores). Oogonia are borne on a single stalk 

cell, in sori or small groups. Antheridia borne in many 

tiered locules that are in whitish sori, surrounded by 3–5 

rows of pigmented multicellular paraphyses.

One species of Rugulopteryx is recognized in the 

northern Gulf of California.

Rugulopteryx okamurae (E. Y. Daws

I.-K. Hwang, W. J. Lee et H.-

FIGURE 68

Dilophus okamurae E. Y. Dawson, 1950a:86, as “D. okamurai”; Dawson, 1961b:388; Hommersand, 1972:70; 

Tseng, 1983:196. pl. 99: fig. 1; Kajimura, 1992:95–97, 

figs. 1–17; González-González et al., 1996:147; Yoshida, 1998:220, pl. 2-8: fig. B, pl. 2-9: figs. A, B; De 

Clerck, 2003:180

Rugulopteryx okamurae (E. Y. Dawson) I.-K. Hwang, 

W. J. Lee et H.-S. Kim in Hwang, Lee, Kim and De 

Clerck 2009:5, figs. 1–49.

Dictyota okamurae (E. Y. Dawson) Hörnig, Schnetter 

et Prud’homme van Reine, 1992a:54; Hörnig et al., 

1993:170; De Clerck, 2003:174; Pedroche et al., 

2008:44.

Dictyota marginata Okamura, 1913:33, pl. 108: fig. 9, pl. 

109: figs. 1–8, nom. illeg., i.e., a later homonym of Dictyota marginata (C. Agardh) Greville, 1830:xliii, which 

= Stoechospermum polypodioides (J. V. Lamouroux) 

J. Agardh, 1848:100 (see De Clerck, 2003:173)].

Dilophus marginatus (Okamura) Okamura, 1915:154 

[Corrigenda], nom. illeg.; Okamura, 1936:167, fig. 

85 (1–3) [a later homonym of Dilophus marginatus

J. Agardh, 1894:91, which is now Rugulopteryx marginata (J. Agardh) De Clerck et Coppejans in De Clerck 

et al., 2006:1286].

Algae of flattened, straplike fronds, up to 5 cm tall, 

and dichotomously to subdichotomously branched; 

attached below by rhizoids. Branches up to 5 mm wide, 

with segments of equal length, or with one segment of a 

pair frequently longer than the other; apices are rounded. 

Medulla of large colorless cells; one-cell layer thick 

toward center (seen in transection) and 2 or more cell 
1 4 2 • 

layers in the margins. Cortex of small cells, of one-cell 

layer, throughout.

Reproduction not seen in Gulf material.

Habitat. On rocks or occasionally entangled

with other algae; low intertidal to shallow subtidal; also 

dredged from 19 m depths.

Distribution. Gulf of California: Puerto

Peñasco; Cabeza Ballena to Bahía de San Lucas. Western 

Pacific coast: China (Tseng, 1983); Taiwan (Lewis and 

Norris, 1987); Japan (Yoshida, 1998); Korea (Lee and 

Kang, 2001).

Type Locality. Enoshima, Kanagawa Prefecture (formerly Sagami Province), Honshu ̄, Japan

Remarks. On the basis of their habit and 

morphological similarities, the northern Gulf specimens 

are referred to Rugulopteryx okamurae. The specific and 

generic status of the northern Gulf material of R. okamurae needs to be verified. Hwang et al. (2009) noted R. 

okamurae to be a subtropical to temperate western Pacific 

species, and did not include Gulf of California material, 

or the type (Dictyota marginata Okamura, 1913) or type 

locality material of Dilophus okamurae in their analyses. 

It would be interesting to study the life history of Gulf R. 

okamurae in culture and using molecular analyses to compare them to Japanese R. okamurae (type locality) and 

south Korean R. okamurae (Hwang et al., 2009) to test 

their phylogenetic relationships.

Although the species epithet has been spelled with 

different endings, the correct ending is –e (McNeill et al., 

2006:Art. 60, Rec. 60C.1a), thus R. okamurae. A novel 

form of vegetative propagation by the development of 

proliferous initials along the surface and margins of its 

fronds was shown in R. okamurae from Japan (Kajimura, 

1992, as Dilophus okamurae).

Spatoglossum Kützing

FIGURE 54G,H

Spatoglossum Kützing, 1843:339.

Algae are erect, flattened fronds, up to 5 cm wide, 

that are subdichotomously to subpalmately divided and 

attached by a fibrous holdfast. The flat fronds are often 

expanded upward and are irregularly to more or less laterally branched up to four orders, with entire, undulate, 

or slightly dentate margins. A midrib is entirely lacking in 

most or only partly extended from the holdfast into the 

lower basal portion. Growth is from marginal meristems 

of small groups of apical cells. Thalli are multilayered, up 

to 10 cell layers in transection, with a medulla and cortex, not clearly demarcated. Medulla is composed of two 

to several rows of colorless large cells that become progressively smaller outward to a cortical layer of smaller 

pigmented cells. Hair tufts arise from depressions that are 

scattered on both surfaces.

Sporangia are scattered over both surfaces of the thallus; borne either completely above or partially embedded 

and partly protruding, or entirely embedded on the surface 

and, produce four aplanospores (tetraspores). Oogonia 

FIGURE 68. Rugulopteryx okamurae: A. Habit (JN-4086). B. Transection; note cortex of a single cell layer and medulla of more than one cell 

layer near margin (JN-3724).
n u m b e r 9 4 • 1 4 3

are solitary or in groups and are usually raised and spread 

over the thallus surfaces. Antheridia are borne in small, 

partially or entirely embedded sori that are scattered over 

both surfaces.

Remarks. Sasaki et al. (1999) used presence 

or absence of sulfuric acid to separate morphologically 

similar species of Spatoglossum and suggested this was a 

useful taxonomic character (Sasaki et al., 2004, 2005).

One species, Spatoglossum howellii Setchell et N. L. 

Gardner (1937), has been reported in the southern Gulf 

of California (Hommersand, 1972). Two species are currently known in the northern Gulf of California.

Key to the Species of Spatoglossum in the n

1a. Narrow lanceolate blades, up to 17 mm wide, with sparsely dentate margins; surface cells irregularly arranged; oogonia 

and antheridia embedded in cortical layer  . . . . . . .

1b. Narrow blades expanding to 2(–3.5) cm wide, with entire margins; surface cells arranged in longitudinal rows; oogonia 

and antheridia projecting above surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. subflabellatum

Spatoglossum lanceolatum E. Y. Daw

FIGURE 69

Spatoglossum lanceolatum E. Y. Dawson, 1954e:328, 

pl. 1: figs. 1, 2, pl. 6; Dawson, 1961b:389; GonzálezGonzález et al., 1996:164; Pedroche et al., 2008:56.

Algae of few to several flat, 3–4 times irregularly, subdichotomously to subpalmately divided fronds, up to 28 

cm tall, above a narrow slightly stupose stipe, up to 10 mm 

long and 1.0–2.0 mm wide; attached below by a discoid 

stupose cushion. Branches narrower near dichotomies, 

widening upward, becoming more or less lanceolate, to 

(3–)5–10(–17) mm broad; margins with irregularly spaced 

dentations; blades 200–250 μm thick. Medulla of (3–)5–7 

irregular layers of large cells, 40–65 μm in diameter; subcortex of partial layers of similarly shaped smaller cells. 

Cortex of quadrangular cells, not projecting, outer walls 

forming a smooth surface; in surface view, cells of irregular 

shape and irregularly arranged (not in longitudinal rows).

Apparently monoecious. Oogonia ovate to hemispherical (or sometimes more or less angular), up to about 

70 μm in diameter; either solitary, in pairs, or in groups 

of 3–4; embedded within cortical layer (barely projecting 

above its suface). Antheridia in sori, 100–200 μm in diameter, embedded in cortical layer and not projecting above 

surface; composed of densely packed cells, most 8–9 μm in 

diameter; formed by periclinal cell divisions into irregular 

tiers within vertical columns, 30–40 μm tall.

Habitat. On rocks, in tide pools, and on tidal

platforms; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Ensenada de San Francisco.

Type Locality. North shore of Ensenada de

San Francisco, near Punta de Las Cuevas, Sonora, Gulf of 

California.

Remarks. Spatoglossum lanceolatum, a Gulf 

endemic species, is distinguished from other species of 

Spatoglossum by its narrower fronds with sparsely dentate margins, irregularly arranged surface cells, and the 

embedded (not projecting above blade surface) reproductive structures.

Spatoglossum subflabellatum E. Y. Da

FIGURE 70

Spatoglossum subflabellatum E. Y. Dawson, 1954e:326, 

pl. 1: figs. 3–5, pl. 5: fig. 2; Dawson, 1961b:389; Dawson, 

1966a:11; Dawson, 1966b:56; Hommersand, 1972:70; 

Espinoza-Ávalos, 1993:333; González-González et al., 

FIGURE 69. Spatoglossum lanceolatum: Part of the type specimen; 

note the irregularly and distantly spaced dentations (EYD-1966, 

after Dawson, 1954e: pl. 6).
1 4 4 • 

1996:164; Mateo-Cid et al., 2006:49; Pedroche et al., 

2008:56.

Spatoglossum cf. schroederi sensu Dawson, 1959a:19, 

fig.1; Dawson, 1961b:389 [non Spatoglossum schroederi (C. Agardh) Kützing, 1859:21, pl. 51].

Algae of few to several flat, narrow, irregularly 

divided or deeply split subflabellate to subpalmate fronds, 

up to 25 cm tall, arising from a stupose basal cushion 

(up to 2 cm tall). Fronds cuneate and narrow at base, 

becoming wider distally to 2(–3.5) cm broad and up to 

180 μm thick; with entire margins. Medulla of 2 irregular 

layers of large thin-walled cells, 40–70 μm in diameter; 

subcortex of partial layers of smaller, similarly shaped 

cells. Cortex a single layer of pigmented cells, 20–30 μm 

in diameter, with projecting rounded outer walls; in surface view, cells arranged in longitudinal rows and about 

twice as long as wide.

Sporangia unknown. Dioecious. Oogonia either solitary or in rows of 2–4 elevated on thallus surface; ovate, 

up to 70 μm long, borne on a modified cortical cell. 

Antheridia elevated in sori, 150–400 μm in diameter, scattered on thallus surface in groups; about 50 μm tall by 25 

μm wide; each antheridium about 2.5 μm in diameter in 

vertical and horizontal rows.

Habitat. On rocks and in crevices and tide

pools; mid intertidal to shallow subtidal, down to 6 m 

depths.

Distribution. Gulf of California: Puerto

Peñasco to Punta Los Frailes.

Type Locality. Dredged 11–18 m depths,

near Punta de Las Cuevas, Ensenada de San Francisco, 

Sonora, Gulf of California.

Remarks. Spatoglossum subflabellatum is a 

Gulf endemic. The more recent collections of Dawson 

(1966a, 1966b) that he identified as S. subflabellatum

included specimens that were much broader than those 

originally described (Dawson, 1954e). Thus, this broadly 

defined species concept is in need of critical study and 

may include more than one species. Further, Dawson 

(1966b:56) observed that the Isla San Pedro Nolasco specimens were intermediate morphologies between S. subflabellatum and S. howellii Setchell et N. L. Gardner (1937) 

and suggested that S. subflabellatum probably reflects 

“progressive speciation in the specialized environment of 

the northern Gulf of California.”

Desmares

Desmarestiales Setchell et N. L. Gardner, 1925:554.

Algae are heteromorphic, with large sporophytes that 

are cylindrical, compressed, flattened, or bladelike, and 

may be entire or branched. Growth is trichothallic-like 

and initiated by individual filaments with bidirectional 

intercalary divisions. The resulting structure is pseudoparenchymatous, with a cortex developed from the descending rhizoidal filaments growing from the bases of lateral 

hairs. Cells have several discoid plastids and no pyrenoids.

Life histories, where known, are hetermorphic. The 

large sporophytes develop unilocular sporangia that produce zoospores. The gametophytes are microscopic filaments, and sexual reproduction is oogamous.

There is one family represented in the northern Gulf 

of California.

Desmares

Desmarestiaceae (Thuret) Kjellman, 1880:10.

Sporophytes are erect cylindrical to flattened thalli, 

which are sparingly to abundantly branched or rarely 

entire and attached to the substratum by a discoid holdfast. Branching is opposite or alternate and generally pinnate. A pseudoparenchymatous cortex surrounds a single 

FIGURE 70. Spatoglossum subflabellatum: Habit (JN-5902).
n u m b e r 9 4 • 1 4 5

filament, which initiates axial trichothallic-like growth of 

the bidirectional intercalary meristem. Some species have 

persistent or deciduous hairs on their margins that are 

irregularly scattered or in transverse whorls.

Unilocular sporangia are solitary or grouped in sori 

and produced by transformation of superficial cells of the 

thallus or of assimilatory filaments. Gametophytes are 

monoecious or dioecious, microscopic filamentous algae. 

Oogonia are usually produced on the filaments of a separate thallus, and each produces a single egg. Antheridia 

develop on filaments, and each antheridium produces a 

single biflagellate antherizoid.

One genus occurs in the northern Gulf of California, with its species apparently restricted to the Islas de 

la Cintura.

Desmarestia J. V. Lamou

Desmarestia J. V. Lamouroux, 1813:43.

Sporophytic thalli are macroscopic, erect, with a 

single cylindrical or compressed main axis that is ligulate 

to broadly bladelike and usually pinnately or oppositely 

branched in most species or are simple and unbranched in 

a few, and attached below by a discoid holdfast. If the thallus is branched, the branches are cylindrical, compressed 

or flat, with opposite or alternate branchlets. Trichothallic growth occurs at the apices of the axis and branches. 

The main axial filament is surrounded by a thick medulla 

of large, colorless cells developed by downward growing 

hyphae. The cortical layer is composed of small cells that 

become progressively smaller toward the thallus surface. 

Cells have lenticulate plastids and vacuoles (in many species these contain sulfuric acid) and lack pyrenoids. Phaeophycean hairs are absent.

Unilocular sporangia are isolated and scattered over 

the surface cells, developing from the transformation of 

cortical cells, or on stalk cells in small sori along with 

one- to two-celled paraphyses. Gametophytic thalli, little 

known in nature, are microscopic, irregularly branched filaments, and endozooic or endophytic. Gametophytes may 

be monoecious or dioecious, and are dimorphic in cell size 

of the filaments. Elongated oogonia develop on the cells 

of filamentous thalli of larger diameter; each oogonium 

produces a single oocyte. Antheridia develop on cells of 

filamentous thalli of narrower diameter, are usually borne 

in clusters, and produce motile spermatozoids.

Remarks. In collecting species of Desmarestia, 

care should be taken in placing it with other specimens. 

Many species of Desmarestia are known to contain sulfuric 

acid in their cell vacuoles (Blinks, 1951; Eppley and Bovell, 

1958; Sasaki et al., 2004), which is probably a herbivore 

deterrent (Anderson and Velimirov, 1982). The vacuolar 

sap, with a pH 0.44 to 2.0 (Schiff, 1962), will discolor or 

damage your other collections. McClintock et al. (1982) 

showed the acidic or nonacidic condition to be species specific, and the analyses of Peters et al. (1997) revealed that 

sulfuric acid vacuoles were strong phylogenetic characters.

Carbohydrates were elucidated in some species of 

Desmarestia (Carlberg et al., 1978). The sexual hormone 

desmarestene and its role in sperm release and sperm 

attraction in the sexual reproduction of Desmarestia has 

been described by Müller and Lüthe (1981) and Müller 

et al. (1982). Filamentous gametophytes of Desmarestia,

although known from culture studies (e.g., Kornmann, 

1962; Nakahara, 1984; Peters and Müller, 1986b), are 

apparently rare in nature and have not been found in the 

Gulf. Filamentous gametophytes have been found in the 

tissue of a Pacific Northwest sea pen, Ptilosarcus gurneyi

(Anthozoa; Pennatulacea), and were cultured and grew to 

be sporophytes of Desmarestia (Dube and Ball, 1971). In 

the Antarctic, endophytic, monoecious filamentous gametophytes of Desmarestia were identified in the red alga 

Curdiea racovitzae Hariot (Moe and Silva, 1989).

Solely on the basis of vegetative morphology, the 

Gulf-endemic species D. mexicana and D. filamentosa

were placed in synonomy with D. ligulata and D. viridis, respectively (Chapman, 1972b). Noting that vegetative characters may be of secondary importance and that 

reproductive structures are known for only a few of the 

species, Moe and Silva (1977) have questioned the conclusions of Chapman (1972a, 1972b).

Comparative morphological studies of vegetative characters and reproductive structures and life history culture 

studies, such as those of Peters and Müller (1986b) and 

Anderson and Bolton (1989), are needed for the northern Gulf taxa. These studies could test another morphological feature; e.g., Garbary (1978) suggested the surface 

characteristics of macroalgae may also be of taxonomic 

value, and Oates and Cole (1990b) found distinctive cell 

wall ornamentation on two species of Desmarestia from 

the northeast Pacific. Critical studies should also include 

DNA investigations to clarify the taxonomic status and 

phylogenetic relationships of the Gulf taxa.

Two taxa, apparently endemic, are known in the 

northern Gulf of California, and both contain sulfuric 

acid. While acknowledging the northern Gulf specimens 

are vegetatively similar to some species from the Pacific 

coast, they are ecologically and geographically isolated, 

with a restricted distribution to the cooler, subtidal waters 
1 4 6 • 

in the vicinity of Isla Ángel de la Guarda and adjacent areas 

and Isla San Esteban. Therefore, for now, two species are 

recognized, one of these as a subspecies to reflect its possible relationship to a Pacific coast species.

Key to the Species of Desmarestia in th

1a. Thalli cylindrical to subcylindrical and narrow, less than 3 mm wide; oppositely branched to 3–5 orders 

. . . . . . . 

1b. Thallus flat, of large, broad blades, greater than 10 mm wide; either unbranched or sparsely, oppositely branched to 1–2 

orders  . . . . . . . . . . . 

Desmarestia filamentosa E. Y. Daw

FIGURES 71, 72

Desmarestia filamentosa E. Y. Dawson, 1944:236, pl. 

76; Dawson, 1961b:392; González-González et al., 

1996:144.

Desmarestia viridis sensu Chapman, 1972b:225, fig. 9 

[in part, with reference to Gulf Desmarestia filamentosa only]; Abbott and Hollenberg, 1976:225, fig. 

187; Norris and Bucher, 1976:6; González-González 

et al., 1996:145; Pacheco-Ruíz and Zertuche-González, 

1996b:171; Pacheco-Ruíz et al., 2008:204; Pedroche et 

al., 2008:78 [non Desmarestia viridis (O. F. Müller) J. 

V. Lamouroux, 1813:45; basionym: Fucus viridis O. F. 

Müller, 1782:5].

Algae yellow-brown when fresh, pale green on drying, up to 92 cm tall; main axis cylindrical to subcylindrical, (1–)2–3 mm in diameter, percurrent and oppositely 

branched to 3–4(–5) orders; arising above a stipe, up to 

5 cm long by 1–2 mm in diameter, and attached below 

by a small discoid holdfast. Branches mostly cylindrical 

and gradually reduced in size upward; ultimate branchlets 

80–110 μm in diameter; apices with trichothallic filament 

with cortication below the meristematic zone (Figure 72A).

Reproduction not seen in Gulf of California specimens.

Habitat. On rocks, sand/shell bottom; subtidal, (3–)10–23 m depths

Distribution. Gulf of California: Isla Mejía

and Puerto Refugio, Isla Ángel de la Guarda to Bahía de 

Los Ángeles; Isla San Esteban.

Type Locality. Puerto Refugio, Isla Ángel de

la Guarda, Las Islas de la Cintura, Gulf of California.

Remarks. Apparently a northern Gulf 

endemic, Desmarestia filamentosa is easily distinguished 

from the very large, flat, Gulf D. munda subsp. mexicana

by its much smaller, narrower subcylindrical axes and 

branches. Originally known only from the dredged type 

specimen (Dawson, 1944), numerous specimens of D. filamentosa were found in our diving off Roca Blanca, Puerto 

Refugio (Norris and Bucher, 1976, as “D. viridis”).

Dawson (1944) noted D. filamentosa was close to D. 

media (cf. Setchell and Gardner, 1925) and maybe to D. 

pacifica Setchell et N. L. Gardner (1924b, 1925). Chapman (1972b) considered D. filamentosa to be a synonym 

of D. viridis, a North Atlantic species that was subsequently reported on the Pacific coast from Alaska to Baja 

California (cf. Abbott and Hollenberg, 1976; Scagel et al., 

1989). However, Abbott and Hollenberg (1976:222) suggested that most central California specimens identified as 

“D. viridis” were probably D. kurilensis Yamada (1935). 

Given the questionable taxonomic status of some species 

of Desmarestia in general (see “Remarks” under Desmarestia) and the ecological and geographical isolation of the 

Gulf taxon, it seems best to consider it a separate species, 

as D. filamentosa, until it can be further evaluated and 

its systematic status and phylogenetic relationships can 

be tested.

Desmarestia munda Setchell et N. L. Gardn

subsp. mexicana (E. Y. Daws

J. N. Norris, comb. nov.

FIGURE 73

Desmarestia mexicana E. Y. Dawson, 1944:236, pl. 

77; Dawson, 1961b:393; González-González et al., 

1996:145; Pedroche et al., 2008:78.

Desmarestia ligulata sensu Chapman, 1972a:2, 17 [in 

part, with reference to Gulf Desmarestia mexicana

only]; Abbott and Hollenberg, 1976:222; Norris and 

Bucher, 1976:6; González-González et al., 1996:144; 

Pacheco-Ruíz et al., 2008:204 [non Desmarestia ligulata (Lightfoot) J. V. Lamouroux, 1813:25; Fucus ligulatus Lightfoot, 1777b:946].
n u m b e r 9 4 • 1 4 7

FIGURE 71. Desmarestia filamentosa: Habit, specimen from the type locality (JN-5271).
1 4 8 • 

Algae large, up to 150 cm in length; of 1 or more 

thin, flat blades, up to 85 cm long and widening gradually upward from its basal stipitate region, to 8–20 cm 

in width. Thalli arising from a small, conical holdfast 

with a slender, compressed stipe, (2–)3–4 mm wide, that 

extends into main axis, 3–4 mm wide, bearing a terminal 

primary blade. Main axis may remain unbranched or with 

opposite lateral blades. Primary and lateral blades broadly 

lanceolate and unbranched or may become sparsely and 

oppositely branched with secondary blades formed on the 

margin of its lower portion. Blades with a slender, visible midrib and faint opposite lateral veins that gradually 

disappear or occasionally continue to become midrib of 

marginal blades. Blade margins coarsely serrate; spinelike outgrowth tapering gradually, with a terminal tooth 

pointed upward.

Reproduction not observed in Gulf of California 

specimens.

Habitat. On rocks, sand/shell bottom; subtidal, 9–23 m depths

Distribution. Gulf of California: Puerto

Refugio, Isla Ángel de la Guarda to Isla Estanque; Punta 

la Gringa, Bahía de Los Ángeles.

Type Locality. Puerto Refugio, Isla Ángel de

la Guarda, Las Islas de la Cintura, Gulf of California.

Remarks. Desmarestia munda subsp. mexicana was originally known from a single dredged collection in the northern Gulf (Dawson, 1944, as “D. 

mexicana”). Desmarestia munda subsp. mexicana is easily recognized by its very large size and flat, long, and 

broad, thin blades; diving surveys at its type locality 

showed it to be rather abundant at 15–23 m depths (Norris and Bucher, 1976).

Although Desmarestia mexicana was considered by 

Chapman (1972a) to be one of the synonyms of a broadly 

redefined D. ligulata, Silva (1979:336) noted that taxonomic treatment is not convincing. The northern Gulf 

specimens show some similarities with larger specimens 

of the Pacific coast D. munda Setchell et N. L. Gardner (1924b; Dawson, 1945c), a species recorded from 

Alaska to central Baja California (Setchell and Gardner, 

1925:567, pl. 89; Smith, 1944:121, pl. 17: fig. 1; Dawson, 

1951:52; Dawson et al., 1960a:30, pl. 7: fig. 1). However, 

Gulf specimens of D. munda subsp. mexicana differ from 

D. munda subsp. munda in having much broader, longer 

blades (up to 20 cm wide and to 85 cm long) that either 

lack secondary blades or have secondary blades that are 

sparse and opposite off the margins of the primary blade 

(Figure 73) or lateral blades. Therefore, it seems best to 

recognize the Gulf taxon as a subspecies to reflect its possible relationship to the Pacific D. munda, until its systematic and phylogenetic status can be tested.

Spo

Sporochnales Sauvageau, 1926:364.

The sporophyte of these algae is a large erect macrophyte. Members usually have prominent, slender, terete 

or compressed main axes that are moderately to densely 

branched. The axes and branches are pseudoparenchymatous in structure and have characteristic terminal and/

or lateral tufts of simple filaments. Growth occurs in the 

meristematic zone in apices of the axes and branches and 

in the intercalary meristem of each trichothallic filament 

of the terminal tufts. The cortical layer is composed of 

pigmented cells with discoid plastids and no pyrenoids. 

In contrast, the gametophytes, where known, are microscopic filamentous algae.

FIGURE 72. Desmarestia filamentosa: A. Apex of a branch showing single trichothallic filament above the meristematic zone (arrow), 

below which the cortical envelope develops. B. Upper portion 

of a branch with opposite initials of lateral branches developing 

(JN-5271).
n u m b e r 9 4 • 1 4 9

FIGURE 73. Desmarestia munda subsp. mexicana: Portion of thallus showing main axis with opposite lateral blades and primary blade with 

secondary blades developed oppositely along its margin (JN-5283).
1 5 0 • 

Life histories are heteromorphic. Reproduction in 

the sporophytes is by meiospores produced in unilocular meiosporangia. The gametophytes are monoecious or 

dioecious, and reproduction is oogamous (Caram, 1965; 

Müller et al., 1985).

The order is represented by one family in the northern 

Gulf of California.

Spo

Sporochnaceae Greville, 1830:36.

Members of this family have a branched main axis that 

is usually very distinct. The axis and branches bear short, 

usually stalked, determinate branchlets that bear a conspicuous terminal tuft of filaments. Growth is trichothallic in the 

axis and branches from an apical meristem located below 

the tuft of filaments, and in each of these filaments, growth 

is from an intercalary meristem just above their base. Anatomical structure is initially uniaxial in the youngest growing portions, which soon become surrounded by developing 

cells to form a pseudoparenchymatous medulla of large colorless cells and an outer cortex of small pigmented cells.

Unilocular sporangia produced by the macroscopic 

sporophyte are superficial on the cortex or borne laterally 

off clustered paraphyses. These produce meiospores that 

when released, germinate and develop into gametophytes. 

Sexual reproduction is oogamous. Gametophytes are 

microscopic, branched uniseriate filaments that produce 

oogonia and antheridia.

One genus of the family is found in the northern Gulf 

of California.

Sporochnus C. Agar

Sporochnus C. Agardh, 1817:xii.

Algae (sporophyte) have an erect cylindrical axis with 

few to many lateral branches, branched up to several 

orders and attached below by a fibrous holdfast. The main 

axis and branches are slender, threadlike to wiry, more 

or less indeterminate in growth, and terminate in a tuft 

of trichothallic filaments. Numerous determinate tufted 

branchlets, of more or less similar length, are borne on the 

axis and all orders of branches. These determinate branchlets in most species are stalked (pedicel) below a thicker 

fertile portion (sporangial receptacle) with a terminal tuft 

of trichothallic filaments, although in a few species they 

are without stalks (non-pedicellate).

The large sporophytes produce unilocular sporangia in a layer encircling the fertile portion of determinate 

branchlets below their terminal tuft (see, e.g., Womersley, 1987:fig. 103G). Unilocular sporangia are borne 

laterally on unbranched or branched paraphyses that 

have an enlarged spherical to subspherical terminal cell. 

Gametophytic thalli are microscopic branched filaments 

and are monoecious or dioecious. Sexual reproduction is 

oogamous.

Remarks. Caribbean species of Sporochnus

have been shown to produce some bioactive secondary 

metabolites, sporochnols A, B, and C (Shen et al., 1993). 

The natural products of the Gulf of California species 

should also be investigated.

Two species are found with a restricted distribution in 

the northern Gulf of California.

Key to the Species of Sporochnus in the n

1a. Ultimate tufted branchlets long, up to 9.0 mm long; sporangial receptacles with stalks 1.0–3.0(–4.0) mm in length 

. . . . . . 

1b. Ultimate tufted branchlets much shorter, mostly 0.5–1.0 mm long; sporangial receptacles mostly lacking stalks or, if 

present, minute, less than 240 μm in length  . . . . . .

Sporochnus anomalus (Pallas) M. J. Wynn

FIGURES 74, 75A,B

Fucus anomalus Pallas, 1766:199, pl. 14: figs. 24, 24*[sic].

Sporochnus anomalus (Pallas) M. J. Wynne, 2003:78, 

fig. 1.

Sporochnus gaertnera C. Agardh, 1820:150; C. Agardh, 

1824:259; Montagne, 1846:27.

Fucus gaertnera Gmelin, nom. illeg., 1768:164, pl. XIX: 

figs. 1, 1a.

Sporochnus pedunculatus sensu Dawson, 1951:56 

[non Sporochnus pedunculatus (Hudson) C. Agardh, 

1820:149; basionym: Fucus pedunculatus Hudson, 

1778: 587].

Sporochnus bolleanus sensu Dawson et al., 1960b:14; 

Dawson, 1961b:392; Norris and Bucher, 1976:4, figs. 

3, 4; González-González et al., 1996:165; Pacheco-Ruíz 
n u m b e r 9 4 • 1 5 1

FIGURE 74. Sporochnus anomalus: Habit, branches with numerous stalked, tufted branchlets (JN-5264).
1 5 2 • 

and Zertuche-González, 1996b:171; Pacheco-Ruíz et 

al., 2008:204; Pedroche et al., 2008(in part; Gulf of 

California records only):76 [non Sporochnus bolleanus

Montagne, 1856:393].

Algae (sporophyte) erect, up to 44 cm tall, of terete 

branched axes; lateral branches up to 38 cm long. Determinate tufted branchlets on axes and branches usually 

numerous and up to 9.0 mm in length; composed of a long 

stalk, (1.0–)2.0–3.0(–5.0) mm long and up to 200 μm in 

diameter, elongated cylindrical receptacle, 1.0–2.25 mm 

long, and a terminal cluster of filaments, 3.0–5.0(–6.0) 

mm long, extending from an apical pit.

Habitat. On rocks and shells; subtidal,

20–30 m depths.

Distribution. Gulf of California: Roca

Blanca, Puerto Refugio (Isla Ángel de la Guarda) to Bahía 

de Los Ángeles. Pacific coast: Isla Guadalupe; Punta 

Abreojos to Punta Pequeña, Baja California Sur.

Type Locality. Not given by Pallas (1766);

Mediterranean was listed by Gmelin (1768).

Remarks. The northern Gulf of California 

specimens of Sporochnus anomalus have measurements 

intermediate between two other species: larger than those 

of S. pedunculatus (Hudson) C. Agardh (1820) and smaller 

than those of S. bolleanus Montagne (1856) (cf. Taylor, 

1960; Earle, 1969). Earle (1969) has noted that in the 

northeastern Gulf of Mexico certain forms of S. bolleanus

have branchlets shorter than usual for this species, making it sometimes difficult to separate from S. pedunculatus

FIGURE 75. Sporochnus anomalus: A. Portion of a branch with determinate branchlets, each showing the sterile, long stalk, a wider fertile portion (receptacle), and an apical tuft of filaments (JN-5264). B. Overall length variations of determinate lateral branchlets, with stalk, receptacle 

(sporangial sori surrounding branch), and terminal tuft of assimilatory filaments (JN-5264).
n u m b e r 9 4 • 1 5 3

where the two occur together. The northern Gulf Sporochnus is tentatively referred to S. anomalus, until critical studies of northern Gulf specimens can be made to determine 

their taxonomic status and phylogenetic relationships.

Sporochnus neushulii J. N. Norris, sp. nov.

FIGURE 76A–E

Latin Description. Algae usque ad 31 cm

longae, maiores teretes axes cum ramis plerumque alternis et lateralibus, usque ad 2.5 cm longis; ab imo affixae 

cum parvo discoidi hapterone. Axes et rami ferentes parvos et numerosos (plerumque denses) ramulos, radialiter 

dispositos, breves, determinatos, caespitosos, (0.25–)0.5–

1.0(–2.0) mm longos, 200–600(–750) μm in diametro, 

plerumque non-pedicellatos praecipue in altis partibus 

thalli, vel, quamvis rarum (ut videtur), si pedicelli adsunt, 

maxime minuti tantum usque ad 80(–240) μm longi et 

usque ad 125 μm in diametro. Sporangialia receptacula 

cylindricalia, magis vel minus ovoidea, cum terminalibus 

caespitibus pilorum. Uniloculares sporangiae, elongatae, 

latae lateraliter super paraphyses, quorum terminales cellulae largae et sphaericae usque sub-sphaericae sunt.

Algae (sporophyte) up to 31 cm tall, main terete axes 

with usually alternate, lateral branches, up to 12.5 cm 

long; attached below by a small discoid holdfast. Axes 

and branches bearing numerous (often dense), radially 

arranged, short, determinate, tufted branchlets, (0.25–)

0.5–1.0(–2.0) mm in length, 200–600(–750) μm in diameter; mostly without a stalk (pedicel), especially in upper 

portions of thallus, or, although apparently rare, with very 

minute stalks, only up to 80(–240) μm long and up to 125 

μm in diameter.

Sporangial receptacles cylindrical, sometimes more 

or less ovoid, with terminal tuft of filaments from apical 

pit. Unilocular sporangia, clavate, 14–22 μm in length and 

3–4 μm in diameter, borne laterally on paraphyses, with 

enlarged, spherical to subspherical terminal cell, 12–15

(–18) μm in diameter.

Gametophytes unknown.

Habitat. On rocks, subtidal; 20–30 m depths

Distribution. Gulf of California: Puerto

Refugio, Isla Ángel de la Guarda.

Holotype. US Alg. Coll.-158492, 21 Apr

1974, coll. J. N. Norris & K. E. Bucher, JN-5262; isotypes: MICH; US Alg. Coll.

Type Locality. Off west side of Roca Blanca,

Puerto Refugio, Isla Ángel de la Guarda, Las Islas de la 

Cintura, Gulf of California.

Etymology. The species is named in honor 

of the late Michael Neushul Jr. (University of California, 

Santa Barbara), professor, mentor, and friend, in recognition of his numerous contributions to marine botany 

(Lewis et al., 1994; Reed et al., 1994), for his teaching and 

encouragement to students, and particularly for his support of my Gulf of California research. He was inspired 

early on by E. Yale Dawson to study marine algae and 

become a scuba diver, and he also collected and published 

papers with Dawson on algae of southern California and 

Pacific Mexico.

Remarks. Sporochnus neushulii is an endemic 

species, thus far only known from Puerto Refugio, Isla 

Ángel de la Guarda. The northern Gulf specimens somewhat resemble S. apodus Harvey (1859) in habit (cf. 

Womersley, 1967, 1987). The two differ in the nature of 

their sporangial receptacles. There is a complete absence 

of pedicels for the receptacles of S. apodus (Harvey, 1859: 

pl. XCII; Womersley, 1987: 278, fig. 101C–E; Brostoff, 

1984, tbl. 1), whereas not all the receptacles of S. neushulii are nonpedicellate because some are on very minute 

pedicels. Their receptacles also differ in size and shape: in 

S. apodus they are 0.3–2.0(–4.0) mm in length and 200–

300 μm in diameter, broadest at their base, and slightly 

tapered upward; in contrast, they are mostly shorter in S. 

neushulii, 0.5–1.0(–2.0) mm in length and usually wider in 

diameter, 200–600(–750) μm, and cylindrical (sometimes 

ovoid) and of similar diameter throughout. These two are 

also widely geographically separated from each other. 

Sporochnus neushulii is apparently restricted to the north 

end of Isla Ángel de la Guarda (29°32′N, 113°33′W), 

and S. apodus is known from Tasmania (type locality: 

George Town) and southern Australia (South Australia to Victoria; Womersley, 1987) (range approximately 

33°–38.4°S,134°–147°E). Thus, for now, they are recognized as different species.

Sporochnus neushulii also superficially resembles some 

S. pedunculatus (Hudson) C. Agardh (1820; type locality: 

England). However, S. neushulii differs from S. pedunculatus primarily by its sporangial receptacles, which usually lack stalks or have stalks, when present, that are very 

minute (less than 240 μm long), and its sporangial receptacles are of similar or longer length, up to 2.0 mm. Sporochnus pedunculatus, from England (Fletcher, 1987) and 

the Mediterranean Sea, has sporangial receptacles that are 

slightly shorter in length, 0.5–1.0 mm, and borne on distinctive long stalks, up to 1 mm in length (Wynne, 2003).

Sporochnus pedunculatus has been reported on 

the Pacific coast from Santa Catalina Island, California 

Channel Islands (Mower and Widdowson, 1969, as “S. 
1 5 4 • 
n u m b e r 9 4 • 1 5 5

apodus”), to Laguna Ojo de Liebre (Scammon’s Lagoon), 

Baja California Sur (Dawson et al., 1960b), and Isla Guadalupe (Setchell and Gardner, 1930; Stewart and Stewart, 

1984). In contrast to S. neushulii, southern California S. 

pedunculatus (Abbott and Hollenberg, 1976) has much 

longer stalks (0.75–2.5 mm long) and longer (up to 2 

mm long) and slightly narrower (up to 500 μm in diameter) sporangial receptacles. Finally, S. pedunculatus as 

reported in the western Atlantic (Taylor, 1960) and Gulf 

of Mexico (Earle, 1969) also has longer stalks (1–2 mm 

long) and generally smaller (0.75–1.25 mm; rarely to 2.0 

mm long) and somewhat narrower (up to 500 μm in diameter) sporangial receptacles. Further studies and molecular 

analyses will be helpful in testing the taxonomic status and 

phylogenetics of the northern Gulf S. neushulii and California and Pacific Baja California S. pedunculatus.

Ralfsia

Ralfsiales Nakamura ex Lim et Kawai in Lim et al., 

2007:464.

Ralfsiales Nakamura, 1972:152, nom. invalid.

Algae are crustose in at least one stage of their life 

history or have a crustose form only in the early stages 

of development (i.e., Analipus Kjellman, 1889; Wynne, 

1971). The initial development of the thallus is discshaped in all members. Cells have one to several chloroplasts without pyrenoids.

Unilocular sporangia (zoidangia) are terminal or lateral and either with or without paraphyses. Plurilocular 

sporangia (zoidangia) are intercalary with terminal sterile 

cells. The Rubisco large subunit gene sequence is distinctive (Lim et al., 2007).

* Contributed by Daniel León-Álvarez and James N. Norris; D. León-Álvarez: Sección de algas del Herbario de la Facultad de Ciencias, Universidad Nacional Autónoma de México, 

Apartado Postal 70–592, 04510 Mexico D.F., Mexico.

Remarks. Although the Ralfsiales was first 

proposed by Nakamura (1972), the name lacked the 

required Latin description (see McNeill et al., 2006:Art. 

36.2). Silva and de Reviers (2000) and León-Álvarez and 

Norris (2004) noted that since then, the name, although 

not validly published, had been used in the literature by 

many phycologists. The ordinal name was only recently 

validly published as Ralfsiales Nakamura ex Lim et Kawai 

(in Lim et al., 2007).

Originally, Nakamura (1972) proposed the order to 

include three families: Ralfsiaceae Farlow (1881), Lithodermataceae Hauck (1884), and Nemodermataceae J. 

Feldmann (1937). Subsequently, another family, the 

Mesosporaceae Tanaka and Chihara (1982), was added. 

Recently, the monotypic Nemodermataceae based on 

Nemoderma Kuckuck (1912), was removed from the 

Ralfsiales to a new order, the Nemodermatales Parente, 

R. A. Fletcher, F. Rousseau et N. Phillips (in Phillips et 

al., 2008).

There is one family represented in the northern Gulf 

of California.

Ralfsi

Ralfsiaceae Farlow, 1881:17, 86.

