Smithsonian contributions to the marine sciences

The Red Alga Polystphonia

(Rhodomelaceae) in the
orthern Gulf of California

comme SHOLLENBERG “= Ses |

Ge. caer

= 5 a
gis ie

sda othe
PPS

NABN(O) 0:5

SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

Emphasis upon publication as a means of “‘diffusing knowledge’ was expressed
by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph
Henry outlined a program that included the following statement: “‘It is proposed to
publish a series of reports, giving an account of the new discoveries in science, and
of the changes made from year to year in all branches of knowledge.”’ This theme
of basic research has been adhered to through the years by tl.ousands of titles issued
in series publications under the Smithsonian imprint, commencing with Smithsonian
Contributions to Knowledge in 1848 and continuing with the following active series:

Smithsonian Contributions to Anthropology
Smithsonian Contributions to Astrophysics
Smithsonian Contributions to Botany
Smithsonian Contributions to the Earth Sciences
Smithsonian Contributions to the Marine Sciences
Smithsonian Contributions to Paleobiology
Smithsonian Contributions to Zoology
Smithsonian Studies in Air and Space
Smithsonian Studies in History and Technology
In these series, the Institution publishes small papers and full-scale monographs
that report the research and collections of its various museums and bureaux or of
professional colleagues in the world cf science and scholarship. The publications are
distributed by mailing lists to libraries, universities, and similar institutions throughout
the world.
Papers or monographs submitted for series publication are received by the
Smithsonian Institution Press, subject to its own review for format and style, only
through departments of the various Smithsonian museums or bureaux, where the

manuscripts are given substantive review. Press requirements for manuscript and art
preparation are outlined on the inside back cover.

S. Dillon Ripley
Secretary
Smithsonian Institution

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES e NUMBER 1

The Red Alga Polysiphoma
(Rhodomelaceae) in the
Northern Gulf of California

George 7. Hollenberg

and fames N. Norris

ISSUED
pea

SMITHSONIAN INSTITUTION PRESS
City of Washington
1977

ABSTRACT

Hollenberg, George J., and James N. Norris. The Red Alga Polysiphonia
(Rhodomelaceae) in the Northern Gulf of California. Smithsonian Contributions
to the Marine Sciences, number 1, 21 pages, 10 figures, 1977—Taxonomic studies
of Polysiphonia show 14 species to be present in the northern Gulf of California,
Mexico. One of these, P. sphaerocarpa var. cheloniae is described herein as a
new variety, and is found growing exclusively on the green sea turtle, Chelonia
mydas L. This and three other species are recorded for the first time in the Gulf
of California. The distribution of six other species is extended.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded
in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Seascape along the
Atlantic coast.

Library of Congress Cataloging in Publication Data

Hollenberg, George J

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

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

Bibliography: p.

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

1. Polysiphonia. 2. Algae—Mexico—California, Gulf of. I. Norris, James N., joint author.
II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to the marine
sciences ; no. 1.

QK569.R37H64 589'.41 76-608299

Contents

Page
Jn ErOCINGMOM “so 65.66.58 6m dS a OD Oa ne aes HIE eee err enCe aeeaeeeaare
iK@y (@ UNS STSTIES Oi JHGIN WHOIS 6066 eco soso nap Onn obb do poose ooh eaueS 2
Oly st PIL OnCOMfusaatAOlleENbeE oir eos aise -o os psee oo. eaienle oie) 2
OMe PILOMIANLACCIAISS171 A EIOMEMDELS ila. © cles): ss als oe we ees wie so oes 4
Polysiphonia johnstonii Setchell & Gardner var. johnstonii ............. 4
Polysiphonia johnstonii var. concinna (Hollenberg) Hollenberg ......... 7
olsipwoniaynasoni,. setchelll % Gardner 2. ss. 252. oe ae 8
Polysiphonia mollis J. D. Hooker & Harvey in Harvey ................. 9
Eolsrphonia pacijica var. delicatula Hollenberg .-......:.-.....5.0... 10
malsipwoniauapaniculata Ntontagpme) 5)... ee. sles sess ee cle ee ee 1]
EROUY SUPILONMIDSAUATICT IFA ATI OL: ats Acs, Sie ste a Saya s sj sincste 6 oieteisis oe Sloe wales 12
Polysiphonia scopulorum var. villum (J. G. Agardh) Hollenberg ......... 14
RaGUY SEP LOMIGESTIT PLCXPEAOUEMDELE) Vici x oc. Se Hcreos ine eS eres eevee ve one we Ass
eG MICOS OILONCIISIS EAOMENMDCL GE ya\cte elas = is sels oe ose soe eee 16
Polysiphonia sphaerocarpa var. cheloniae, new variety ................-. 16
PONS MOORID GUAGES os 4 Sagon ec aa po Ace eRe cae a eer ae 17
Literate CHIGGL: 5 a5 Falco es 8 as COINS Gone HEEL OC eras eee nrae 19
ERC 5.0 6 08'8 50:60 00.0 0 HEIDE SE SE Oe Ocenia eet re saa eee 21