Algae are discoid to spreading prostrate crusts or 

cushionlike, with smooth to convoluted surfaces. Crustose 

thalli growing on hard substrata are usually without rhizoids on their ventral surface and may be firmly attached 

or easily detached. Internally, crustose members are composed of a basal layer of horizontal radiating filaments, 

one to several cell layers thick, and middle to upper layers 

of assurgent to erect filaments produced by the basal cells. 

These layers may be loosely (e.g., Hapalospongidion De 

A. Saunders, 1899), moderately (e.g., Petroderma Kuckuck, 1897, and Ralfsia confusa Hollenberg, 1969), or 

closely packed (e.g., members of Ralfsia subgen. Ralfsia 

[= Batters, 1902:42, as “R. subgen. Euralfsia”]). Growth 

is from terminal cells of the horizontal and erect filaments 

or, in a few, intercalary from basal meristems of cortical 

FIGURE 76. Sporochnus neushulii sp. nov., holotype (JN-5262; US Alg. Coll.-158492). A. Habit of holotype specimen. B. Portion of a branch 

with short, radially arranged, determinate, tufted branchlets. C. Example of a typical non-pedicellate determinate branchlet, completely covered 

by sporangial paraphyses from apical tuft to parent branch. D. Very minute, sterile stalk, as found on only a few of the determinate tufted 

branchlets. E. Portion of a receptacle showing a group of paraphyses, each with terminal subspherical cell (arrowhead), and developing laterally 

borne, unilocular sporangia (arrow).
1 5 6 • 

filaments. Cells have cup-shaped or plate-shaped chloroplasts without pyrenoids.

Reproductive structures are lateral, terminal, or intercalary on the reproductive filament and are usually associated with surrounding paraphyses or vegetative filaments 

(but lacking in a few) (León-Álvarez and Norris, 2004). 

Life histories, where known, are isomorphic, with unilocular or plurilocular sporangia (zoidangia) on separate 

individuals.

Remarks. The Ralfsiaceae had been placed in 

the Ectocarpales (e.g., Wynne, 1981; Nelson, 1982; Scagel et al., 1989; Schneider and Searles, 1991; Silva et al., 

1996a), or in the Myrionemataceae (e.g. Loiseaux, 1986), 

but on the basis of rDNA studies, Tan and Druehl (1994) 

concluded the family did not belong in the Ectocarpales, 

and Draisma et al. (2003) stated molecular sequences of 

Ralfsia were needed to clarify their systematic and phylogenetic relationships. The ordinal taxonomic placement 

of the Ralfsiaceae was recently resolved in the molecular phylogeny of Lim et al. (2007), which supported the 

Ralfsiales. Taxonomic studies of members of the family 

include those of Tanaka and Chihara (1980a, 1980b, 

1980c, 1981a, 1981b, 1982).

Riosmena-Rodríquez and Paul-Chávez (1997) have 

reported two genera in the southern Gulf from La Paz: 

Endoplura Hollenberg (1969) with E. aurea Hollenberg; 

and Pseudolithoderma Svedelius (in Kjellman and Svedelius 1910) with P. nigrum Hollenberg (1969).

Two genera, including one of uncertain taxonomic 

status, are known in the northern Gulf of California.

Key to the Genera of Ralfsiaceae in the no

1a. Thalli low-growing clumps of overlapping linear, mostly horizontal, short branches. . . . . . .

1b. Thalli crustose; flat, discoid to spreading, sometimes overlapping; of 2–3 layers: a basal layer of 1 or more horizontal filaments and 1–2 upper layers of assurgent or vertical filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ralfsia

Hapterophycus Setchell et N. L. Gardn

in Setchell

Hapterophycus Setchell et N. L. Gardner in Setchell, 

1912:233.

Algae are low growing and spreading, but only slightly 

crustose, with linear, mostly horizontal, short branches. 

Medulla has a middle layer of horizontal filaments of 

elongated cells that have with downwardly directed filaments and assurgent filaments that end with adjoined erect 

filaments. Cryptostomata are present. Cells have a single 

parietal chloroplast. Unilocular sporangia are in broad 

sori; sessile at the base of multicellular paraphyses. Gametangia not confirmed.

Remarks. The generic status of Hapterophycus is in need of investigation. In culture studies, 

Kogame (1996) has shown that Japanese specimens identified as “H. canaliculatus” were a sporophytic phase 

in the life history of a species of Scytosiphon C. Agardh 

(1820) and, on the basis of their morphological similarity 

to the generitype (H. canaliculatus Setchell et N. L. Gardner in Setchell, 1912), concluded them to be Scytosiphon 

canaliculatus (Setchell et N. L. Gardner) Kogame, thus 

reducing Hapterophycus to a synonym of Scytosiphon 

(Scytosiphonaceae). However, in culture studies of southern California H. canaliculatus (type locality: San Pedro, 

California) neither Hollenberg (1941) nor Wynne (1969) 

observed erect thalli, and Wynne concluded there was a 

“direct type” of life history with a macroscopic alga with 

unilocular sporangia. Hollenberg (1941; see also Abbott 

and Hollenberg, 1976) concluded Hapterophycus should 

be in the Ralfsiaceae, close to Ralfsia. The question of 

its generic and familial placement and if the algae from 

California and Japan are the same species will need to be 

resolved by molecular and phylogenetic analyses.

There is one endemic species in the northern Gulf of 

California.

Hapterophycus anastomosans E. Y. Da

FIGURE 77A,B

Hapterophycus anastomosans E. Y. Dawson, 1966b:56, 

figs. 2, 6A; Hollenberg, 1969:301; Wynne, 1969:7; 

González-González et al., 1996:152; Pacheco-Ruíz et 

al., 2008:204; Pedroche et al., 2008:26.

Algae form extensive, entangled mats, up to 40 cm or 

more in width; consisting of horizontal to partially erect, 

short, rigid (haptera-like) branches, subdichotomously 

to irregularly branched. Branches subcylindrical to compressed, up to 1.0 mm wide and to 15 mm long, simple to 

palmately divided terminally, with blunt, broadly rounded 
n u m b e r 9 4 • 1 5 7

apices; entangled and fused at most lateral or terminal 

contacts by anastomosing growth, attachments give rigidity to thallus. Medulla in transection of compact, more 

or less isodiametric, angular cells of irregular size, 20–40 

μm in diameter; in longitudinal section, central portion of 

irregularly oblong cells and outward cells in a “fountain 

pattern” toward the apices and cortical surfaces. Cortex 

of sparsely branched, anticlinal rows of 7–8 cells, 6–7 μm 

in diameter.

Reproduction not known.

Habitat. On rocks, mid intertidal

Distribution. Gulf of California: Isla San

Lorenzo; Isla Raza.

Type Locality. Isla Raza, Islas de la Cintura,

Gulf of California.

Remarks. Apparently a Gulf endemic, the 

taxonomic status of H. anastomosans is in need of investigation (see also Remarks above under genus). Until the 

generic and specific status of Gulf H.? anastomosans can 

be tested by comparative morphological, life history, and 

molecular analyses with the generitype H. canaliculatus

Setchell et N. L. Gardner (in Setchell 1912) (type locality 

San Pedro), and those reported from southern California 

(Abbott and Hollenberg, 1976), Pacific Baja California 

(Dawson, 1951), and Isla Guadalupe (Stewart and Stewart, 1984), the Gulf of California taxon is referred to this 

genus with a query.

Ralfsia Berkeley in J. E. Smith et J. C. Sower

Ralfsia Berkeley in J. E. Smith et J. C. Sowerby, 1843:pl. 

2866.

Algae are crustose and loosely to firmly adherent 

(without rhizoids) to the substratum or attached by rhizoids. Initially, crusts are more or less disc shaped,but 

later spread and may become slightly lobed at the margins. Few to several crusts often grow close to each other, 

and each may become thicker by overgrowth of overlapping margins. Anatomically, there are two or three layers: a lower or basal layer of one to several horizontally 

radiating filaments, either with or without rhizoids from 

FIGURE 77. Hapterophycus anastomosans: Habit (holotype, EYD-26110; US Alg. Coll.-37228.). A. Portion of air-dried holotype specimen. B. 

Liquid-preserved portion of type specimen showing the entangled, fused, short branches that form the low-growing mat.
1 5 8 • 

its lowermost cells, and upper and/or middle layers developed from basal filaments that ascend to form more or 

less vertical, moderately to firmly adjoined, unbranched to 

sparingly branched filaments of similar length. Cells of the 

outer surface layer usually have one chloroplast per cell. 

Hairs are grouped in hair pits within the upper surface 

layers.

Reproductive structures are unilocular or plurilocular on separate algae. Unilocular structures are lateral or 

terminal and sessile or on one to few stalk cells (pedicel), 

usually associated with paraphyses, and grouped in superficial sori. Plurilocular structures are slender, of one to two 

rows of locules, with one to two sterile terminal cells, and 

grouped within the crust.

Remarks. Life history studies of the species of 

Ralfsia thus far cultured are heteromorphic (with mostly 

asexual phases, but sexual phases have been reported for 

some in culture studies), with an erect bladelike thallus 

(such as Petalonia) and a prostrate crustose Ralfsia or 

“Ralfsia-like” phase (e.g., Wynne, 1969, 1972; Nakamura and Tatewaki, 1975; Fletcher, 1978). Loiseaux 

(1968) found two species of Ralfsia to be apomeiotic with 

apparent isomorphic life histories. Culture and molecular 

studies of Gulf species should prove interesting and help 

elucidate their taxonomic status.

One species, Ralfsia hesperia Setchell et N.L. Gardner (1924b), has been reported in the southern Gulf 

(Carballo et al. 2002). In addition to those species known 

in the northern Gulf, there are at least two other species 

of “Ralfsia-like” crusts occurring on mid-intertidal rocks 

in the upper Gulf at Puerto Peñasco (Dawson, 1966a), but 

the lack of fertile material has left them undetermined.

Three species and one uncertain record are reported in 

the northern Gulf of California.

Key to the Species of Ralfsia in the nor

1a. Crusts of 3 distinct layers: basal layer of 2–4 horizontal rows of basal filaments, upper medulla layer of assurgent rows of 

branching filaments, and multicellular cortical layer; unilocular sporangia on (3–)4–5(–6) celled pedicel . . . R. hancockii

1b. Crusts of 2 layers of filaments: a basal layer and an upper layer  

2a. Upper layer of more or less assurgent filaments developed from basal layers; crusts usually more than 300 μm thick; mostly 

sessile sporangia  . . . . . 

2b. Upper layer of erect, vertical rows of filaments above a thin basal layer of 1(–2) horizontal rows of basal filaments; crusts 

mostly less than 200 μm thick; unilocular sporangia on 1- to 6-celled pedicels . . . . . . . . . . . . . . . . . . . . . . . . . R. confusa

Ralfsia confusa Hollenber

FIGURE 78C

Ralfsia confusa Hollenberg, 1969:291, figs. 1–12; Abbott 

and Hollenberg, 1976:164, fig. 131; Stewart and Stewart, 1984:141; Dreckmann et al., 1990:26; Mateo-Cid 

and Mendoza-González 1992:23; León-Álvarez and 

González-González, 1993:461, 464; Mateo-Cid et al., 

1993:49; González-González, 1993:443; León-Tejera 

et al., 1993:200; León-Tejera and González-González, 

1993:497; Servière-Zaragoza et al., 1993:482; MateoCid and Mendoza-González 1994a:50; MendozaGonzález et al., 1994:110; González-González et al., 

1996:158; Rodríguez-Morales and Siqueiros-Beltrones 

1999:22; Mateo-Cid et al., 2000a:67; Cruz-Ayala et al., 

2001:190; Mateo-Cid and Mendoza-González 2003:21; 

Pedroche et al. 2008:28.

Ralfsia californica sensu Dawson, 1944:232 [in part; see 

Hollenberg, 1969:293] [non Ralfsia californica Setchell 

et N. L. Gardner, 1924b:2].

Algae, light to medium brown crusts, often coalescent, mostly 5–10 mm wide, and relatively thin, usually 

less than 200 μm thick and 150–250 μm thick in fertile 

portions. Basal layers of 1–2 horizontally elongate cells, 

13.5–24.0 μm long and 9–12 μm in diameter. Upper layer 

of erect, vertical (not assurgent) filaments of 10–15 cells, 

5–6 μm in diameter. Cells with a single chloroplast. Hair 

pits frequent.

Unilocular and plurilocular sporangia on separate 

thalli. Unilocular sporangia, 70–90 μm long by 20–25 μm 

in diameter, sessile or borne on pedicels of 1–2(–6) cells; 

associated with paraphyses of (3–)10–12 cells, 75–100 μm 

in length and 6–8 μm in diameter. Plurilocular sporangia 

mostly in uniseriate rows, 30–45(–60) μm in length and 5 

μm in diameter, with a single sterile terminal cell, 7.5–10 

μm long and mostly 5 μm in diameter.

Habitat. On rocks; upper to low intertidal

Distribution. Gulf of California: Guaymas

to Cabo Pulmo. Pacific coast: Washington; southern California to Baja California; Isla Guadalupe; Isla Socorro 

(Islas Revillagigedo); Sinaloa to Oaxaca.
n u m b e r 9 4 • 1 5 9

FIGURE 78. Species of Ralfsia. A, B. Ralfsia hancockii: A. Section of thallus showing assurgent branched rows of cells and unilocular sporangia 

on five-celled stalk and associated paraphyses. B. Habit, surface view of crust in the intertidal, showing the smooth surface and concentric lines 

of an orbicular crust. C. Ralfsia confusa: internal structure (three-dimensional depiction) showing vertical and horizontal growth of margin (left) 

and middle region showing vertical rows of cells, hairs in pits, and unilocular sporangia with associated paraphyses (right). D. Ralfsia pacifica: 

section through sterile crust showing filaments curving upward from basal layer. Illustrations A, C, and D drawn by Adrián Ramírez, UNAM; 

photo B taken by D. León-Álvarez.
1 6 0 • 

Type Locality. Corona del Mar, Orange

County, California.

Remarks. In describing Ralfsia confusa from 

southern California to Pacific Baja California, Hollenberg 

(1969:293) also suggested some of the Dawson collections 

(1944, as “R. californica”) were probably this species; 

however, these thin Gulf specimens were not reproductive 

and thus are tentatively referred to R. confusa.

Ralfsia hancockii E. Y. Daw

FIGURE 78A,B

Ralfsia hancockii E. Y. Dawson, 1944:223, pl. 31: figs. 6, 

7, pl. 54: fig. 2; Dawson, 1954a:3; Dawson, 1961b:387; 

León-Álvarez and González-González, 1993:461, 464; 

León-Álvarez and González-González, 1995:358, figs. 

3, 4; González-González et al., 1996:158; L. AguilarRosas et al., 2000:132; Mateo-Cid et al., 2000a:67; 

León-Álvarez and González-González, 2003:614, figs. 

2–12; López et al., 2004:10; Mateo-Cid et al., 2006:49, 

57; Servière-Zaragoza et al., 2007:8; Pedroche et al., 

2008:29.

Ralfsia expansa sensu Børgesen 1912a:123; Børgesen 

1914:189 (see: León-Álvarez and González-González 

2003:614);.Chávez 1972:268; Tanaka and Chihara, 1980b:231, figs. 1B, 2C,D; González-González, 

1993:443; León-Tejera et al., 1993:200 [non Ralfsia 

expansa (J. Agardh) J. Agardh, 1848:63].

Crusts light-brown to reddish brown, orbicular, up to 

7 cm in diameter, upper surface smooth with radial and 

concentric growth lines or rugose to verrucose and with 

a conspicuous margin; vegetative portions108–306 μm 

thick; reproductive portions 198–360 μm thick. Crusts 

firmly attached to substratum by rhizoids. Growth mainly 

unilateral; in thick portions, filaments may grow downward, tending toward bilateral symmetry. Base of 2–4 

horizontal layers of thick-walled, elongate basal cells, 

24–42 μm long and 10.5–15.0 μm in diameter; curving 

upward. Upper layer of assurgent, branching rows of cells; 

middle cells 22–31.5 μm long by 10.5–18 μm in diameter; 

outer cells of 4–7 layers of anticlinal cells, 6–10 μm long 

by 6–8.5 μm in diameter.

Unilocular sporangia, in sori sparsely scattered over 

thallus surface; ovoid, ellipsoid to clavate, 62.5–93 μm in 

length and 22–43 μm in diameter; terminal on a narrow 

stalk of 4–5 cells, 7.5–13 μm in diameter. Paraphyses slender of 10–18 cells, 100–150 μm long; basal cells 9–18 μm 

long by 3.8 μm in diameter; upper cells 6–12 μm long by 

6–8 μm in diameter. Plurilocular sporangia not known for 

northern Gulf specimens.

Habitat. Mid intertidal; on rocks and in tide

pools; forming small continuous or isolated patches and 

often associated with crustose coralline algae.

Distribution. Gulf of California: San Felipe

to Puertecitos; Punta Arena to San José del Cabo. Pacific 

coast: Isla San Benedicto (Islas Revillagigedo); Nayarit to 

Oaxaca; Japan.

Type Locality. Southern end of San José del

Cabo, Baja California Sur.

Remarks. Studies of León-Álvarez and 

González-González (1995, 2003) have clarified Ralfsia 

hancockii, and although some morphologically intergrading forms are found, two different morphological forms 

are recognized. One form is a light-yellowish-brown to 

dark-brown crust, 165–495 μm thick, with smooth upper 

surface without growth lines and sometimes greenish; 

strongly adhering to substratum by scattered rhizoids or 

without rhizoids (difficult to detach); anatomically tending toward unilateral symmetry. The other form is a 

light-brown to greenish crust, 108–750 μm thick; upper 

surface usually irregular, rough or sometimes verrucose, 

and with radial and/or concentric lines; partially attached 

to substratum (easily detached); anatomically of variable 

symmetry, usually tending toward irregular bilateral symmetry. Structures tentatively identified as “plurilocular 

sporangia” were found on Pacific Mexico R. hancockii 

(León-Álvarez and González-González, 2003), but thus 

far, no plurilocular reproductive structures have been 

reported in northern Gulf specimens.

Ralfsia pacifica Hollenber

FIGURE 78D

Ralfsia pacifica Hollenberg in G. M. Smith, 1944:95, pl. 

12: figs. 4–6; Dawson, 1944:222; Dawson, 1961b:387; 

Smith, 1969:95, pl. 12: figs. 4–6; Hollenberg, 1969:296, 

figs. 17–18; Abbott and Hollenberg, 1976:167, fig. 135; 

Devinny 1978:358; Pacheco-Ruíz, 1982:70; Dreckmann 

et al., 1990:26; González-González, 1993:443; LeónÁlvarez and González-González, 1993:461; ServièreZaragoza et al. 1993:485; Mateo-Cid et al., 1993:49; 

González-González et al., 1996:158; Pacheco-Ruíz and 

Zertuche-González, 1996b:171; Riosmena-Rodríguez 

et al., 1998:45; L. Aguilar-Rosas et al., 2000:132; Mateo-Cid et al., 2000a:68; López et al., 2004:10; ServièreZaragoza et al., 2007:8; Pacheco-Ruíz et al., 2008:203; 

Pedroche et al., 2008:30.
n u m b e r 9 4 • 1 6 1

Ralfsia occidentalis Hollenberg in W. R. Taylor, 1945:81; 

Salcedo-Martínez et al., 1988:82.

Ralfsia verrucosa sensu Setchell and Gardner, 1925:497 

[non Ralfsia verrucosa (Areschoug) Areschoug in E. 

Fries, 1845:124; basionym: Cruoria verrucosa Areschoug, 1843:264].

Crusts, irregularly circular, 2–10 cm broad, 0.4–1.0 

mm thick, olive-brown to dark-brown, upper surface 

often with radial and concentric growth lines or ridges 

and firmly adhering to substratum without rhizoids. Basal 

layer filaments curving upward; of cells, 9–16 μm long 

by 11–32(–40) μm wide. Upper layer of forked filaments; 

of cells, mostly 6–8 μm in diameter and 12–24 μm long 

below, near the surface of similar diameter but shorter in 

length 6–15 μm; hair pits frequent.

Unilocular sporangia oblong to clavate, up to 90 μm 

long and to 23 μm wide, usually sessile at base of paraphyses; borne in more or less circular sori, about 1 mm wide. 

Paraphyses of 9–12 cells, up to 150 μm long and up to 9 

μm in diameter. Plurilocular sporangia in groups up to 1 

mm in diameter within crust; uniseriate, up to 90 μm long, 

on cells shorter than broad (to 6 μm in diameter) of erect 

filaments, and with a single terminal sterile cell, 1.5–2.0 

times long as wide.

Habitat. On rocks; mid to low intertidal

Distribution. Gulf of California: Playa

Santa Teresa; Bahía Tepoca to Isla San Esteban; Bahía 

Concepción to Cabo Pulmo; Matzatlán. Pacific coast: 

Alaska to Sinaloa; Isla Socorro (Islas Revillagigedo); 

Nayarit to Oaxaca.

Type Locality. Corona del Mar, Orange

County, California.

Remarks. Although Womersley (1987) suggested that Pacific North America Ralfsia pacifica may not 

be distinct from the Australian R. verrucosa, most continue 

to recognize R. pacifica in Pacific Mexico (e.g., Abbott and 

Hollenberg, 1976; León-Álvarez and González-González, 

1993; L. Aguilar-Rosas et al., 2000). The relationship of 

R. pacifica (type locality: southern California) and the 

Gulf of California R. pacifica to R. verrucosa (Areschoug) 

Areschoug in E. Fries (1845:124; type locality: Bohuslän, 

Sweden) requires further investigation (see also Tanaka 

and Chihara, 1980a, 1980b).

Uncertain Record:

Ralfsia californica Setchell et N. L. Gardn

Ralfsia californica Setchell et N. L. Gardner, 1924b:2; 

Setchell and Gardner, 1925:497, pl. 36: fig. 22; Dawson, 1944:223 [in part]; Dawson, 1961b:386 [in part]; 

González-González et al., 1996:158; Pedroche et al., 

2008:27.

Remarks. The report of Ralfsia californica 

from the northern Gulf is questionable since it was based 

on nonfertile specimens (Dawson, 1944). Culture studies 

of Pacific coast “R. californica” (as interpreted by Wynne, 

1969, 1972) indicate that it is a sporophytic crustose phase 

in the heteromorphic life history of Pacific North American species of Scytosiphonaceae, including Petalonia fascia

(O. F. Müller) O. Kuntze (1898) (see also Nakamura and 

Tatewaki, 1975; Dethier, 1981, 1987; Kogame, 1997a), 

which has been reported from Alaska to Baja California 

and Sinaloa (Scagel et al., 1989; Hansen, 1997; Perdoche 

et al., 2008). Other specimens similar to “R. californica” 

(Wynne, 1969, 1972) or Ralfsia-like crusts (Nakamura, 

1965; Tatewaki, 1966) were found to be a phase in the life 

history of Scytosiphon lomentaria (Lyngbye) Link (1833).

Interestingly, southern California to Pacific Baja California specimens identified as “R. californica” were determined to be a new species, R. confusa Hollenberg (1969; 

Abbott and Hollenberg, 1976). Thus, “R. californica” 

from the Gulf clearly needs reinvestigation. Fertile specimens need to be found, and culture studies and DNA analyses are needed before its identification can be verified.

Cutle

Cutleriales Oltmanns, 1922:109.

Cutleriales Bessey, 1907:289.

Algae are erect or prostrate or partially prostrate 

and erect. Gametophytes may be cylindrical or narrow 

and slightly compressed and variously branched, with all 

branches having an apical tuft of unbranched filaments, 

or flat and fan shaped, with a fringed distal margin of 

monosiphonous unbranched filaments. Life histories are 

isomorphic or heteromorphic. Members of the latter 

group have gametophytes that are erect and a morphologically different macroscopic crustose sporophyte. Erect 

thalli have a trichothallic mode of growth, with intercalary cell division occurring near the base of each distal unbranched pigmented filament. Both gametophytes 

and sporophytes are parenchymatous in structure, with 

cortical layers of small pigmented cells and a medulla of 

irregularly arranged larger, colorless, rounded cells. Cortical cells have numerous discoid plastids and apparently 

lack pyrenoids; however, Hori and Udea (1975) reported 

that although inconspicuous, small pyrenoids may also 
1 6 2 • 

be present. Single or small clusters of phaeophycean hairs 

arising from surface cells are scattered over the thallus 

surface.

Sporophytes (asexual phase) develop unilocular 

sporangia in sori scattered on the blade surface. Each 

sporangium produces 8, 16, or 32 motile meiospores. 

Gametophytes are either monoecious or dioecious, with 

gametangia borne in clusters on lower portions of simple 

or branched fertile filaments (which, in some, may extend 

into hairlike filaments). Sexual reproduction is anisogamous. Female gametangia have 8–16 large locules, while 

male gametangia have numerous smaller locules.

Remarks. Recent study of the phylogenetic 

relationships of brown algae by Phillips et al. (2008:figs. 

3, 4) have revealed the Cutleriaceae and Tilopteridaceae 

to diverge within the same group. Further, they considered 

these two families to be a single order Tilopteridales Kylin 

(1917; = Tilopteridales Bessey, 1907) and suggested that 

ordinal name be used. However, herein the more widely 

used Cutleriales Oltmanns sensu stricto is recognized for 

the order.

There is one family represented in the Gulf of 

California.

Cutle

Cutleriaceae Hauck, 1883:318; Hauck, 1884:403 [see 

Silva, 1980:71].

Cutleriaceae J. W. Griffith et Henfrey, 1856:179.

The family has the characteristics of the order and 

includes three genera: Cutleria Greville (1830), Microzonia J. Agardh (1894), and Zanardinia Nardo ex P. Crouan 

et H. Crouan (1857).

One genus, Cutleria, is represented in the Gulf of 

California.

Cutleria Greville

Cutleria Greville, 1830:xliv, 59.

Most species of Cutleria have an alternation of heteromorphic generations with an erect gametophyte and a 

prostrate crustose sporophyte. The latter was once considered to be a separate genus, Aglaozonia Zanardini 

(1843), until it was recognized as an alternate phase in 

the life history of Cutleria (Falkenberg, 1879; Yamanouchi, 1912; Drew, 1955). Gametophytic thalli of Cutleria

are mostly compressed to flat and are either fan shaped or 

straplike (however, at least one species is cylindrical) and 

may be entire or dichotomously to irregularly branched, 

with laciniated apices, and attached to the substratum by 

rhizoidal outgrowths or discoid holdfasts. Growth is by 

trichothallic meristems near the base of unbranched pigmented filaments that are either along the distal margin 

of flabellate blades or, in other species, terminal on the 

branch apices. These filaments coalesce with each other 

behind the apical tufts, and longitudinal divisions take 

place to form a parenchymatous thallus that develops a 

one- to two-celled cortical layer and a medulla of several 

layers of larger, colorless, rounded cells. Cortical cells contain many discoid chloroplasts without conspicuous pyrenoids. Phaeophycean hairs are usually grouped together in 

small clusters and are scattered over the suface.

Sporophytes of Cutleria (“Aglaozonia-phase”) are 

prostrate and flabellate to irregularly shaped and lobed 

thalli, from 3 to up to 10 cells thick, with tufts of phaeophycean hairs on their upper surface. Crusts are loosely 

attached to the substratum by rhizoidal filaments produced from their lower surface. Growth is by a row of 

meristematic cells along the thallus margin. The medulla is 

composed of irregularly placed large inner cells, and cortex is composed of smaller cells.

The life history, where known, is an alternation of 

heteromorphic generations. Unilocular sporangia of the 

encrusting sporophyte (“Aglaozonia-phase”) are sessile, 

cylindrical, and grouped in clusters on the crust surface. 

Plurilocular sporangia on sporophytic thalli that developed neutral plurispores have been reported in one species, C. cylindrica from Japan (Kitayama et al., 1992). 

Gametophytes of Cutleria are dioecious or monoecious. 

Plurilocular gametangia are typically grouped in superficial tufts on erect blade surfaces. Female gametangia 

are ovoid, with large locules, and produce gametes more 

than twice as long and wide as male gametes. Female 

gametes attract male gametes by the secretion of a sexual 

pheromone (Müller, 1974). Antheridial gametangia are 

elongated, with numerous smaller locules. Parthenogenesis of unfused female gametes and androgenesis of 

unfused male gametes have both been reported (Kitayama et al., 1992).

Remarks. Gametophytes of Cutleria are typically reported to be annual or ephemeral, and sporophytes 

are usually reported to be perennial. The reported dominance in field populations of female gametophytic thalli 

has been partly explained by parthenogenetic females 

(Fletcher, 1987; Womersley, 1987; Kitayama et al., 1992). 

Unfused female gametes may either develop into female 

gametophytes or develop directly into the sporophyte 

(Kitayama et al., 1992; Nagasato et al., 1998).
n u m b e r 9 4 • 1 6 3

There is one species, Cutleria cylindrica Okamura 

(1902b), reported on the Pacific coast from southern California (Hollenberg, 1978; Stewart, 1991) to northern Baja 

California (R. Aguilar-Rosas, 1994).

One endemic species occurs in the Gulf of California.

Cutleria hancockii E. Y. Da

FIGURE 79A–F

Cutleria hancockii E. Y. Dawson, 1944:226, pl. 54: fig. 

1; Dawson, 1953b:3; Dawson, 1961b:387; Dawson, 

1966a:9–10; Hommersand, 1972:70; Norris, 1973:4; 

La Claire and West, 1978:93, figs. 1–31; La Claire and 

West, 1979:247, figs. 1–30; Littler and Littler, 1981:153, 

tbl. 4; Mendoza-González and Mateo-Cid, 1986:420; 

Espinoza-Ávalos, 1993:333; Mendoza-González et 

al., 1994:110; González-González et al., 1996:143; 

Pacheco-Ruíz and Zertuche-González, 1996b:171; 

L. Aguilar-Rosas et al., 2000:132; Mateo-Cid et al., 

2000a:68; Riosmena-Rodriguez et al., 2001:461, fig. 

1; Mateo-Cid et al., 2006:49, 58; Pacheco-Ruíz et al., 

2008:203; Pedroche at al., 2008:58.

Zanardina prototypus sensu Dawson 1941:119 [non Zanardinia prototypus (Nardo) Nardo 1841:189, nom. illeg.; which is now Zanardinia typus (Nardo) P.C. Silva 

in Greuter 2000:2].

Gametophytic algae (Figure 79A–D) erect, light-brown 

to golden-yellow-brown, flat, fan-shaped blades with fringe 

of monosiphonous pigmented filaments along distal margin, blades often becoming laciniate; 1 to several blades per 

clump, up to 15(–22) cm high by (2.5–)10–21 cm wide; 

arising from a flattened, prostrate basal portion attached 

by rhizoids from its undersurface. Trichothallic growth 

occurs at the basal meristem of each monosiphonous 

unbranched filament near the transition from free filaments 

to consolidated blade. Blades parenchymatous below the 

fringed margin, with 1(–2) layer(s) of small pigmented cortical cells. Fronds 100–140(–180) μm thick. Cortical cells: 

in transection, brick-shaped, 6–9(–12) μm long by 5–9 

μm wide; in surface view, quadrate with rounded corners. 

Medulla cells in transection, irregularly arranged, (45–)60–

100(–125) μm long by (17–)35–53 μm wide.

Gametophytes dioecious, gametangial sori in tufts on 

thallus surface, coalescing to form concentric bands on 

mature thalli, on both surfaces of blade. One to a number 

of plurilocular gametangia develop on a fertile filament, a 

greater number of male gametangia in a series than female 

gametangia. Female gametangia develop laterally on fertile 

filaments; initially, a protuberance forms a 2- and then 

4-celled filament that divides twice longitudinally to form 

a gametangium of 4 tiers with 4 locules per tier (16 locules) and matures acropetally. Mature female gametangia 

are ovoid, usually biseriate in lateral view, 40–55 μm long 

by 18–20 μm wide. Antheridial gametangia borne laterally 

(rarely terminal) on fertile filaments; mature antheridial 

gametangia often multiseriate in lateral view, of 16 tiers 

with 4–6 locules per tier (64–96 locules). Antheridial gametangia cylindrical, 40–60 μm long by 7–9 μm wide. The 

uniseriate fertile filament continues on above both types 

of gametangia with a sterile portion that has shorter and 

wider cells (9.5–12 μm in diameter) and more pigmented 

cells than filament cells below it.

Aglaozonia-phase of Cutleria hancockii

Sporophytic thalli (“Aglaozonia-phase”; Figure 79 

E,F) encrusting, brown to olive-brown, loosely attached, 

thin, prostrate blades, multilobed, small, up to 2 cm 

wide; loosely attached by branched, multicellular rhizoids. Lobes 2–5 mm wide, rounded; 70–85(–105) μm 

and often 6–7 cell layers thick. Thallus parenchymatous, 

arising from slightly larger, darker row of meristematic 

apical cells along the crust’s margin. Crust dorsiventral, 

cortical surface cells quadrate, 6–8 μm in diameter, lower 

cortex flanking substrate of larger cells, medulla of several 

slightly uneven layers of larger colorless cells.

Habitat. On rocks or loosely attached to various fleshy algae and corallines; epizoic on mollusk shells

and carapace of black sea turtle (Chelonia mydas agassizii); mid to low intertidal, occasionally in shallow subtidal, to 5 m depths.

Distribution. Gulf of California: Puerto

Peñasco to Isla San Pedro Nolasco; Múlege to Bahía Concepción; Bahía de La Paz to Punta Arena (Baja California 

Sur); Mazatlán, Sinaloa.

Type Locality. Gametophyte: low intertidal,

rocky reef, Isla Turners (off Isla Tiburón), Las Islas de la 

Cintura, Gulf of California. Sporophyte: southwest shore 

of Isla Tiburón (opposite Isla Turners), Las Islas de la Cintura, Gulf of California.

Remarks. Apparently a Gulf of California 

endemic, gametophytes of Cutleria hancockii in the northern Gulf occur seasonally from midwinter to midspring, 

only rarely extending into late spring (during cooler years). 

On the reef platform at Playa Estación, Puerto Peñasco, 

gametophtyes are found in the mid to low intertidal from 

January to June, sometimes with the darker-brown, lobed 

crustose “Aglaozonia-phase.” Cutleria hancockii is easily 
1 6 4 • 
n u m b e r 9 4 • 1 6 5

separated from somewhat similar looking Gulf Padina: 

Cutleria is more delicate, yellowish brown to golden light 

brown, with a fringed margin of monosiphonous filaments, whereas Padina is more rigid and darker brown, 

with an inrolled margin.

Ec

Ectocarpales Setchell et N. L. Gardner, 1922c:403.

Mesogloiales Nägeli, 1847:141, 252.

Ectocarpales Bessey, 1907:288.

Chordariales Setchell et N. L. Gardner, 1925:570.

Dictyosiphonales Setchell et N. L. Gardner, 1925:586.

Scytosiphonales Feldmann, 1949:112.

Gametophytic and sporophytic thalli exhibit a 

great diversity of size and shape. Some are erect tufts of 

unbranched or branched uniseriate filaments that are 

attached by basal rhizoids, and some are filaments that are 

entirely or partly endophytic. Others are cylindrical to compressed, foliose, saccate, or subglobose and may be simple 

and entire or branched, membranous or gelatinous in texture, with surfaces that are smooth, irregular or folded, or 

that may be perforated with holes of various sizes. Internally, the latter group are centrally solid or hollow, with a 

medulla either of colorless filaments surrounding a core of 

uniaxial or multiaxial filaments or of large, colorless parenchymatous cells. The cortex of pigmented filaments or 

small pigmented cells bears the reproductive structures and 

is with or without phaeophycean hairs. Growth is apical, 

intercalary, diffuse, or trichothallic in cortical filaments. 

Cells have a single or few to many ribbon-shaped or discoid 

chloroplasts, with one to several pyrenoids. Ultrastructural 

studies have shown a pedunculate pyrenoid with a cap layer 

in the plastids (Evans, 1966; Hori 1971; Hori and Udea, 

1975), a character unique to the Ectocarpales, and vesiclelike globules (physodes) that may be present or lacking.

Life histories, where known, may be any of four basic 

types: (1) isomorphic generations of sporophytes and 

gametophytes; (2) heteromorphic, with a large sporophyte and microscopic gametophytes; (3) heteromorphic, 

with either gametophytic or sporophytic macrothalli that 

have either a sporophytic or gametophytic crustose phase, 

respectively; or (4) some macrothalli that have direct 

development to the next phase.

Reproductive structures are terminal, lateral, or intercalary and may be either solitary or in series or in sori 

scattered over the thallus surface. Sporangia may be unilocular or plurilocular; both produce zoospores. Also, 

some have been found to have unilocular sporangia that 

produce meiospores, and a few with neutral plurilocular 

sporangia are known. Sexual reproduction is isogamous 

or anisogamous. Gametophytes develop plurilocular or 

unilocular gametangia that produce isogametes or anisogametes. Gametes in some have been shown to be capable 

of parthenogenetic development.

Remarks. The taxonomy of the Ectocarpales 

is still in flux. There has been much debate about the distinctiveness of the orders Chordariales, Dictyosiphonales, 

Mesogloiales, and Scytosiphonales and if they should be 

included in the Ectocarpales. On the basis of molecular 

analyses, many now consider these orders to be part of the 

Ectocarpales (e.g., Rousseau and de Reviers, 1999a; Silva 

and de Reviers, 2000; Peters and Ramírez, 2001; Draisma 

et al., 2003; Cho et al., 2004; Adl et al., 2005; Phillips et 

al., 2008).

There are seven families represented in the northern 

Gulf of California.

FIGURE 79. Cutleria hancockii: A–D. Gametophytes: A. Frond with concentric bands of dense reproductive filaments bearing male gametangia 

(JN-4953). B. Trichothallic growth occurs at meristems near the base of monosiphonous filaments fringing the frond margin (JN-4786). C. 

Fertile filament bearing female gametangia with large locules (JN-6040). D. Fertile filament bearing a row of male gametangia with numerous 

smaller locules (JN-7065). E, F. Sporophytes (Aglaozonia-phase of Cutleria hancockii): E. Surface view of a portion of the crust; note larger, 

darker apical cells lining margin (EYD-27365). F. Transection through dorsiventral crust (JN-3691b).

Key to the FamIlies of Ectocarpales in the n

1a. Algae of uniseriate filaments throughout or basically filamentous (mostly uniseriate, but a few cells may divide longitudinally to become parenchymatous)  

1b. Algae not filamentous; erect, terete to compressed or foliose and simple to branched or saccate or globose . . . . . . . . . . 4

2a. Filaments mostly uniseriate, becoming parenchymatous by a few longitudinal divisions; meristems random; reproductive 

structures developed within vegetative cells. . . . . . . 

2b. Uniseriate filaments throughout; meristems intercalary; reproductive structures lateral or terminal . . . . . . . . . . . . . . . . 3
1 6 6 • 

Acineto

Acinetosporaceae Hamel, 1931b:8 [as “Acinétosporacées”]; Hamel ex J. Feldmann, 1937:110.

Algae are uniseriate filaments that either are sparsely 

and irregularly branched or can be more densely branched 

and of variable branching patterns. Species grow as epiphytes or attached on hard substratum. Branches are often 

formed from the middle of an axial cell (appearing perpendicular to it) and have rounded or tapered and hairlike 

apices; some develop short, curved laterals (crampons) 

that attach to or become entangled with other algae. Intercalary meristematic growth regions are basal, usually near 

the base of laterals, or diffuse. Cells have numerous plastids, each with a pyrenoid.

Reproduction by fragmentation has been documented for 

some species (Amsler, 1984). Ovoid monosporangia, which 

are found in some, produce and release a single nonmotile 

spore. Unilocular and plurilocular sporangia are sessile, shortstalked, or intercalary (formed by direct transformation of 

vegetative cells). Plurilocular structures have been described 

with large locules (zoosporangia) in some species and with 

numerous small locules in others. Plurilocular gametangia, of 

different shapes and sizes, that produce isogametes or anisogametes are also known for many of the species.

Remarks. Life histories for some members of 

the family are not completely understood. For example, 

there is no evidence that meiosis occurs in the unilocular sporangia, nor has sexual reproduction been observed 

in the type of the family (Acinetospora pusilla (Griffiths 

ex Harvey) De Toni, 1895b; = A. crinita (Carmichael) 

Kornmann, 1953). Müller (1986) reported apomeiotic 

unilocular sporangia and suggested that the loss of sexual 

reproduction could allow the establishment of geographically isolated populations and could account for the great 

variability within species of Acinetospora.

The report of Asteronema breviarticulatum

(J. Agardh) Ouriques et Bouzon (2000; basionym: Ectocarpus breviarticulatus J. Agardh 1847) establishes the 

genus Asteronema Delépine et Asensi (1975) in the southern Gulf (Huerta-Múzquiz and Mendoza-González 1985, 

as “E. breviarticulatus”; Riosmena-Rodríguez and PaulChávez 1997; and Mateo-Cid et al. (2000a, as Hincksia breviarticulata (J. Agardh) P. C. Silva (in Silva et al., 

1987). The familial placement of the genus is problematic, 

and it has been placed in the Ectocarpaceae, Scytothamnaceae Womersley (1987) or Acinetosporaceae.

Two genera are known in the northern Gulf of 

California.

3a. Meristems localized or random on axes and branches; ribbonlike plastids with several pyrenoids . . . . . . . Ectocarpaceae

3b. Meristems mostly basal on axes and branches; discoid plastids usually with single pyrenoid. . . . . . . . 

4a. More or less cylindrical, axes and branches, narrow diameter; internally solid parenchyma (not hollow); surface of irregularly arranged cells,and numerous shallow pits with conspicuous tufts of hairs; single plastid with 1 pyrenoid; growth by a 

subapical meristem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chnoosporaceae

4b. Cylindrical to compressed and simple to branched or frondose, saccate, or globose; discoid plastids with pyrenoids; growth 

by diffuse intercalary meristems or single apical cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5a. Gametophytes erect, tubular to compressed, saccate, globose, or foliose, either open flat and entire or clathrate (with 

holes); internally, usually hollow; medulla parenchymatous, cortex of small cells; sporophytes crustose 

. . . . . . . . 

5b. Thalli variable: minute filaments or small to moderate size globose to lobed forms or cylindrical to compressed branched 

forms; internally, not hollow; medulla filamentous; cortex of filaments  

6a. Small, globose, subglobose, or lobed forms; without an evident basal layer. . . . . . . 

6b. Minute filaments, epiphytic or endophtyic, simple to branched with or without a distromatic basal layer or erect, cylindrical to compressed branched forms with a holdfast. . . . . . . 

Key to the Genera of Acinetosporaceae in th

1a. Filaments relatively simple; with branches and reproductive structures below distinct intercalary meristems in lower portion of axes; above the meristematic zone filaments long and unbranched, with distal portion often appearing hairlike. . . . . 

. . . . . 

1b. Filaments repeatedly branched, 2–4 orders; with intercalary meristematic zones and reproductive structures distributed 

throughout main axes and branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hincksi
n u m b e r 9 4 • 1 6 7

Feldmannia Ham

Feldmannia Hamel, 1939b:xli.

Algae are small, usually epiphytic, erect tufts of uniseriate filaments; each of the axes, and branches, has a 

distinct localized intercalary meristematic zone near its 

base; and are attached below by rhizoids or sometimes 

by prostrate filaments. Branches are restricted to below 

the meristematic zone. Above the meristematic zone, the 

filaments are unbranched and become long, with hair-like 

ends. Cells contain several to many discoid chloroplasts 

with one to many pyrenoids.

Life histories, where known, are isomorphic. Reproductive structures are below the meristematic zones. 

Unilocular sporangia are ovoid to spherical. Plurilocular 

sporangia and plurilocular gametangia are multiseriate, 

similar in appearance, and apparently borne on separate 

thalli. Gametes are isogamous.

Remarks. The taxonomic status of Feldmannia has been questioned by some phycologists. For 

example, Knoepffler-Péguy (1974) found some species of 

Acinetospora Bornet (1891) were involved in the life history of some species of Feldmannia and Hincksia and later 

concluded that these genera could not be convincingly 

separated on morphology and may represent a single variable genus (Knoepffler-Péguy, 1977). Later, Kim and Lee 

(1994), recognizing Feldmannia, proposed that the genus

be based solely on the characteristics seen in the illustration of Ectocarpus irregularis Kützing (1855:pl. 62: fig. 

1). Several others have also continued to recognize Feldmannia (e.g., Womersley, 1987; Silva et al., 1987; Silva 

et al., 1996a; Abbott and Huisman, 2004). Pedersen and 

Kristiansen (2001) elucidated generic characters and concluded that Acinetospora, Feldmannia, and Hincksia could 

be reliably distinguished. Life history culture studies and 

molecular analyses on the generitype, Feldmannia lebelii

(Areschoug ex P. Crouan et H. Crouan) G. Hamel (1939a; 

basionym: Ectocarpus lebelii Areschoug ex P. Crouan et H. 

Crouan, 1867), will help clarify its generic status.

One species, Feldmannia hemispherica (De A. Saunders) Hollenberg (1971), has been reported in the southern Gulf (Mateo-Cid and Mendoza-González 1994a).

There is one species known in the northern Gulf of 

California.

Feldmannia irregularis (Kützing) Ham

FIGURE 80

Ectocarpus irregularis Kützing, 1845:234; Kützing, 

1855:19, pl. 62: fig. 1 [note: illustration redrawn by 

Kim and Lee, 1994: fig. 4A]; Dawson 1954a:3; Dawson, 1961b:385; González-González et al., 1996:149; 

Yoshida, 1998:168.

Feldmannia irregularis (Kützing) Hamel, 1939b:xvii, 

fig. 61f; Cardinal, 1964:54, fig. 29A–K; Hollenberg, 

1970:61, fig. 1; Hollenberg, 1971:285; Abbott and 

Hollenberg, 1976:136, fig. 99; Stewart, 1991:39; Kim 

and Lee, 1994:154, figs. 1A,B, 2A–C, 3A–K, 4A,B; 

González-González et al., 1996:296; Abbott and Huisman, 2004:159, fig. 58A; Mateo-Cid et al., 2006:49; 

Servière-Zaragoza et al., 2007:8; Pedroche et al., 

2008:20.

Giffordia irregularis (Kützing) Joly, 1965:72, pl. VIII: figs. 

111–119.

Hincksia irregularis (Kützing) Amsler in Schneider et Searles, 1991:120, figs. 129–131.

Ectocarpus mucronatus De A. Saunders, 1898:152, pl. 19; 

Setchell and Gardner, 1925:429; Dawson, 1944:220; 

Dawson, 1961b:385; Dawson, 1966a:9; GonzálezGonzález et al., 1996:149.

Algae small, slender tufted thalli, up to 3 mm tall, of 

erect, loosely intertwined filaments that become elongated 

FIGURE 80. Feldmannia irregularis: Drawing of type specimen 

(from Kützing, 1855: pl. 62, fig. 1).
1 6 8 • 

and taper slightly toward apices (terminal hairlike appearance due to elongated cells with fewer chloroplasts); 

attached below by a prostrate system of entangled rhizoidal 

filaments. Axes often sparse and irregularly branched several times below the intercalary meristematic zone (meristem shifts upward toward the lower-middle portion of the 

filament as it grows, obscuring its original basal position). 

Cells 18–20 μm in diameter; 0.5–1.0 times as long as wide 

in meristematic zone; 1–3 times as long as wide in lower 

portions and 2–5 times as long as wide in upper portions.

Unilocular sporangia unknown in Gulf material. Plurilocular sporangia sessile on branches below meristematic 

zone; elongate-conical, 90–150 μm long and 20–35 μm in 

diameter, often with attenuated apices.

Habitat. Epizoic on the black sea turtle (Chelonia mydas agassizii; see Parham and Zug, 1996), epiphytic

on Sargassum and probably other algae, and growing on 

rocks; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco; Canal de Infiernillo (between coast of Sonora and 

Isla Tiburón). Pacific coast: British Columbia to Oregon; 

southern California to Baja California; Isla San Benedicto 

(Islas Revillagigedo); Hawaiian Islands; Japan; China.

Type Locality. On Laurencia obtusa, Adriatic Sea

Remarks. Feldmannia irregularis has a wide 

distribution (Guiry and Guiry, 2008), but its generic 

placement has been somewhat uncertain, in part because 

the type specimen of Ectocarpus irregularis Kützing 

(1845) was not preserved and the original description 

was vague (Clayton, 1974). This uncertainty has resulted 

in the species being variously interpreted as belonging to 

other genera; e.g., Womersley (1987) noted that Australian F. irregularis exhibited intermediate characteristics of 

both Feldmannia and Hincksia, and Amsler (in Schneider 

and Searles, 1991) considered it to be Hincksia irregularis. 

However, studies of Kim and Lee (1994) support recognition of Feldmannia irregularis.

Life history culture studies and comparative molecular analyses on Gulf of California specimens referred to F. 

irregularis are needed to elucidate their taxonomic status 

and phylogenetic relationships.

Hincksia J. E. Gr

Hincksia J. E. Gray, 1864:12.

Giffordia Batters, 1893:86; Hamel, 1939b:X, as “Gifforidia (Batters) Hamel.”

Algae are erect, tufts of uniseriate filaments with variable branching patterns, and attached below by rhizoids. 

Branching is to 2–3(–4) orders and unilaterally secund 

or tending to spiral or irregular, alternate, or opposite. 

Growth is diffuse by intercalary cell divisions. Meristems 

are scattered throughout the axes and near the base of 

most branches, with additional meristems distally. Meristematic zones are not always obvious but in some species are very distinct. Cells are more or less isodiametric 

to barrel-shaped and have numerous discoid chloroplasts, 

each with a single pyrenoid. True phaeophycean hairs are 

unknown.

Life histories are mostly of isomorphic alternating 

generations; however, some have repeating generations 

via spores or parthenogenic gametes. Reproductive structures are usually sessile and singular or in a series on the 

branches. Unilocular sporangia are apparently rare. Plurilocular sporangia are formed on the adaxial side of the 

supporting lateral branch and may also occur on the main 

axis. Plurilocular gametangia (where known) are of different sizes and shapes and produce anisogametes.

Remarks. One species is reported in the southern Gulf, Hincksia rallsiae (Vickers) P. C. Silva (in Silva et 

al., 1987), by Mateo-Cid et al. (2000a).

Three species are known in the northern Gulf of 

California.

Key to the Species of Hincksia in the no

1a. Filaments occasionally to frequently oppositely branched; plurilocular sporangia sessile, asymmetrically ovoid 

. . . . . .

1b. Filaments not oppositely branched, mostly dichotomous, alternate, unilaterally secund, or spiral; plurilocular sporangia 

sessile or pedicellate and fusiform or ellipsoidal to cylindrical  

2a. Branching sparse, dichotomous below (nearly at algal host’s surface cells); apical cells blunt; plurilocular sporangia sessile 

or pedicellate, fusiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. bryanti

2b. Branching alternate or unilaterally secund or spiral; apices attenuated; plurilocular sporangia sessile and ellipsoidal to 

cylindrical with rounded apices  . . . . . . 
n u m b e r 9 4 • 1 6 9

Hincksia bryantii (Setchell et N. L. Gardne

J. N. Norris, comb. nov.

FIGURE 81C

Ectocarpus bryantii Setchell et N. L. Gardner, 1924a:720, 

pl. 17: fig. 45; Setchell and Gardner, 1925:440; Dawson, 1944:220; Dawson, 1961b:384; Dawson, 1966a:9; 

Dawson, 1966b:55; Huerta-Múzquiz and MendozaGonzález, 1985:44; Rocha-Ramírez and Siqueiros-Beltrones, 1991:24; Espinoza-Ávalos, 1993:333; GonzálezGonzález et al., 1996:147; Paul-Chávez and RiosmenaRodriguez 2000:146; Mateo-Cid et al., 2006:49; Pedroche et al. 2008:16.

Algae tufted epiphytes, up to 2.5 mm high, of 

intertwined, uncorticated filaments, attached below by 

penetrating rhizoidal filaments. Branching below dichotomous (nearly at surface of algal host cells); above sparse 

with very few short, slightly tapering branchlets terminating in a blunt cell and lacking terminal hairs. Cells 

28–64 μm long by 28–32 μm in diameter. Chloroplasts 

discoid.

Unilocular sporangia unknown. Plurilocular sporangia scattered along filaments; sessile or pedicellate on 1 

cell; narrow to broadly fusiform, 70–100(–140) μm long 

by 25–35 μm wide.

Habitat. Epiphytic on Codium brandegeei and

C. simulans; mid to low intertidal.

FIGURE 81. Two species of Hincksia. A, B. Hincksia granulosa: A. Habit, with lateral branches predominately opposite (from Saunders, 1898: 

pl. 24, fig. 3). B. Plurilocular sporangia sessile, asymmetrically ovoid, borne on lateral branches (from Saunders, 1898: pl. 24, fig. 4). C. Hincksia 

bryantii: Plurilocular reproductive structures; varying from narrow to broadly fusiform and either sessile or borne on one-celled pedicel (after 

Setchell and Gardner, 1924a: pl. 17, fig. 45).
1 7 0 • 

Distribution. Gulf of California: Puerto

Peñasco; Isla Espíritu Santo to Bahía de La Paz.

Type Locality. Epiphytic on Codium brandegeei, La Paz, Baja California Sur, Gulf of California,

Mexico.

Remarks. The discoid chloroplasts and 

branching pattern of Ectocarpus bryantii Setchell et N. 

L. Gardner (1924a) suggest that it more properly belongs 

to Hincksia, and a new combination is proposed. Setchell 

and Gardner (1924a, as E. bryantii) noted that Hincksia 

bryantii could be related to the Gulf Ectocarpus gonodioides but it differed in its mode of branching. Further collections are needed to elucidate the generic placement and the 

phylogenetic relationship of H. bryantii to other species.

Hincksia granulosa (J. E. Smith) P. C. S

FIGURE 81A,B

Conferva granulosa J. E. Smith, 1811:pl. 2351.

Hincksia granulosa (J. E. Smith) P. C. Silva in Silva, Meñez et Moe, 1987:130; Stewart, 1991:39; Kim and Lee, 

1992:252, figs. 7A–D, 8E–H; Hoffmann and Santelices, 

1997:116, fig. 27; Yoshida, 1998:171; R. Aguilar-Rosas et al., 2005b:34; Pedroche et al., 2008:21.

Ectocarpus granulosus (J. E. Smith) C. Agardh, 1824:163; 

C. Agardh, 1828:45; De A. Saunders, 1898:156, pl. 

24: figs. 3–5; Setchell and Gardner, 1925:426; Smith, 

1944:81, pl. 11: figs. 1, 2; Dawson, 1945b:64; Dawson, 

1960a:36; Dawson, 1961b:385; Dawson, 1962c:278; 

Devinny, 1978:358; Mendoza- González and MateoCid, 1986:420; González-González et al., 1996:149.

Giffordia granulosa (J. E. Smith) G. Hamel 1939b:xv, fig. 

61E; Cardinal, 1964:39, figs. 19A–J, 20A–J; Abbott and 

Hollenberg, 1976:140, fig. 103; Fletcher, 1981:211; Pacheco-Ruíz, 1982:70; Mendoza-González and MateoCid, 1986:420; Santelices, 1989:267, pl. DD: figs. 9–11; 

González-González et al., 1996:151.

Algae tufted, up to 2.5 cm high, intertwined filaments; 

lower axes of filaments corticated, upper portions uncorticated; attached below by penetrating rhizoidal filaments. 

Lateral branches predominately opposite, with short ultimate branchlets; branches and branchlets taper distally 

and terminate in a blunt apical cell and lack terminal hairs. 

Cells mostly shorter than wide, 35–80 μm long, 50–80(–

100) μm in diameter; with numerous discoid chloroplasts.

Unilocular sporangia unknown in Gulf material 

(reported elsewhere to be globose). Plurilocular sporangia adaxial along branches and branchlets; sessile, broadly 

ovate, asymmetrical, 45–60(–100) μm long by 40–60 μm 

in diameter.

Habitat. Epiphytic on Codium, also probably

on other algae and on rocks; intertidal.

Distribution. Gulf of California: Isla Pelícano, Bahía Kino. Pacific coast: Alaska to Baja California;

Chile; Japan.

Type Locality. Brighthelmston (Brighton)

and Shoreham, Sussex, England.

Remarks. Description of Hincksia granulosa

is based primarily on Abbott and Hollenberg (1976) since 

I have not seen the Sonoran specimens reported by Mendoza-González and Mateo-Cid (1986).

Hincksia mitchelliae (Harvey) P. C. Si

FIGURE 82

Ectocarpus mitchelliae Harvey, 1852:142, pl. 12G: figs. 

1–3; D. A. Saunders, 1898:153, pl. 21: figs. 1, 2; Setchell and Gardner, 1925:428; Setchell and Gardner, 

1930:140; Dawson, 1944:220; Dawson, 1954:5; Dawson, 1961b:385; Dawson, 1966a:9.

Hincksia mitchelliae (Harvey) P. C. Silva in Silva, Meñez et 

Moe, 1987:73; R. Aguilar-Rosas and Machado-Galindo, 

1990:188; Stewart, 1991:39; Kim and Lee, 1992:246, 

figs. 3A–D, 5A–D; González-González, 1993:443; 

León-Tejera and González-González, 1993:497; Stout 

and Dreckmann, 1993:6; Mendoza-González et al., 

1994:110; González-González et al., 1996:153; Mendoza-González and Mateo-Cid, 1996:70, 85, pl. 18: 

figs. 75–77; Hoffman and Santelices, 1997:119, fig. 28; 

Yoshida, 1998:171; L. Aguilar-Rosas et al., 2000:132; 

León-Tejera and González-González, 2000:328; Mateo-Cid and Mendoza-González, 2003:21; Abbott and 

Huisman, 2004:162, fig. 59C; R. Aguilar-Rosas et al., 

2005b:34; Mateo-Cid et al., 2006:49; Dreckmann et al., 

2006:154; Servière-Zaragoza et al., 2007:8; Pedroche et 

al., 2008:22.

Giffordia mitchelliae (Harvey) Hamel, 1939a:66; Hamel, 

1939b:XIV, fig. 61c,d; Cardinal, 1964:45, fig. 23A–H; 

Huerta-Múzquiz and Tirado-Lizárraga, 1970:127; Abbott and Hollenberg, 1976:143, fig. 105; L. Aguilar-Rosas, 1982:84; Schnetter and Bula Meyer, 1982:55, pl. 

8: figs. C, D; Stewart and Stewart, 1984:141; HuertaMúzquiz and Mendoza-González, 1985:44; MendozaGonzález and Mateo-Cid, 1985:24; Mendoza-González 

and Mateo-Cid, 1986:420; Sánchez-Rodríguez et al., 

1989:40; Mateo-Cid et al., 1993:49; González-González 

et al., 1996:296.
n u m b e r 9 4 • 1 7 1

Algae tufted, 1–4 cm tall, often densely branched in 

mid to upper portions; attached to host by creeping rhizoidal filaments. Filaments of main axis up to 50 μm in diameter; branching alternate or unilaterally secund or spiral; 

apices often attenuated. Intercalary meristematic zones 

scattered along axes and near base of branches. Cells 1–3 

times longer than wide; 20–120 μm long by 20–40 μm in 

diameter; with numerous discoid plastids.

Unilocular sporangia not seen in Gulf material 

(reported elsewhere as sessile, ovoid to elliptical, 40–45 

μm long by 15–20 μm in diameter). Plurilocular sporangia sessile on main axes and adaxial on lateral branches, 

often in short series (2–5); more or less cylindrical, tapering slightly with rounded apices, (40–)60–150 μm long by 

30–45 μm in diameter; with 1–3(–4) locules in each horizontal row (seen in lateral view).

Habitat. Epiphytic on Sargassum and Ralfsia,

occasionally on other algae, and epizoic on the black sea 

turtle, Chelonia mydas agassizii (see Parham and Zug, 

1996); mid intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Canal de Infiernillo (between Isla Tiburón and 

Campo Ona, Sonora); Bahía Concepción to San José del 

Cabo. Pacific coast: central California to Sinaloa; Isla San 

Benedicto (Islas Revillagigedo); Michoacán to Chiapas; 

Colombia; Chile; Hawaiian Islands; Taiwan; Korea; Japan.

Type Locality. Nantucket, Nantucket

County, Massachusetts.

Remarks. The northern Gulf collection of 

Hincksia mitchelliae (US Alg. Coll.-158145), growing on 

the carapace of a black sea turtle in the Canal de Infiernillo 

(between Campo Ona, Sonora, and Isla Tiburón), extends 

its known distribution south from Puerto Peñasco (Dawson, 1966a). Southern Gulf collections of Dawson (1959a, 

as “E. sp. aff. E. mitchelliae”) were also referred to this 

species, but with doubt.

Chnoo

Chnoosporaceae Setchell et N. L. Gardner, 1925:400, 

552.

Algae are composed of terete to compressed axes and 

usually subdichotomously divided branches above a crustose or discoid holdfast. Internally they have a solid, pseudoparenchymatous structure of elongate, colorless cells in 

the medulla that is developed from a subapical meristem 

below the outer cortical layer. The cortex is composed of 

one to two layers of small, pigmented, irregularly arranged 

cells and numerous shallow pits with conspicuous tufts of 

colorless hairs (cryptoblasts). Each cortical cell has a single chloroplast with one pyrenoid.

Life histories are apparently heteromorphic but are 

not fully known for most of the species. An independent 

crustose phase that repeated itself was described by Fotos 

(1981), and a heteromorphic life history was reported in 

culture studies of C. implexa J. Agardh (1848) from Japan 

(Kogame, 2001). Unilocular and plurilocular reproductive structures, both sporangia (zoidangia), are known in 

the sporophytic crustose phase of one species. Plurilocular 

structures are found in the erect phase and either develop 

in association with the colorless hairs or sometimes may 

be separate from these hairs.

Remarks. Although some consider the Chnoosporaceae to be a synonym of the Scytosiphonaceae (e.g., 

Kogame, 2001), others continue to recognize it as a distinct family (e.g., Silva et al., 1996a; Yoshida, 1998; 

Abbott and Huisman, 2004), primarily on the basis of 

its members having vegetative cells that contain a single 

chloroplast with one pyrenoid. Molecular analysis of C. 

pacifica J. Agardh (lectotype of the genus) will help elucidate the taxonomic status and phylogenetic relationships 

of this monotypic family.

FIGURE 82. Hincksia mitchelliae: Filaments with intercalary meristem (arrow) at base of branch and several sessile, ellipsoid to cylindrical plurilocular sporangia with rounded apices, borne on adaxial 

side of branches, often arranged in short series (JN-4774).
1 7 2 • 

The family’s single genus, Chnoospora, occurs in the 

northern Gulf of California.

Chnoospora J. Agar

Chnoospora J. Agardh, 1847:7.

Gametophytes are erect tufts, clumps, or partly 

decumbent mats, and composed of one to several, more 

or less terete to compressed axes arising from a crustose 

holdfast. Axes are dichotomously, subdichotomously, or 

irregularly branched. Growth is in the subapical meristematic row of cells. Anatomically, the medulla is solid, 

and pseudoparenchymatous, composed of large and small 

colorless medullary cells, and the cortex is of one to two 

layers of smaller pigmented cells. The surface layer is of 

irregularly arranged cortical cells and numerous shallow 

pits from which groups of colorless hairs extend. Each 

cortical cell has a single chloroplast and a pyrenoid.

Unilocular and plurilocular structures (zoidangia) are 

developed on discoid sporophytes for at least one species (Kogame, 2001) and have been reported as “densely 

packed around groups of hairs” to form “more or less 

continuous sori” (Lawson and John, 1982:134). Plurilocular gametangia that produce isogametes are developed on 

the erect thalli in superficial sori or may be associated with 

colorless hairs on the outer surfaces.

Remarks. The generitype of Chnoospora was 

not specifically designated by Agardh (1847), and Setchell 

and Gardner (1925) were apparently the first to choose a 

lectotype, C. pacifica J. Agardh, for the genus.

Reproductive structures in Chnoospora were first 

described by Barton (1898) and Børgesen (1924) and later 

more completely described by Fotos (1981) and Kogame 

(2001), who also found a repeating crustose phase in culture studies. In the most complete life history study, erect 

dioecious gametophytes of Japanese C. implexa produced 

isogametes in plurilocular gametangia that formed zygotes, 

which grew into crustose sporophytes. Under long-day 

photoperiod conditions the latter developed plurilocular 

zoidangia that produced sporophytic crusts repeatedly, 

whereas under short-day regimes the crusts developed unilocular zoidangia that grew into the dioecious, erect gametophytes (Kogame, 2001). Some strains of cultured C. 

implexa exhibited a direct life history with gametophytic 

thalli produced without sporophytes (Kogame, 2001).

Three other species are reported in the southern Gulf 

and Pacific Mexico: C. implexa J. Agardh (1848) by Dawson (1959a, 1959b, 1960c,1961b); C. minima (K. Hering) Papenfuss (1956) by Chávez B. (1972), Pedroche 

and González-González (1981), Huerta-Múzquiz and 

Mendoza-González (1985), León-Tejera and GonzálezGonzález (1993), Stout and Dreckmann (1993), MateoCid et al. (1993), and Servière-Zaragoza et al. (1993); 

and C. pannosa J. Agardh (1848) from Isla Guadalupe 

(Setchell and Gardner, 1930; Stewart and Stewart, 1984).

One species is reported in the northern Gulf of 

California.

Chnoospora pacifica J. Agar

FIGURE 83A–C

Chnoospora pacifica J. Agardh, 1847:7; Kützing, 1859:36, 

pl. 86: fig. 1; Setchell and Gardner, 1924a:728; Setchell 

and Gardner, 1925:553; Okamura, 1936:236; Dawson, 

1944:234; Taylor, 1945:82; Dawson, 1954a:4, pl. 3: 

fig.1; Huerta-Múzquiz and Tirado-Lizárraga, 1970:127; 

Huerta-Múzquiz, 1978:339; Mendoza-González and 

Mateo-Cid, 1986:421; Salcedo-Martínez et al., 1988:82; 

León-Tejera and González-González, 1993:497; Servière-Zaragoza et al., 1993:482; González-González et 

al., 1996:141; Hernández-Herrera et al., 2005:146.

Chnoospora fastigiata α [var.] pacifica (J. Agardh) J. Agardh, 

1848:171–172; Børgesen, 1924:263, figs. 11, 12.

Chnoospora minima sensu Dawson 1959b:8; Mateo-Cid 

et al. 1993:49; Pedroche et al. 2008:63 [non Chnoospora minima (K. Hering) Papenfuss 1956:69].

Algae erect, up to 6 cm tall, of few to many more 

or less cylindrical to compressed branched axes, up to 2 

mm in diameter, arising from a basal crustose holdfast. 

Branched dichotomously to subdichotomously; branches 

of more or less the same diameter throughout, tapering to 

slightly smaller apices. Surface with irregularly arranged 

cortical cells and scattered, noticeable groups of colorless hairs, up to 3 mm long, extending from shallow pits. 

Medulla of thick-walled, small to large, colorless cells; 

cortex of 1–2 layers of smaller pigmented cells.

Plurilocular uniseriate and biseriate structures, up to 

80 μm long, develop in long sori on branch surfaces.

Habitat. Intertidal

Distribution. Gulf of California: Bahía San

Pedro (north of San Carlos); Bahía Concepción to Bahía de 

La Paz. Pacific coast: Isla San Benedicto (Islas Revillagigedo); 

Isla María Magdalena (Las Islas Tres Marías); Nayarit to 

Oaxaca; Hawaiian Islands; Rapa Nui (Easter Island).

Type Locality. San Agustín, Oaxaca, Mexico (Agardh, 1847, “St. Augustin, Stilla Oceanen [Pacific

Ocean]”).
n u m b e r 9 4 • 1 7 3

Remarks. Described from southern Oaxaca, 

Pacific Mexico (Agardh, 1847), Chnoospora pacifica was 

first reported in the southern Gulf by Setchell and Gardner 

(1924a). Subsequently, Dawson (1944) noted its absence 

from further Gulf collections and suggested it may be a 

strictly tropical species, limited to the southernmost Gulf 

(Dawson, 1961b). More recently, C. pacifica was reported 

from Sonora in the northern Gulf by Mendoza-González 

and Mateo-Cid (1986); although I have not seen their 

specimens, Chnoospora is very distinctive, with characteristic scattered pits in the thallus surface from which tufts 

of colorless hairs extend (Figure 83B,C).

Some have considered the Pacific Mexico C. pacifica to 

be a synonym of the South African C. minima (K. Hering) 

Papenfuss (1956; cf. Levring, 1938:22, fig. 10, pl. 4: fig. 

12, as “C. pacifica”; basionym: Fucus minimus K. Hering, 

1841). Others have continued to recognize them as separate species; e.g., Servière-Zaragoza et al. (1993) reported 

both occurring in Jalisco and Nayarit, Pacific Mexico. 

Until the types and type locality materials of C. pacifica

(generitype) and C. minima have been reexamined and 

tested using molecular and phylogenetic analyses and 

compared to the Gulf of California C. pacifica, it seems 

best to recognize them as separate species.

FIGURE 83. Chnoospora pacifica, EYD-12121: A. Habit, air-dried herbarium specimen. B. Liquid-preserved specimen showing scattered 

groups of hairs from shallow pits. C. Close-up of clumps of hairs extending from superficial pits.
1 7 4 • 

Chord

Chordariaceae Greville, 1830:44.

Dictyosiphonaceae Kützing, 1849:484 as “Dictyosiphoneae.”

Elachistaceae Kjellman, 1890:41.

Myrionemataceae Nägeli, 1847:145, 252 as “Myrionemeae.”

Myriotrichiaceae Kjellman, 1890:46.

Punctariaceae (Thuret) Kjellman, 1880:9; Punctarieae Thuret in Le Jolis, 1863:14, 21.

Spermatochnaceae Kjellman, 1890:32.

Algae are microscopic or macroscopic, erect or partly 

erect, and variable in size and shape, ranging from (1) simple to branched filaments or tufts, (2) cushion-shaped to 

irregularly lobed forms, or (3) cylindrical to compressed, 

simple, entire, or branching forms, attached by basal filaments. Some are membranous or gelatinous in texture. 

Some of the species are filamentous or have a filamentous 

form in their life history. Other species are structurally 

differentiated, with a medulla composed of filaments that 

surround a central axis (core) of one or more colorless filaments, each of which grows from an apical cell; internally, 

they may be either solid or hollow when mature. Cells 

divide radially from the central medullary filaments and 

give rise to whorls of outer assimilatory filaments. These 

assimilatory filaments bear reproductive structures, and 

hairs or paraphyses may also be present or absent. Growth 

may be apical, marginal, diffuse, or intercalary in cortical 

filaments. Cells have one to several discoid, elongate, or 

irregular chloroplasts and usually one pyrenoid.

Life histories, where known, are mostly heteromorphic, usually with a macroscopic pseudoparenchymatous 

sporophyte of uniaxial or multiaxial structure and a usually smaller or microscopic filamentous gametophytic 

phase. Sporophytes produce unilocular sporangia, which 

are borne at the base of paraphyses. Plurilocular structures are uniseriate or multiseriate and may be sessile or 

pedicellate or borne at the base of paraphyses or on special 

branches growing from the outer cortical cells. Gametophytes are filamentous and produced unilocular or plurilocular gametangia. Gametes are usually of unequal size.

Remarks. On the basis of molecular phylogenies, Peters and Ramírez (2001) considered the Chordariaceae to include several families (see those listed above as 

synonyms). In addition to the genera known in the northern Gulf, another, Haplogloia Levring (1939), has been 

reported in the southern Gulf by Mateo-Cid et al. (1993, 

as H. andersonii (Farlow) Levring).

There are three genera represented in the northern 

Gulf of California.

Key to the Genera of Chordariaceae in the 

1a. Erect, terete, narrow axis and branches (up to 1.0 mm in diameter), up to 20 cm or more in length; more or less alternately 

to irregularly branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemacystu

1b. Much smaller, microscopic to minute uniseriate filaments, less than 3 mm tall and 5–15 μm in diameter  

2a. Sparingly branched filaments, up to 0.5 mm tall, above a discoid base of laterally adjoined prostrate filaments which become mostly distromatic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hecatonem

2b. Uniseriate filaments entirely to partly endophytic, up to 3 mm in length; without a discoid base . . . . . . . . . Streblonema

Hecatonema Sauvageau

Hecatonema Sauvageau, 1897:248.

Algae are minute, discoid to irregularly shaped or 

cushion-shaped filamentous, crustose thalli. Structually, 

they are composed of a mostly distromatic basal layer of 

laterally adjoined, radiating filaments. Most cells of these 

basal filaments grow parallel to the substratum or, if epiphytic, on the surface of the host. Longitudinal divisions 

of the basal filament cells occur inconsistently, thus forming a partly distromatic disc; other cells of the filaments 

may divide in an intercalary manner or marginally. Some 

of the lower filament cells may produce rhizoids that penetrate the cells of the host alga; the upper cells develop 

an erect, simple, unbranched or branched filament or an 

endogenous hair or a hyaline paraphysis-like structure 

(ascocyst) or a reproductive structure. Cells contain one to 

several discoid chloroplasts and one pyrenoid.

Unilocular sporangia, where known, are usually sessile or sometimes pedicellate on the basal layer. Plurilocular sporangia are multiseriate, variable in shape, sessile, 

and terminal or lateral.

Remarks. The taxonomic status and placement of the genus Hecatonema needs further investigation. Culture studies have shown some Hecatonema-like 
n u m b e r 9 4 • 1 7 5

microthalli to produce macrothalli that are other genera, 

e.g., Asperococcus Lamouroux (1813), Desmotrichum 

Kützing (1845), Myriotrichia Harvey (1835), or Punctaria

Greville (1830) (see Loiseaux, 1967, 1969; Clayton, 1974; 

Lockhart, 1982; Fletcher, 1984; Pedersen, 1984); therefore, some phycologists have placed Hecatonema in the 

Punctariaceae (Thuret) Kjellman (1880; e.g., Christensen, 

1980; Fletcher, 1987; Womersley, 1987; Wynne, 1998a). 

However, other species of Hecatonema have been shown 

to have an isomorphic life history and do not develop macrothalli in culture (e.g., Clayton, 1974; Pedersen, 1984). 

These species have been placed in the Myrionemataceae 

(e.g., Abbott and Hollenberg, 1976; Schneider and Searles, 1991; Silva et al., 1996a). Most recent investigations 

consider both the Myrionemataceae and Punctariaceae as 

members of a more broadly defined Chordariaceae (e.g., 

Peters and Ramírez, 2001; Wynne, 2005).

Easily overlooked, future studies of these microscopic 

to minute filamentous brown algae will probably reveal 

more species in the Gulf of California.

Currently there is one species known in the northern 

Gulf of California.

Hecatonema streblonematoides

(Setchell et N. L. Gardner) Lois

FIGURE 84A,B

Compsonema streblonematoides Setchell et N. L. Gardner, 1922a:353, pl. 35: fig. 4; Setchell and Gardner, 

1925:481; Dawson, 1961b:391.

FIGURE 84. Hecatonema streblonematoides: Upper portions of filaments from a dense tuft (JN-4033). A. Two filaments with terminal plurilocular gametangia (arrows). B. Plurilocular gametangium with short lateral protuberances (arrow).
1 7 6 • 

Hecatonema streblonematoides (Setchell et N. L. Gardner) 

Loiseaux, 1970b:253, figs. 2, 3; Abbott and Hollenberg, 1976:162, fig. 130; Stewart, 1991:40; GonzálezGonzález et al., 1996:153; Pedroche et al., 2008:62.

Compsonema myrionematoides Setchell et N. L. Gardner, 1922a:361, pl. 36: fig. 1; Setchell and Gardner, 

1925:483; Smith, 1944:109; Dawson, 1961b:390.

Compsonema nummuloides Setchell et N. L. Gardner, 

1922a:359, pl. 35: figs. 5, 6; Setchell and Gardner, 

1925:475; Dawson, 1961b:391.

Compsonema pusillum Setchell et N. L. Gardner, 

1922a:356, pl. 37: fig. 3; Setchell and Gardner, 

1925:477, pl. 54: fig. 3; Smith, 1944:108; Dawson, 

1961b:391.

Compsonema ramulosum Setchell et N. L. Gardner, 

1922a:362, pl. 39: figs. 1–5; Setchell and Gardner, 

1925:486; Smith, 1944:111; Dawson, 1961b:391.

Compsonema secundum Setchell et N. L. Gardner, 

1922a:361, pl. 37: figs. 1, 2; Setchell and Gardner, 

1925:483, pl. 54: figs 1, 2; Smith, 1944:109, pl. 15: 

fig. 4; Dawson, 1961b:391; Servière-Zaragoza et al., 

1993:482; González-González et al., 1996:374; Pedroche et al., 2008:69.

Compsonema secundum f. terminale Setchell et N. L. 

Gardner, 1922a:366, pl. 37: figs. 4, 5; Setchell and 

Gardner, 1925:484, pl. 54: figs. 4, 5; MendozaGonzález and Mateo-Cid, 1986:420; Servière-Zaragoza 

et al., 1993:482; González-González et al., 1996:143, 

374; Yoshida, 1998:269; Mateo-Cid et al., 2006:58; Pedroche et al., 2008:69.

Compsonema speciosum f. piliferum Setchell et N. L. 

Gardner, 1922a:356, pl. 38: figs. 1, 2; Setchell and 

Gardner, 1925:485; Dawson, 1961b:391.

Hecatonema clavatum Setchell et N. L. Gardner, 

1922a:378, pl. 40: figs. 1–4; Setchell and Gardner, 

1925:491; Dawson, 1961b:391.

Hecatonema variabile Setchell et N. L. Gardner, 

1922a:377, pl. 41: figs. 1–12; Setchell and Gardner, 

1925:490; Smith, 1944:113; Dawson, 1961b:391.

Algae minute tufts form a discoid to irregular cushion, 2–10 mm in diameter; basal layer mostly distromatic, 

and 1 cell layer at margins; attached below by rhizoids; 

erect filaments unbranched, mostly 100–500 μm tall 

(older specimens reported up to 1000 μm tall and with 

secund branching). Filaments of cylindrical cells, 5–7 μm 

in diameter basally and 10–12(–15) μm above. Cells with 

(1–)2–4(–5) chloroplasts. Endogenous hairs, up to 7 μm in 

diameter; rare in juveniles, abundant in mature specimens. 

Ascocysts rare, sessile or lateral on erect filaments, 20–50 

μm long by 10–15 μm in diameter.

Unilocular sporangia erect on basal filaments, sessile, lateral or pedicellate; 40–80 μm long by 15–30 μm 

in diameter. Plurilocular structures very variable in shape 

and position; sessile, 40 μm long and 12 μm in diameter, 

or pedicellate, 40–180 μm long by 12–17 μm in diameter, 

or lateral or in long terminal rows on erect filaments, up 

to 400 μm in length (sometimes resembling a cockscomb; 

see Setchell and Gardner, 1925:pl. 54: fig. 1).

Habitat. Epiphytic on Padina durvillei and P.

caulescens; mid to low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Bahía Kino. Pacific coast: Alaska to central 

California; Baja California; Jalisco to Nayarit; Japan.

Type Locality. On pneumatocyst of Nereocystis luetkeana, Tomales Bay, Marin County, California

Remarks. Hecatonema streblonematoides has 

been recorded from the Sonora coast at Puerto Peñasco 

(Loiseaux, 1970b) and in the vicinity of Bahía Kino at Isla 

Pelícano and Segundo Cerro Prieto (Mendoza-González 

and Mateo-Cid, 1986, as “Compsonema secundum f. 

terminale”). The form of H. streblonematoides we have 

collected (Playa Hermosa, vicinity of Puerto Peñasco, 

JN-4033; US Alg. Coll. slide 5350) closely resembles those 

originally described as Compsonema secundum (see Setchell and Gardner, 1925:pl. 54: fig. 2).

Smith (1944, as “C. secundum”) noted its sporangia 

were so variable that it was impossible to distinguish the 

type species of C. secundum from the forma, C. secundum

f. terminale. On the basis of culture and morphological 

studies, Loiseaux (1970b) concluded H. streblonematoides

was a single species that included C. secundum and several 

other northeastern Pacific taxa previously placed in Compsonema or Hecatonema. The species encompassed within 

the broadly taxonomically defined H. streblonematoides

(Loiseaux, 1970b) need to be molecular phylogenetically 

and morphologically tested.