ili

ey

The Red Alga Polystphonia
(Rhodomelaceae) in the
Northern Gulf of California

George 7. Hollenberg
and James N. Norris

Introduction

The earliest records of Polysiphonia Greville
(1824:308) in the Gulf of California were provided
by Setchell and Gardner (1924), who described three
new species. Later, Dawson (1944), in his investiga-
tions on marine algae of the Gulf of California,
recorded seven species, including a new one. A de-
tailed study of the taxa of Polysiphonia from the
Gulf of California and Pacific coast of Baja Cali-
fornia was first presented by Hollenberg (1961).
These three papers constitute the basis for our
knowledge of Polysiphonia in the Gulf of California.

Members of the genus Polysiphonia (Rhodome-
laceae, Ceramiales) are distinguished by several im-
portant taxonomic characters: the number of peri-
central cells; presence or absence of cortication; the
manner of origin of rhizoids; the manner of origin
of branches; the nature and position of trichoblasts;
and the nature of spermatangial branchlets. ‘These
features have been pointed out in previous investi-
gations by Hollenberg (1942a; 1942b; 1944; 1961;
1968a).

Herein we present our studies of recent intertidal
and subtidal collections of Polysiphonia from the

George J. Hollenberg, Department of Biology, University of
Redlands, Redlands, California 92373. James N. Norris, De-
partment of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

northern Gulf of California. These collections were
obtained by the junior author during field expedi-
tions in 1972-1974 on the Gulf coasts of Baja Cali-
fornia and Sonora. In addition, the following collec-
tions were utilized during the course of this study:
those of the senior author from Puerto Penasco,
Sonora, in 1967; some additional material of the
late Dr. E. Yale Dawson; and those of the junior
author and Katina Bucher from the cruise of the
R/V Dolphin to Las Islas de la Cintura (Midriff
Islands) (Norris and Bucher, 1976).

Collectors are designated as follows: JN for J. N.
Norris; H for G. J. Hollenberg; D for E. Y. Dawson;
and KB for Katina E. Bucher. The numbers cited
correspond to the collector’s field notebooks. Speci-
mens are being deposited in the following herbaria:
first set, U.S. National Herbarium, Smithsonian In-
stitution (US); duplicate material to University of
California, Berkeley (UC); Phycology Herbarium,
University of Arizona (ARIZ); Universidad Na-
cional Autonoma de México (MEXU). Herbaria
housing the holotypes are abbreviated in accordance
with Holmgren and Keuken (1974). Geographical
locales under “Specimens Studied”’ are listed from
north to south.

ACKNOWLEDGMENTS.—The Latin diagnosis was
kindly provided by Dr. Hannah T. Croasdale
(Dartmouth College). Field studies for collections
were made possible through the support of the Na-

tional Science Foundation (research grant GB-28623)
to Dr. M. Neushul and J. Norris (Marine Science
Institute, University of California, Santa Barbara).
The junior author also wishes to thank again the
University of Arizona for use of their Laboratorio
de Biologia Marina, Puerto Penasco, Sonora, while
he served as resident Marine Biologist /Station Di-
rector of the facility. Collections by Norris and
Bucher from Las Islas de la Cintura were made
possible only through the generosity of Dr. William
Fenical (Scripps Institution of Oceanography), Chief
Scientist on the R/V Dolphin cruise of April 1974.

SMITHSONIAN CONTRIBUTIONS FO THE MARINE SCIENCES

Collections from the sea turtles were obtained by
Dr. Richard S. Felger (Arizona-Sonora Desert Mu-
seum, Tucson, Arizona) and Edward W. Moser
(Summer Institute of Linguistics, University of
North Dakota). Further collections were given to
Norris by Dr. Felger, Professor Richard Evans
Schultes (Director, Botanical Museum of Harvard
University), and Dr. Andrew T. Weil (Botanical
Museum of Harvard University). Finally, we wish
to thank Drs. Isabella A. Abbott, Robert W.
Hoshaw, and Katina Bucher for reading the manu-
script and offering suggestions for its improvement.