Nemacystus Derbès et Solier

Nemacystus Derbès et Solier, 1850:269.

Algae are erect, lax, with cylindrical narrow axes, 

400–1000 μm in diameter, usually irregularly branched, 

and attached by a small discoid holdfast. The single axial 

filament of the medulla (usually evident in the upper portions) grows by divisions of a dome-shaped apical cell. 

Surrounding the axial filament are large pseudoparenchymatous cells that develop from filaments growing downward from the base of the assimilatory filaments. The 

outermost pseudoparenchymatous cells bear pigmented 

assimilatory filaments that are simple or branched at their 
n u m b e r 9 4 • 1 7 7

bases, straight or curved, and up to 20 cells in length. Colorless phaeophycean hairs develop from basal cells of the 

assimilatory filaments, and together with the assimilatory 

filaments, they cover the surface, giving it a hairy or furry 

appearance under low magnification.

Life histories, where known (e.g., Migita and Yotsui, 

1972; Yotsui, 1980), are either direct from spores of plurilocular sporangia or heteromorphic, with macroscopic 

sporophytes and microscopic gametophytes. Sporophytes 

produce two kinds of sporangia: unilocular, ovoid to 

pyriform sporangia that are borne at base of assimilatory 

filaments and plurilocular sporangia that are uniseriate 

and borne on simple or sometimes branched pedicels at 

the base of assimilatory filaments (Yotsui, 1976, 1980). 

Microscopic “streblonematoid” gametophytes, known 

in some species (e.g., N. decipiens (Suringar) Kuckuck, 

1929), bear few-celled, uniseriate gametangia that produce isogametes (Peters and Müller, 1986a; Peters, 1987).

There is one species, originally described from the 

southern Gulf, and known to occur throughout the Gulf 

of California.

Nemacystus brandegeei (Setchell et N. 

Gardner) Kylin

FIGURE 85A–C

Meneghiniella brandegeei Setchell et N. L. Gardner, 

1924b:5; Setchell and Gardner, 1925:549, pl. 47: fig.11, 

pl. 49: fig.16.

Nemacystus brandegeei (Setchell et N. L. Gardner) Kylin, 1940:49; Dawson, 1944:232; Dawson, 1949:230; 

Dawson, 1959a:19, fig. 2; Dawson, 1961b:392; Hommersand, 1972:70; Norris, 1973:6; Huerta-Múzquiz 

and Mendoza-González, 1985:46; González-González 

et al., 1996:155; de Jong and Prud’homme van Reine, 1997:6; de Jong, 1999:88; L. Aguilar-Rosas et al., 

2000:132; Mateo-Cid et al., 2006:58; Pacheco-Ruíz et 

al., 2008:204; Pedroche et al., 2008:62.

Haplogloia andersonii sensu Norris, 1973:6; PachecoRuíz et al., 2008:204 [non Haplogloia andersonii (Farlow) Levring, 1939:50; basionym: Mesogloia andersonii

Farlow, 1889:6].

Algae soft, dark-brown, slender cylindrical main axes, 

irregularly and alternately branched, mostly up to 20 cm 

(to 1.0 m) long and up to 500–600(–1000) μm in diameter. 

Main axis and branches develop from a single apical cell; 

initial axial filament more evident in upper portions. Internally of large, thin-walled, colorless, pseudoparenchymatous cells; innermost up to 100 μm wide by up to 400 μm 

long; outward mostly 45 μm by 125 μm; outermost cells 

36 μm by 65 μm. Assimilatory filaments up to 140(–160) 

μm in length, composed of 6–12 cells with discoid plastids; basal cells smaller, 5–7 μm in diameter, distal cells 

relatively larger and moniliform, 7–10(–13) μm in diameter. Phaeophycean hairs with a basal meristem; 3–8 μm 

in diameter.

Unilocular sporangia ovoid, 41 μm long by 32 μm in 

diameter. Plurilocular sporangia on single or branched 

pedicels; mostly uniseriate, up to 50 μm long, and up to 

10 μm in diameter.

Habitat. Epiphytic or entangled with various algae, notably Sargassum; low intertidal to shallow

subtidal.

Distribution. Gulf of California: Puerto

Peñasco to La Paz. Pacific coast: west side of Laguna Ojo 

de Liebre (Scammon’s Lagoon), opposite Isla La Concha 

(Baja California Sur).

Type Locality. La Paz, Baja California Sur,

Gulf of California, Mexico.

Remarks. In the field, Nemacystus brandegeei 

is recognized by its thin cylindrical axes, which are mostly 

irregularly branched (usually less than 600 μm in diameter), and slippery texture.

Streblonema Derbès et Solier

Streblonema Derbès et Solier, 1851:100.

Algae are minute, uniseriate, branched filaments, up 

to 3 mm in height. They may be entirely endophytic, growing between cells of the host, or partly endophytic, with 

both creeping and erect portions above the host’s surface. 

Growth is mostly diffuse. Cells have one to several bandshaped or discoid plastids and are without pyrenoids. Colorless hairs are usually present.

Reproductive structures are terminal or lateral on 

erect or creeping filaments, mostly extending above surface cells of its host. Unilocular structures are only known 

in a few species. Plurilocular sporangia are cylindrical or 

ovoid and uniseriate or multiseriate and occur either on 

the same thallus or on separate thalli. Gametophytes are 

apparently unknown.

Remarks. Many species of Streblonema are 

problematic or poorly known, and studies have revealed 

the genus to be paraphyletic. Some, but apparently not 

all, may be distinct independent species (Pedersen 1984). 

These species of Streblonema may have evolved by complete suppression of the macrothallus, or their sexual 

reproduction has been lost.

Streblonema microthalli have been found physically 

connected to a macrothallus. Pedersen (1978), studying the 
1 7 8 • 

FIGURE 85. Nemacystus brandegeei: A. Habit (JN-3985), epiphytic on Sargassum. B. Transection of a liquid-preserved specimen showing 

single axial filament cell and pseudoparenchymatous cortex, grading from larger inner cells to smaller outer cells that bear the assimilatory 

filaments. C. Furry external appearance of axes and branches derived from well-developed outer layer of free assimilatory filaments and phaeophycean hairs. B, C. EYD-25847.
n u m b e r 9 4 • 1 7 9

life history of Myriotrichia clavaeformis Harvey (1834), 

found its prostrate system was probably S. sphaericum

(Derbès et Solier) Thuret in Le Jolis (1863), the possible 

generitype of Streblonema. Culture studies have shown 

other Streblonema to be a microthallus phase in the heteromorphic life history of other species of brown macroalgae. 

For example, Streblonema-like microthalli were found to 

be life history phases of Litosiphon laminariae (Lyngbye) 

Harvey (1849) (Pedersen, 1981, as Litosiphon pusillus), 

Sphaerotrichia divaricata (J. Agardh) Kylin (1940) (Ajisaka and Umezaki, 1978), and Stilophora tenella (Esper) 

P. Silva (in Silva et al., 1996a; = S. rhizodes (C. Agardh) J. 

Agardh, 1841) (Peters and Müller, 1986a; Peters, 1987).

One species of Streblonema may be responsible for an 

algal disease. The endophytic infestations that result in loss 

of quality of Undaria (human food, known as wakame) in 

Japan are caused by S. aecidioides (Rosenvinge) Fosile in 

DeToni (1895b) (Yoshida and Akiyama, 1979) (note that 

molecular evidence places S. aecidioides in a group of kelp 

endophytes, Laminariocolax Kylin, 1947a; Burkhardt and 

Peters, 1998).

Three species of Streblonema are reported from Pacific 

Baja California (see L. Aguilar-Rosas, 1985; R. AguilarRosas and M. Aguilar-Rosas, 1986) and Isla Guadalupe 

(Dawson, 1961b; Stewart and Stewart, 1984).

One species and one undetermined species are 

recorded from the northern Gulf of California.

Streblonema transfixum Setchell et N. 

Gardner

FIGURE 86

Streblonema transfixum Setchell et N. L. Gardner, 

1922b:391; Setchell and Gardner, 1925:446; Dawson, 1945c:28; Abbott and Hollenberg, 1976:156, fig. 

121; R. Aguilar-Rosas and M. Aguilar-Rosas, 1986:18; 

Scagel et al., 1989:139; González-González et al., 

1996:165; Pedroche et al., 2008:24.

Algae minute filaments; lower portion, partly endophytic of irregularly branched filaments, 4–5 μm diameter, and upper portion of erect, unbranched filaments. 

Upper filaments short, of squarish to slightly rectangular 

cells, (5–)7–15 μm long and 7–10 μm diameter. Cells with 

band-shaped chloroplast.

Unilocular sporangia unknown. Plurilocular sporangia terminal on filaments; cylindroconical, 40–60 μm long 

and up to 15 μm wide; with locules in both longitudinal 

and transverse rows.

Habitat. Partly endophytic-epiphytic on

Gracilaria textorii; shallow subtidal, 10 m depths.

Distribution. Gulf of California: Isla Coronado (Isla Smith), Bahía de Los Ángeles. Pacific coast:

British Columbia to Oregon; central California to northern Baja California.

Type Locality. On Desmarestia herbacea,

San Pedro, Los Angeles County, California.

Remarks. Gulf specimens from Isla Coronado (JN-4411c; US Alg. Coll. slide 4257) are tentatively 

referred to Streblonema transfixum and represent a new 

record for the Gulf of California. Previously, S. transfixum

has been known on Desmarestia (Smith, 1944; R. Aguilar-Rosas and M. Aguilar-Rosas, 1986), and Gracilaria

would be an unusual host for this species. Life history and 

FIGURE 86. Streblonema transfixum: Habit, uniseriate unbranched 

filaments; one filament bearing a terminal plurilocular structure 

(arrow) with locules in longitudinal and transverse rows; epiphytic 

on Gracilaria textorii (JN-4411c).
1 8 0 • 

molecular studies are needed to confirm the identification 

of this species in the northern Gulf of California.

Streblonema sp.

Streblonema sp. of La Claire II and West, 1977:127, figs. 

1–4.

Microscopic, endophytic, and partly emerged irregularly branched filaments above surface cells of host, Cutleria hancockii. Reproduction not observed.

Habitat. Endophytic or epiphytic on gametophytes of Cutleria hancockii; mid to low intertidal

Distribution. Gulf of California: Punta Pelícano (vicinity of Puerto Peñasco), Sonora

Remarks. Filaments of a Streblonema sp. A 

were collected on gametophytes of Cutleria hancockii in 

the upper Gulf (La Claire II and West, 1977). Spores of 

the Streblonema were isolated and grown in culture studies, and ultrastructural study revealed viruslike particles 

(previously unknown in brown algal vegetative cells) 

within their cytoplasm (La Claire II and West, 1977). 

Studies of future collections, as well as life history culture and molecular analyses, are needed to identify the 

species.

Ec

Ectocarpaceae C. Agardh, 1828:9.

Members of this family are generally small or occasionally microscopic, branched, uniseriate filaments. The 

erect filaments can be corticated or uncorticated throughout and are usually attached by a disc-shaped base or a 

creeping rhizoidal holdfast, but some may penetrate the 

host or substrata. The branches are most often uniseriate, 

but in a few they are biseriate. Cells have a few to many 

parietal or axially elongated, band-shaped plastids and 

one to several pyrenoids.

Reproductive structures, unilocular or plurilocular 

sporangia and plurilocular gametangia, are borne laterally 

on branches or are intercalary in some and are formed 

by transformation of vegetative cells. Plurilocular and unilocular structures sometimes occur on the same thallus. 

Sporangia produce meiospores (zoospores). Gametangia 

produce isogametes or anisogametes.

One genus is represented in the northern Gulf of 

California.

Ectocarpus Lyngb

Ectocarpus Lyngbye, 1819:130.

Algae are erect, uniseriate filaments that are sparsely 

to densely irregularly branched and basally attached by 

rhizoids and/or prostrate filaments. The main axis may be 

uncorticated or, in some, corticated by descending rhizoidal filaments. Growth is diffuse by intercalary meristems 

(among the vegetative cells). Axis and branches often terminate in elongated terminal cells or colorless “pseudohairs.” 

Cells have one to several parietal, band-shaped or ribbonlike chloroplasts and usually several pyrenoids per plastid.

Life histories are isomorphic or heteromorphic with 

only slightly different generations; however, a few species 

are known to be only asexual. Reproductive structures are 

similar in appearance and sessile or pedicellate. Unilocular 

and plurilocular sporangia can be borne on the same alga 

or on separate thalli. Unilocular sporangia are globose or 

ellipsoidal, usually sessile, or terminal on short laterals. 

Plurilocular sporangia are usually terminal but, in some, 

may develop from vegetative cells below a pseudohair. 

Plurilocular gametangia are borne on separate thalli, and 

the gametes are isogamous.

Remarks. Ectocarpus is worldwide in distribution, with species that often exhibit wide phenotypic plasticity. The relationship of the species of Ectocarpus recorded in 

the Gulf to other known species is in need of critical study, 

including comparative morphological and life history studies and DNA analyses. Further field exploration will probably yield additional species in the Gulf of California.

Three species are reported in the southern Gulf 

(Huerta-Múzquiz and Mendoza-González 1985; MateoCid et al., 2000a; Cruz-Ayala et al., 2001): Ectocarpus 

corticulatus De A. Saunders (1898), E. chantrasoides

Setchell et N. L. Gardner (1922c) and E. simulans Setchell 

et N. L. Gardner (1922c).

Seven species are recorded in the northern Gulf of 

California.

Key to the Species of Ectocarpus in the n

1a. Filaments corticated basally; distally narrowing in diameter; cells more or less barrel-shaped; plurilocular sporangia more 

or less cylindrical with a conical upper portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. acutus

1b. Filaments not corticated basally; cells not barrel-shaped; plurilocular sporangia not cylindrical  
n u m b e r 9 4 • 1 8 1

Ectocarpus acutus Setchell et N. L. Gardn

FIGURE 87A,B

Ecotcarpus acutus Setchell et N. L. Gardner, 1922c:404, 

pl. 48: figs. 36–39, pl. 49: figs. 40, 41; Setchell and 

Gardner, 1925:415; Smith, 1944:80, pl. 10: fig. 1; Dawson, 1961b:383; Abbott and Hollenberg, 1976:123, 

fig. 83; Pacheco-Ruíz, 1982:70; Rocha-Ramírez 

and Siqueiros-Beltrones, 1991:24; Mateo-Cid and 

Mendoza-González, 1991:23; Mateo-Cid and Mendoza-González, 1994b:42; González-González et al., 

1996:147; Hoffmann and Santelices, 1997:110, fig. 25; 

Rodríguez-Morales and Siqueiros-Beltrones, 1999:17, 

22; Paul-Chávez and Riosmena-Rodríguez, 2000:146; 

Mateo-Cid et al., 2006:57; Servière-Zaragoza et al., 

2007:3, 8; Pedroche et al., 2008:15.

Algae filamentous, up to 5 cm tall; usually densely 

branched; attached by prostrate, entangled filaments (not 

penetrating deeply into host). Main filaments corticated 

below by rhizoidal filaments growing into holdfast and 

not corticated above; branching mostly alternate below 

and secund above; with branches gradually narrowing and 

becoming attenuated distally. Cells of filaments more or 

less barrel-shaped; in lower to middle portions (16–)24–

120 μm long and 40–60 μm in diameter; with few irregular band-shaped chloroplasts, each with several pyrenoids.

Unilocular sporangia unknown. Plurilocular sporangia on 1–2 celled pedicels; more or less cylindrical with a 

conical upper portion, 100–150(–230) μm long and 20–35 

μm in diameter.

FIGURE 87. Ectocarpus acutus: A. Portion of a filament showing 

the barrel shape of the cells with irregular band-shaped chloroplasts 

(from Setchell and Gardner, 1922b: pl. 48, fig. 38). B. Upper portion 

of filament showing branches attenuated toward acute apices (from 

Setchell and Gardner, 1922b: pl. 48, fig. 39).

2a. Upper branches usually in secund pattern; plurilocular sporangia up to 500 μm (or more) in length, with a sterile apex or 

long extension  . . . . . . .

2b. Upper branches usually not in secund series; plurilocular sporangia shorter and without sterile apices  

3a. Two distinct kinds of reproductive structures on same thallus; plurilocular structures: one of multiseriate rows of locules 

and the other of a uniseriate row of large locules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. commensalis

3b. Different kinds of reproductive structures on separate thalli: unilocular and plurilocular sporangia and plurilocular 

gametangia  

4a. Reproductive structures borne on both prostrate and erect filaments  . . . . 

4b. Reproductive structures restricted to erect filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5a. Epiphytic on Codium. . . . . . . .

5b. Growing on rocks or crustose coralline algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6a. Thalli small, filaments infrequently branched, about 20 μm in diameter, apices usually blunt; plurilocular sporangia sessile, 

to 175 μm long, without apical extensions. . . . . . .

6b. Thalli larger, filaments abundantly branched, up to 50 μm in diameter; plurilocular sporangia pedicellate, to 400 μm long, 

with apical extension of cells  . . . . . . 
1 8 2 • 

Habitat. Epiphytic, usually on brown algae;

intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco; Isla Espíritu Santo to Bahía de La Paz. Pacific 

coast: British Columbia to Baja California Sur; Colima; 

Chile.

Type Locality. Carmel, Monterey County,

California.

Ectocarpus commensalis 

Setchell et N. L. Gardn

FIGURE 88A

Ectocarpus commensalis Setchell et N.L. Gardner, 

1922c:407, pl. 48: figs. 32–35; Setchell and Gardner, 1925:424; Smith, 1944:86; Dawson, 1961b:384; 

González-González et al., 1996:147; Mateo-Cid et al., 

2006:57; Pedroche et al., 2008:16.

Ectocarpus dimorphus P. C. Silva nom. nov., 1957:42; 

Dawson, 1961b:384; Silva, 1979:332.

Ectocarpus variabilis (De A. Saunders) G. M. Smith, 

nom. illeg., 1942:647, figs. 1–4 [note Ectocarpus variabilis Vickers, 1905:59 is an earlier homonym]; Smith, 

1944:85, pl. 10: figs. 2–4; Dawson, 1945b:64; Dawson, 

1951:52; Dawson, 1954b:111; González-González et 

al., 1996:149.

Ecotcarpus confervoides f. variabilis De A. Saunders, 

1898:155, pl. 23: figs 1–10; Setchell and Gardner, 

1925:414, pl. 46: fig. 7; Dawson, 1944:220. 

Algae epiphytic, forming tufts up to 1.0 cm tall. Erect 

filaments uncorticated, sparingly and predominately alternate branched; upper ends attenuated, with round apices; 

prostrate filaments irregularly branched, attached to but 

not penetrating host cells. Cells of erect filaments 10–75(–

90) μm long by (10–)15–30 μm in diameter, with bandshaped chloroplasts.

Plurilocular structures of two kinds, both pedicellate 

(1–3 celled stalk), and usually borne on separate branches 

of same individual: (1) multiseriate, 10–15 locules wide, 

fusiform to ovoid, 75–180 μm long by 15–35 μm in diameter; each locule about 0.2 μm in diameter; and (2) a uniseriate row, 70–200 μm in length; of 2–15 barrel-shaped 

large locules, each 15–35 μm in diameter.

Habitat. Epiphytic on Sargassum and probably other algae; low intertidal

Distribution. Gulf of California: Isla Turners (off south end of Isla Tiburón). Pacific coast: British

Columbia to Baja California Sur; Nayarit.

Type Locality. Not stated by Saunders

(1898); Point Pinos, Pacific Grove, Monterey County, 

California, given by Smith (1944).

Remarks. Description of Ectocarpus commensalis is based primarily on Dawson (1944, as “E. confervoides f. variabilis”) and Smith (1944, as “E. variabilis”). 

Although considered to be conspecific with E. parvus by 

Abbott and Hollenberg (1976), Silva (1979, as “E. dimorphus”) noted that the epiphytic E. commensalis should 

probably be separated from the epilithic E. parvus.

Ectocarpus gonodioides

Setchell et N. L. Gardn

FIGURE 88B

Ectocarpus gonodioides Setchell et N. L. Gardner, 

1924a:721, pl. 17: fig. 44; Setchell and Gardner, 

1925:439; Dawson, 1944:221; Smith, 1944:86; Dawson, 1961b:384; Dawson, 1966a:9; Dawson, 1966b:55; 

Abbott and Hollenberg, 1976:126, fig. 87; GonzálezGonzález et al., 1996:148; Pacheco-Ruíz and ZertucheGonzález, 1996b:171; Mateo-Cid et al., 2006:49; 

Pacheco-Ruíz et al., 2008:203; Pedroche et al., 2008:16.

Algae minute tufts, up to 550 μm high, epiphytic, 

attached by rhizoidal filaments deeply penetrating into the 

host. Erect filaments near base, almost colorless, sparsely 

branched at host’s surface; upper filaments simple, mostly 

becoming attenuated toward apices. Cells 1–2 times as 

long as wide; lower cells 18–24 μm in diameter; upper 

cells 10–14 μm in diameter. Chloroplasts not adequately 

described in Gulf specimens.

Unilocular sporangia unknown. Plurilocular gametangia pedicellate, on 1–2 celled stalk, borne near base of 

filaments; narrowly fusiform, up to 125 μm long by 20–28 

μm wide.

Habitat. Epiphytic on Codium macdougali

(= C. brandegeei) and C. cuneatum (= C. simulans); mid 

to low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Bahía de Los Ángeles; Bahía Agua Verde to 

Cabo Pulmo. Pacific coast: central California.

Type Locality. Isla Coronado (Isla Smith),

Las Islas de la Cintura, Gulf of California.

Remarks. Setchell and Gardner (1924a:722), 

in describing the Gulf E. gonodioides, stated “the chromatophores are too much disorganized for characterization.” Recording this species from Carmel, central 

California, Smith (1944:86) noted it differed somewhat 
n u m b e r 9 4 • 1 8 3

FIGURE 88. Four species of Ectocarpus. A. Ectocarpus commensalis: Filaments bearing both multiseriate and uniseriate plurilocular sporangia 

(from Saunders, 1898: pl. 23, figs. 1, 9). B. Ectocarpus gonodioides: Tuft of filaments showing plurilocular reproductive structures borne right 

above the rhizoidal portion that penetrates the host (drawn from type specimen, I. M. Johnston-47e; Setchell and Gardner, 1924a: pl. 17, fig. 

44). C. Ectocarpus hancockii: Portion of a filament bearing a sessile, plurilocular sporangium (drawn from type specimen, EYD-[1940]133, 

Dawson, 1944: pl. 31, fig. 4). D. Ectocarpus sonorensis: Portion of a filament bearing a narrow, elongate plurilocular sporangium with sterile 

apical cells (drawn from type specimen, Drouet & Richards-3281, Dawson, 1944: pl. 31, fig. 5).
1 8 4 • 

from those of the Gulf of California type locality and 

described its plastids as band-shaped, and later, Abbott 

and Hollenberg (1976:126) noted there were few per cell. 

Until further collections of Gulf type locality material are 

made and its plastids and mode of growth are described 

and tested with molecular analyses, it is best to retain the 

species here (see also “Remarks” under E. hancockii).

Ectocarpus hancockii E. Y. Da

FIGURE 88C

Ectocarpus hancockii E. Y. Dawson, 1944:222, pl. 31: fig. 

4; Dawson, 1961b:385; Espinoza-Ávalos, 1993:333; 

González-González et al., 1996:149; L. Aguilar-Rosas 

et al., 2000:132, 134; Pedroche et al., 2008:18.

Algae tufted, 4–5 mm high, saxicolous or epiphytic 

on crustose coralline algae, attached by a mat of cells and 

basal filaments. Erect filaments infrequently branched, 

about 20 μm in diameter, apices usually blunt. Cells 

30–60 μm in length and up to 20 μm in diameter. Plastids 

probably band-shaped.

Unilocular sporangia unknown. Plurilocular sporangia, abundant, lateral, sessile, 125–175 μm long by 28–32 

μm in diameter.

Habitat. On rocks and on crustose coralline

algae; mid intertidal.

Distribution. Gulf of California: El Faro de

San Felipe (lighthouse of San Felipe); Isla Turners.

Type Locality. Reef off Isla Turners (off

southeast end of Isla Tiburón), Las Islas de la Cintura, 

Gulf of California.

Remarks. The generic placement of the Gulfendemic Ectocarpus hancockii remains uncertain, as characters necessary for generic separation, such as the plastids 

or type of growth, were not given in the original description (Dawson, 1944). However, in Dawson’s illustration 

(1944, pl. 31: fig. 4) the plastids appear to be mostly bandshaped, and thus, for now, this species is retained in Ectocarpus. Further study, including DNA analyses, on future 

collections from the type locality should lead to a better 

understanding of this species.

Ectocarpus parvus (De A. Saunders) Hollen

FIGURE 89

Ectocarpus siliculosus var. parvus De A. Saunders, 

1898:153, pl. 22: figs. 1–9.

Ectocarpus parvus (De A. Saunders) Hollenberg, 1971:283; 

Abbott and Hollenberg, 1976:126, fig. 89 [in part]; L. 

Aguilar-Rosas, 1982:31; Pacheco-Ruíz, 1982:70; Mendoza-González and Mateo-Cid, 1985:24; MendozaGonzález and Mateo-Cid, 1986:420; Stewart, 1991:38; 

Servière-Zaragoza et al., 1993:482; R. Aguilar-Rosas 

and M. Aguilar-Rosas, 1994:517; Mendoza-González 

et al., 1994:109; González-González et al., 1996:149; 

Mendoza-González and Mateo-Cid, 1996:69, 85, pl. 

17: figs. 69–72; Cruz-Ayala et al., 2001:190; Dreckmann et al., 2006:154.

Ectocarpus confervoides f. parvus (DeA. Saunders) Setchell et N. L. Gardner, 1922c:414; Setchell and Gardner, 

1925:413; Dawson, 1961b:384.

Ectocarpus confervoides f. pygmaeus sensu Setchell and 

Gardner, 1925:415 [non E. confervoides f. pygmaeus

(Areschoug) Kjellman, 1890:77; basionym: Ectocarpus 

pygmaeus Areschoug in Kjellman, 1872:85].

Ectocarpus confervoides var. pygmaeus sensu Hollenberg 

and Abbott, 1968:1237 [non E. confervoides var. pygmaeus (Areschoug) Rosenvinge, 1893:883; the basionym, E. pygmaeus Areschoug in Kjellman, 1872:85, is 

a synonym of Ectocarpus fasciculatus Harvey, 1841:40 

(see Russell, 1966:270)].

Algae erect, forming short, filamentous tufts, 5–10 

mm high. Filaments uncorticated, 20–30 μm in diameter; 

simple or sparingly, more or less alternately branched; 

attenuated at branch apices. Cells of filaments 25–75 μm 

in length, 20–30 μm in diameter; upper cells shorter than 

lower cells. Chloroplasts irregularly band-shaped.

Unilocular and plurilocular sporangia not reported in 

northern Gulf material.

Habitat. Growing on rocks; low intertidal

Distribution. Gulf of California: Puerto

Peñasco to Isla Pelícano, Bahía Kino; Bahía de La Paz. 

Pacific coast: Alaska to Baja California Sur; Sinaloa to 

Chiapas.

Type Locality. On “sand-colored rocks,”

San Pedro, Los Angeles County, California (Saunders, 

1898).

Remarks. The relationship of Ectocarpus 

parvus with E. commensalis is problematic. While recognizing E. parvus as a separate taxon (including those 

misidentified as “E. confervoides var. pygmaeus”), Scagel 

et al. (1989) noted its taxonomic status, as well as that of 

other morphologically similar taxa, is uncertain. Future 

collections will be helpful in understanding the degree of 

variation and the status of this taxon within the Gulf of 

California. Specimens of “E. confervoides f. pygmaeus” 
n u m b e r 9 4 • 1 8 5

as reported from Sonora (Mendoza-González and MateoCid, 1986) and Nayarit (León-Tejera et al., 1993) may 

also be this species.

Ectocarpus siliculosus (Dillwyn) Lyngb

Conferva siliculosa Dillwyn, 1809: 69, Suppl. pl. E.

Ectocarpus siliculosus (Dillwyn) Lyngbye, 1819:131, 

pl. 43B,C; Setchell and Gardner, 1925:410; Doty, 

1947:31; Dawson, 1961b:385; Abbott and Hollenberg, 

1976:128, fig. 90; Silva, 1979:332; Servière-Zaragoza 

et al., 1993:482; Mateo-Cid and Mendoza-González 

1991:23; Mateo-Cid and Mendoza-González 1992:23; 

Mateo-Cid and Mendoza-González 1994b:42; 

González-González et al., 1996:376; Kitayama, 

1996:224, figs. 2, 3; Yoshida, 1998:166; L. Aguilar-Rosas et al., 2000:132, L. Aguilar-Rosas et al., 2002:235; 

Pedroche et al., 2008:18.

Algae forming tufts, up to 2.0 cm tall, composed of 

uniseriate, uncorticated filaments; branching subdichotomous in lower portion and alternately secund above; 

larger branches 45–60 μm in diameter, ultimate branchlets 10–25 μm in diameter. Cells (0.6–)1–2 times as long 

as wide; with ribbon-shaped chloroplasts and several 

pyrenoids.

Plurilocular sporangia conical, usually on short 

pedicels (also reported sessile), 90–500(–600) μm long, 

15–25(–40) μm in diameter; with sterile tip or sometimes 

with hairlike multicellular extension.

Habitat. On rocks, limestone, and clay, in tide

pools, or epiphytic on Codium; often becomes free-floating; high to mid intertidal.

Distribution. Gulf of California: El Tornillal to Puertecitos. Pacific coast: Alaska to Baja California

Sur; Nayarit to Colima, Peru; Chile; China; Japan.

Syntype Localities. Cromer in Norfolk

and Hastings in East Sussex, England (Silva et al., 1996a).

Remarks. Ectocarpus siliculosus has been 

recorded in the northern Gulf by L. Aguilar-Rosas et al. 

(2000, 2002); description is based on Abbott and Hollenberg (1976).

FIGURE 89. Ectocarpus parvus: Habit, portion of a filament showing terminal and lateral plurilocular sporangia (drawn from type 

specimen, W.A. Setchell-1213, Saunders, 1898: pl. 22, fig. 1).
1 8 6 • 

Ectocarpus sonorensis E. Y. Daw

FIGURE 88D

Ectocarpus sonorensis E. Y. Dawson, 1944:221, pl. 31: 

fig.5; Dawson, 1961b:385; Espinoza-Ávalos, 1993:333; 

González-González et al., 1996:149; Pedroche et al., 

2008:20.

Algae filamentous, light green-brown, up to 10 cm tall, 

abundantly branched, with main filaments up to 50 μm in 

diameter, and branches attenuating distally from 40 μm to 

12 μm in diameter; secondary and other order branches, 

slender, somewhat cylindrical, and tapering upwards to 

a sharp point; branches bearing numerous short, spinelike secondary branchlets near apices. Cells thin-walled, 

1.25–1.50 times as long as wide; with pale chloroplasts 

apparently lining the wall.

Unilocular sporangia unknown. Plurilocular sporangia borne on 2-celled pedicels, slender, up to 400 μm long 

by 20–25 μm in diameter, with an apical, pointed extension of vegetative cells, 30–120 μm long.

Habitat. On rocks; intertidal

Distribution. Gulf of California: Guaymas

Type Locality. “On rocks in tidal stream,” 3

km east of Guaymas, Sonora, Gulf of California (Dawson, 

1944).

Remarks. Apparently a Gulf of California 

endemic, Ectocarpus sonorensis is known only from the 

type collection.

Pylai

Pylaiellaceae P. M. Pedersen, 1984:50, as “Pilayellaceae.” 

Algae are filamentous, with erect and sometimes lowgrowing portions, attached by rhizoids. Filaments are 

mostly uniseriate, becoming parenchymatous by longitudinal divisions of the cells.

Sporangia are formed by the transformation of vegetative cells.

Remarks. Pedersen (1984) considered the Pylaiellaceae to be close to some families of the orders Dictyosiphonales and Tilopteridales, noting that the more or less 

frequent formation of longitudinal walls in Pylaiella justifies 

its position among the parenchymatous brown algae. However, Wynne (1986) questioned the taxonomic significance 

of this character and placed Pylaiella in the Ectocarpaceae 

and, later (Wynne, 2005), within the Acinetosporaceae 

(both Ectocarpales). Herein the Pylaiellaceae is recognized 

as a family within the Ectocarpales (Silva et al., 1996a).

Pylaiella Bory de Saint-Vincen

Pylaiella Bory de Saint-Vincent, 1823:393.

Algae are filamentous, composed of creeping and 

erect portions, which may be simple or more or less opposite, alternate, or irregularly branched, and attached by 

rhizoids. Filaments are initially uniseriate and without 

a terminal hair; later, these filaments may be frequently 

divided by longitudinal walls (some interpret this as 

becoming parenchymatous) and taper upward into pseudohairs (note that true phaeophycean hairs are absent). 

Growth is by intercalary divisions, which are generally diffuse throughout the thallus or sometimes localized. Cells have numerous discoid plastids and a single 

pyrenoid.

Thalli are isomorphic, sometimes with both unilocular and plurilocular structures on the same individual. 

Plurilocular and unilocular sporangia are barrel-shaped or 

subspherical, are intercalary in catenate series with lateral 

openings, and are formed by direct transformation of vegetative cells. Plurilocular gametangia are oblong to more 

or less cylindrical.

Remarks. The original spelling, Pylaiella

Bory de Saint-Vincent (1823) has been conserved over 

“Pilayella” (Silva et al., 1999; Compère, 2003). Species 

of Pylaiella exhibit a wide range of phenotypic plasticity; thus, the species concepts are in need of testing and 

clarification.

Although species of Pylaiella are not well known in 

the Gulf, large loose-lying mats of another species, P. littoralis (Linnaeus) Kjellman (1872), have created environmental problems on the northeastern coast of the United 

States (Wilce et al., 1982).

One undetermined species is now recorded from the 

northern Gulf of California.

Pylaiella sp.

Alga minute and filamentous. Main axes branched 

irregularly alternate.

Sporangia not seen.

Habitat. In a tide pool, entangled with Antithamnionella spirographidis; low intertidal

Distribution. Gulf of California: Punta Pelícano, Puerto Peñasco

Remarks. A minute specimen of Pylaiella (JN3979) establishes the genus as occurring in the northern 

Gulf. Unfortunately, the material is not fertile; thus, the 

species cannot be determined. It is noted herein to call its 

presence to the attention of future collectors.
n u m b e r 9 4 • 1 8 7

Leathesi

Leathesiaceae Farlow, 1881:16, 79.

Corynophlaeceae Oltmanns, 1922:23 [see Silva, 1980: 

71, 73].

Sporophytic algae are cushionlike, globose, or hemispherical, some becoming irregularly globular, with 

smooth to convoluted surfaces and without a distinct basal 

layer. Assimilatory filaments are anticlinally arranged, 

with intercalary growth. Cells of assimilatory filaments 

contain several chloroplasts, each with a pyrenoid.

Life histories, where known, are heteromorphic with 

a sporophytic macrothallus and a haploid filamentous 

microthallus. Sporophytes may have unilocular and/or 

plurilocular sporangia on the same or separate thalli. Presumed gametophytes are filamentous, microscopic thalli, 

with uniseriate gametangia.

Remarks. Some taxonomists consider the 

Leathesiaceae to be synonymous with the family Chordariaceae on the basis of morphological and life history similarities. Molecular phylogenetic analyses of the generitype 

Leathesia tuberiformis S. F. Gray (1821; which is now 

Leathesia marina (Lyngbye) Decaisne, 1842, basionym 

Chaetophora marina Lyngbye 1819; L. difformis (Linnaeus) J. E. Areschoug, 1847; see Pedroche et al., 2008) 

will help resolve the familial status.

One of the genera, Myriactula Kuntze (1898), is represented by two poorly known endemic species in the southern Gulf: Myriactula johnstonii (Setchell et N. L. Gardner) 

Feldmann (1945; basionym: Gonodia johnstonii Setchell 

et N. L. Gardner 1924a); and M. marchantiae (Setchell 

et N. L. Gardner) Feldmann (1945; basionym: Gonodia 

marchantiae Setchell et N.L. Gardner 1924a).

There is one genus recorded in the northern Gulf of 

California.

Leathesia S. F. Gray

Leathesia S. F. Gray, 1821: 279, 301.

Algae are globular to hemispherical or irregularly 

convoluted and internally solid or initially solid and later 

*Contributed by Luis E. Aguilar-Rosas, Raúl Aguilar-Rosas, 

and James N. Norris; L. Aguilar-Rosas: Instituto de Investigaciones Oceanológicas, Universidad Autónoma de Baja California, 

Km 107 carretera Tijuana-Ensenada, 22830 Ensenada, Baja California, Mexico; R. Aguilar-Rosas: Facultad de Ciencias Marinas, Universidad Autónoma de Baja California.

becoming hollow. Species are epiphytic or growing on 

solid substratum. Growth is by intercalary cell divisions. 

Internal structure is of two kinds of filaments: inner 

medullary filaments that are loosely arranged, dichotomously to trichotomously branched, and composed of 

large, elongated, colorless cells; and outer multicelled 

assimilatory filaments that are composed of cylindrical 

pigmented cells that terminate with a large globular cell. 

Cells of assimilatory filaments contain several chloroplasts with pyrenoids. Phaeophycean hairs, often common, are among the assimilatory filaments and project 

above the surface layer.

The life history is heteromorphic, with plurispores 

of sporophytic macrothalli producing filamentous microthalli and unfused gametes of the latter developing gametophytes (e.g., Kylin, 1933; Dangeard, 1969). Unilocular 

and plurilocular sporangia develop on the basal cells of 

the assimilatory filaments, sometimes on the same individual. Unilocular sporangia are ovoid to ellipsoid and 

are usually separated from each other. Plurilocular sporangia with locules in uniseriate rows are usually grouped 

together.

Remarks. Two taxonomic sections of the 

genus were recognized by Inagaki (1958). Leathesia

sect. Leathesia includes members that are hollow, with a 

reticulated medullary layer, and Leathesia sect. Primariae

includes those that are solid, with densely packed medullary filaments. Species of the latter section resemble the 

genus Corynophlaea Kützing (1843) in their inner anatomy; however, they are different genera. Ajisaka (1984) 

cultured one member, L. japonica Inagaki (1958), and 

reported the occurrence of filamentous thalli that resembled Polytretus reinboldii (Reinke) Sauvageau (1900; 

=Botrytella reinboldii (Reinke) Kornmann et Sahling, 

1988) and formed plurilocular sporangia with several lateral release pores in its life history.

One species has been reported in the northern Gulf of 

California.

Leathesia nana Setchell et N. L. Gardn

FIGURE 90A–C

Leathesia nana Setchell et N. L. Gardner, 1924b:3; 

Setchell and Gardner, 1925:511, pl. 43: fig. 67; Smith, 

1944:115, pl. 15: fig. 1; Dawson, 1958:65; Dawson, 

1961b:391; Hollenberg, 1971:285, fig. 3; Abbott and 

Hollenberg, 1976:177, fig. 143; Pacheco-Ruíz, 1982:70; 

Tseng, 1983:174, pl. 88: fig. 1; R. Aguilar-Rosas et 

al., 1984a:152; R. Aguilar-Rosas et al., 1984b:161; 

R. Aguilar-Rosas et al., 1990:123; González-González 
1 8 8 • 

et al., 1996:154; L. Aguilar-Rosas et al., 2000:127, 136; 

Mateo-Cid et al., 2006:58; Pedroche et al., 2008:60.

Algae small, more or less globose or hemispherical, 

1.0–3.0(–5.0) mm in diameter; mostly with a smooth 

surface and usually not hollow; epiphytic on Sargassum. 

Assimilatory (cortical) filaments of 3–4(–5) elongate cells, 

(6–)7–8 μm in diameter, with a larger, subspherical terminal cell, 10 μm in diameter. Medullary filaments of 

larger colorless cells. Hairs not observed in Gulf specimens 

(reported as present or absent for the species).