Key to the Species of Polysiphonia

THE SPECIES ARE ARRANGED ALPHABETICALLY IN THE TEXT

J: Pericentrall-cells4(Section Oligosiphomia) e-c---..--.cscsssssrsssceccesscsvsterseconrastsesnsser tapeteesrsesnacenttameententes 2
Pericentral cells more than 4 (Section Polysiphonia) .........0.0..cccccecceee cece cece ceeeeeseneneeseneeeseeseneeees 11

2. Trichoblasts or scar-cells in general, regularly 1 per S€GMENt ........... cesses este ees esesesesesteneseseseees 3
Trichoblasts or scar-cells lacking or not regularly 1 per segMeNt ........::cc ccc eects eseseseseseeeaeeeeeees 9

3. Branching dichotomous, throughout <.-.2...c.:.scs-s-nscerusneseeressstessssesonsracasss2sotcsucussscoseet seam ates P. masonii
Branching lateral, not strictly dichOtomGus) 2iive!cssecssscscescsecstserstscesscsses sosusseucorscesearene tease nea 4

4 GTOWANG TON SeATtUTtLES ssc tes ee ee etscscersrereetere est P. sphaerocarpa var. cheloniae, new variety
NOt growin Ee KOMUSeANtUTElES| coke. ccccwrscsscestetettee fesse nstnansuar suse Me ereee nae tana atar eae aean nanan manatee ereemeeten 5

5. Branches arising from a primary branch of a trichoblast (axillary to a trichoblast) ................
Ee beech gnc da cu eien tee Eades de eaaseya eta cocsnce be ce due see best eee aed ca cute a OMe gee R WBE A aataet DMCR EE ER Een P. flaccidissima
Branches arising from an entire trichoblast priMOrdiuM .........c.cc cscs ecseses eee eeeereseseseeeeneneneeeeee 6

6.- Epiphytic, mostly less tham);2 cml bighy esses sresesccsscecessestererssesstscsoscvesesststecrastetsetaaness P. savatieri
Wesurally motiepiphy tice cece escccosccececereroresscsseetecsccacseuste rows ntscor esac ereratncac nesters ote enn erp t

7. Segments in main branches mostly shorter than their diameter or sometimes as long ..............
wvedesenecsssieSvotcavrevatero¥lshusseusyisissacsnssurveau(Survecoutsasterecesssnstesdritneossecesuvetosrkistacseicewoaeiovosemenc erate P. simplex
Segments in main branches mostly 1.5 times as long as their diameter or longer .................. 8

8. Trichoblasts usually well developed; spermatangial stichidia arising as a primary branch of
EY oe 000) 0) ESL rere socee eee eee ee reer pect rseacpeacpar had ss sooc CaP ASCOEC sata oan cncn seo oasbotasooaocasos06020a00000008 P. mollis
Trichoblasts very rudimentary; spermatangial stichidia arising from an entire trichoblast

FONG Bets CO BND TT Oty ear ee setae eee cca orca nee Ce po re eR Sheba Cte cee Beopo ee coneto Polysiphonia species

9. Thalli delicate, mostly intertidal, commonly forming mats or furry patches .........ccee
Sesessarderegsseseetiaea ges svesusaevs stvassistes totonststen ate ctee west rnin aoueaeape nese ats seca rose P. scopulorum var. villum

- Thalli not as delicate, mostly subtidal, not forming mats or patches .......ssssssssuseeeeeseeee 10
10. Thalli mostly less than 2 cm high; trichoblasts infrequent to rare ....P. pacifica var. delicatula
Thalli up to 4 cm high, of a terracotta color; trichoblasts numerous, arising at irregular
PS 1 CSc Bas acca ee. oC CAFE ESOC SE CD SGEBOIISOB CODD P. sonorensis

Lz) )Pericentrall=cells) 5G) scree ceseca sess teo ecto se ese recte eee eos ene te ere eee rea 12
erate a el CaN GC) Oye TIVE) crcatercscocetoscocacaaceorcacaeaca aoco; aces osaccdcéd+co00-FeSnocacosancoraacococceceacacosoacatcaccee20c0 13

12. Thalli occasionally epiphytic, mostly 5 cm or more high ............ P. johnstonii var. johnstonii
Thalli usually epiphytic, mostly less than 5 cm high ..............0. P. johnstonii var. concinna

13S halligmostly;lessithane2eemishig hikes nesses scccs ee ectacserecceete et carter etraceereanece ce steerer P. confusa
Thalli mostly much largers to 20) craVMighy eset sns-cscectocasscsressevbectesctve eseecuccertoresscorsess P. paniculata

Polysiphonia confusa Hollenberg

Polysiphonia confusa
Dawson, Acelto, and Foldvik, 1964:87, pl. 33p.—Abbott and
Hollenberg, 1976:696, fig. 643.

Polysiphonia inconspicua sensu Hollenberg,

Polysiphonia inconspicua Reinsch, 1888:146].

Ficure |

Hollenberg, 1961:350, pl. 1; fig. 5.—

1944:479 [not

DescriPpTION.—Thalli 0.8-1.5(-3.0) cm high, with
erect axes mostly assurgent from prostrate branches
100-175 wm in diameter, composed of segments

NUMBER 1

Pas

a
\ |
ee.N
‘on &
Tea
100 pm*
& We a 4 a. 2)

5 | " B
" et

om Se

Figure 1.—Polysiphonia confusa: A, upper portion of thallus (JN-5577); B, cystocarpic plant
(JN-3835).

about as long as the diameter or somewhat longer,
attached by numerous unicellular rhizoids, arising
1 to several per segment as cells cut off from the
proximal ends of the pericentral cells, and fre-
quently having digitate apices; erect branches
60-150 »m in diameter, the segments 1.0—2.5 times
as long; lateral branches arising exogenously in con-
nection with trichoblasts at irregular intervals; peri-
central cells 8-10, without cortication; trichoblasts
to 1 mm long, with | or 2 dichotomies, arising 1 per
segment in spiral sequence with 14 divergence, soon
deciduous, leaving persistent scar-cells.

Tetrasporangia, 60-80 ym in diameter, in long
spiral series in slightly swollen segments of ultimate
and subultimate branches; cystocarps globular, to
225 um in diameter; spermatangial plants unknown.

Type-Locatity.—Mid-intertidal rocks, Corona
del Mar, Orange County, California.

Ho.otyPe.—H-3285, 24 October 1942, in her-
barium of G. J. Hollenberg (now at US).

GULF OF CALIFORNIA DiIsTRIBUTION.—Mid-
intertidal regions; epizoic on sand dollars, epiphytic
on Codium and growing on hard substrate of buoy;
Puerto Pefiasco to Desemboque de San Ignacio.

PaciFic Coast DistRisuTION.—U.S.A.: southern
California. Mexico: northern Baja California.

SPECIMENS STUDIED.—sONORA: JN-5577, sterile, growing on a
sand dollar, Encope grandis, Bahia Cholla near Punta Peli-
cano, legit J. N. and K. B., 25 March 1974; JN-3835, cysto-
carpic, on NASA buoy, anchored off Laboratorio de Biologia
Marina, Playa Estacién, Puerto Penasco, legit J. N., 22
March 1973; JN-5158, growing on Codium sp., Desemboque
de San Ignacio, legit R. E. Schultes, R. S. Felger, andwAtmlr
Weil, 3 April 1974.

ReMARKsS.—Previously known on the Pacific coast
from southern California and northern Baja Cali-

4

fornia (Hollenberg, 1961:351), this taxon is now
recorded in the Gulf of California. It is apparently
rare in the northern Gulf; only three collections
have come to our attention. Although none of these
specimens is from the habitat described as charac-
teristic of P. confusa Hollenberg, size and structural
features leave little room for doubt as to the identity
as given.

Polysiphonia flaccidissima Hollenberg
FIGURE 2E

Polysiphonia flaccidissima Hollenberg, 1942b:783, figs. 8, 19;
1961:351, pl. 2: fig. 2; 1968a:63, figs. 2a, 11.—Dawson,
Acelto, and Foldvik, 1964:88, pl. 788—Abbott and Hollen-
berg, 1976:688, fig. 634.