Reproductive structure not observed in upper Gulf of 

California specimens.

Habitat. Epiphytic on Sargassum sinicola and

S. herporhizum; intertidal.

Distribution. Gulf of California: Puerto

Peñasco; Playa El Coloradito. Pacific coast: northern 

Washington to Cabo Colnett, Baja California; China.

Type Locality. Monterey, Monterey County,

California.

Remarks. The northern Gulf Leathesia nana

were found on brown algae, species of Sargassum (L. 

Aguilar-Rosas et al., 2000; Mateo-Cid et al., 2006). This 

is a new host, as central California L. nana are known 

as epiphytes on the sea grass Phyllospadix (Setchell and 

Gardner, 1924b, 1925; Smith, 1944) and on the red algae 

Neorhodomela larix (Turner) Masuda (1982; = Rhodomela larix (Turner) C. Agardh, 1822) and Odonthalia 

floccosa (Esper) Falkenberg (1901), both Rhodomelaceae 

(Sparling, 1971; Abbott and Hollenberg, 1976).

Although reproductive structures were not seen 

in upper Gulf specimens, two kinds of sporangia are 

reported in California L. nana (Hollenberg, 1971; Abbott 

and Hollenberg, 1976): plurilocular sporangia, uniseriate, 

of 4–6 locules, 20–30 μm long and 3.5–5.0 μm in diameter, borne at base of 3- to 4-celled paraphyses, with a 

globular terminal cell only slightly larger than lower cells 

(Setchell and Gardner, 1925); and unilocular sporangia 

25–40(–50) μm long and 10–20 μm in diameter, borne at 

base of 4- to 6-celled assimilatory filaments with a large 

FIGURE 90. Leathesia nana: A. Several thalli epiphytic on a leaflike blade of Sargassum sinicola. B. Habit of several small globose thalli. C. 

Transection of thallus showing three- to five-celled assimilatory filaments with a larger, subspherical, terminal cell (CMMEX-6447).
n u m b e r 9 4 • 1 8 9

globular terminal cell (larger than those that bear plurilocular sporangia; Hollenberg, 1971:fig. 3). Specimens of 

L. nana with plurilocular sporangia are known only on 

Phyllospadix, whereas those with unilocular sporangia 

are almost exclusively on Neorhodomela and Odonthalia

(Abbott and Hollenberg, 1976).

Webber (1981, as “L. difformis”) suggested L. nana

was a growth-form of L. marina (Lyngbye) Decaisne, and 

Scagel et al. (1989, as “L. difformis”), considering the 

study of Novaczek (1987), reduced L. nana to a synonym 

of L. marina. Comparative molecular and morphological studies of L. marina (type locality: Båstad, Sweden), 

central California “L. difformis sensu Smith (1944) and 

Abbott and Hollenberg (1976),” L. nana (type locality: 

Monterey), and the northern Gulf L. nana are needed to 

elucidate the taxonomic status and phylogenetic relationships of these four.

Scyto

Scytosiphonaceae Farlow, 1881:15, 62.

Algae of this family exhibit a wide variety of forms and 

can be tubular, compressed, or flattened and elongated, 

cylindrical, foliose, spherical, or saccate. The habit of 

these algae can be entire, irregularly shaped, or branched. 

Structurally, they are multiaxial and pseudoparenchymatous, with a medulla of large colorless cells and a cortical 

layer of small pigmented cells, and can be either solid or 

hollow centrally. Growth in most of the members is intercalary or from subapical cells. Cells have a large plastid 

with a pyrenoid, and vesicle-like globules (physodes) may 

be present or lacking. Phaeophycean hairs are present and 

either isolated or grouped in superficial pits.

Life histories, where known, are heteromorphic. 

Reproductive structures are in sori. Sporophytes produce 

unilocular and plurilocular sporangia. Sexual reproduction is by unilocular gametangia and uniseriate or multiseriate plurilocular gametangia that produce isogametes or 

ansiogametes.

Remarks. The report of two members of the 

family in the northern Gulf (Mendoza-González and 

Mateo-Cid, 1986), Petalonia binghamiae (J. Agardh) 

Vinogradova (1973; = Endarachne binghamiae J. Agardh 

1896) and Petalonia fascia (O. F. Müller) Kuntze (1898), 

needs to be confirmed.

Four genera of the Scytosiphonaceae are known in the 

northern Gulf of California.

Key to the Genera of Scytosiphonaceae in the 

1a. Minute to small epiphytes or endophytes; a disclike basal layer of monostromatic filaments, with erect, short, simple to 

laterally branched filaments  . . . . 

1b. Large, erect, and centrally hollow (not filamentous) and may be cylindrical and tubular or compressed, or globose or saccate, or open and sheetlike with or without holes (netlike)  

2a. Thalli either long, narrow cylindrical axes and branches, up to 3 mm in diameter, or low-growing clumps of more or less 

cylindrical to compressed entangled branches, up to 6 mm in diameter  . . . . . 

2b. Thalli globose or saccate or open sheetlike or netlike  

3a. Thalli may be spherical to irregularly globose or saccate or more or less tubular irregularly lobed branches or open sheetlike 

without holes  . . . . 

3b. Thallus three-dimensional “net” of small to large holes, usually torn open and forming netlike sheets  . . . . . H

Colpomenia (Endlicher) Derbès et Solie

Colpomenia (Endlicher) Derbès et Solier, 1851:95.

Asperococcus β. Colpomenia Endlicher, 1843:26.

Algae are more or less globose or saccate and hollow, occasionally forming prostrate expanses or open 

expanded sheets. The surface may be smooth, lobed, or 

constricted, or may be heavily convoluted and with prominent protuberances. Some may become torn or open and 

flattened with age. The medulla is composed of large 

colorless cells, the innermost medullary layer facing the 

open hollow cavity. The cortex is generally one layer of 

small pigmented cells and has scattered tufts of hairs with 

basal meristems.

Plurilocular gametangia are borne in superficial sori 

associated with paraphyses, scattered over the thallus 

surface.

Four species of Colpomenia are recognized in the 

northern Gulf of California. Another, C. peregrina Sauvageau (1927), has been reported in the southern Gulf 

(Cruz-Ayala et al., 2001).
1 9 0 • 

Colpomenia phaeodactyla

M. J. Wynne et J. N. Norris

FIGURE 91A

Colpomenia phaeodactyla M. J. Wynne et J. N. Norris, 1976:5, figs. 4, 5a,b, 9a, 11c; Littler and Littler, 

1981:151, tbl. 3, figs. 4, 5, 7; Norris, 1985:209, fig. 16.3 

(bottom); Santelices, 1989:289, pl. GG: fig. 7; Yoshida 

et al., 1995:280; González-González et al., 1996:142; 

Pacheco-Ruíz and Zertuche-González, 1996b:171; 

Hoffmann and Santelices, 1997:159, fig. 40; Yoshida, 

1998:305, fig. 2-21G; Mateo-Cid et al., 2006:49, 58; 

Pacheco-Ruíz et al., 2008:204; Pedroche et al., 2008:64.

Colpomenia sinuosa f. deformans sensu Setchell and Gardner, 1924a:726, pl. 19: figs. 61, 62; Setchell and Gardner, 1925:542; Dawson, 1949:239 [in part; Gulf of California specimens only]; Dawson, 1944:233; Dawson, 

1966a:11; Wynne, 1972:137, figs. 1, 17, 18; Wynne, 

1973:141 [non Colpomenia sinuosa f. deformans Setchell et N. L. Gardner, 1903:242; which is Colpomenia 

bullosa (De A. Saunders) Yamada, 1948:6].

Colpomenia bullosa sensu Norris, 1973:6; Pacheco-Ruíz 

et al., 2008:204 [non Colpomenia bullosa (De A. Saunders) Yamada, 1948:6; basionym: Scytosiphon bullosus

De A. Saunders, 1898:163].

Algae usually forming clusters of long, hollow sacs, 

golden brown (darker when fertile), 8–15(–25) cm long, 

0.8–1.5(–5.5) cm wide, arising from a compressed base 

attached to the substrate. Sacs flaccid and delicate, elongated and tapering toward base; hollow throughout; thin, 

70–88(–110) μm thick. Medulla 2–3 cells thick, of large 

colorless cells. Outer cortex a single layer of small cuboidal cells, each with 1 chloroplast; tufts of colorless hairs 

rare. Subcortex also a single layer of slightly larger cells.

Unilocular sporangia unknown. Plurilocular structures, uniseriate and biseriate, 28–38 μm long; forming 

dense, extensive sori over surface of erect sacs; paraphyses 

rare (if present, more or less same height as plurilocular 

structures).

Habitat. On tops of rocks and tidal platforms;

mid to low intertidal.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas. Pacific coast: Bahía San Bartolomé 

to Bahía Asunción (Baja California Sur); Costa Rica; central Chile; Japan.

Type Locality. Playa Estación, Puerto

Peñasco, Sonora, Gulf of California.

Remarks. Named for the brown finger-like 

appearance of its elongated sacs, Colpomenia phaeodactyla is an ephemeral species and can be relatively common 

at some locales in the upper Gulf from December to May. 

It is quite flaccid, lying on the substratum when exposed 

during low tides, and during high tides it is submerged and 

vertical, with the hollow portions appearing to be at least 

partly filled with air.

Ramírez and Rojas V. (1991) considered Chilean 

records identified as “C. phaeodactyla” (Santelices et al., 

1989; Ramírez and Santelices, 1991) to be C. durvillei

(Bory de Saint-Vincent) Ramírez (see Wynne, 1999, for 

clarification of status of this taxon). However, Hoffmann 

and Santelices (1997) recognized C. phaeodactyla from 

central Chile.

Key to the Species of Colpomenia in the no

1a. Thalli consisting of erect, clustered, fingerlike sacs arising from a common prostrate holdfast; wall of sacs thin, delicate, 

usually less than 100 μm thick  . . . . . . .

1b. Thalli not fingerlike sacs, but range from spherical, balloonlike to irregularly shaped, low-growing or sessile expanses or 

clumps of branched thalli; thicker and firmer, >200 μm in thickness  

2a. Thalli forming a spreading clump of adherent, irregular lobed branches; smooth surface . . . . . . . . . . . . . . . . . C. ramos

2b. Thalli globose or expanded sacs (occasionally open and flat sheets); lacking definite branches; smooth, convoluted, or 

tuberculate surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Thalli with tuberculate, warty surface; of a crisp, rigid texture; thickness exceeding 500 μm (often up to 1.0 mm); plurilocular structures up to 25 μm long  . . . . . . 

3b. Thalli with smooth or convoluted surface; flaccid, never tuberculate or rigid; thinner in transection, 300–500 μm thick; 

plurilocular structures 40 or more μm long  

4a. Thallus hollow, vesicle-like or globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sinuos

4b. Thallus open, broad or expanded thin sheets  . . .
n u m b e r 9 4 • 1 9 1

Colpomenia ramosa W. R. Taylor

FIGURE 91B

Colpomenia ramosa W. R. Taylor, 1945:84, pl. 6, fig. 2; 

Dawson, 1949:228, 250; Dawson, 1951:52; Dawson, 

1952:431; Dawson, 1961b:394; Wynne and Norris, 

1976:11, fig. 8a,b; Mendoza-González and Mateo-Cid, 

1985:24; Ramírez and Rojas V., 1991:16, figs. 4, 8; 

Martínez-Lozano et al., 1991:23; Mendoza-González 

et al., 1994:110; González-González et al., 1996:142; 

Pacheco-Ruíz and Zertuche-González, 1996b:171; 

Mateo-Cid and Mendoza-González, 1997:59, pl. 7: 

figs. 28, 29, pl. 9: fig. 39; Mateo-Cid and MendozaGonzález, 2003:22; Pacheco-Ruíz et al., 2008:204; Pedroche et al., 2008:65.

Rosenvingea intricata sensu Dawson, 1944:233 [in part], 

pl. 52: fig. 1 [non Rosenvingea intricata (J. Agardh) 

Børgesen, 1914:26].

Algae forming adherent clumps, up to 4 cm broad and 

to 2 cm tall, with several areas of attachment. Thallus of 

congested, hollow, more or less cylindrical to compressed 

axes, irregularly subdichotomously to polychotomously 

FIGURE 91. Two species of Colpomenia. A. 

Colpomenia phaeodactyla: Habit of several 

fingerlike hollow sacs (JN-4759). B. Colpomenia ramosa: Upper portion of thallus showing 

branching pattern (JN-4697).
1 9 2 • 

branched; broader basally, becoming narrower distally; 

terminal branch divisions cylindrical, short, 1–2 mm in 

diameter by 2–3 mm long, with rounded ends. In transection, outer wall 200–500 μm thick, of 6–8 cell layers; cells 

larger inward toward hollow cavity, becoming smaller 

outward to small-celled cortex.

Plurilocular sporangia in sori, uniseriate, with 10–12 

locules, 35–40 μm long.

Habitat. Often entangled with other algae;

low intertidal to shallow subtidal.

Distribution. Gulf of California: Isla Willard, Bahía San Luis Gonzaga to Bahía Tepoca; Isla Espiritu Santo to Bahía de La Paz. Pacific coast: Baja California

to Sinaloa; Guerrero to Oaxaca; Costa Rica; Galápagos 

Islands; Chile.

Type Locality. Bahía Sur, Isla Cedros, Baja

California, Pacific Mexico.

Colpomenia sinuosa (Mertens ex Roth)

Derbès et Solier

FIGURES 5A, 92A

Ulva sinuosa Mertens ex Roth, 1806:327, pl. 12: figs. a–c.

Colpomenia sinuosa (Mertens ex Roth) Derbès et Solier, 1851:95; Derbès and Solier, 1856:11, pl. 22: figs. 

18–20; Saunders, 1898:164, pl. 32: figs. 7, 8; Setchell 

and Gardner, 1925:539, 541, pl. 45: figs. 82–86; Dawson, 1944:232; Dawson, 1951:52; Dawson, 1959a:19; 

Dawson et al., 1960a:30, pl. 9: fig. 1, 2; Dawson, 

1962b:199, fig. 77; Dawson, 1966a:10; Earle, 1972:85; 

Clayton, 1975: 187, tbl. 1, figs. 5–7, 12, 13; Abbott 

and Hollenberg, 1976:204, fig. 168; Wynne and Norris, 

1976:2, figs. 1, 2a,b, 11a; L. Aguilar-Rosas, 1982:31; 

Parsons, 1982:291, figs. 2, 3, 11, 12; Schnetter and Bula 

Meyer, 1982:65, pl. 10: fig. N; Stewart and Stewart, 

1984:141; L. Aguilar-Rosas et al., 1985:125; HuertaMúzquiz and Mendoza-González, 1985:46; MendozaGonzález and Mateo-Cid, 1985:24; Sánchez-Rodríguez 

et al., 1989:40; Santelices, 1989:280, pl. FF: fig. 3; R. 

Aguilar-Rosas et al., 1990:123; Martínez-Lozano et 

al., 1991:23; Ramírez and Rojas V., 1991:16, figs. 2, 

6; Rocha-Ramírez and Siqueiros-Beltrones, 1991:24; 

Stewart, 1991:44; Mateo-Cid et al., 1993:50; R. Aguilar-Rosas and M. Aguilar-Rosas, 1994:518; Mendoza-González et al., 1994:110; González-González et 

al., 1996:143; Pacheco-Ruíz and Zertuche-González, 

1996b:171; Hoffmann and Santelices, 1997:156, fig. 

39; Yoshida, 1998:305, fig. 2-21 H; Mateo-Cid et al., 

2000a:68; Cruz-Ayala et al., 2001:190; Abbott and 

Huisman, 2004:179, fig. 67A,B; R. Aguilar-Rosas et al., 

2005b:34; Hernández-Herrera et al., 2005:146; MateoCid et al., 2006:49, 58; Servière-Zaragoza et al., 2007:8; 

Pacheco-Ruíz et al., 2008:204; Pedroche et al., 2008:66.

Algae globose or vesicle-like, hollow, golden- to 

medium-brown, flaccid and slippery, up to 14 cm in diameter; 200–500 μm thick; becoming compressed, irregularly convoluted and expanded; attached by rhizoids on 

lower surface. Medulla and subcortex of 4–6 layers of 

FIGURE 92. Two species of Colpomenia. A. Colpomenia sinuosa: Habit, showing smooth, slightly convoluted surface (JN-4580). B. Colpomenia tuberculata: Habit, photo of live specimen showing protuberances (from Puerto Peñasco, Sonora, Mexico).
n u m b e r 9 4 • 1 9 3

large, irregularly shaped cells; gradually becoming smaller 

toward outer surface layer of small cuboidal cells.

Unilocular sporangia unknown. Plurilocular sporangia uniseriate (to partially biseriate), up to 40 μm long, 

in sori covered with a cuticle, clustered often around an 

invaginated tuft of hairs. Paraphyses usually longer than 

plurilocular sporangia, to 55 μm long.

Habitat. On rocks and tidal platforms or epiphytic

on various algae; low intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto Peñasco to

Bahía Agua Verde; Bahía de La Paz to Cabo San Lucas. 

Pacific coast: southern California to Baja California Sur; 

Sinaloa to Jalisco; Isla Clarión (Islas Revillagigedo); Hawaiian Islands; Panama; Colombia; Chile.

Type Locality. “Location uncertain” (Parsons,

1982); given as near Cádiz, Spain (Silva et al., 1996a).

Remarks. In the northern Gulf of California, specimens of Colpomenia sinuosa were found only with 

plurilocular sporangia (Wynne and Norris, 1976). Interestingly, the life history of C. sinuosa in the Azores 

was found to be highly variable (Toste et al., 2003b) 

with three different kinds of life histories: (1) asexual 

plurispores from gametophytes that developed directly 

into new gametophytes; (2) heteromorphic (involving an 

alternation of a macrothallus and a filamentous prostrate 

thalli), and either diplohaplontic or monophasic life history; and (3) isomorphic, monophasic life histories.

Colpomenia sp.

Colpomenia sinuosa? of Wynne and Norris, 1976: 5, fig. 

3 [“open expanded sheet form”].

Colpomenia sinuosa f. expansa sensu Norris 1973:6; Pedroche et al. 2008:67 [non Colpomenia sinuosa f. expansa De A. Saunders 1898:164; which is now Colpomenia expansa (De A. Saunders) Y.-P. Lee 2008:116]. 

Algae thin, broadly expanded open sheets, up to 25 

cm wide, with smooth surfaces and margins.

Reproductive structures not seen.

Habitat. On rocks; shallow subtidal, 4–16 m

depths.

Distribution. Gulf of California: Puerto

Calamajue (Bahía de Calamajue) to Bahía San Francisquito; Isla San Esteban; Puerto Escondido to Isla Carmen.

Remarks. Relatively large specimens of open 

expanded sheets of Colpomenia sp. A, were collected in the 

Gulf (Norris, 1973:6; Wynne and Norris, 1976:5, as C. cf. 

sinuosa). These may possibly be undescribed or another 

taxon or form of Colpomenia. Although somewhat similar to the Gulf of California C. sinuosa f. expanissima

Setchell et N. L. Gardner (1924a:726; Dawson, 1944, as 

“C. sinuosa f. tuberculata”), these Gulf specimens have 

smooth surfaces and lack the minute, spinelike projections of C. sinuosa f. expanissima. Further collections are 

needed for study to clarify the taxonomic status of these 

Gulf open-sheet specimens.

Colpomenia tuberculata De A. Saunde

FIGURE 92B

Colpomenia tuberculata De A. Saunders, 1898:164, pl. 

32: figs. 1–3; Wynne, 1972:137, fig. 2; Wynne and Norris, 1976:8, figs. 6, 7a,b, 11b; Abbott and Hollenberg, 

1976:204, fig. 169; Littler and Littler, 1981:150, tbls. 

2–4, figs. 3–7; Stewart, 1982:54; Huerta-Múzquiz and 

Mendoza-González, 1985:46; Norris, 1985:209, fig. 16.3 

(top); Santelices, 1989:281, pl. FF: fig. 2; Sánchez-Rodríguez et al., 1989:40; R. Aguilar-Rosas et al., 1990:123; 

Ramírez and Rojas V., 1991:16, figs. 3, 7; Stewart, 

1991:44; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:32; Martínez-Lozano et al., 1991:23; Mateo-Cid 

et al., 1993:50; González-González et al., 1996:143; 

Pacheco-Ruíz and Zertuche-González, 1996b:171; 

Hoffmann and Santelices, 1997:162, fig. 41; RiosmenaRodríguez et al., 1998:45; Mateo-Cid et al., 2000a:68; 

Cruz-Ayala et al., 2001:190; Abbott and Huisman, 

2004:180, fig. 67C,D; Mateo-Cid et al., 2006:49, 58; 

Pacheco-Ruíz et al., 2008:204; Pedroche at al., 2008:67.

Colpomenia sinuosa f. tuberculata (De A. Saunders) 

Setchell et N. L. Gardner, 1903:242; Setchell and Gardner, 1924a:725; Setchell and Gardner, 1925:541; Dawson, 1944:233; Dawson, 1959a:19; Dawson, 1966a:11; 

Norris, 1973:6; González-González et al., 1996:143.

Algae olive- to dark-brown globose or hemispherical, 

up to 15 cm in diameter, to irregularly shaped, compressed 

expanses, up to 24 cm wide; rigid and crisp, 0.6–1.5 mm 

thick; surface coriaceous, deeply convoluted and sometimes folded, covered with blunt tubercles (protuberances), 1.0–10 mm high and 1.0–10 mm wide. Cortex of 

4–5 small cuboidal cells; hairs arising in tufts from deep 

invaginations; subcortex and medulla of 4–8 large colorless cells, becoming increasingly smaller outward.

Unilocular sporangia unkown. Plurilocular sporangia, 

uniseriate (occasionally with biseriate portions), with 5–7 
1 9 4 • 

locules, 16–23 μm long; in extensive sori over entire thallus. Paraphyses slightly shorter than plurilocular sporangia.

Habitat. On rocks and tidal platforms; occasionally epiphytic on other algae; mid to low intertidal

Distribution. Gulf of California: Puerto

Peñasco to San José del Cabo. Pacific coast: southern California to Baja California Sur; Sinaloa; Hawaiian Islands; 

Ecuador; Chile.

Type Locality. Near San Pedro, Los Angeles

County, California.

Compsonema Kuckuck

Compsonema Kuckuck, 1899:58.

Algae discoid, composed of a monostromatic basal 

layer of irregular contorted filaments and upper erect 

filaments that are simple and unbranched or branched 

laterally.

Plurilocular sporangia multiseriate, borne on erect 

filaments.

Remarks. Compsonema has been placed in the 

Ectocarpaceae or the Myrionemataceae (now = Chordariaceae) (e.g. Abbott and Hollenberg, 1976). The generic 

status of Compsonema has been questioned. For example, 

Loiseaux (1970a) concluded one species of Compsonema

was a phase in the life history of a Scytosiphon, and on 

the basis of studies of the generitype, Compsonema gracile

Kuckuck (1899), Pedersen and Sokhi (1990) placed the 

genus in the Scytosiphonaceae. Further studies may reveal 

some species of Compsonema in the Gulf of California to 

be microthallus phases in the life history of other brown 

algae.

One species, Compsonema intricatum Setchell et 

N. L. Gardner (1922a), is reported in the southern Gulf 

(Mendoza-González et al. 1994).

Two species are currently known in the northern Gulf 

of California.

Key to the Species of Compsonema In the no

1a. Plurilocular sporangia long and narrow, 24–30 μm long and 6–8 μm in diameter  . . . . .

1b. Plurilocular sporangia longer and much wider, 60–130 μm long and 18–28 μm in diameter. . . . . . . . . . . . . . . C. serpen

Compsonema immixtum

Setchell et N. L. Gardn

Compsonema immixtum Setchell et N. L. Gardner, 

1924a:724, pl. 17: fig. 49; Setchell and Gardner, 

1925:487; Dawson, 1944:222; Dawson, 1961b:390; 

González-González et al., 1996:143; Pacheco-Ruíz et 

al., 2008:204; Pedroche et al., 2008:69.

Algae minute, epiphytic; lower portion of prostrate 

basal filaments, creeping among the plurilocular reproductive structures of the host alga; upper portion with 

numerous erect, short unbranched filaments, most bearing 

plurilocular structures.

Unilocular sporangia not known. Plurilocular sporangia (?), narrow, ellipsoid, 24–30 μm long, and 6–8 μm 

in diameter; terminal on erect, short filaments (extending 

above the plurilocular structures of the host alga).

Habitat. Epiphytic on Colpomenia phaeodactyla; intertidal

Distribution. Gulf of California: Isla

Partida.

Type Locality. Isla Partida, Islas de la Cintura, Gulf of California

Remarks. A poorly known species, Setchell and Gardner (1924a) commented that Compsonema 

immixtum was close to both Myrionema Greville (1827) 

and Compsonema and placed it in the latter genus on 

the basis of its plurilocular structures. Abbott and Hollenberg (1976:160) stated that in their opinion, several of 

the Setchell and Gardner (1922) species of Compsonema, 

including the Gulf of California C. immixtum, did not 

merit recognition. However, it is included herein in hopes 

that when it is once again collected, its taxonomic status, 

life history, and phylogenetic relationships will be investigated and its unique habit and association with Colpomenia phaeodactyla will be elucidated.

Compsonema serpens Setchell et N. L. Gardn

FIGURE 93

Compsonema serpens Setchell et N. L. Gardner, 1922a:363, 

pl. 39: fig. 7; Setchell and Gardner, 1925:480; Smith, 

1944:110, pl. 15: fig. 3; Dawson, 1960b:391; Norris and 

Abbott, 1972:89; Abbott and Hollenberg, 1976:160, 

fig. 128; Mendoza-González and Mateo-Cid, 1992:21; 

Servière-Zaragoza et al., 1993:482; González-González 
n u m b e r 9 4 • 1 9 5

et al., 1996:374; L. Aguilar-Rosas et al., 2000:133; Pedroche et al., 2008:70.

Algae minute, epiphytic, lower portion of irregularly 

branched prostrate filaments with a few endophytic rhizoids attached to host; above erect, unbranched (rarely 

subulate) filaments, up to 550 μm tall. Erect filaments narrower in diameter at base and near the apices. Cells of 

basal filaments 5–9 μm in diameter; wider in midportions, 

10–12(–17) μm in diameter.

Unilocular sporangia not known. Plurilocular sporangia, conical, 60–130 μm long, 18–28 μm in diameter; terminal on erect pedicel of several cells above basal filaments.

Habitat. Epiphytic and endophytic on Padina

durvillei; intertidal.

Distribution. Gulf of California: Playa El

Coloradito to Puertecitos. Pacific coast: British Columbia; 

northern to central California; Jalisco.

Type Locality. Cypress Point, Monterey

County, California.

Remarks. Compsonema serpens was reported 

in the northern Gulf by L. Aguilar-Rosas et al. (2000); 

description of the species is based on Abbott and Hollenberg (1976).

Hydroclathrus Bory de Saint-Vincen

Hydroclathrus Bory de Saint-Vincent, 1825:419.

Algae are a three-dimensional net, often becoming 

torn, perhaps from grazing or physical stress, to form 

small- to large-holed, netlike sheets. Initially saccate, the 

thalli become irregularly shaped and convoluted with 

perforations of varying sizes that sometimes may join 

together to form larger holes. Outer cortical cells are small 

and pigmented, and large colorless medullary cells of up 

to several layers thick are above a usually hollow center. 

Hairs are borne in shallow depressions scattered over the 

thallus surface.

Life history is a heteromorphic alternation of the perforated thallus that produces plurilocular structures and a 

filamentous phase with both plurilocular and unilocular 

structures; the latter are known only in culture (Toste et 

al., 2003a).

One species is reported in the northern Gulf of 

California.

Hydroclathrus clathratus (C. Agardh) M. 

FIGURE 94

Encoelium clathratum C. Agardh, 1823:412.

Hydroclathrus clathratus (C. Agardh) M. Howe,1920:590; 

Setchell and Gardner, 1924a:727; Setchell and Gardner, 1925:543; Dawson, 1944:324; Dawson, 1959a:19; 

Dawson, 1961b:395; Dawson, 1962b:197–198, figs. 

74, 229; Abbott and Hollenberg, 1976:206, fig.170; 

L. Aguilar-Rosas, 1982:30, 31; Stewart and Stewart, 

1984:141; Huerta-Múzquiz and Mendoza-González, 

1985:46; Mendoza-González and Mateo-Cid, 1985:24; 

Mendoza-González and Mateo-Cid, 1986:421; Sánchez-Rodríguez et al., 1989:40; R. Aguilar-Rosas et 

al., 1990:123; Oates and Cole, 1990a:12–16; Stewart, 1991:46; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:32; Martínez-Lozano et al., 1991:23; Mateo-Cid 

et al., 1993:50; González-González et al., 1996:297; 

Pacheco-Ruíz and Zertuche-González, 1996b:171; Kogame, 1997b:228, figs. 7–14; Yoshida, 1998:306; CruzAyala et al., 2001:190; Kraft and Abbott, 2003:246, 

figs. 2–11; Abbott and Huisman, 2004:183, fig. 68C,D; 

FIGURE 93. Compsonema serpens: Erect filaments, with terminal 

plurilocular reproductive structures, arising from prostrate filaments 

(drawn from type specimen, N. L. Gardner-4684, Setchell and 

Gardner, 1922a: pl. 39, fig.7).
1 9 6 • 

R. Aguilar-Rosas et al., 2005b:34; Pacheco-Ruíz et al., 

2008:204; Pedroche et al., 2008:70.

Hydroclathrus cancellatus Bory de Saint-Vincent,

1825:419; Setchell and Gardner, 1924a:727.

Algae netlike, initially more or less globose, then 

becoming irregularly shaped and convoluted with numerous irregularly shaped, variously sized holes; often torn 

into sheets; somewhat coarse and slippery; attached by 

groups of rhizoids along the lower surface. Surface perforations from less than 1 mm in diameter to holes over 

3 cm in width; larger holes often with inrolled margins; 

surface distance between the perforations usually less than 

2 mm. Colorless hairs in clusters of 3–15 within shallow 

pits, scattered over and extended above thallus surface. 

Thallus thickness from less than 200 μm to over 900 μm. 

Medulla of up to 6 layers of thin-walled, subspherical to 

ovoid colorless medullary cells, 60–320 μm, around hollow center. Cortex of 1–2 layers of pigmented cells, 6–7.5 

μm by 7–17 μm; surface layer in transection papillate, of 

domed cells, 10–15(–20) μm tall.

Plurilocular sporangia 5–7 μm by 10–12(–25) μm, in 

2–3 rows of 3–4 locules; in irregularly shaped sori, usually 

near hair tufts. Sori scattered and grow separately, spread 

and coalesce, sometimes covering much of the surface.

Habitat. On rocks or occasionally epiphytic or

entangled on other algae; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Punta

Perla, northeast end of Isla Tiburón to Bahía de Los 

Ángeles; Bahía Concepción to Bahía de La Paz. Pacific 

coast: southern California to Laguna Ojo de Liebre (Scammon’s Lagoon), Baja California Sur; Ecuador; Chile; 

Hawaiian Islands; China; Japan; Vietnam.

Type Locality. “Uncertain” (see Silva et al.,

1996a:631). Lectotype with locality of Belle-Île, Brittany, 

France, was selected by Howe (1920); however, neither 

Hydroclathrus clathratus or mollusk shells found with the 

lectotype collection are known from this locality (Hamel, 

1937; Silva et al., 1996a; Kraft and Abbott, 2003). Other 

syntype locales of the basionym, Encoelium clathratum

C. Agardh (1823), were Rauki, Waigeo Island, Moluccas, 

Indonesia; and Shark Bay, Western Australia.

Remarks. Generally considered a warm-water 

species, Hydroclathrus clathratus was reported in the northern Gulf by Mendoza-González and Mateo-Cid (1986) and 

Pacheco-Ruíz and Zertuche-González (1996b). This description of H. clathratus is after Kraft and Abbott (2003).

Rosenvingea Børgesen

Rosenvingea Børgesen, 1914:22.

Algae are terete to compressed, erect but mostly lax. 

They are branched alternately, oppositely, subdichotomously, or irregularly. Internally, they are hollow, except 

near the base, where they are filled with rhizoidal filaments. Growth is intercalary and apical. A few layers of 

pseudoparenchymatous cells face the hollow center and 

FIGURE 94. Hydroclathrus clathratus: Habit, of a sheetlike thallus with numerous small to large holes (US Alg. Coll.-36619).
n u m b e r 9 4 • 1 9 7

outwardly join the surface layer of small pigmented cells. 

Phaeophycean hairs are distributed over the thallus.

Plurilocular gametangia are borne in sori with groups 

of hairs. Sori are scattered over the thallus surface.

Remarks. The erect, cylindrical species of 

Rosenvingea are problematic, with species largely separated by branching pattern. Characters of vegetative 

anatomy and reproductive structures need to be clarified for type species, and comparative morphological and 

DNA analyses are needed to define species limits. Further 

studies of Gulf of California specimens, including fertile 

material and DNA analyses, are needed to assess their 

taxonomic status in comparison to the western Atlantic 

taxa.

Three species are known in the northern Gulf of 

California.

Key to the Species of Rosenvingea in the n

1a. Algae clumps of low-growing mats of entangled branches; branches more or less cylindrical to compressed, 2–6 mm in 

diameter; attached by rhizoids from branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. intricata

1b. Algae erect; of narrow, terete axes and branches, less than 2 mm in diameter; attached by single holdfast  

2a. Thalli sparsely and irregularly branched; up to 1 mm in diameter; thin in transection (3–4 cell layers)  . . .  R. an

2b. Thalli mostly oppositely or occasionally irregularly branched; mostly 1.5 to 2.5 mm in diameter; thicker in transection (4–6 

cell layers)  . . . . . .

Rosenvingea antillarum

(P. Crouan et H. Crouan) M. J. Wy

Stilophora antillarum P. Crouan et H. Crouan in Schramm et Mazé, 1865:2; Schramm and Mazé, 1866:3; 

Mazé and Schramm, 1878:116.

Rosenvingea antillarum (P. Crouan et H. Crouan) M. 

J. Wynne, 1997:334, fig. 1; L. Aguilar-Rosas et al., 

2002:235; Pedroche et al., 2008 (in part; excluding records of R. floridana):72.

Algae erect, composed of terete, internally hollow, 

axes and branches, up to 25 cm long, and 0.5–1.0 mm in 

diameter. Branching sparse and irregular, 2–3 orders.

Plurilocular reproductive structures uni- to biseriate, 

of 6–7 locules; 32–36 μm long and 4–6 μm in diameter; 

with enlarged cells (“apparently paraphyses”).

Habitat. Intertidal

Distribution. Gulf of California: El Tornillal to Punta Gorda, Sonora

Type Locality. Le Moule, Grande-Terre,

Guadeloupe, French West Indies.

Remarks. Rosenvingea antillarum was 

reported in the upper Gulf by L. Aguilar-Rosas et al. 

(2000); since I have not seen their specimens, the description is based on Wynne (1997).

Rosenvingea floridana 

(W. R. Taylor) W. R. Taylor

FIGURE 95A,B

Cladosiphonia? floridana W. R. Taylor, 1928:113, pl. 15: 

figs 10–14.

Rosenvingea floridana (W. R. Taylor) W. R. Taylor, 

1955:72, pl. IV; Taylor, 1960:262, pl. 29: figs.7, 8; Abbott and Hollenberg, 1976:202, fig. 165; Mateo-Cid 

and Mendoza-González, 1997:59, pl. 8: figs. 34, 35, pl. 

9: fig. 38.

Rosenvingea antillarum sensu Pedroche et al., 2008 (in 

part; only records of R. floridana):72 [non Rosenvingea antillarum (P. Crouan et H. Crouan) M. J. Wynne, 

1997:334].

Rosenvingea aff. sanctae-crucis sensu Norris and Bucher, 

1976:7, fig. 5a,b; Pacheco-Ruíz et al., 2008:204; Pedroche et al., 2008:72 [non Rosenvingea sanctae-crucis

Børgesen, 1914:22, figs. 14–17].

Algae composed of terete, internally hollow axes and 

branches, up to 55 cm long and (0.5–)1.5–2.5 mm in diameter. Branching often predominately opposite, sometimes 

irregular, 2–3(–4) orders; branches narrowed at their base. 

Transection with internal hollow cavity; outer wall composed of 4–5(–6) cell layers, 90–120 μm thick.

Reproductive structures not observed in Gulf material.

Habitat. On rocks or occasionally entangled

with other algae; low intertidal to shallow subtidal, 3–9 

m depths.

Distribution. Gulf of California: El Tornillal to Punta Gorda; Puerto Refugio, Isla Ángel de la

Guarda; Punta la Gringa, Bahía de Los Ángeles. Pacific 

coast: Santa Catalina Island (California Channel Islands); 

Oaxaca.

Type Locality. East Cay, Dry Tortugas

Islands (Dry Tortugas National Park), Florida.

Remarks. The northern Gulf specimens are 

tentatively referred to Rosenvingea floridana. Wynne 

(1997) considered R. floridana to be a synonym of 
1 9 8 • 

R. antillarum. Although both are terete species, R. antillarium has narrower axes (up to 1.0 mm in diameter), is 

thinner (up to 60 μm thick), has 3–4 cell layers, and is 

sparsely and irregularly branched (Wynne, 1997:334, fig. 

1). Our Gulf specimens (Norris and Bucher, 1976) have 

wider axes (1.5–2.5 mm in diameter), are thicker (90–

120 μm), have 4–6 cell layers, and are mostly oppositely 

branched (a character that can be seen in Taylor, 1955:pl. 

IV) to occasionally irregularly branched; and thus, Gulf 

specimens are closer to the characteristics of R. floridana

sensu stricto (see Taylor, 1955:72). A terete specimen of 

Dawson (1944, as “R. intricata”) from Isla Estanque may 

also belong here.

Rosenvingea intricata (J. Agardh) Børges

FIGURE 96

Asperococcus intricatus J. Agardh, 1847:7.

Rosenvingea intricata (J. Agardh) Børgesen, 1914:26; Taylor, 1928:111, pl. 15: figs. 15–17; Dawson, 1944:233 [in 

part]; Taylor, 1945:83 [as “R. intricata prox.”]; DawFIGURE 95. Rosenvingea floridana

(JN-5471): A. Habit. B. Portion of tubular thallus showing opposite branching 

(arrow heads).
n u m b e r 9 4 • 1 9 9

son, 1959a:19; Taylor, 1960:262, pl. 36, fig. 2; Dawson, 1961b:395; Wynne and Norris, 1976:12; HuertaMúzquiz and Mendoza-González, 1985:46; RochaRamírez and Siqueiros-Beltrones, 1991:32; ServièreZaragoza et al., 1993:482; Mateo-Cid et al., 1993:50; 

González-González et al., 1996:298, 380; Mateo-Cid et 

al., 2000a:68; Cruz-Ayala et al., 2001:190; Abbott and 

Huisman, 2004:185, fig. 69A–C; Servière-Zaragoza et 

al., 2007:8; Pacheco-Ruíz et al., 2008:204; Pedroche et 

al. 2008:72.

Algae forming low-growing clumps of more or less 

cylindrical to compressed, crisp, hollow branches, up 

to 30 cm long, and subdichotomously to irregularly 

branched; attached by groups of rhizoids at several places 

along the lower branches or sometimes unattached and 

free-floating. Branches (2–)4–6 mm in diameter, becoming narrower distally to ultimate branches; apices mostly 

rounded. Transection of 3–4 cell layers, less than 100 

μm thick. Cells in surface view in more or less longitudinal series or irregularly arranged. Phaeophycean hairs in 

groups scattered over thallus surface.

Plurilocular structures, biseriate, in small, discrete sori 

scattered over thallus surface.

Habitat. On rocks or shells, in tide pools, or

free-floating; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Punta

Bufeo to Isla Ángel de la Guarda; Bahía Concepción to 

Punta Arena. Pacific coast: Isla Clarión and Isla Socorro 

(Islas Revillagigedo); Guerrero to Oaxaca; China; Japan; 

Hawaiian Islands; Ecuador.

Type Locality. Veracruz, Veracruz (estado),

Gulf of Mexico, Mexico.