DescripTion.—Epiphytic or saxicolous algae 10-
25 mm high, erect from creeping basal branches
attached by numerous unicellular rhizoids, which
are cut off as separate cells from the pericentral
cells; main erect axes 70-80 »m in diameter, with
segments mostly 1-2 times as long as the diameter;
branches of several to many orders, arising exoge-
nously from a primary branch of a trichoblast,
mostly at intervals of 6 segments or less, narrowed
gradually below and above, at first usually strict
and curving toward the parent branch; pericentral
cells 4, totally ecorticate, with thin hyaline walls;
trichoblasts simple or mostly with a single dichot-
omy, tapering to slender apices, arising 1 per -seg-
ment in spiral sequence, with 14 divergence, soon
deciduous, leaving persistent scar-cells.

Tetrasporangia 50-70 ym in diameter, in spiral
series in smaller branches, somewhat distending the
segments; cystocarps globular; spermatangial sti-
chidia arising as a primary branch of a trichoblast,
with 1-2 celled sterile apices.

Typre-LocaLtiry.—On a coralline alga, rocky
point, Laguna Beach, Orange County, California.

Ho.LotypPr.—H-2269, 1 January 1938, in her-
barium of G. J. Hollenberg (now at US).

GULF OF CALIFORNIA DisTRIBUTION.—Intertidal
to subtidal, on rocks or occasionally epiphytic;
Guaymas to Isla Espiritu Santo.

PaciFic Coast Distrisution.—U.S.A.: southern
California; Hawaiian Islands. Mexico: northern
Baja California; Revilla Gigedo Archipelago; Guer-
rero. Costa Rica. Panama. Peru.

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

SPECIMENS STUDIED.—SONORA: JN-4267, tetrasporangial, and
JN-4270, on Padina, from 1.5-6.1 m depth, Nueva Guaymas,
legit Dale Evensen, 13 Oct 1973.

Remarks.—In the northern Gulf this species is
apparently rare, restricted in distribution to Guay-
mas and vicinity, Sonora. Across the Gulf it has
been recorded from Santa Rosalia, and in the
southern Gulf from Puerto Escondido and Isla
Espiritu Santo (Hollenberg, 1961:352).

Polysitphonia johnstonii Setchell & Gardner
var. johnstonii

FIGURES 2B, 3

Polysiphonia johnstonii Setchell & Gardner, 1924:767.—
Hollenberg, 1944:475; 1961:357, pl. 2: figs. 4, 5——Dawson,
1944:329; 1949:237; 1951:53, 57; 1954b:160; 1959:32; 1966:29.
—Dawson, Neushul, and Wildman, 1960a:72; 1960b:26.—
Norris, 1972:16.—Abbott and Hollenberg, 1976:699, fig. 649.

Polysiphonia marchantae Setchell & Gardner, 1924:768, pl.
49a.

Polysiphonia sinicola Setchell & Gardner, 1924:769.

Polysiphonia acuminata sensu Dawson, 1950:68; 1954b:159
[non Polysiphonia acuminata Gardner, 1927:100}].

Polysiphonia richardsoni sensu Segi, 1951:253.—Dawson,
1954b:160 [in part, with reference to Gulf of California
specimens cited by Setchell and Gardner, 1924:768 as
P. marchantae only; non Polysiphonia richardsoni W. J-
Hooker ex Harvey in W. J. Hooker, 1833:333].

Description.—Thalli relatively rigid, 5-12 cm
high, dull red in color, nearly black on drying,
attached by numerous unicellular rhizoids, which
are cut off as separate cells from the pericentral
cells of a limited prostrate base; primary erect
branches several, relatively distinct, to 1 mm in
diameter, repeatedly branched, the branches spirally
arranged, arising at a narrow angle, which widens
as the branch matures; ultimate branches usually
relatively close and dense; segments in main
branches 1.5 times as long as the diameter or
shorter, with 5-6 pericentral cells, totally ecorticate;
trichoblasts once or twice forked, arising | per seg-
ment in spiral sequence, with 14 or 1% divergence,
soon deciduous, leaving persistent scar-cells; branches
replacing trichoblasts in origin at irregular in-
tervals.

Tetrasporangia 1(—2) per segment, 70-90 pm in
diameter; cytocarps globular, 450-500 ym in diam-

NUMBER 1

aN

99 Old eaaos SCS
agQaoe

eS
IQ) oe
p WO — es

FicurE 2.—a, Polysiphonia johnstonii var. concinna, branch tip of a tetrasporic plant (after
Hollenberg, 1944); 8, P. johnstonii var. johnstonii, apical portion of an axis showing trichoblasts
and characteristically incurved tips (after Hollenberg, 1961); c, P. sonorensis, branch tip of the
holotype (after Hollenberg, 1961); bp, P. masonii, upper portion of a tetrasporic plant (after
Hollenberg, 1961); ©, P. flaccidissima, branch tip (after Hollenberg, 1942b).