Remarks. In the Gulf, Rosenvingea intricata seems to prefer calmer protected habitats, bays, and 

lagoons, often living in large depressions or holes in sand 

bottoms. Large, broad specimens, similar to those illustrated by Dawson (1944:pl. 52, fig. 1, as “R. intricata”) 

from Isla Willard, Bahía Tepoca and Isla Espiritu Santo, 

were recognized to be Colpomenia ramosa W. R. Taylor 

by Wynne and Norris (1976).

Fu

Fucales Bory de Saint-Vincent, 1827c:62.

Fucales Kylin, 1917:309.

Algae of this order are all macroscopic, often very 

large and rather sturdy in structure, flattened to cylindrical, and branched. Branching in flattened forms is dichotomous (e.g., Fucus, common in temperate regions). Others, 

such as Sargassum (abundantly represented in the Gulf 

and widespread in tropical to temperate areas worldwide), 

are variously branched but mostly radial about a central 

cylindrical axis. Growth in most is initiated by a single 

apical cell, usually sunken within an apical pit, and diffuse 

in a few members. Divisions from the apical cell result in 

a parenchymatous structure in many species; only a few 

have a filamentous structure. Cells have several discoid 

plastids and no pyrenoids.

Asexual reproduction is generally absent in the Fucales. 

The majority have only a parenchymatous sporophyte. 

Sexual reproduction is mostly oogamous, but at least 

one species is anisogamous. Members are monoecious or 

dioecious, with reproductive structures developed within 

conceptacles. In some monoecious species the reproductive structures of both sexes are produced within the same 

conceptacle. Vegetative division of the initial cell produces 

a layer of cells which line the conceptacle. The upper lining 

produces hairs which project through the opening of the 

cavity (ostiole), and the basal layer develops paraphyses. 

The oogonia are borne on the conceptacle wall, usually 

on a stalk cell, and the antheridia are on branched filaments. Antheridia normally produce 64–128 biflagellated, 

pyriform-shaped antherizoids. The number of eggs produced by the oogonia is variable. The primary nucleus in 

the oogonium undergoes meiotic divisions followed by a 

single mitotic division of each of the four nuclei produced. 

This results in eight haploid eggs (e.g., as in Fucus), but in 

others, four, six, or seven of them degenerate.

Remarks. On the basis of DNA sequence data, 

Rousseau and de Reviers (1999b) proposed a phylogeny 

of the Fucales that included additional families, merged 

FIGURE 96. Rosenvingea intricata: Upper portion of clump showing compressed tubular branches that narrow in diameter toward 

tips (US Alg. Coll.-163441).
2 0 0 • 

the Cystoseiraceae with the Sargassaceae, and expanded 

the ordinal characters.

One family is represented in the northern Gulf of 

California.

Sarga

Sargassaceae Kützing, 1843:349, 359.

Cystoseiraceae Kützing, 1843:349, 353, as “Cystosireae.”

Members of this family are among the most highly 

morphologically differentiated algae. They have a basal 

attachment that is distinct from the erect axis or stemlike 

parts. The stipe and primary axes are cylindrical, are usually branched, and bear leaflike blades with a basal pedicel. Blades are with or without a midrib or cryptostomata 

(sterile pits) and have smooth, dentate, or serrate margins. 

Vesicles (flotation structures) are usually present and are 

borne laterally, terminally, or within short branchlets.

Reproductive structures, oogonia and antheridia, are 

contained within monoecious or dioecious conceptacles in 

regular branches or in specialized receptacle branchlets. 

Each oogonium develops a single egg, and the antheridia 

produce numerous antherizoids.

There is one genus, Sargassum, well represented in 

northern Gulf and widely distributed throughout the Gulf 

of California.

Sargassum C. Agar

Sargassum C. Agardh, 1820:1.

Algae are light to dark brown and morphologically 

differentiated into holdfasts, stipes, terete to compressed 

axes and branches, leaflike blades, buoyant vesicles, and 

reproductive branchlets (receptacles). Primary axes have 

few to numerous, radially or distichously arranged lateral 

branches of determinate or indeterminate growth and are 

usually above a stipe that is attached below to the substratum by a holdfast that is discoid, crustose, conical, or 

lobed or with protuberances or rhizoidal to rhizomatous. 

Ultimate branchlets are leaflike blades that may be simple 

or divided, subcylindrical, filiform, or compressed to flattened and broad, with entire, dentate, or serrate margins 

and with or without a midrib or cryptostomata (sterile 

pits). Vesicles (usually present) are subspherical to ovoid 

above a pedicel and with or without terminal extensions. 

Growth is from a three-sided cell in an apical depression. 

Internally, the stipe and branch axes are composed of a 

central medulla of elongate cells, surrounded by a core of 

isodiametric cells outward to the meristoderm (surface tissue that acts as meristem with cells often dividing).

Reproductive receptacles (specialized branchlets) 

develop in axils of lateral branches, ultimate branchlets, 

and leaflike blades and are single or grouped in clusters. 

Receptacles are simple or branched and can be terete, 

angular, or compressed, with a smooth, verrucose, or 

spinous surface. Conceptacles within the receptacles are 

ostiolate and contain either oogonia or antheridia or both.

Remarks. Species of Sargassum are worldwide 

in tropical to temperate seas and are often quite large, 

10–100 cm in length (some may be up to 3 m or more tall). 

In the Gulf of California, species of Sargassum comprise 

the bulk of the biomass of fleshly macroalgae, providing 

habitat for diverse communities of epiphytic algae and a 

canopy for understory algae, invertebrates (Brusca, 1980), 

and fishes (Thomson and Lehner, 1976; Thomson et al., 

2000). Gulf species of Sargassum, such as S. sinicola, 

could be important sources for alginates, and it has been 

suggested they could be harvested in spring, the season of 

maxium biomass and high yield (Rodríguez-Montesinos 

et al., 2008).

In addition to those of the northern Gulf, there are 

six other species reported in the southern Gulf (Dawson 

1959a; Huerta-Múzquiz and Mendoza-González 1985; 

Rocha-Ramírez and Siqueiros-Beltones 1990, 1991; 

Martínez-Lozano et al. 1991, Riosmena-Rodríguez et al.

2005): Sargassum cylindrocarpum Setchell et N. L. Gardner (1924a), S. horridum Setchell et N. L. Gardner (1924a); 

S. howelli Setchell (1937b); S. liebmanni J. Agardh (1847); 

S. pacificum Bory de Saint-Vincent (1828), and S. palmeri

Grunow (1915).

A Japanese species, S. muticum (Yendo) Fensholt 

(1955), introduced in British Columbia in the 1940s (Scagel, 1956, 1957) now ranges on the Pacific coast from 

southeast Alaska to Baja California Sur (Abbott and Hollenberg, 1976; Silva, 1979; Norton, 1981; L. Aguilar-Rosas 

et al., 1982; Deysher and Norton, 1982; R. Aguilar-Rosas 

et al., 1984a, 1984b; R. Aguilar-Rosas and L. AguilarRosas, 1985, 1993; Scagel et al., 1989; R. Aguilar-Rosas 

and Machado-Galindo, 1990; Espinoza, 1990b; RochaRamírez and Siqueiros-Beltrones, 1991; Stewart, 1991; R. 

Aguilar-Rosas et al., 2005b). Riosmena-Rodríguez et al. 

(1992) have suggested the invasive S. muticum in Pacific 

Baja California could continue its expansion southward 

and may colonize into the Gulf of California.

Eight species, two subspecies, and one form of Sargassum are found in the northern Gulf of California.
n u m b e r 9 4 • 2 0 1

Sargassum subgen. Sargassum

Remarks. Dawson (1944, as “S. subgen. 

Eusargassum”) considered all the Gulf of California species of Sargassum to be in S. subgen. Sargassum. However, Phillips (1995:118, tbl. 1) noted their subgeneric 

placement to be “unknown.” Thus, as has been previously 

noted, there is still much work to be done.

The four morphological groups of Dawson (1944)

for the Gulf of California species of Sargassum are herein 

described and designated by the name chosen by him for 

the principal or most representative species of each group. 

Recognizing that many of the Gulf species of Sargassum

are problematic, poorly known, or inadequately defined, it 

is helpful in attempting to identify specimens to first associate them with the suite of characters of these morphological groups.

Sargassum sect. johnstonii E. Y. Dawson e

N. Norris, sect. nov.

“The Johnstonii Group” of Dawson, 1944:239, pls. 32, 

33.

Latin Description. Sectio cum sequentibus

characteribus: laminae folio similes, maxime angustae, 

sub-cylindraceae usque ad compressae, sine costa, et cryptostomata frequenter praesentia; vesiculae ellipticae cum 

longis apiculatis extensionibus.

The section is characterized by leaf-like blades that 

are very narrow, subcylindrical to compressed, without a 

midrib, and with cryptostomata usually present; vesicles 

are elliptical and with long apiculate extensions.

Type Species  Sargassum johnstonii Setchell e

N. L. Gardner, 1924a:737, pl. 20: fig. 72, pl. 21: fig. 80.

Key to the Species of Sargassum in the 

1a. Leaf-like blades without a midrib; holdfast small, discoid,conical, or irregularly discoid to spreading crust . . . . . . . . . . 2

1b. Leaf-like blades either with inconspicuous or distinct midribs; holdfast not crustose. . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

2a. Leaf-like blades subcylindrical to slightly compressed, long and very narrow, 10–22 mm long by 1.0–1.5 mm wide; vesicles 

with long apiculate extensions; holdfast disc-shaped to spreading crust . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. johnstonii

2b. Leaf-like blades wider and mostly asymmetrical  

3a. Upper leaf-like blades, to 8 mm long by 4–7 mm wide; some almost as wide as long; discoid holdfast  . . . . . S. 

3b. Upper leaf-like blades larger, mostly longer than 12 mm in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4a. Axes branched; leaf-like blades 12–16 mm long; vesicles subspherical to ellipsoidal, smooth, with short apiculate extensions; holdfast presumably discoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. acinacifolium

4b. Axes unbranched; upper leaf-like blades mostly 20–25 mm long; vesicles ellipsoidal with long, usually dentate, foliaceous 

extensions; holdfast knotty, of short protuberances. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sonorense

5a. Leaf-like blades variable in shape and size, with an inconspicuous midrib; 5.0–12.5 mm long: smallest blades symmetrical 

and dentate; midsized blades asymmetrical, as wide as long, and partly dentate; and largest blades symmetrical, elongated, 

and dentate; holdfast discoid  . . . . . . 

5b. Leaf-like blades symmetrical, longer than wide, with a conspicuous midrib; holdfast of long branching rhizomes or woody 

and conical or knotty with short protuberances  

6a. Several thalli arising from long, branched, narrow in diameter rhizomes that are attached below at various points by haptera; leaf-like blades usually without cryptostomata (or rarely with only a few cryptostomata)  

6b. Thallus arising from a single woody, discoid to conical holdfast or a coarse, knotty holdfast; leaf-like blades with conspicuous to inconspicuous cryptostomata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

7a. Leaf-like blades mostly smaller, 15–25 mm by 2–5 mm; with a midrib (often less distinct or not evident toward apices) and 

slightly and irregularly dentate or uneven margins; without or with inconspicuous cryptostomata  . . . . . S. 

7b. Leaf-like blades mostly larger, 30–60(–110) mm by 4–10 mm; with a distinct midrib and serrate margins of regularly, evenly 

spaced, pronounced dentation; mostly without cryptostomata (or rarely a few cryptostomata on only a few blades) 

. . . . . . . 

8a. Thalli usually lax; axes smooth; holdfast coarse, knotty, with short, thick protuberances  . . . . . .  S. sinicola s

8b. Thalli usually not as lax; axes muricate; holdfast conical, woody . . . . . . . . . . . . . . . . . . . . . . . . . S. sinicola subsp. camouii
2 0 2 • 

There is one species, and one form in the northern 

Gulf of California.

Sargassum johnstonii Setchell et N. L. Gardn

FIGURES 6, 97A,B

Sargassum johnstonii Setchell et N. L. Gardner, 1924a:737, 

pl. 20: fig. 72, pl. 21: fig. 80; Setchell and Gardner, 

1925:719; Dawson, 1944:240, pl. 23: figs. 1–15, pl. 

33: figs. 1, 2, 17–22; Dawson, 1961b:400; Dawson, 

1966a:12; Norris, 1973:7, pl. 13; Huerta-Múzquiz, 

1978:337; Littler and Littler, 1981:151, tbl. 3, figs. 4, 

5, 7; McCourt, 1984a:142; McCourt, 1984b:141; Mendoza-González and Mateo-Cid, 1985:26; MendozaGonzález and Mateo-Cid, 1986:421; Rocha-Ramírez 

and Siqueiros-Beltrones, 1990:19; Rocha-Ramírez 

and Siqueiros-Beltrones, 1991:26; Espinoza-Ávalos, 

1993:333; Phillips, 1995:118; González-González et 

al., 1996:161; Pacheco-Ruíz and Zertuche-González, 

1996b:171; L. Aguilar-Rosas et al., 2002:235; MateoCid et al., 2006:50, 58; Pacheco-Ruíz et al., 2008:205; 

Pedroche et al., 2008:95.

Sargassum guardiense Setchell et N. L. Gardner, 

1924a:723, pl. 19: fig. 64; Setchell and Gardner, 

1925:713; González-González et al., 1996:160.

Algae with several erect axes, up to 1(–2) m long, 

arising from a perennial, parenchymatous, disc-shaped 

to spreading crustose holdfast. Axes and branches terete 

and smooth. Leaf-like blades narrow, lanceolate to linear, 

slightly compressed to flattened, up to 10 mm long and 

0.5–1.5 mm wide; margins smooth to sparsely dentate; 

without a midrib and with few to many cryptostomata. 

Vesicles on short pedicels, subspherical to elliptical, up 

to 4 mm long and 1 mm wide; apiculate with foliaceous 

crests or extensions, up to 5 mm long.

Receptacle branchlets narrow in diameter, branching up to 4 orders, dentate in varying degrees. Antheridial 

receptacles smaller in diameter than oogonial receptacles.

Habitat. On rocks; high to low intertidal;

occasionally shallow subtidal.

Distribution. Gulf of California: El Tornillal to Bahía de La Paz. Pacific coast: Baja California Sur

Type Locality. Isla Jorge (George’s Island),

off Sonora, Gulf of California.

Remarks. The distinctive crustlike, spreading 

holdfast of Sargassum johnstonii and its narrow (up to 

1.5 mm wide), lanceolate to linear leaflike blades make 

it one of the Gulf species easiest to recognize in the field. 

Although generally considered a Gulf-endemic species, S. 

johnstonii has also been reported from India (Thivy and 

Chauhan, 1964; Silva et al., 1996).

Sargassum johnstonii f. gracile

Setchell et N. L. Gardn

FIGURE 98

Sargassum johnstonii f. gracile Setchell et N. L. Gardner, 1924a:738, pl. 21: fig. 76; Setchell and Gardner, 

1925:720; Dawson, 1944:241, pl. 33: figs. 3–16; Dawson, 1961b:400; Norris, 1973:7; González-González et 

al., 1996:161; Mateo-Cid et al., 2006:50; Pedroche et 

al. 2008:96.

Sargassum johnstonii f. laxius Setchell et N. L. Gardner, 

1924a:737, pl. 21: figs. 75, 81; Setchell and Gardner, 

1925:719; Norris, 1973:7; González-González et al., 

1996:161.

Although very similar to Sargassum johnstonii f. johnstonii, this form, S. johnstonii f. gracile, differs in being 

more slender, lax, and longer, with fewer branches; having 

leaf-like blades less closely spaced and longer, up to 22.5 

mm long, usually filiform and not as flattened, with abundant conspicuous cryptostomata; and having vesicles with 

long, terete apicula.

Habitat. Often floating, in drift, or cast ashore;

probably low intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas.

Type Locality. Cast ashore, Guaymas,

Sonora, Gulf of California.

Remarks. Setchell and Gardner (1924a) noted 

the type specimen lacked basal parts.

Sargassum sect. lapazeanum 

E. Y. Dawson ex J. N. Norris, sect. n

“The Lapazeanum Group” of Dawson, 1944:241, pls. 

34–37.

Latin Description. Sectio cum sequentibus

characteribus: laminae folio similes planae, expansae et 

assymetricae, cum costa absenti vel aliquando presenti sed 

inconspicua, afixae cum hapterone rhizomatozo.

The section is characterized by leaf-like blades that are 

flattened, expanded, and asymmetrical, without a midrib 
n u m b e r 9 4 • 2 0 3

FIGURE 97. Sargassum johnstonii: A. Upper portion of a branch showing the narrow, elongate leaf-like blades, elliptical vesicles with apiculate 

extensions, and receptacle branchlets (EYD-27204). B. Habit, multiple axes arising from a crustose holdfast (JN-4559).
2 0 4 • 

or occasionally, if present, inconspicuous, and attached by 

rhizomatous holdfast.

Type Species  Sargassum lapazeanum Setchel

et N. L. Gardner, 1924a:733, pl. 20: fig. 74.

There are four species found in the northern Gulf of 

California.

Sargassum lapazeanum 

Setchell et N. L. Gardn

FIGURE 99A,B

Sargassum lapazeanum Setchell et N. L. Gardner, 

1924a:734, pl. 20: fig.74; Setchell and Gardner, 

1925:714; Dawson, 1944:243, pl. 34: figs. 1–34; Dawson, 1961b:400; Norris, 1973:7; Huerta Muzquiz, 

1978:337; Huerta-Múzquiz and Mendoza-González, 

1985:46; Mendoza-González and Mateo-Cid, 1986:421; 

Rocha-Ramírez and Siqueiros-Beltrones, 1990:18; Rocha-Ramírez and Siqueiros-Beltrones, 1991:26; Martínez-Lozano et al., 1991:23; Mateo-Cid et al., 1993:50; 

Phillips, 1995:118; González-González et al., 1996:161; 

Núñez-López and Casas-Valdez, 1997:19; Paul-Chávez 

and Riosmena-Rodríguez, 2000:137; Cruz-Ayala et al., 

2001:190; Rivera and Scrosati, 2006:178; Rivera and 

Scrosati, 2008:45–49; Pacheco-Ruíz et al., 2008:205; 

Pedroche et al., 2008:96.

FIGURE 98. Sargassum johnstonii f. gracile: Habit, isotype specimen (D. R. Marchant-28a; US Alg. Coll.-56958).
n u m b e r 9 4 • 2 0 5

Sargassum bryantii Setchell et N. L. Gardner, 1924a:733, 

pl. 21: fig. 83.

Sargassum insulare Setchell et N. L. Gardner, 1924a:735, 

pl. 20: figs. 67, 68, pl. 21: fig. 78; Setchell and Gardner, 

1925:717.

Erect axes, up to 1 m or more long, above a terete stipe, 

up to 2 cm long; with 5–7 terete, smooth primary branches 

from which several long, slender secondary branches arise; 

attached by a parenchymatous disc, formed by fusing of 

short haptera. Leaf-like blades, variable in shape, 5–20 

FIGURE 99. Sargassum lapazeanum: A. Habit. B. Close-up showing smooth axes and small leaflike blades with dentate margins and prominent 

cryptostomata. A, B. Isotype specimen (D. R. Marchant-21; US Alg. Coll.-56961).
2 0 6 • 

FIGURE 100. Sargassum acinacifolium: A. Upper portion of a branch showing a leaf-like blade with coarsely dentate portion, several vesicles 

with apiculate extensions, and receptacle branchlets (drawn from type, Setchell and Gardner, 1924a: pl. 21, fig. 82). B, C. Isotype specimen (T. 

S. Brandegee-2; UC 221158).
n u m b e r 9 4 • 2 0 7

mm long, with a short pedicel, without or sometimes with 

an inconspicuous midrib, and few to abundant conspicuous cryptostomata. Smallest blades more or less symmetrical with dentate margins; midsized blades asymmetrical, 

often as wide as long in midportions, with crenate to 

dentate lower margins and often smooth upper margins; 

upper blades symmetrical, larger, often widest toward apices, and with dentate margins. Vesicles among receptacles, 

spherical to subspherical, ellipsoidal, up to 2 mm long, 

usually with a short, small apical blade-like extension.

Receptacles intermixed with blades and vesicles; on 

short pedicel at base of a blade; spinulose, branched 2–3 

times, up to 7 mm long.

Habitat. On rocks; mid intertidal to shallow

subtidal.

Distribution. Gulf of California: Bahía San

Francisquito to Cabo San Lucas.

Type Locality. La Paz, Baja California Sur,

Gulf of California.

Sargassum acinacifolium 

Setchell et N. L. Gardn

FIGURE 100A–C

Sargassum acinacifolium Setchell et N. L. Gardner, 

1924a:732, pl. 21: fig. 82; Setchell and Gardner, 

1925:713; Dawson, 1944:244, pl. 37: figs. 1–26; Dawson, 1961b:399; Dawson, 1966a:12; De Lara Isassi, 

1992:22; González-González et al., 1996:159; MateoCid et al., 2006:50; Pedroche et al., 2008:92.

Algae erect with terete, smooth axes and branches; leaflike blades asymmetrical, mostly 12–16 mm long, without 

a midrib and originally described without cryptostomata 

(see Remarks below); upper side (adaxial) of blades with 

concave, smooth margins and lower side and apical margins coarsely dentate; holdfast unknown (“probably a parenchymatous disc;” see Dawson, 1944:244). Vesicles on a 

short pedicel among or at base of receptacles; subspherical to ellipsoidal, 1.5–2.5 mm in diameter, smooth, with 

short, simple apicula extending to about half of vesicle’s 

length.

Oogonial receptacles nearly cylindrical and branched 

2–3 times, smooth or slightly dentate toward apices (Dawson, 1944:fig. 18). Antheridial receptacles branched 2–3 

times and dentate (Dawson, 1944:figs. 17, 24).

Habitat. On rocks and tidal platforms; mid

intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas.

Type Locality. “Guaymas (?),” Sonora, Gulf

of California (Setchell and Gardner, 1924a).

Remarks. Sargassum acinacifolium is apparently a northern Gulf endemic. The basal parts of the type 

specimen are unknown (Setchell and Gardner, 1924a), 

although Dawson (1944) has suggested they are probably 

a parenchymatous disc. Although the leaf-like blades were 

described as being without cryptostomata (Setchell and 

Gardner, 1924a), illustrations of S. acinacifolium by Dawson (1944:pl. 37: figs. 2–3, 19–26; the latter figures were 

drawn from the type specimen) show some to have cryptostomata, thus sharing that character with S. lapazeanum

and S. sonorense. Until S. acinacifolium can be adequately 

described and molecularly and phylogenetically analyzed, 

it remains a poorly known species.

Sargassum macdougalii E. Y. Da

FIGURE 101A,B

Sargassum macdougalii E. Y. Dawson, 1944:245, pl. 35: 

figs. 1–8, pl. 36: figs. 1–7, pl. 37: figs. 27–30; Dawson, 

1959a:19; Dawson, 1961b:400; Dawson, 1966b:59; 

Huerta-Múzquiz and Mendoza-González, 1985:46; Espinoza-Ávalos, 1993:333; Phillips, 1995:118; GonzálezGonzález et al., 1996:161; Pacheco-Ruíz and ZertucheGonzález, 1996b:171; Pacheco-Ruíz et al., 2008:205; 

Pedroche et al., 2008:97.

Several erect terete, smooth axes, up to 50 cm long, 

arising from a slender stipe, to 1 cm long, attached below 

by a discoid holdfast. Leaf-like blades asymmetrical, without a midrib and lacking cryptostomata; mostly up to 8 

mm long by (2.5–)4–6(–7) mm wide (sometimes almost 

as wide as long); upper blades usually larger, longer than 

wide, up to 15(–20) mm long. Blade margins sharply denticulate (usually more prominent at apices); lower portion 

of upper side of blade concave and smooth. Vesicles ellipsoidal, with foliar apical extensions (usually about as long 

as length of vesicle).

Receptacles unknown.

Habitat. On rocks; mid intertidal to shallow

subtidal.

Distribution. Gulf of California: Puerto Libertad to Guaymas; Puerto Escondido to Bahía de La Paz

Type Locality. Puerto Libertad, Sonora,

Gulf of California.

Remarks. Sargassum macdougalii is said to 

differ from S. lapazeanum primarily by its longer vesicles 

with elongated foliaceous extensions and larger upper 

leaflike blades, up to 15(–20) mm long (Dawson, 1944). 
2 0 8 • 

FIGURE 101. Sargassum macdougalii: A. Habit (JN-5975). B. Upper portion of an axis showing characters of asymmetrical leaflike blades and 

some vesicles (JN-5944).
n u m b e r 9 4 • 2 0 9

Both S. macdougalii and S. lapazeanum are attached by 

parenchymatous discoid holdfasts. The nature of the holdfast helps separate them in the field from S. sonorense, 

which has a knotty holdfast with short protuberances.

Sargassum sonorense E. Y. Daw

FIGURE 102

Sargassum sonorense E. Y. Dawson, 1960a:36; Dawson, 

1961b:400; Phillips, 1995:118; González-González et 

al., 1996:163; Mateo-Cid et al., 2006:50; Pedroche et 

al., 2008:102.

Sargassum asymmetricum E. Y. Dawson, nom. illeg.,

1944:244, pl. 36: figs. 8–18; Dawson, 1966a:12; 

González-González et al., 1996:159 [non Sargassum asymmetricum Yamada, 1942:563, figs. 30, 31; 

Yamada, 1944:3 (which is now a synonym of Sargassum alternato-pinnatum Yamada, 1942; see Yoshida, 1988; Noro and Nanba, 1990; Ajisaka et al., 

1994)].

FIGURE 102. Sargassum sonorense: Habit of young thallus showing asymmetrical leaflike blades and holdfast of short protuberances (arrow) 

(JN-5205).
2 1 0 • 

Algae erect of terete, smooth primary axes, unbranched, 

up to 60 cm long, arising above a slender stipe, up to 1 cm 

long, attached below by a small, knotty holdfast of short 

protuberances, up to 1.5 cm wide. Leaf-like blades, mostly 

asymmetrical, resulting from a usually smooth, concave 

upper side; without a midrib; with small, inconspicuous 

cryptostomata; blade margins dentate. Upper blades 20–25 

mm long, sometimes more or less lobed; lowermost blades 

usually smaller, 3–6 mm long, and variable in shape: from 

lanceolate with blunt apices to more or less triangular. Vesicles ellipsoid, usually with dentate foliaceous extensions, 

1–3 times longer than vesicle.

Oogonial receptacles unknown. Antheridial receptacles, up to 1.0 cm long, branched 2–4 times, upper portions dentate.

Habitat. On rocks; low intertidal to shallow

subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas.

Type Locality. On shore, southern end of

Isla Tiburón (opposite Isla Turners), Las Islas de la Cintura, Gulf of California.

Remarks. Sargassum sonorense can be separated from other morphologically similar members of this 

section primarily by its holdfast, which is knotty, with 

short protuberances, in contrast to the parenchymatous 

discoid holdfast found in the other species of this group. In 

addition to its holdfast, S. sonorense differs from similar S. 

acinacifolium by its larger blade sizes. These two are both 

problematic species, and their relationship needs further 

study.

Originially described as Sargassum asymmetricum

E. Y. Dawson (July 1944), it was the second use of the 

name “asymmetricum” in Sargassum. The earliest use 

of the name was S. asymmetricum Yamada (1942; April 

1944) from Japan. Therefore, since S. asymmetricum E. Y. 

Dawson is a later homonym, a replacement name, S. sonorense, was given for the northern Gulf of California species by Dawson (1960a).

Sargassum sect. sinicola E. Y. Dawson ex J.

Norris, sect. nov.

“The Sinicola Group” of Dawson, 1944:245, pls. 38–39.

Latin Description. Sectio cum sequentibus

characteribus: laminae folio similes, longiores quam latae, 

symmetricae, cum costa, margines dentatae usque ad serratae, cryptostomata presentia, saepe abundantia; vesiculae 

sub-sphaericae usque ad sphaericae; hapteron ligneum, 

conicum vel nodosum cum parvis protuberationibus.

The section is characterized by leaf-like blades that 

are longer than wide, symmetrical, with a midrib, dentate to serrate margins, and cryptostomata present, often 

abundant; vesicles are subspherical to spherical; holdfasts 

are woody, conical, or knotty, with short protuberances.

Type Species  Sargassum sinicola Setchell e

N. L. Gardner, 1924a:736, pl. 20: fig. 73.

Remarks. There are two subspecies of Sargassum sinicola known in the northern Gulf of California. 

Another member of this section, S. horridum Setchell et 

N. L. Gardner (1924a), is known from the southern Gulf 

(Dawson, 1944, 1959.; Pedroche et al., 2008).

Sargassum sinicola Setchell et N. L. Gardn

subsp. sinicola

FIGURES 103, 104, 105A

Sargassum sinicola Setchell et N. L. Gardner, 1924a:736, 

pl. 20: fig. 73; Setchell and Gardner, 1925:718; Dawson, 1944:247, pl. 35: fig. 10, pl. 38: figs. 5–11, pl. 39: 

figs. 1–11; Dawson, 1959a:19; Dawson, 1961b:400; 

Dawson, 1966a:13; Norris, 1973:8; Huerta-Múzquiz, 

1978:337; Stewart and Stewart, 1984:142; McCourt, 

1984b:141; Huerta-Múzquiz and Mendoza-González, 

1985:46; Mendoza-González and Mateo-Cid, 1985:26; 

Mendoza-González and Mateo-Cid, 1986:421; Ortega 

et al., 1987:77; Espinoza-Ávalos and Rodríguez-Garza, 

1985:115, fig. 1; Espinoza-Ávalos and RodríguezGarza, 1987:183; Sánchez-Rodríguez et al., 1989:41; 

Espinoza-Ávalos and Rodríguez-Garza, 1989:141; Espinoza, 1990a:1; Rocha-Ramírez and Siqueiros-Beltrones, 

1990:20 [in part]; Rocha-Ramírez and Siqueiros-Beltrones, 

1991:26; Martínez-Lozano et al., 1991:23; Mateo-Cid 

et al., 1993:51; Phillips, 1995:118; González-González 

et al., 1996:163; Pacheco-Ruíz and Zertuche-González, 

1996b:171; Riosmena-Rodríguez et al., 1998:45; MateoCid et al., 2000a:68; Cruz-Ayala et al., 2001:190; L. Aguilar-Rosas et al., 2002:235; Mateo-Cid et al., 2006:50, 58; 

Pacheco-Ruíz et al., 2008:205; Rodríguez-Montesinos et 

al., 2008:33; Pedroche et al., 2008:100.

Sargassum polyacanthum f. americanum Setchell et N. L. 

Gardner, 1924a:736; Setchell and Gardner, 1925:718; 

González-González et al., 1996:162.

Algae up to 60 cm long; of smooth axes and branches 

above a short stipe, attached below by a knotty, coarse 
n u m b e r 9 4 • 2 1 1

holdfast with short, thick protuberances. Leaf-like 

blades, long and narrow, very variable in size, 30–60(–

90) mm long and (2–)3–5 mm wide; with a midrib, 

inconspicuous cryptostomata, and irregularly dentate 

margins. Vesicles smooth, mostly spherical, up to 6 mm 

in diameter, with a pedicel, and sometimes with a small 

terminal apicula.

Receptacle branchlets forked up to 4 times, subcylindrical, smooth, with acuminate apices, and scattered 

among vesicles in upper portions of the thallus. Oogonial 

FIGURE 103. Sargassum sinicola subsp. sinicola: Habit; note holdfast of rhizomelike protuberances (arrow) (EYD-21568).
2 1 2 • 

receptacles generally shorter and more robust than the 

antheridial receptacles.

Habitat. On rocks and tidal platforms; usually

low intertidal to shallow subtidal, occasional in mid intertidal tide pools.

Distribution. Gulf of California: Piedras de

La Salina to Cabo San Lucas. Pacific coast: Isla Guadalupe; Baja California Sur.

Type Locality. Eureka, near La Paz, Baja

California Sur, Gulf of California.

FIGURE 104. Sargassum sinicola subsp. sinicola: Habit, smooth axes and long, leaflike blades with irregularly dentate margins and vesicles 

with a pedicel (JN-3119).
n u m b e r 9 4 • 2 1 3

Remarks. The most common of the species of 

Sargassum in the northern Gulf, S. sinicola usually reaches 

its largest size in spring. Sargassum sinicola subsp. sinicola is recognized by its characteristic coarse, knotty holdfast, smooth branch axes, leaf-like blades that are always 

longer than wide and with dentate margins, a percurrent 

midrib and cryptostomata, and its round, smooth vesicles. Unusual reproductive forms of S. sinicola, with both

oogonial and antheridial conceptacles in the receptacle 

branchlets, were reported from Puerto Peñasco by Dawson (1944:pl. 39: figs. 6, 7).

Some specimens of S. sinicola can be confused with 

the southern Gulf S. horridum. Characters pointed out 

by Rocha-Ramírez and Siqueiros-Beltrones (1991) that 

distinguish S. horridum are spinose or terminally dentate 

receptacles of oogonial thalli, strongly muricate stipes and 

axes, leaf-like blades that are wavy or ruffled. Both S. sinicola subsp. sinicola and S. horridum primarily differ from 

S. sinicola subsp. camouii in having holdfasts of short, 

rhizome-like protuberances.

Another species, Sargassum californicum (Grunow) 

Setchell (1937b; basionym S. fuliginosum var.? californica 

Grunow, 1916), was listed as a synonym of S. sinicola by 

Dawson (1944). Since it was described from the Pacific 

coast (Grunow, 1916: 173, type locality: “ad litora Californiae”) and was subsequently reported in the Gulf of 

California (Setchell and Gardner, 1924a, as “S. paniculatum”), it has remained a poorly known and largely 

overlooked species (e.g., not mentioned by Abbott and 

Hollenberg, 1976, or Phillips, 1995). Specimens of “S. 

paniculatum” sensu Setchell and Gardner (1924a, 1925, 

1930; non S. paniculatum J. Agardh. 1848) from the Gulf, 

along with those from Isla Guadalupe (Pacific Mexico), 

were also referred to S. californicum by Setchell (1937b) 

when he made the taxonomic combination. Shortly after 

this, Dawson (1944) considered S. californicum (Grunow) 

Setchell (1937b) to be a synonym of the Gulf S. sinicola

and also included “S. paniculatum” sensu Setchell and 

Gardner (1924a, 1925, 1930). Since then, this taxonomic 

interpretation of S. sinicola has been followed (e.g., Dawson, 1961b; Stewart and Stewart, 1984; Rocha-Ramírez 

and Siqueiros-Beltrones, 1990; González-González et 

al., 1996). Although presumably somewhat similar to S. 

sinicola, the Pacific coast S. californicum appears to differ in having slightly muricate axes, leaf-like blades that 

lack a midrib, and subspherical to ovate vesicles, some 

slightly pointed, on relatively long pedicels (cf. Grunow, 

1916; Setchell and Gardner, 1925:pl. 46: fig. 5). Thus, 

the morphology and relationship of the little-known California S. californicum to the Gulf S. sinicola needs to be 

reinvestigated. Either these two are taxonomically the 

same species, as proposed by Dawson (1944), or S. californicum is a separate species.

Sargassum sinicola subsp. camouii

(E. Y. Dawson) J. N. Norris et Yensen, comb.

FIGURES 105B, 106

Sargassum camouii E. Y. Dawson, 1944:248, pl. 35: figs. 

9, 11–13; Dawson, 1961b:399; Dawson, 1966a:13; 

Norris, 1973:12, 20; McCourt, 1984a:142; McCourt, 

1984b:141; Huerta-Múzquiz and Mendoza-González, 

1985:46; Mendoza-González and Mateo-Cid, 1986:421; 

Sánchez-Rodríguez et al., 1989:41; De Lara Isassi, 

1992:22; Mateo-Cid et al., 1993:51; González-González 

et al., 1996:159; Mateo-Cid et al., 2006:50, 58.

Sargassum sinicola var. camouii (E.Y. Dawson) J. N. Norris et Yensen in Pedroche et al., 2008:101.

Algae erect and rigid, up to 60 cm tall, axes rough, 

usually muricate, and densely branched; attached by solid, 

woodlike holdfast, initially discoid, becoming conical. 

Leaf-like blades lanceolate, mostly 30–60 mm long and 

3–6 mm wide; with a midrib, dentate margins, and numerous cryptostomata (not always as evident on dried specimens). Vesicles spherical, smooth, 3–4.5 mm in diameter.

Oogonial receptacles, more or less cylindrical, 

branched 1–3 orders, and somewhat spiny.

Habitat. On rocks and tidal platforms and in

tide pools; mid intertidal to shallow subtidal.

Distribution. Gulf of California: Puerto

Peñasco to Isla San Esteban; Bahía Concepción to Bahía 

de La Paz. Pacific coast: Bahía Sebastián Vizcaíno to Bahía 

Magdalena, Baja California Sur.

Type Locality. South end of Isla Tiburón,

Las Islas de la Cintura, Gulf of California.

Remarks. The new combination Sargassum 

sinicola subsp. camouii is proposed to reflect its possible 

relationship to S. sinicola. Although similar in most characteristics with S. sinicola subsp. sinicola, S. sinicola subsp. 

camouii is generally darker in color and is distinguished by 

its woody conical holdfast (see Dawson, 1944:pl. 35: fig. 

9), rough to muricate primary axis, mostly muricate upper 

axes, and a more rigid habit. Critical studies, including 

DNA analyses, are needed to test its taxonomic and phylogenetic status.

Unfortunately, the type specimen lacked a holdfast 

(Dawson, 1944); it was the only specimen saved when 

all others were lost on a boat trip returning from Isla 
2 1 4 • 

Tiburón, and the description of the holdfast was based on 

field notes and another (paratype) specimen from another 

locality, “a bay, 10 miles northwest of Guaymas” (probably Bahía San Carlos).

Sargassum sect. herporhizum 

E. Y. Dawson ex J. N. Norris, sect. n

“The Herporhizum Group” of Dawson, 1944:249, pl. 40: 

figs. 1–26.

Latin Description. Sectio cum sequentibus

characteribus: laminae folio similes cum costa, cum cryptostomatibus absentibus vel, si praesentibus, inconspicuis; 

vesiculae cum apiculata extensione; hapteron cum lungis, 

expansis, saepe intricatis rhizomatibus.

The section is characterized by leaf-like blades that 

have a midrib and are without cryptostomata or, if present, with inconspicuous cryptostomata; vesicles with an 

apiculate extension; holdfast of long, spreading, often 

entangled rhizomes.

Type Species  Sargassum herporhizum Setchell et N. L. Gardner, 1924a:739, pl. 20: figs. 69–7

Remarks. Currently there are two species 

from the northern Gulf of California and another, Sargassum liebmannii J. Agardh (1847, 1889), from Pacific 

Mexico (Baja California to Oaxaca; González-González et 

al., 1996) that belong to this section.

Sargassum herporhizum 

Setchell et N. L. Gardn

FIGURES 107, 108A,B

Sargassum herporhizum Setchell et N. L. Gardner, 

1924a:739, pl. 20: figs. 69–71; Setchell and Gardner, 

1925:720; Dawson, 1944:249, pl. 40: figs. 1–12; Dawson, 1961b:400; Dawson, 1966a:13; Norris, 1973:7; 

Huerta-Múzquiz, 1978:337; McCourt, 1984b:141; 

FIGURE 105. Comparison of Sargassum sinicola: A. Sargassum sinicola subsp. sinicola, showing smooth axes, irregular dentate margin of 

leaflike blades, vesicles, and developing receptacles (JN-3119). B. Sargassum sinicola subsp. camouii, showing muricate axes, regular dentate 

margins of leaflike blades, and vesicles (JN-3250).
n u m b e r 9 4 • 2 1 5

Rocha-Ramírez and Siqueiros-Beltrones, 1990:19; De 

Lara Isassi, 1992:22; Espinoza-Ávalos, 1993:333; Phillips, 1995:118; González-González et al., 1996:160; 

Pacheco-Ruíz and Zertuche-González, 1996b:171; 

Cruz-Ayala et al., 2001:190; Mateo-Cid et al., 2006:50, 

58; Pacheco-Ruíz et al., 2008:205; Pedroche et al., 

2008:93.

Algae up to 30 cm long, with several erect terete, 

smooth axes, alternately to irregularly branched, often 

FIGURE 106. Sargassum sinicola subsp. camouii: Habit; note insert (lower right) showing conical holdfast attached to rock (JN-6064).
2 1 6 • 

whorled, above entangled rhizomes. Primary axis initially arising from a small discoid base that develops 

numerous, branching, narrow in diameter (less than 3 

mm), repent cylindrical rhizomes that are attached at 

various points by small haptera; several erect secondary 

axes arise along horizontal rhizomes as they lengthen. 

Leaf-like blades, long and narrow, 15–25 mm long and 

2–5 mm wide, with a midrib (sometimes less distinct or 

faint on uppermost blades); cryptostomata absent or few 

and inconspicuous; margins vary, uneven or slightly to 

FIGURE 107. Sargassum herporhizum: Habit, showing basal attachment of entangled rhizomes (JN-4963).
n u m b e r 9 4 • 2 1 7

irregularly dentate. Vesicles spherical to ovoid, mostly 

smooth, up to 3 mm in diameter; on pedicels; occasionally apiculate.

Dioecious; receptacle branchlets scattered among vesicles, narrow, subcylindrical, up to 10 mm long, branched, 

on short pedicel, with conspicuous conceptacles.

Habitat. On rocks and tidal platforms and in

tide pools; mid intertidal to shallow subtidal, down to 6 

m depths.

Distribution. Gulf of California: Puerto

Peñasco to Bahía de La Paz. Pacific coast: Jalisco; Galápagos Islands.

Type Locality. Isla San Jorge (George’s

Island), off Sonora, Gulf of California.

Remarks. Seasonal observations of Sargassum 

herporhizum at Playa Estacíon, Puerto Peñasco, showed 

it to be one of the three dominant species of Sargassum, 

along with S. johnstonii and S. sinicola subsp. camouii

(see also McCourt, 1984a, 1984b). Some intertidal specimens can be difficult to separate from S. brandegeei, but 

S. herporhizum has smaller, narrower leaf-like blades with 

less distinct midribs and margins that are only slightly 

and irregularly dentate or sometimes uneven (see also 

“Remarks” for S. brandegeei).

Sargassum brandegeei 

Setchell et N. L. Gardn

FIGURE 109A–C

Sargassum brandegeei Setchell et N. L. Gardner, 

1924a:736, pl. 21: fig. 79; Setchell and Gardner, 

1925:718; Dawson, 1944:249, pl. 40: figs. 13–26; 

Taylor, 1945:120 [with a query]; Dawson, 1959a:19; 

Dawson, 1961b:399; Dawson, 1966a:12; Norris, 

1973:7; Huerta-Múzquiz, 1978:337, 338; Pedroche 

and González-González, 1981:67; Rocha-Ramírez 

and Siqueiros-Beltrones, 1990:18–19 [in part, only S. 

brandegeei]; Servière-Zaragoza et al., 1993:482; Phillips, 1995:118; González-González et al., 1996:159; 

Pacheco-Ruíz et al., 2008:204; Pedroche et al., 2008:93.

Algae up to 30 cm long, of 1 to several erect, terete, 

smooth axes, branching alternately to irregularly. Primary 

axis initially from a discoid base that develops numerous, 

FIGURE 108. Sargassum herporhizum: A. Young thallus arising from basal rhizomes (JN-5870b). B. Upper portion of thallus showing long, 

thin leaflike blades, with faint midrib and serrate margins, pedicellate vesicles, and receptacles (JN-3117).
2 1 8 • 

FIGURE 109. Sargassum brandegeei: A. Habit, showing developing basal rhizomes (JN-6034). B, C. Smooth axes, with long and comparatively 

wider leaflike blades, with midribs and moderate to deeply serrate margins (JN-4829).
n u m b e r 9 4 • 2 1 9

irregularly branched, narrow in diameter, decumbent rhizomes that become entangled and attach by haptera at 

various points. Leaf-like blades, longer than wide, mostly 

30–60(–110) mm long and 4–10 mm wide; with a midrib and without cryptostomata (note some observed with 

very few cryptostomata); margins regularly, moderately to 

deeply, serrate. Vesicles spherical to subspherical, (2–)3–5 

mm in diameter, smooth, and usually apiculate or sometimes with a foliaceous extension; borne on pedicels (more 

or less equal to or longer than their diameter) at base of 

leaflike blades or scattered among receptacles.

Receptacle branchlets relatively short, up to 20 mm 

long, branched mostly to 2 orders, smooth, and with 

blunt, rounded apices.

Habitat. On rocks and tidal platforms and

in tide pools; low intertidal to shallow subtidal to 8 m 

depths.

Distribution. Gulf of California: Puerto

Peñasco to Guaymas. Pacific coast: Jalisco; Galápagos 

Islands (?).

Type Locality. Cast ashore, probably Guaymas, Sonora, Gulf of California (Setchell and Gardner

1924a).

Remarks. Northern Gulf specimens generally 

agree with Sargassum brandegeei as described, including 

that most have leaf-like blades that lack cryptostomata 

(Setchell and Gardner, 1924a; Dawson, 1944), but a few 

specimens were observed to also have a small number 

of scattered cryptostomata on some of their leaf-like 

blades.

Dawson (1966a) observed that some of the intertidal 

S. brandegeei were not always clearly distinct from S. herporhizum. However, the size of the leaf-like blades and the 

nature of their margins distinguish these two in the field. 

Sargassum brandegeei has larger, longer, and comparatively wider blades, with a distinct midrib, and margins 

of regularly, evenly spaced, pronounced serration; in contrast, S. herporhizum has smaller, narrower blades, with 

less distinct midribs (sometimes very faint or not apparent 

on blades of the upper thallus and not always evident on 

the upper portion of some individual blades; see Figure 

108B), and margins that are uneven or slightly, irregularly 

dentate.

Uncertain Record:

Sargassum agardhianum Farlow ex J. Agar

Sargassum agardhianum Farlow ex J. Agardh, 1889:93; 

38, pl. 16:fig. 1; Mendoza-González and Mateo-Cid, 

1986:421; Martínez-Lozano et al., 1991:23; GonzálezGonzález et al., 1996:159; Pedroche et al., 2008(in part; 

Gulf of California records only):93.

Remarks. Described from San Diego, California, Sargassum agardhianum (J. Agardh, 1889) is known 

on the Pacific coast from southern California to central 

Baja California (Dawson et al., 1960a; Abbott and Hollenberg, 1976; Pacheco-Ruíz, 1982; Stewart and Stewart, 

1984; R. Aguilar-Rosas et al., 1990; Stewart, 1991; Phillips, 1995). The reported presence of this species in the 

northern Gulf of California needs to be verified by reexamining specimens identified as S. agardhianum (Mendoza-González and Mateo-Cid, 1986; Martínez-Lozano 

et al., 1991).

Excluded Records

Laminar

Laminariales Migula, 1908:173, 243.

Laminariales Kylin, 1917:308.

Remarks. In 25 years of collecting in the Gulf 

of California, Dawson himself did not collect any members of the Laminariales. Since then, neither I nor others 

have encountered or reported any Laminariales in field 

collecting and diving surveys in the northern Gulf.

Alar

Alariaceae Setchell et N. L. Gardner, 1925:591.

Egregia Arescho

Egregia Areschoug, 1876:66.

Egregia laevigata Setchell

Egregia laevigata Setchell, 1896:44; Dawson, 1946a:83; 

Dawson, 1946b:182; Dawson, 1947:43; Dawson, 

1949:241; González-González et al., 1996:149; Pedroche et al., 2008:79.

Remarks. Two beach drift specimens of Egregia laevigata reputed to be from Bahía de Los Ángeles 

were recorded in the northern Gulf (Dawson, 1946a). 

Later, Dawson (1949b) noted E. laevigata had been erroneously reported in the Gulf of California.
2 2 0 • 

Lessonia

Lessoniaceae Setchell et N. L. Gardner, 1925: 621.

Macrocystis C. Agar

Macrocystis C. Agardh, 1820:46.

Macrocystis pyrifera (Linnaeus) C. Aga

Fucus pyriferus Linnaeus, 1771:311.

Macrocystis pyrifera (Linnaeus) C. Agardh, 1820:47; 

Setchell and Gardner, 1924a:728; Dawson, 1944:237; 

González-González et al., 1996:154; Pedroche et al., 

2008 (in part; only Gulf of California record):85.

Remarks. A single specimen, about 3 ft. long, 

apparently adrift, was caught on [the] log-line of [the] vessel between Isla Espiritu Santo and La Paz and “brought 

aboard the vessel, but no specimens were found growing attached within the Gulf” (Setchell and Gardner, 

1924a:728; Dawson, 1944). Since then, there has never 

been any published reports of attached Macrocystis, 

nor any further reports of drift specimens in the Gulf of 

California.
I express my appreciation to my professors and mentors of the marine algae, 

Isabella Aiona Abbott (Stanford University), Michael Neushul, Jr. (University of California, Santa Barbara [UCSB]), Robert W. Hoshaw (University 

Arizona), and George T. Oberlander and Harry D. Thiers (San Francisco 

State University), and to Demorest Davenport and Alfred W. Ebeling (UCSB) 

for their continued encouragement, advice, and support throughout my Gulf of 

California research. Special thanks are owed to initiators of the Puerto Peñasco 

project, the marine scientists of the University of Arizona, Robert W. Hoshaw 

(McCourt, 1993), Donald A. Thomson, J. Laurens Barnard (Brusca, 1993), and 

John R. Hendrickson (Owens, 2003). I appreciated the use of Laboratorio de 

Biología Marina in Puerto Peñasco (shared facility of the Universidad de Sonora 

and University of Arizona) while I was serving as University Arizona Resident 

Marine Biologist and Station Director. I particularly thank Carlos Peña, Jose 

Ramon Infante-León, Ramon Durazo, Fernando Montanez, Argelia González, 

Manuel Esper-Huerta, Christine A. Flanagan, and Richard C. Brusca; and also 

those at the Peñasco Experimental Unit of the University of Arizona Environmental Research Laboratory (ERL), notably David Moore (Station Manager), 

Carl N. Hodges (Director, ERL), Carl Hodge, and Jim Ure. I thank Agustin 

Cortéz C., Eugenio Zamorano and his family, and the many people of Puerto 

Peñasco for making me welcome while I lived in Mexico.

During the course of this floristic study, my travel, fieldwork, and scuba 

diving was supported by grants from the National Science Foundation (NSFBMS-73-07000) and NOAA (04-5-158-20 8/75); later support was received from 

the American Philosophical Society, the National Museum of Natural History, 

Smithsonian Institution, and NSF-PEET grant DEB-0328491. Help in designing 

and equipping a four-wheel-drive truck for off-road use and land-based scuba 

operations came from Vic Hickey, Bill Issacson, and especially Gary Haden 

(Ventura, California), all of whom developed and allowed the rigorous testing 

of off-road prototype equipment. Advice and assistance on setting up the truck 

for land-based scuba and field work that could accomodate four divers (and 

dogs, Cyril and Beau) and all the collection, scuba, photographic, and camping 

gear along with food, water, and fuel came from Whitney Baines (UCSB), Craig 

Acknowledgments
2 2 2 • 

Askin (Tucson), and Norm Lammer (Marine Science Institute, UCSB), who also constructed much of the specialized 

field equipment.

Thanks are owed to William Fenical (UCSD-SIO, 

Chief Scientist) for the opportunity to participate on 

National Science Foundation-sponsored cruises of the 

Scripps Institution of Oceanography R/V Dolphin to 

Gulf of California and the OR/V Alpha Helix to islands 

of Pacific Mexico. He and Robert Jacobs (UCSB), Valerie Paul (Smithsonian Marine Station at Fort Pierce), 

William Gerwick (UCSD, SIO), and Oliver McConnell 

(the latter three were students of Bill Fenical at the time) 

provided many hours of collecting, lab work on natural 

product chemistry, and interesting discussions; I am grateful to have shared many adventures with them. Thanks 

are also owed to Dennis Divins (UCSB) for scuba training and support and continued friendship. The field work 

and collecting could not have been completed without 

the help of scuba diving partners Katina Bucher Norris, 

William Fenical, Howard Sleeper, William Gerwick, John 

Paul, George Boehlert, David Coon, David Moore, Jerry 

D. Kudenov, Mark Helvey, Whitney Baines, Valerie Paul, 

Bruce Howard, and Oliver McConnell, all of whom made 

one or more of the land-based and/or ship-based surveys 

and assisted with logistics and collecting. It is a pleasure 

to thank Richard S. Felger (University Arizona), Richard 

Evans Schultes (Harvard University), and Andrew T. Weil 

(University Arizona) for their gifts of algae collected in 

Sonora; David G. Lindquist (University North Carolina 

Wilmington; Howe, 2001) for algal specimens from El 

Tornillal, Rocas Consag, and Caleta Santa Maria; Nicolas 

P. Yensen for his collection from Isla San Pedro Nolasco; 

Mark and Diane Littler (NMNH), fellow marine botanists, 

divers, and “Baja-proven travelers” for their collections 

of algae and hours of discussion; Max H. Hommersand 

(University North Carolina) and Makoto Yoshizaki (Toho 

University, Japan) for sharing their collections and traveling with my group when we met in Bahía de Los Ángeles 

during a trip to Baja California; and Nonie McKibbin and 

Dale Evanson (Tucson, Arizona) for their collections of 

Sonoran algae. My appreciation goes to those who made 

Tucson a home during my trips to University of Arizona, 

particularly Craig Askin, Richard S. Felger, Ronald and 

Jan Pulliam, Robert and Ruth Hoshaw, and Alexander 

“Ike” and Jean Straub Russell. I thank Ike Russell, who 

generously flew my team in his small aircraft to locales 

in Sonora and to Isla Ángel de la Guarda, enabling us to 

collect and work with the Seri (including one trip with 

Clinton J. Dawes, University South Florida, and Donald P. 

Cheney, Northeastern University), and who also once flew 

in critical parts to get us on the road again when our vehicle was broken down in a remote location. I am indebted 

to Edward and Mary Beck Moser, who assisted in numerous ways, including sharing their home in El Desemboque 

(El Desemboque de San Ignacio; Haxöl Ihoom of the 

Seri) during our Sonora coast trips. I am also indebted to 

the Seri fishermen of El Desemboque and Punta Cheuca 

for guiding and providing boats for scuba diving. Their 

knowledge of locales, sea turtle foraging sites, and offshore reefs of Isla Tiburón and coastal Sonora yielded 

numerous collections, and their understanding of the sea 

conditions in this region kept us safe more than once. The 

assistance of Glenn and Martha Vargas (Thermal, California) enabled my team to scuba dive on many of the islands 

in the vicinity of Bahía de Los Ángeles, and I thank them 

for their generous hospitality and numerous boat trips. I 

again extend my thanks to Donald and Isabella Abbott 

for sharing their Pacific Grove home and providing access 

to the Hopkins Marine Station laboratory to study Gulf 

seaweeds; to William and Fran Fenical and Joan G. and 

James Stewart for letting us stay with them while studying at Scripps Institution of Oceanography, La Jolla; and 

to Gary and Mary Haden for providing accommodations 

during trips to Ventura and UCSB.

I am grateful for the taxonomic expertise, discussions, 

and comments of Isabella A. Abbott, Michael Neushul, 

Richard E. Norris, and Paul C. Silva, and of Michael J. 

Wynne on Colpomenia, he and William Randolph Taylor (University Michigan) on Sporochnus, Daniel LeónÁlvarez (UNAM) on the Ralfsiales, Luis Aguilar-Rosas 

and Raúl Aguilar-Rosas (CMMEX) on Gulf Leathesia 

nana, Heroen Verbruggen (University Ghent) for reviewing and discussion on the species of Codium. I am also 

grateful to Frederik Leliaert (University Ghent) for reviewing “Willeella-Phyllodictyon,” Naomi Phillips (Arcadia University) for comments on the classification of the 

brown algal orders and their evolutionary relationships, 

and Nicolas P. Yensen (University Arizona) for discussions on Gulf Sargassum species. My appreciation goes to 

Dan H. Nicolson (NMNH) and Larry J. Dorr (NMNH) 

for advice on solving numerous nomenclatural problems, 

to Alain Touwaide (NMNH) for the Latin descriptions of 

new taxa, to Francisco F. Pedroche, Luis Aguilar-Rosas, 

Raúl Aguilar-Rosas, and Rafael Riosmena-Rodríguez for 

help on obtaining important literature and the loan of 

specimens, and finally, to Paul M. Peterson (Botany Editor; NMNH) for his encouragement throughout the submission and review process.

I thank those who have prepared or helped with 

illustrations: Susan Manchester (Monterey, California) 
n u m b e r 9 4 • 2 2 3

for drawing many of the algae, Ilka C. Feller (NMNH; 

SERC) for line drawings of Sargassum, Adrián Ramírez 

(UNAM) for the species of Ralfsia, and Alice R. Tangerini 

(Botanical Illustrator, NMNH) for the northern Gulf of 

California map and the comparative diagrammatic illustrations of Ulva and the Gulf genera of the Dictyotaceae. 

Most of the algal and habitat photographs were taken by 

me. Additionally, I am grateful not only for the help of 

many others but for the photographs and digital images 

of selected herbarium specimens they skillfully provided, 

including Alison Habel (Photographic Illustrator, University of Arizona), Chip Clark (NMNH Scientific Photographer), Harvey Kupferberg (NMNH volunteer), and Ingrid 

Lin (NMNH Botany); I am grateful to Scott Whittaker 

(NMNH) for assistance on digital photomicrographs and 

Kathy Ann Miller (Herbarium, UC Berkeley) for kindly 

(and rapidly) providing the isotype image of Sargassum 

acinacifolium. I thank Roger H. Hill (USAF) for obtaining 

the NASA Gemini V photo of the northern Gulf of California and NASA for permission for its use herein. Thanks 

go to Chip Clark (NMNH) for consultation and advice on 

digital photography, 35mm photographic slide digitizing, 

and his appreciation of margaritas. The photos, digital 

images, and line drawings were competently prepared for 

this study by Robert H. Sims (NMNH), whose research 

assistance is much appreciated.

To my wife, Katina “Katy” Bucher Norris (NMNH 

Botany), it is impossible to express all my thanks for her 

companionship and friendship. She has participated in 

every phase of this research: planning and carrying out 

the field work, scuba diving, collecting and studying Gulf 

of California marine algae, research collaboration, curatorial assistance, taking the digital photomicrographs of 

algal images used, and reviewing the various stages of the 

manuscript throughout this project. It is with Katy that I 

have shared many great adventures of diving and traveling 

through the pristine desert, beaches, reefs, and islands of 

the Gulf of California. To our sons, James “Jesse” Alexander and Shaun William, I give thanks and appreciation for 

their interest in Baja California, scuba diving, and listening to many tales of the Gulf of California forays.

Many people were kind enough to review and discuss 

various parts of the manuscript as it developed, and I particularly thank Robert W. Hoshaw and Jeanine L. Olsen 

(University of Groningen), for reviews and comments on 

earlier drafts of the greens, and Isabella A. Abbott for 

comments on the browns. I also extend sincere thanks to 

the reviewers of the final draft of the manuscript—Luis 

Ernesto Aguilar-Rosas and Raúl Aguilar-Rosas (Universidad Autónoma de Baja California, Ensenada), David 

L. Ballantine (University of Puerto Rico, Mayagüez), 

Suzanne L. Fredericq (University of Louisiana Lafayette), 

Rafael Riosmena-Rodríguez (Universidad Autónoma de 

Baja California Sur, La Paz), and Richard M. McCourt 

(National Science Foundation)—for their thoughtful and 

constructive suggestions and comments that led to its 

improvement. Despite all this input and help, I alone am 

responsible for any errors.

Appendix 1: 

List of New Taxa and Combinations

I. CHLOR

Ctenocladales: Ulvellaceae

Epicladia condensata (Setchell et N. L. Gardner) J. N. Norris, comb. nov.

Epicladia mexicana (Setchell et N. L. Gardner) J. N. Norris, comb. nov.

II. P

Desmarestiales: Desmarestiaceae

Desmarestia munda Setchell et N. L. Gardner subsp. mexicana (E. Y. Dawson) 

J. N. Norris, comb. nov.

Sporochnales: Sporochnace

Sporochnus neushulii J. N. Norris, sp. nov.

Ectocarpales: Acinetosporacea

Hincksia bryantii (Setchell et N. L. Gardner) J. N. Norris, comb. nov.

Fucales: Sargassacea

Sargassum sect. johnstonii E. Y. Dawson ex J. N. Norris, sect. nov.

Sargassum sect. lapazeanum E. Y. Dawson ex J. N. Norris, sect. nov.

Sargassum sect. sinicola E. Y. Dawson ex J. N. Norris, sect. nov.

Sargassum sect. herporhizum E. Y. Dawson ex J. N. Norris, sect. nov.

Sargassum sinicola Setchell et N. L. Gardner subsp. camouii (E.Y. Dawson) 

J. N. Norris et Yensen, comb. nov.

I. CHLOR

Ctenocladales: Ulvellaceae

Entocladia marchantiae

Epicladia condensata

Epicladia mexicana

Cladophorales: Cladophoraceae

Cladophora macdougalii

Cladophora tiburonensis

Siphonocladales: Valoniace

Valoniopsis hancockii

Bryopsidales: Codiaceae

Codium brandegeei

II. P

Ishigeales: Ishigeaceae

Ishige sinicola

Dictyotales: Dictyotaceae

Spatoglossum lanceolatum

Spatoglossum subflabellatum

Desmarestiales: Desmarestiaceae

Desmarestia filamentosa

Desmarestia munda subsp. mexicana

Appendix 2: 

Endemic Taxa of the Gulf of California 

that Occur in the Northern Gulf
2 2 8 • 

Sporochnales: Sporochnace

Sporochnus neushulii

Ralfsiales: Ralfsiaceae

Hapterophycus anastomosans

Cutleriales: Cutleriaceae

Cutleria hancockii

Ectocarpales: Acinetosporacea

Hincksia bryantii

Ectocarpales: Chordariaceae

Nemacystus brandegeei

Ectocarpales: Ectocarpaceae

Ectocarpus hancockii

Ectocarpus sonorensis

Ectocarpales: Scytosiphonacea

Compsonema immixtum

Fucales: Sargassacea

Sargassum johnstonii f. gracile

Sargassum lapazeanum

Sargassum acinacifolium

Sargassum macdougalii

Sargassum sonorense

Sargassum brandegeei
I. CHLOR

Entocladia marchantiae (Figure 9A): Epiphytic on Gracilaria textorii, north

end of Isla Coronado, Bahía de los Ángeles, Baja California, Coll. H. Sleeper 

and M. Helvey (JN-4411b; US Alg. Coll. slide 4257). 

Entocladia polysiphoniae (Figure 9B): Endophytic on Polysiphonia, El Desemboque, Sonora, Coll. R. E. Schultes, R. S. Felger, and A. T. Weil (JN-5190; US 

Alg. Coll.-157757 and slide 5344). 

Entocladia sp. (Figure 9C): Epiphytic on Cladophora, Islas de los Gemelos, 

Bahía de los Ángeles, Baja California, Coll. H. Sleeper and M. Helvey (JN4492b; US Alg. Coll. slide 4154).

Entocladia viridis (Figure 9D,E): Epiphytic on Chaetomorpha aerea, El Desemboque, Sonora, Coll. R. E. Schultes, R. S. Felger, and A. T. Weil (JN-5180b; 

US Alg. Coll. slide 4144).

Stromatella monostromatica (Figure 11A,B): Epiphytic on Polysiphonia, Isla la

Ventana, Bahía de los Ángeles, Baja California, Coll. J. N. Norris, M. Helvey, 

and H. Sleeper (JN-4442b; US Alg. Coll. slide 4160).

Ulvella lens (Figure 12A): Epiphytic on Cladophora graminea, channel between

Isla Mejia and Isla Ángel de la Guarda, Puerto Refugio, Isla Ángel de la 

Guarda, Coll. E. Y. Dawson (EYD[1940]-271; US Alg. Coll. slide 4165). 

Ulvella setchellii (Figure 12B): Epiphytic on Cladophora cf. microcladioides, 

Bahía Aqua Dulce, Isla Tiburón, Coll. E. Y. Dawson (EYD-46-980; US Alg. 

Coll. slide 892).

Ulva californica (Figure 14A): Vicinity of Punta Peñasco, Sonora, Coll. E. Y.

Dawson (EYD-27407; US Alg. Coll.-55027). 

Ulva californica (Figure 14B): Isla Willard, N end of island, Bahía de San Luis 

Gonzaga, Baja California, Coll. J. N. Norris and K. E. Bucher (JN-5425; US 

Alg. Coll.-157778).

Appendix 3: 

Algal Specimens Illustrated: 

Species Locality Information and Herbarium
2 3 0 • 

Ulva lactuca (Figure 15): La Paz, Baja California Sur, 

Coll. O. Holguín Q. (ChP-1606; US Alg. Coll.-36624).

Ulva lobata (Figure 16): Punta la Gringa, Bahía de los 

Ángeles, Baja California, Coll. J. N. Norris (JN-6103a; 

US Alg. Coll.-157781).

Ulva nematoidea (Figure 17A): Ensenada, Bahía de Todos Santos, Baja California (Pacific Mexico), Coll. E. Y.

Dawson (EYD-10421; US Alg. Coll.-41727). 

Ulva nematoidea (Figure 17B): San Isidro, Baja California (Pacific México), Coll. Clair Etienne (US Alg. 

Coll.-8799).

Ulva rigida (Figure 18): Puerto Refugio, Isla Ángel de la 

Guarda, Coll. J. N. Norris and K. E. Bucher (JN-5742; 

US Alg. Coll.-157779).

Ulva acanthophora (Figure 19A): Punta la Gringa, Bahía

de los Ángeles, Baja California, Coll. J. N. Norris and 

K. E. Bucher (JN-5455; US Alg. Coll.-157716). 

Ulva acanthophora (Figure 19B): Estuary, Bahía de los 

Animas, Baja California, Coll. G. Bockus (JN-383; US 

Alg. Coll.-89466).

Ulva clathrata (Figure 20): Bahía Cholla, vicinity of 

Puerto Peñasco, Sonora, Coll. J. N. Norris and K. E. 

Bucher (JN-4994; US Alg. Coll.-157727).

Ulva flexuosa (Figure 21A,B): Epizoic on carapace of female black sea turtle (Chelonia mydas agassizii), vicinity of Campo Ona, Canal de Infiernillo, Sonora, Coll. R.

S. Felger, E. Moser, and G. Mendez (JN-4790; US Alg. 

Coll.-157732 and slide 4172).

Ulva intestinalis (Figure 22): El Tornillal, Sonora, Coll. D. 

G. Lindquist (JN-3327; US Alg. Coll.-157741).

Ulva linza (Figure 23): Playa Hermosa, Puerto Peñasco, 

Sonora, Coll. J. N. Norris (JN-3694; US Alg. Coll.-

157743).

Chaetomorpha antennina (Figure 24A): Near Bahía Sa

Carlos, Ensenada de San Francisco, Sonora, Coll. E. Y. 

Dawson (EYD-11023; US Alg. Coll.-39950). 

Chaetomorpha aerea (Figure 24B): El Desemboque, Sonora, Coll. R. E. Schultes, R. S. Felger, and A. T. Weil

(JN-5180a; US Alg. Coll. slide 4144).

Cladophora graminea (Figure 28A): West side of larg

rock, Rocas Consag, Coll. D. G. Lindquist (JN-3118; 

US Alg. Coll.-157544).

Cladophora hesperia (Figure 29A): Puerto Lobos o

Bahía Tepoca, Sonora, Coll. E. Y. Dawson (EYD-832; 

US Alg. Coll.-30072 and slide 4170).

Cladophora microcladioides (Figure 30B): Bahía Kino,

Sonora, Coll. E. Y. Dawson (EYD-2928; US Alg. Coll.-

30073 and slide 4166).

Phyllodictyon pulcherrimum (Figure 33): East side of

Punta Concepción, Baja California Sur, Coll. J. Bleck 

(EYD-25872; US Alg. Coll.-36522).

Phyllodictyon robustum (Figure 34A): Isla San Pedr

Nolasco, Coll. E. Y. Dawson (EYD-18555; US Alg. 

Coll.-5603).

Valoniopsis pachynema (Figure 36C): Photo of living

clump from Puerto Peñasco, Sonora.

Bryopsis pennata (Figure 37): South of rocky shore fronting Laboratorio de Biología Marina, vicinity of Punta 

Peñasco, Sonora, Coll. A. E. Dennis (EYD-27451; US 

Alg. Coll.-40783).

Halicystis-phase (Figure 39A,B): Isla Willard, Bahía d

San Luis Gonzaga, Baja California, Coll. J. N. Norris 

and K. E. Bucher (JN-5732; US Alg. Coll.-157764).

Codium amplivesiculatum (Figure 40): Isla San Esteban,

Coll. J. N. Norris, with J. Paul and K. Robertson (JN5700; US Alg. Coll.-157600).

Codium brandegeei (Figure 42): “Codium macdougaliiform” of C. brandegeei, Playa Estación, Puerto Peñasco,

Sonora, Coll. J. N. Norris and R. W. Hoshaw (JN-5020; 

US Alg. Coll.-157629).

Codium simulans (Figure 43): SE side of Isla San Esteban,

Coll. J. N. Norris, with J. Paul and K. Robertson (JN5713; US Alg. Coll.-157676).

Codium sp.? (Figure 44): cf. Codium cuneatum-form

of C. simulans, off west side of Roca Blanca, Puerto 

Refugio, Isla Ángel de la Guarda, Coll. J. N. Norris, 

with J. Paul and K. Robertson (JN-5286; US Alg. Coll.-

157616).

Caulerpa mexicana f. pectinata (Figure 45): Playa Estación, Puerto Peñasco, Sonora, Coll. N.J. Lang (JN5861; US Alg. Coll.-157510)

Caulerpa sertularioides (Figure 46A): N side of bay, Bahí

Bocochibampo, Guaymas, Sonora, Coll. M. Helvey 

(JN-4471; US Alg. Coll.-157509). 

Caulerpa sertularioides (Figure 46B): Bahía de San Carlos, Nueva Guaymas, Sonora, colI. D. Evanson (JN4277; US Alg. Coll.-157507)

Caulerpa vanbosseae (Figure 47A): El Desemboque d

San Ignacio, Sonora, Coll. J. N. Norris, K. E. Bucher 

and D. Coon (JN-6562; US Alg. Coll.-157529). 

Caulerpa vanbosseae (Figure 47B): Punta Pelícano, vicinity of Puerto Peñasco, Sonora, Coll. J. N. Norris and K.

E. Bucher (JN-4981; US Alg. Coll.-157521).
n u m b e r 9 4 • 2 3 1

Halimeda discoidea (Figure 48B,C): 3 miles [4.8 km]

south of Punta Mangles, Baja California Sur, Coll. J. 

Bleck (EYD-25921; US Alg. Coll.-36518 and slide 8477 

and slide 8478).

Acetabularia caliculus (Figure 49A): Bahía Cholla, vicinity of Punta Peñasco, Sonora, Coll. A. E. Dennis (EYD27569; US Alg. Coll.-40937 and vial 3442, jar 406)

Acetabularia caliculus (Figure 49B): Bahía Cholla, vicinity of Puerto Peñasco, Sonora, Coll. R. W. Hoshaw (US

Alg. Coll.-95320 and slide 5533).

II. P

Ishige sinicola (Figure 50A): Specimens, from left to right

La Mona, Bahía de los Ángeles, Baja California, Coll. 

J. N. Norris (JN-2980; US Alg. Coll.-158148); Puerto 

Calamajué, Baja California, Coll. J. N. Norris and K. E. 

Bucher (JN-4621; US Alg. Coll.-158165); and Playa Estación, Puerto Peñasco, Sonora, Coll. J. N. Norris and 

K. E. Bucher (JN-4166; US Alg. Coll.-158160). 

Ishige sinicola (Figure 50B): Northwest of rock window

on shore, Puerto Refugio, Isla Ángel de la Guarda, Coll. 

J. N. Norris and K. E. Bucher (JN-5824; US Alg. Coll.-

158176 and slide 8481).

Sphacelaria rigidula (Figure 51A): Playa Hermosa, Puerto

Peñasco, Sonora, Coll. J. N. Norris and K. E. Bucher 

(JN-4041; US Alg. ColI.-158531 and slide 4201). 

Sphacelaria rigidula (Figure 51B): Bahía San Francisquito, Baja California, Coll. J. N. Norris (JN-3271; US 

Alg. Coll.-158528 and slide 4194).

Sphacelaria rigidula (Figure 51C): Playa Hermosa, Puerto 

Peñasco, Sonora, Coll. J. N. Norris (JN-3715; US Alg. 

Coll.-158514 and slide 4196).

Sphacelaria tribuloides (Figure 53A–C): Playa Hermosa,

Puerto Peñasco, Sonora, Coll. J. N. Norris and K. E. 

Bucher (JN-4038; US Alg. Coll.-158525 and slide 4198 

and slide 4199).

Dictyopteris undulata (Figure 55): Off west side of Roca

Blanca, Puerto Refugio, Isla Ángel de la Guarda. Coll. J. 

N. Norris (JN-5801; US Alg. Coll.-157994).

Dictyota concrescens (Figure 56A): Punta Hughes, vicinity of Bahía Magdalena, about 31⁄2 miles [5.6 km

southeast of Cabo San Lazaro, Baja California Sur (Pacific Mexico), Isotype (W. R. Taylor 34-598; US Alg. 

Coll.-56360). 

Dictyota concrescens (Figure 56B): North side of Puerto

Calamajué, Baja California, Coll. J. N. Norris and K. E. 

Bucher (JN-4695; US Alg. Coll.-158074).

Dictyota coriacea (Figure 57): Off southeast end of Isla

San Esteban, Coll. K. E. Bucher (JN-5522; US Alg. 

Coll.-158202).

Dictyota crenulata (Figure 58A): Playa between San Jos

del Cabo and Cabo San Lucas, Baja California Sur, 

Coll. J. N. Norris and H. Sleeper (JN-4091a; US Alg. 

Coll.-158007). 

Dictyota crenulata (Figure 58B): Playa between San Jose

del Cabo and Cabo San Lucas, Baja California Sur, 

Coll. J. N. Norris and K. E. Bucher, and H. Sleeper (JN4114; US Alg. Coll.-158009).

Dictyota dichotoma var. intricata (Figure 59A): Cabez

Ballena, Baja California Sur, Coll. J. N. Norris and H. 

Sleeper (JN-4087; US Alg. Coll.-158034). 

Dictyota dichotoma var. intricata (Figure 59B): In front

of Laboratorio de Biología Marina, Playa Estación, 

Puerto Peñasco, Sonora, Coll. J. N. Norris (JN-5630; 

US Alg. Coll.-158042). 

Dictyota dichotoma var. intricata (Figure 59C): Playa

between San Jose del Cabo and Cabo San Lucas, Baja 

California Sur, Coll. J. N. Norris and H. Sleeper (JN4094; US Alg. Coll.-158016).

Dictyota flabellata (Figure 60): Playa Hermosa, Puerto

Peñasco, Sonora, Coll. J. N. Norris and K. E. Bucher 

(JN-3474; US Alg. Coll.-158049).

Dictyota stolonifera (Figure 61A): Playa Estación, Puert

Peñasco, Sonora, Coll. J. N. Norris and K. E. Bucher 

(JN-3865a; US Alg. Coll.-158031). 

Dictyota stolonifera (Figure 61B): Playa between San Jose

del Cabo and Cabo San Lucas, Baja California Sur, 

Coll. J. N. Norris, K. E. Bucher and H. Sleeper (JN4113a; US Alg. Coll.-158017). 

Dictyota stolonifera (Figure 61C): C. In front of marine

laboratory, vicinity of Puerto Peñasco, Sonora, Coll. E. 

Y. Dawson (EYD-27409; US Alg. Coll.-40641).

Dictyota vivesii (Figure 62): Punta Prieta, outer Bahía Topolobampo, Sinaloa, Coll. E. Y. Dawson (EYD-10938;

US Alg. Coll.-39951).

Padina caulescens (Figure 63A,B): Islas de los Gemelos

Bahía de los Ángeles, Baja California, Coll. J. N. Norris, H. Sleeper and W. Baines (JN-4748; US Alg. Coll.-

158312).
2 3 2 • 

Padina crispata (Figure 64A–C): Isla Grande, Guerrer

(Pacific Mexico), Coll. E. Y. Dawson (EYD-20957; US 

Alg. Coll.-6160).

Padina durvillei (Figure 65): Punta Pelícano, Puerto Peñasco, Sonora, Coll. M. and D. Littler, s.n. (US Alg.

Coll.-163415).

Padina gymnospora (Figure 66): Playa Arenosa, east of

Punta Pelícano, Puerto Peñasco, Sonora, Coll. J. N. Norris and K. E. Bucher (JN-3425b; US Alg. Coll.-158298).

Padina mexicana var. mexicana (Figure 67): Photo of live

specimen from Playa Estación, Puerto Peñasco, Sonora.

Rugulopteryx okamurae (Figure 68A): Cabeza Ballena,

Baja California Sur, Coll. J. N. Norris and H. Sleeper 

(JN-4086; US Alg. Coll.-158127). 

Rugulopteryx okamurae (Figure 68B): Playa Estación, 

Puerto Peñasco, Sonora, Coll. J. N. Norris (JN-3724; 

US Alg. Coll.-158125 and slide 4204).

Spatoglossum subflabellatum (Figure 70): Punta Lobos, 

north side of point, vicinity of Puerto Lobos, Sonora, 

Coll. J. N. Norris (JN-5902; US Alg. Coll.-158512).

Desmarestia filamentosa (Figure 71): Off west side of

Roca Blanca, Puerto Refugio, Isla Ángel de la Guarda, 

Coll. J. N. Norris (JN-5271; US Alg. Coll.-157950). 

Desmarestia filamentosa (Figure 72A,B): Off west side o

Roca Blanca, Puerto Refugio, Isla Ángel de la Guarda, 

Coll. J. N. Norris, with J. Paul and K. Robertson (JN5271; US Alg. Coll.-157950 and slide 4183).

Desmarestia munda subsp. mexicana (Figure 73): Off

west side of Roca Blanca, Puerto Refugio, Isla Ángel de 

la Guarda, Coll. J. N. Norris (JN-5283; US Alg. Coll.-

157944).

Sporochnus anomalus (Figure 74): Off west side of Roca 

Blanca, Puerto Refugio, Isla Ángel de la Guarda, Coll. J. 

N. Norris (JN-5264; US Alg. Coll.-158494).

Sporochnus anomalus (Figure 75A,B): Off west side of

Roca Blanca, Puerto Refugio, Isla Ángel de la Guarda, 

Coll. J. N. Norris (JN-5264; US Alg. Coll.-158494; US 

Alg. Coll. vial 5824, jar 568, and slide 4210).

Sporochnus neushulii J. N. Norris sp. nov. (Figure 7

E): Off west side of Roca Blanca, Puerto Refugio, Isla 

Ángel de la Guarda, Coll. J. N. Norris (A–E. Holotype: 

JN-5262, US Alg. Coll.158492; B. US Alg. Coll. vial 

5823, jar 568; C. US Alg. Coll. slide 4208; D. US Alg. 

Coll. slide 4209; and E. US Alg. Coll. slide 8474).

Hapterophycus anastomosans (Figure 77A,B): Isla Raza

Gulf of California (A,B. Holotype: EYD-26110; US Alg. 

Coll.-37228 and vial 2479, jar 296).

Cutleria hancockii (Figure 79A): Gametophyte, Playa

Estación, Puerto Peñasco, Sonora, Coll. J. N. Norris 

and K. E. Bucher (JN-4953; US Alg. Coll.-209237). 

Cutleria hancockii (Figure 79B): Gametophyte, epizoic 

on carapace of female black sea turtle (Chelonia mydas 

agassizii), vicinity of Campo Ona, Canal de lnfiernillo, 

Sonora, Coll. R. S. Felger and E. Moser (JN-4786; US 

Alg. Coll.-157912 and slide 4182). 

Cutleria hancockii (Figure 79C): Gametophyte, Puerto 

Lobos, Sonora, Coll. J. N. Norris and M. Helvey (JN6040; US Alg. Coll.-157937 and slide 8471). 

Cutleria hancockii (Figure 79D): Gametophyte, Camp 

Dolar, Cabo Tepopa, Sonora, Coll. J. N. Norris (JN7065; US Alg. Coll.-157942 and slide 8473). 

Cutleria hancockii (Figure 79E): Sporophyte (“Aglaozonia-phase”), Punta Pelícano–Playa Arenosa area, vicinity of Puerto Peñasco, Sonora, Coll. E. Y. Dawson 

(EYD-27365; US Alg. Coll.-40713 and slide 8469). 

Cutleria hancockii (Figure 79F): Sporophyte (“Aglaozonia-phase”), epiphytic on Sargassum, Playa Hermosa, 

Puerto Peñasco, Sonora, Coll. J. N. Norris (JN-3691b; 

US Alg. Coll. slide 4158).

Hincksia mitchelliae (Figure 82): Epizoic on black sea

turtle (Chelonia mydas agassizii) carapace, vicinity of 

Campo Ona, Canal de Infiernillo, Sonora, Coll. R. S. 

Felger and E. Moser (JN-4774; US Alg. Coll.-158145 

and slide 4223).

Chnoospora pacifica (Figure 83A–C): Binners Cove, Isl

Socorro, Revillagigedo Arch. (Pacific Mexico), Coll. E. 

Y. Dawson (EYD-12121; US Alg. Coll.-5794 and vial 

784, jar B88).

Hecatonema streblonematoides (Figure 84A,B): Play

Hermosa, Puerto Peñasco, Sonora, Coll. K. Bucher and 

J. N. Norris (JN-4033; US Alg. Coll. slide 5350).

Nemacystus brandegeei (Figure 85A): Epiphytic on Sargassum, Playa Hermosa, Puerto Peñasco, Sonora

Coll. J. N. Norris and K. E. Bucher (JN-3985; US Alg. 

Coll.-158192). 
n u m b e r 9 4 • 2 3 3

Nemacystus brandegeei (Figure 85B,C): Isla San Jose,

Baja California Sur (EYD-25847; US Alg. Coll.-36453; 

B. US Alg. Coll. slide 8479 and C. US Alg. Coll. jar 269, 

vial 2271).

Streblonema transfixum (Figure 86): Epiphytic on 

Gracilaria textorii, north end of Isla Coronado, Bahía 

de los Ángeles, Baja California, Coll. H. Sleeper and M. 

Helvey (JN-4411c; US Alg. Coll. slide 4257).

Leathesia nana (Figure 90A–C): Epiphytic on leaflik

blade of Sargassum sinicola, El Coloradito, Baja California, Coll. R. Aguilar-Rosas, L. Aguilar-Rosas, L. E. 

Mateo-Cid, C. Mendoza-Gonzalez and C. E. GutierrezVillaseñor (BC95-S/N3; CMMEX-6447).

Colpomenia phaeodactyla (Figure 91A): Playa Estación

Puerto Peñasco, Sonora, Coll. J. N. Norris and K. E. 

Bucher (drawing of JN-4759; US Alg. Coll.-157841). 

Colpomenia ramosa (Figure 91B): North side of Puerto

Calamajué, Baja California, Coll. J. N. Norris and K. E. 

Bucher (drawing of JN-4697; US Alg. Coll.-157854).

Colpomenia sinuosa (Figure 92A): Playa Estación, Puert

Peñasco, Sonora, Coll. J. N. Norris and K. E. Bucher 

(drawing of JN-4580; US Alg. Coll.-60062). 

Colpomenia tuberculata (Figure 92B): Photo of living

specimen from Playa Estación, Puerto Peñasco, Sonora.

Hydroclathrus clathratus (Figure 94): Isla Pichilingue,

La Paz, Baja California Sur, Coll. O. Holguin Q. (PhP1536; US Alg. Coll.-36619).

Rosenvingea floridana (Figure 95A,B): Punta la Gringa,

Bahía de los Ángeles, Baja California, Coll. J. N. Norris 

and K. E. Bucher (JN-5471; US Alg. Coll.-158327).

Rosenvingea intricata (Figure 96): Punta Bufeo, Baja California, Coll. M. and D. Littler, s.n. (US Alg. Coll.-163441).

Sargassum johnstonii (Figure 97A): Bahía Cholla, vicinity of Punta Peñasco, Sonora, Coll. E. Y. Dawson

(EYD-27204). 

Sargassum johnstonii (Figure 97B): Playa Estación, Puerto 

Peñasco, Sonora, Coll. J. N. Norris and K. E. Bucher 

(JN-4559; US Alg. Coll.-158429). 

Sargassum johnstonii f. gracile (Figure 98): Guaymas, Sonora, Coll. Daisy R. Marchant (Isotype, D. R. Marchant-28a; US Alg. Coll.-56958).

Sargassum lapazeanum (Figure 99A,B): La Paz, Baja

California Sur, Coll. Daisy R. Marchant (Isotype, D. R. 

Marchant-21; US Alg. Coll.-56961).

Sargassum acinacifolium (Figure 100B,C): Cast ashore, 

Guaymas (?), Sonora (Isotype, T. S. Brandegee-2; 

UC-221158).

Sargassum macdougalii (Figure 101A): Cabo Tepoca,

Puerto Lobos, Sonora, Coll. J. N. Norris (JN-5975; US 

Alg. Coll.-158398). 

Sargassum macdougalii (Figure 101B): Punta Lobos, 

Cabo Tepoca, SE side of Puerto Lobos, Sonora, Coll. J. 

N. Norris (drawing of JN-5944; US Alg. Coll.-158397).

Sargassum sonorense (Figure 102): Northeast side of cove, 

Isla San Pedro Nolasco, Coll. N. P. Yensen (JN-5205; 

US Alg. Coll.-158399).

Sargassum sinicola subsp. sinicola (Figure 103): Isla San 

Esteban, Coll. E. Y. Dawson (EYD-21568; US Alg. 

Coll.-6284).

Sargassum sinicola subsp. sinicola (Figure 104): West 

side of large rock, Rocas Consag, colI. D. G. Lindquist 

(JN-3119; US Alg. Coll.-158447).

Sargassum sinicola subsp. sinicola (Figure 105A): West

side of large rock, Rocas Consag, colI. D. G. Lindquist 

(drawing of JN-3119; US Alg. Coll.-158447). 

Sargassum sinicola subsp. camouii (Figure 105B): Playa 

Hermosa, Puerto Peñasco, Sonora, Coll. J. N. Norris and K. E. Bucher (drawing of JN-3250; US Alg. 

Coll.-158450).

Sargassum sinicola subsp. camouii (Figure 106): Playa 

Estación, Puerto Peñasco, Sonora, Coll. J. N. Norris 

and K. E. Bucher (JN-6064; US Alg. Coll.-158477).

Sargassum herporhizum (Figure 107): Playa Estación, 

Puerto Peñasco, Sonora, Coll. J. N. Norris and K. E. 

Bucher (JN-4963; US Alg. Coll.-158382).

Sargassum herporhizum (Figure 108A): Playa Estación,

Puerto Peñasco, Sonora, Coll. J. N. Norris and K. E. 

Bucher (JN-5870b; US Alg. Coll.-158346). 

Sargassum herporhizum (Figure 108B): West side of large 

rock, Rocas Consag, Coll. D. G. Lindquist, drawing of 

(JN-3117; US Alg. Coll.-158356).

Sargassum brandegeei (Figure 109A): North side of point

Punta Lobos, Puerto de Lobos, Sonora, coIl. M. Helvey 

(JN-6034; US Alg. Coll.-158392).

Sargassum brandegeei (Figure 109B,C): Punta Robinson, 

west side about 2.4 km east of Puerto Libertad, Sonora, 

Coll. J. N. Norris and K. B. Bucher (JN-4829; US Alg. 

Coll.-158377).

Abbott, I. A. 1966. Elmer Yale Dawson (1918–1966). Journal of Phycology 2(4): 

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Benthic Chlorophyta, Including Pseudochlorodesmis abbreviata (Gilbert) n. comb. 

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Index of Scientific Names

Names of all taxa recognized in the northern Gulf of California are shown in bold 

type. Higher taxa (e.g., orders and families) that are synonyms or have a distribution 

outside of the northern Gulf are in non-bold, small capital letters. Names of genera 

and species that are synonyms or have a distribution outside of the northern Gulf are in 

non-bold italics.

Page numbers in bold type refer to northern Gulf taxa descriptions. Page numbers 

in non-bold type refer to basionyms or synonyms or to taxa mentioned in the text or in 

discussion, e.g., Remarks. Page numbers in non-bold italics to pages with the figure(s) of 

the genus or species.

Acanthocodium fragile, 92

Acetabularia, 102

	 Acetabularia caliculus, 103, 103

Acetabularia crenulata, 102

Acetabularia farlowii, 102

Acetabularia parvula, 102

Acetabularia pusilla, 102

Acetabularia schenckii, 102

Acetabulariaceae, 102

Acinetospora, 166, 167

Acinetospora crinita, 166

Acinetospora pusilla, 166

Acinetosporaceae, 166, 186

Acrochaete, 30

Acrochaete codicola, 26

Acrochaete marchantiae, 22

Acrochaete viridis, 25

Aglaozonia, 162

Aglaozonia-phase of Cutleria, 163, 164

Alariaceae, 219

Anadyomena, 67, 68

Anadyomenaceae, 67

Analipus. 155

Annulina albida, 55

Archaeplastida, 20

Asperococcus, 175

Asperococcus intricatus, 198

Asperococcus β. Colpomenia, 189
2 7 2 • 

Asteronema, 166

Asteronema breviarticulatum, 166

Batophora, 101

Batophora oerstedii, 101

Blidingia, 32

Blidingia marginata, 32

Blidingia minima, 32

Boodlea, 68, 70

Boodlea composita, 68

Boodleaceae, 67, 68

Botrytella reinboldii, 187

Bryopsidaceae, 78

Bryopsidales, 77, 78

Bryopsidella, 79

Bryopsidineae, 78

Bryopsidophyceae, 20, 77, 78

Bryopsis, 79, 83

	 Bryopsis corticulans, 79, 80

	 Bryopsis hypnoides, 79, 80

Bryopsis muscosa, 79

Bryopsis pachynema, 76

	 Bryopsis pennata var. minor, 79, 81, 82

	 Bryopsis pennata var. pennata, 79, 80, 

81, 82, 85

Bryopsis pennatula, 81, 82

Bryopsis plumosa, 80

Bryopsis plumosa var. pennata, 81

	 Bryopsis salvadoreana, 79, 82

Canistrocarpus, 118

Canistrocarpus crispatus, 122

Carpopeltis sinicola, 109

Caulerpa, 94, 95

Caulerpa antoensis, 95

Caulerpa arenicola, 95

Caulerpa ashmeadii, 98

Caulerpa crassifolia f. pectinata, 95

	 Caulerpa mexicana f. pectinata, 95, 95, 

96

Caulerpa pectinata, 95

Caulerpa peltata, 95

Caulerpa pinnata f. pectinata, 95, 96

Caulerpa racemosa, 95

Caulerpa racemosa var. chemnitzia, 95

Caulerpa racemosa var. turbinata, 95

Caulerpa racemosa var. uvifera, 95

	 Caulerpa sertularioides, 95, 96, 97

Caulerpa sertularioides f. brevipes, 96

Caulerpa sertularioides f. longiseta, 96

	 Caulerpa vanbosseae, 95, 97, 98

Caulerpaceae, 78, 94

Caulerpales, 78

Caulerpella, 94

Chaetomorpha, 50, 53, 64

	 Chaetomorpha aerea, 51, 52, 54

Chaetomorpha aerea f. linum, 54

	 Chaetomorpha antennina, 51, 52, 52, 53

	 Chaetomorpha bangioides, 51, 53, 53

Chaetomorpha javanica, 50

Chaetomorpha linoides, 50

	 Chaetomorpha linum, 51, 53, 54

Chaetomorpha minima, 50

Chaetomorpha pachynema, 50

Chaetomorpha spiralis, 50

Chaetomorpha tortuosa, 66

Chaetophorales, 21

Chaetophora marina, 187

Chamaedoris, 70

Chlorobionta, 20

Chlorodesmis hildebrandtii, 94

Chloropelta, 32

Chlorophyceae, 77

Chlorophyta, 19, 20, 78

Chloroplastida, 19

Chnoospora, 172

Chnoospora fastigiata α [var.] pacifica, 

172

Chnoospora implexa, 171, 172

Chnoospora minima, 172, 173

	 Chnoospora pacifica, 171, 172, 173

Chnoospora pannosa, 172

Chnoosporaceae, 166, 171

Chordariaceae, 166, 174, 175, 187, 194

Chordariales, 108, 165

Chromavelolata, 106

Chromista, 106

Cladophora, 50, 54

	 Cladophora albida, 55, 56, 61

Cladophora bertolonii var. hamosa, 56

Cladophora brachyclados, 61, 62

	 Cladophora columbiana, 55, 56, 57

Cladophora crispata, 54

Cladophora delicatula, 61

Cladophora densa, 60

Cladophora expansa, 63

Cladophora fasicularis, 63

Cladophora flexuosa, 63, 64

Cladophora glomerata, 54

Cladophora glomerata var. crassior, 54

Cladophora gracilis, 63

	 Cladophora graminea, 55, 57, 58, 63

	 Cladophora hesperia, 55, 58, 59

Cladophora insignis, 54

Cladophora lehmanniana, 54

	 Cladophora macdougalii, 13, 55, 57, 59, 

60

Cladophora magdalenae, 56

Cladophora media, 53

Cladophora membranacea, 68

	 Cladophora microcladioides, 55, 56, 60, 

61

	 Cladophora montagneana, 55, 60, 61

Cladophora ovoidea, 60

Cladophora pacifica, 53

Cladophora pellucida, 57

	 Cladophora prolifera, 55, 62

Cladophora rivularis, 54

Cladophora rudolphiana, 62

Cladophora rugulosa, 62

Cladophora sakaii, 60

	 Cladophora sericea, 55, 60, 62

Cladophora stimpsonii, 54

	 Cladophora tiburonensis, 55, 56, 63

Cladophora trichotoma, 56

Cladophora uncinella, 63

Cladophora utriculosa, 54

	 Cladophora vagabunda, 55, 63, 64

Cladophoraceae, 50

Cladophorales, 49, 50, 67, 74

Cladophorophyceae, 20, 49

Cladophoropsis, 67, 68, 70, 76

	 Cladophoropsis gracillima, 68, 69, 69

	 Cladophoropsis macromeres, 68, 69, 70

Cladophoropsis membranacea, 68

Cladophoropsis mexicana, 72

Cladophoropsis pulcherrima, 70

Cladophoropsis robusta, 72, 73, 74, 76

Cladosiphonia? floridana, 197

Codiaceae, 78, 85

Codiales, 78

Codium, 86, 87, 89, 90

	 Codium amplivesiculatum, 86, 87, 88,

89

Codium anastomosans, 90, 91

	 Codium brandegeei, 14, 86, 87, 89, 90, 91

Codium cervicorne, 89, 90, 91

Codium conjunctum, 91

Codium cuneatum, 90, 91, 92

Codium cylindricum, 87

Codium decorticatum, 87

Codium dichotomum, 87, 91

Codium fernandezianum, 87

Codium fragile, 92

Codium isabelae, 90

Codium latum subsp. palmeri, 91

Codium longiramosum, 87

Codium macdougalii, 14, 90

Codium magnum, 87

Codium palmeri, 91

Codium picturatum, 86

Codium reductum, 90

Codium setchellii, 86

	 Codium simulans, 11, 86, 89, 90, 92

Codium sp. A, 92, 93

Codium tomentosum, 90, 91

Codium unilaterale, 91

Colpomenia, 12, 189, 190, 193

Colpomenia bullosa, 190

Colpomenia durvillei, 190

Colpomenia expansa, 193

Colpomenia peregrina, 189

	 Colpomenia phaeodactyla, 8, 190, 191, 

194

	 Colpomenia ramosa, 190, 191, 191, 199

	 Colpomenia sinuosa, 8, 11, 190, 192, 

192, 193
n u m b e r 9 4 • 2 7 3

Colpomenia sinuosa f. deformans, 190

Colpomenia sinuosa f. expanissima, 193

Colpomenia sinuosa f. expansa, 192

Colpomenia sinuosa f. tuberculata, 193

	 Colpomenia sp. A, 190, 193

	 Colpomenia tuberculata, 8, 190, 192,

193

Compsonema, 176, 189, 194

Compsonema gracile,194

	 Compsonema immixtum, 194

Compsonema intricatum, 194

Compsonema myrionematoides, 176

Composonema nummuloides, 176

Compsonema pusillum, 176

Compsonema ramulosum, 176

Compsonema secundum, 176

Compsonema secundum f. terminale, 176

	 Compsonema serpens, 194, 195

Compsonema speciosum f. piliferum, 176

Compsonema streblonematoides, 175

Conferva aerea, 51

Conferva albida, 55

Conferva antennina, 52

Conferva brachyclados, 61

Conferva clathrata, 41

Conferva crispata var. brachyclados, 62

Conferva expansa, 63

Conferva fascicularis, 63

Conferva flexuosa, 44, 63

Conferva gracilis, 63

Conferva granulosa, 170

Conferva hamosa, 56

Conferva implexa, 65

Conferva linum, 53

Conferva media, 53

Conferva pellucida, 57

Conferva prolifera, 62

Conferva riparia, 64

Conferva rudolphiana, 62

Conferva sericea, 62

Conferva siliculosa, 185

Conferva tortuosa, 66

Conferva trichotoma, 56

Conferva vagabunda, 63

Corynophlaea, 187

Corynophlaeceae, 187

Cruoria verrucosa, 161

Ctenocladales, 20, 21

Cutleria, 162, 165

Cutleria cylindrica, 163

	 Cutleria hancockii, 8, 163, 164

Cutleriaceae, 162

Cutleriales, 107, 161, 162

Cystoseiraceae, 200

Dasycladaceae, 101, 102

Dasycladales, 101, 102

Dasycladophyceae, 20, 101

Derbesia, 79, 82, 83, 84, 85

Derbesia hollenbergii, 83

	 Derbesia marina, 83, 84, 85

Derbesia sp., 83, 84, 85

	 Derbesia sp. A, 83, 84

Derbesia tenuissima, 82, 85

Derbesia turbinata, 83

Derbesia vaucheriaeformis, 84

Derbesiaceae, 78, 82

Desmarestia, 13, 145, 146

	 Desmarestia filamentosa, 145, 146, 147,

148

Desmarestia kurilensis, 146

Desmarestia ligulata, 145, 146, 148

Desmarestia media, 146

Desmarestia mexicana, 145, 146, 148

Desmarestia munda, 148

Desmarestia munda var. munda, 148

	 Desmarestia munda subsp. mexicana, 

146, 148, 149

Desmarestia pacifica, 146

Desmarestia viridis, 145, 146

Desmarestiaceae, 144

Desmarestiales, 107, 144

Desmotrichum, 175

Dictyerpa, 133

Dictyopteris, 118, 119, 120

Dictyopteris delicatula, 119

Dictyopteris johnstonei, 122

Dictyopteris membranacea, 119

Dictyopteris polypodioides, 119, 121

Dictyopteris repens, 119

	 Dictyopteris undulata, 8, 119, 121, 122

Dictyopteris zonarioides, 121

Dictyosiphonaceae, 174

Dictyosiphonales, 165, 186

Dictyosphaeria, 67, 74

Dictyosphaeria australis,74

Dictyosphaeria cavernosa, 74

Dictyosphaeria versluysii, 74

Dictyota, 117, 118, 119, 120, 122, 123, 126, 

130, 131, 133

Dictyota bartayresiana, 122

Dictyota binghamiae, 125, 126

Dictyota cervicornis, 122

Dictyota ceylanica, 128

	 Dictyota concrescens, 123, 124

	 Dictyota coriacea, 8, 120, 123, 124, 125, 

126, 130

	 Dictyota crenulata, 123, 126, 127

Dictyota dichotoma, 123, 128, 130, 132, 

133

	 Dictyota dichotoma var. intricata, 123, 

128, 129

Dictyota divaricata, 128

	 Dictyota flabellata, 123, 124, 128, 130, 

133

Dictyota friabilis, 123, 124

Dictyota hesperia, 130

Dictyota johnstonii, 124, 126, 130

Dictyota marginata, 141, 142

Dictyota okamurae, 141

Dictyota pinnata, 118

Dictyota pulchella, 128

	 Dictyota stolonifera, 123, 130, 131

	 Dictyota vivesii, 13, 123, 132, 132, 133

Dictyota volubilis, 123

Dictyotaceae, 109, 117, 118, 120, 126

Dictyotales, 7, 108, 117

Dictyoteae, 118

Dilophus, 118, 132, 133

Dilophus crenulatus, 126

Dilophus flabellatus, 128

Dilophus gunnianus, 118

Dilophus marginatus, 141

Dilophus okamurae, 141, 142

Dilophus pinnatus, 118

Dilophus stoloniferus, 131

Diplura, 108

Ectocarpaceae, 166, 180, 186, 194

Ectocarpales, 107, 108, 109, 156, 165, 

186

Ectocarpus, 180, 183

	 Ectocarpus acutus, 180, 181, 181

Ectocarpus breviarticulatus, 166

Ectocarpus bryantii, 14, 169, 170

Ectocarpus chantrasoides, 180

	 Ectocarpus commensalis, 181, 182, 183, 

184

Ectocarpus confervoides f. parvus, 184

Ectocarpus confervoides f. pygmaeus, 184

Ectocarpus confervoides f. variabilis, 182

Ectocarpus confervoides var. pygmaeus,

184

Ectocarpus corticulans, 180

Ectocarpus dimorphus, 182

Ectocarpus fasciculatus,184

	 Ectocarpus gonodioides, 170, 181, 182, 

183

Ectocarpus granulosus, 170

	 Ectocarpus hancockii, 181, 183, 184

Ectocarpus irregularis, 167, 168

Ectocarpus lebelii, 167

Ectocarpus mitchelliae, 170

Ectocarpus mucronatus, 167

	 Ectocarpus parvus, 181, 182, 184, 185

Ectocarpus pygmaeus, 184

	 Ectocarpus siliculosus, 181, 185

Ectocarpus siliculosus var. parvus, 184

Ectocarpus simulans, 180

	 Ectocarpus sonorensis, 181, 183, 186

Ectocarpus variabilis, 182

Ectochaete ramulosa, 30

Egregia, 219

Egregia laevigata, 219

Elaschistaceae, 174

Encoelium clathratum, 195, 196

Endarachne binghamiae, 189
2 7 4 • 

Endoderma vagans, 30

Endoderma viride, 25

Endoderma viride f. major, 23

Endoplura, 156

Endoplura aurea, 156

Enteromorpha, 32, 33, 41

Enteromorpha acanthophora, 41

Enteromorpha angusta, 34

Enteromorpha clathrata, 41

Enteromorpha clathrata var. crinita, 43

Enteromorpha compressa, 44

Enteromorpha compressa var. f­lexuosa,

Enteromorpha crinita, 43

Enteromorpha flexuosa, 44

Enteromorpha intestinalis, 46

Enteromorpha intestinalis var. tubulosa, 

46, 49

Enteromorpha lingulata, 45

Enteromorpha linza, 47

Enteromorpha marchantiae, 46

Enteromorpha marginata, 32

Enteromorpha micrococca, 32

Enteromorpha minima, 32

Enteromorpha muscoides, 43

Enteromorpha plumosa, 43

Enteromorpha prolifera, 48, 49

Enteromorpha prolifera subsp. radiata, 

48, 49

Enteromorpha prolifera var. flexuosa, 45

Enteromorpha radiata, 47, 49

Enteromorpha ramulosa, 41, 43

Enteromorpha ramulosa var. 

acanthophora, 41

Enteromorpha tubulosa, 46, 49

Entocladia, 21, 22, 25

Entocladia codicola, 26

Entocladia condensata, 25

	 Entocladia marchantiae, 22, 23, 24

Entocladia mexicana, 26

	 Entocladia polysiphoniae, 22, 23, 24

Entocladia robustum, 23

	 Entocladia sp. A, 22, 23, 24, 25

Entocladia velutina, 25

	 Entocladia viridis, 22, 23, 24, 25

Epicladia, 21, 22, 25

	 Epicladia condensata, 22, 25, 26

	 Epicladia mexicana, 22, 26, 26

Ernodesmis, 74

	 Ernodesmis verticillata, 75, 75

Feldmannia, 166, 167, 168

Feldmannia hemispherica, 167

	 Feldmannia irregularis, 167, 167, 168

Feldmannia lebelii, 167

Fucales, 7, 108, 199

Fucophyceae, 106

Fucus, 106, 199

Fucus anomalus, 150

Fucus gaertnera, 150

Fucus ligulatus, 146

Fucus minimus, 173

Fucus pedunculatus, 150

Fucus pyriferus, 220

Fucus sertularioides, 96

Fucus viridis, 146

Gastridium ovale, 84

Geppella, 94

Geppella? decussata, 94

Geppella mortensenii, 94

Giffordia, 168

Giffordia granulosa, 170

Giffordia irregularis, 167

Giffordia mitchelliae, 170

Glossophora, 118, 122

Glossophora coriacea, 124

Glossophorella, 118, 122

Glossophorella coriacea, 124

Gonodia, 187

Gonodia johnstonii, 187

Gonodia manchantiae, 14, 187

Halicystaceae, 82

Halicystidaceae, 82

Halicystis, 82, 84, 85

Halicystis ovalis, 84, 85

Halicystis parvula-phase, 85

	 Halicystis-phase, 82, 83, 84, 85, 85

Halicystis-phase of Derbesia marina, 84, 

85

Halimeda, 99

	 Halimeda discoidea, 99, 100

Halimeda opuntia, 99

Halimedaceae, 78, 94, 98

Halimedales, 77

Halimedineae, 78, 92, 94

Haliseris polypodioides, 121

Haliseris undulata, 121

Hapalospongidion, 155

Haplogloia, 174

Haplogloia andersonii, 174, 177

Hapterophycus, 156

	 Hapterophycus anastomosans, 156, 157

Hapterophycus canaliculatus, 156, 157

Hecatonema, 174, 175, 176

Hecatonema clavatum, 176

	 Hecatonema streblonematoides, 175, 

175, 176

Hecatonema variabile,176

Heterokonta, 106

Heterokontophyta, 106

Hincksia, 166, 167, 168, 169

Hincksia breviarticulata, 166

	 Hincksia bryantii, 168, 169, 169, 170

	 Hincksia granulosa, 168, 169, 170

Hincksia irregularis, 167, 168

	 Hincksia mitchelliae, 168, 170, 171

Hincksia rallsiae, 168

Hydroclathrus, 189, 195

Hydroclathrus cancellatus,196

	 Hydroclathrus clathratus, 195, 196

Ishige, 108, 109

Ishige foliacea, 108, 109, 111

Ishige okamurae, 108, 109, 111

	 Ishige sinicola, 108, 109, 110, 111

Ishigeaceae, 108

Ishigeales, 108

Laminaria, 106

Laminariales, 7, 13, 219

Laminariocolax, 179

Leathesia, 187

Leathesia difformis, 187, 189

Leathesia japonica, 187

Leathesia marina, 187, 189

	 Leathesia nana, 187, 188, 189

Leathesia sect. Leathesia, 187

Leathesia sect. Primariae, 187

Leathesia tuberiformis, 187

Leathesiaceae, 166, 187

Lessoniaceae, 220

Lithodermataceae, 155

Litosiphon laminariae, 179

Litosiphon pusillus, 179

Lobophora, 118

Lobophora variegata, 118

Lola, 64

Lola implexa, 65

Lola lubrica, 66

Lola tortuosa, 66

Macrocystis, 106, 220

Macrocystis pyrifera, 220

Melanophyceae, 106

Meneghiniella brandegeei, 177

Mesogloia andersonii, 177

Mesogloiales, 165

Mesosporaceae, 155

Microdictyon, 67, 68

Microzonia, 162

Myriactula, 187

Myriactula johnstonii, 187

Myriactula marchantiae,187

Myrionema, 194

Myrionemataceae, 156, 174, 175, 194

Myriotrichia, 175

Myriotrichia clavaeformis, 179

Myriotrichiaceae, 174

Nemacystus, 174, 176

	 Nemacystus brandegeei, 14, 177, 178

Nemacystus decipiens, 177

Nemoderma, 155

Nemodermataceae, 155

Nemodermatales, 155

Neomeris, 101
n u m b e r 9 4 • 2 7 5

Neomeris annulata, 101

Neurocarpus undulata, 121

Neurocarpus zonarioides, 121, 122

Newhousia, 106

Ochlochaete dendroides, 30

Pachydictyon, 118, 122, 126

Pachydictyon binghamiae, 125, 126

Pachydictyon coriaceum, 8, 124, 126

Padina, 7, 8, 106, 119, 120, 133, 165

Padina antillarum, 135

Padina boergesenii, 138

	 Padina caulescens, 134, 135

Padina concrescens, 133

	 Padina crispata, 134, 135, 136, 139

	 Padina durvillei, 118, 134, 136, 137, 

138

	 Padina gymnospora, 134, 138, 139

	 Padina mexicana var. mexicana, 133, 

139, 140, 141

	 Padina mexicana var. erecta, 134, 140, 

141

Padina ramonribae, 133

Padina sanctae-crucis, 133

Padina tetrastromatica, 135, 136

Padina vickersiae, 138

Parvocaulis, 102

Parvocaulis parvulus, 102

Parvocaulis pusilla, 102

Petalonia binghamiae, 189

Petalonia fascia, 161, 189

Petroderma, 155

Phaeophila, 30

	 Phaeophila dendroides, 30, 31

Phaeophila engleri, 30

Phaeophila ramulosa, 30

Phaeophila vagans, 30

Phaeophila viridis, 25

Phaeophilaceae, 30

Phaeophilales, 20, 30

Phaeophyceae, 106, 107

Phycoseris lobata, 36, 37

Phyllodictyon, 67, 68, 70, 74

Phyllodictyon anastomosans, 70

	 Phyllodictyon pulcherrimum, 70, 71

	 Phyllodictyon robustum, 70, 72, 73, 74, 

76

Pilayella, 186

Pocockiella variegata, 118

Polyopes sinicola, 109

Polyphysaceae, 102

Polytretus reinboldii, 187

Pringsheimia marchantiae, 14, 22

Pringsheimiella marchantiae, 22

Protista, 106

Protoctista, 106

Pseudolithoderma, 156

Pseudolithoderma nigrum, 156

Pseudostruvea, 72

Pseudostruvea robusta, 72

Pseudulvella, 28

Pseudulvella americana, 28

Pseudulvella applanata, 28

Punctaria, 175

Punctariaceae, 174, 175

Pylaiella, 186

Pylaiella littoralis, 186

	 Pylaiella sp. A, 186

Pylaiellaceae, 165, 186

Ralfsia, 156, 157, 159

Ralfsia californica, 158, 160, 161

	 Ralfsia confusa, 155, 158, 159, 160, 161

Ralfsia expansa, 160

	 Ralfsia hancockii, 158, 159, 160

Ralfsia hesperia, 158

Ralfsia occidentalis, 161

	 Ralfsia pacifica, 158, 159, 160, 161

Ralfsia verrucosa, 161

Ralfsia subgen. ralfsia, 155

Ralfsiaceae, 108, 155, 156

Ralfsiales, 107, 108, 155, 156

Rhipiliopsis mortensenii, 94

Rhizoclonium, 50, 64

Rhizoclonium hieroglyphicum, 64

Rhizoclonium implexum, 65, 66

Rhizoclonium kerneri, 65

Rhizoclonium kochianum, 65

Rhizoclonium lubricum, 66

	 Rhizoclonium riparium, 64, 65, 66

Rhizoclonium riparium f. validum, 66

	 Rhizoclonium riparium var. 

implexum, 64, 65, 66

	 Rhizoclonium tortuosum, 64, 66, 66

Rosenvingea, 189, 196, 197

Rosenvingea aff. santae-crucis, 197

	 Rosenvingea antillarum, 197, 198

	 Rosenvingea floridana, 197, 198

	 Rosenvingea intricata, 191, 197, 198, 

199

Rugulopteryx, 118, 119, 120, 132, 141

Rugulopteryx marginata, 141

	 Rugulopteryx okamurae, 141, 142

Sargassaceae, 200

	 Sargassum, 7, 8, 13, 106, 118, 138, 199, 

200, 201, 213, 217

	 Sargassum acinacifolium, 201, 206,

207, 210

Sargassum agardhianum, 219

Sargassum alternato-pinnatum, 209

Sargassum asymmetricum, 209, 210

	 Sargassum brandegeei, 14, 201, 206, 

217, 218, 219

Sargassum bryantii, 205

Sargassum californicum, 213

Sargassum camouii, 213

Sargassum cylindrocarpum, 200

Sargassum fuliginosum var.? californica, 

213

Sargassum guardiense, 202

	 Sargassum herporhizum, 201, 214, 216, 

217, 219

Sargassum horridum, 200, 210, 213

Sargassum howelli, 200

Sargassum insulare, 205

	 Sargassum johnstonii, 12, 13, 201, 202, 

203, 217

	 Sargassum johnstonii f. gracile, 202, 

204

Sargassum johnstonii f. laxius, 202

	 Sargassum lapazeanum, 201, 204, 205, 

207, 209

Sargassum liebmanni, 200, 214

	 Sargassum macdougalii, 14, 201, 207, 

208, 209

Sargassum muticum, 200

Sargassum pacificum, 200

Sargassum palmeri, 200

Sargassum paniculatum, 213

Sargassum polyacanthum f. americanum, 

210

	 Sargassum sect. herporhizum, 214

	 Sargassum sect. johnstonii, 201

	 Sargassum sect. lapazeanum, 202

	 Sargassum sect. sinicola, 210

	 Sargassum sinicola subsp. camouii,

201, 213, 214, 215, 217

	 Sargassum sinicola subsp. sinicola,

200, 201, 210, 211, 212, 213, 214

Sargassum sinicola var. camouii, 213

	 Sargassum sonorense, 201, 207, 209, 

209, 210

	 Sargassum subgen. Sargassum, 201

Scytosiphon, 156, 194

Scytosiphon bullosus, 190

Scytosiphon canaliculatus, 156

Scytosiphon lomentaria, 161

Scytosiphonaceae, 156, 161, 166, 171, 

189, 194

Scytosiphonales, 165

Scytothamnaceae, 166

Siphonales, 78

Siphonocladaceae, 67, 74

Siphonocladales, 49, 50, 67, 74

Siphonocladus, 67, 68, 74

Siphonocladus pusillus, 74

Siphonocladus wilbergii, 74

Spatoglossum, 118, 119, 120, 142, 143

Spatoglossum cf. schroederi, 144

Spatoglossum howellii, 143, 144

	 Spatoglossum lanceolatum, 143, 143

	 Spatoglossum subflabellatum, 143, 144

Spermatochnaceae, 174

Sphacelaria, 111, 112, 115

Sphacelaria brevicornis, 112
2 7 6 • 

	 Sphacelaria californica, 112, 114, 115, 

116

Sphacelaria cornuta, 112

Sphacelaria didichotoma, 112

Sphacelaria divaricata, 112

Sphacelaria furcigera, 113, 114

	 Sphacelaria hancockii, 112, 114, 115, 

115, 116

Sphacelaria mexicana, 116

Sphacelaria novae-hollandica, 114

Sphacelaria plumula var. californica, 114, 

115

	 Sphacelaria rigidula, 112, 113, 114

Sphacelaria sp., 112

Sphacelaria subfusca, 114

Sphacelaria subgen. Propagulifera, 111, 

112

	 Sphacelaria subgen. Propagulifera

sect. Furcigerae, 112

	 Sphacelaria subgen. Propagulifera

sect. Tribuloides, 114, 116

	 Sphacelaria tribuloides, 112, 114, 116, 

117

Sphacelariaceae, 111

Sphacelariales, 107, 111

Sphacella, 111

Sphaerotrichia divaricata, 179

Sporochnaceae, 150

Sporochnales, 107, 148

Sporochnus, 13, 150

	 Sporochnus anomalus, 150. 151, 152, 

153

Sporochnus apodus, 153, 155

Sporochnus bolleanus, 150, 152

Sporochnus gaertnera, 150

	 Sporochnus neushulii, 150, 153, 154, 

155

Sporochnus pedunculatus, 150, 152, 153, 

155

Stilophora antillarum, 197

Stilophora rhizodes, 179

Stilophora tenella, 179

Stramenopiles, 105

Streblonema, 174, 177, 179, 180

Streblonema aecidioides, 179

	 Streblonema sp. A, 180

Streblonema sphaericum, 179

	 Streblonema transfixum, 179, 179

Stromatella, 22, 26, 27

	 Stromatella monostromatica, 27, 27

Struvea, 68, 70, 72

Struvea anastomosans, 70

Struvea delicatula, 70

Struvea pulcherrima, 70

Struvea ramosa, 70

Struvea sp., 68

Struveopsis, 70, 72

Struveopsis robusta, 72

Taonia, 118

Taonia lennebacherae, 118

Tilopteridaceae, 162

Tilopteridales, 162

Udoteaceae, 78, 94

Ulothrix, 20

Ulotrichales, 20

Ulva, 12, 31, 32, 33, 34

	 Ulva acanthophora, 33, 41, 42

Ulva angusta, 34, 35

	 Ulva californica, 33, 34, 35

	 Ulva clathrata, 34, 41, 43, 44

	 Ulva compressa, 33, 44, 47

Ulva costata, 37, 39

Ulva dactylifera, 39

Ulva dichotoma, 132

Ulva expansa, 33

Ulva fasciata, 33

Ulva fasciata f. costata, 37

Ulva fasciata f. expansa, 33

Ulva fasciata f. lobata, 37

Ulva fasciata f. taeniata, 39

	 Ulva flexuosa, 34, 44, 45, 49

	 Ulva intestinalis, 33, 46, 47

	 Ulva lactuca, 33, 35, 36

Ulva lactuca var. rigida, 40

	 Ulva linza, 33, 47, 48

	 Ulva lobata, 33, 36, 37

Ulva muscoides, 43

	 Ulva nematoidea, 33, 37, 38, 39

Ulva paradoxa, 43

Ulva pertusa, 33

Ulva prolifera, 49

	 Ulva radiata, 34, 41, 47–49

Ulva ramulosa, 41, 43

	 Ulva rigida, 33, 39, 40

Ulva scagelii, 34

	 Ulva sect. enteromorpha, 31, 32, 33, 41

	 Ulva sect. ulva, 31, 32, 33, 34, 41

Ulva sinuosa, 192

Ulva taeniata, 34, 39

Ulvaceae, 31

Ulvales, 20, 21, 31

Ulvella, 21, 27, 28

Ulvella applanata, 28

	 Ulvella lens, 28, 29

Ulvella monostromatica, 27

	 Ulvella setchellii, 28, 29, 29, 30

Ulvellaceae, 21

Ulvophyceae, 20

Undaria, 179

Valonia, 67, 74, 75, 76

Valonia macrophysa,75

Valonia pachynema, 76

Valonia pusilla, 74

Valonia verticillata, 75

Valoniaceae, 67, 74, 75

Valoniopsis, 72, 74, 75, 76, 77

	 Valoniopsis hancockii, 76, 77

	 Valoniopsis pachynema, 72, 76, 77

Vaucheria, 86, 106

Vaucheria marina, 83

Vaughaniella, 118, 133

Willeella, 72

Willeella mexicana, 72, 73, 74

Zanardinia, 162

Zanardina prototypus, 163

Zanardina typus, 163

Zonaria, 118

Zonaria dichotoma var. intricata, 128

Zonaria gymnospora, 138

Zonarieae, 118
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journals, use the parentheses system for volume(number):pagination [e.g., “10(2):5–9”]. Do not use “et al.”; all authors

editors should be included in reference citations. In titles, 

capitalize first word, last word, first word after colon, and all 

other words except articles, conjunctions, and prepositions. 

Examples of the most common types of citations are provided 

in the SISP Manuscript Preparation and Author Style Guide. 

For questions regarding the guidelines, please email SISP at 

schol.press@si.edu. 

requirements for smithsonian series publi