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

FicurE 3.—Polysiphonia johnstonii var. johnstonii: a, B, habits of two specimens showing the
divergent manner of branching (JN-3201).

NUMBER 1

eter, nearly sessile; spermatangial stichidia arising
as a primary branch of a trichoblast, 100-140 by
30-40 pm in length and diameter, respectively.

Typr-LocaLtity.—On Gracilaria sp., Isla San

Esteban, Gulf of California, Mexico.

Hototyre.—Ivan M. Johnston 110, April 1921
(CAS 1361, now at UC).

GULF OF CALIFORNIA DIsTRIBUTION.—Intertidal
and subtidal, on rocks or epiphytic; Puerto Penasco
to La Paz.

PaciFic Coast DistrRipuTion.—Santa Catalina
Island, California; northern Baja California; Si-
naloa.

SPECIMENS STUDIED.—soONORA: JN-3102, rocky intertidal,
Punta Pelicano, 30 Jun 1972; JN-3260, on Sargassum sp. on
rock platform with sand areas, Playa Hermosa, Puerto Pejiasco,
legit J. N. and K. B., 26 Jul 1972; JN-4040, spermatangial,
intertidal rock platform, Playa Hermosa, Puerto Pefiasco, legit
J. N. and K. B., 3 May 1973; JN-5150, beach drift after strong
winds, Desemboque de San Ignacio, legit R. E. Schultes, R. S.
Felger, and A. T. Weil, 3 Apr 1974. BAJA CALIFORNIA DEL
NorTE: JN-5462, 1-7 m depth, Punta la Gringa, Bahia de
Los Angeles, legit J. N. and K. B., 28 Apr 1974; JN-3029,
5 m depth, Punta la Gringa, Bahia de los Angeles, 22 May
1972; JN-3201, and JN-3209 (a slender form), estuary, Bahia
de las Animas, legit J. N. and G. Boehlert, 23 May 1972. Las
ISLAS DE LA CINTURA: JN-5280, tetrasporangial and cystocarpic,
10 m depth, W side of Roca Blanca, Puerto Refugio, Isla
Angel de la Guarda, legit J. N., J. Paul, and K. Robertson,
21 Apr 1974; JN-5314, intertidal to 10 m depth, Puerto
Refugio, Isla Angel de la Guarda, legit J. N. and K. B.,
21 Apr 1974; JN-5369, 5-12 m depth off small islet, Puerto
Refugio, Isla Angel de la Guarda, 21 Apr 1974; JN-5762,
intertidal, rocky shore NE side of Puerto Refugio, Isla Angel
de la Guarda, 23 Apr 1974; JN-5483, 3-10 m depth, S end of
Isla Estanque, legit J. N. and K. B., 27 Apr 1974; JN-5508 and
JN-5616A, 3-12 m depth, S$ end of Isla Estanque, legit J. N.
and K. B., 27 Apr 1974; JN-5561, tetrasporangial, 3-8 m depth,
rocky shore SE end of Isla San Esteban, 25 Apr 1974; JN-5528,
cystocarpic, 3-5 m depth, off SE end of Isla San Esteban,
26 Apr 1974; JN-5707, epiphytic on Laurencia sinicola, and
JN-5709, 3-8 m depth, SE end of Isla San Esteban, 25 Apr
1974; JN-5518, 1-5 m depth, SE end of Isla San Esteban, legit
K. B., 26 Apr 1974.

Remarks.—One of the most frequently encoun-
tered species of Polysiphonia in the northern Gulf,
its distribution is now extended northward to
Puerto Pefiasco from Puerto Libertad (Hollenberg,
1961:358). In the northern Gulf, specimens are
usually epiphytic and commonly sterile.

Polysiphonia johnstonii var. concinna
(Hollenberg) Hollenberg

Ficurss 2A, 4D

Polysiphonia concinna Hollenberg, 1944:474, fig. 10.—Dawson,
1951:53, 56; 1954b:159; 1959:32.

Polysiphonia johnstonii var. concinna (Hollenberg) Hollen-
berg, 1961:358, pl. 3: fig. 4—-Abbott and Hollenberg, 1976:
699, fig. 650.

DescripTion.—This variety is similar to P. john-
toni var. johnstonii in detailed features, but
smaller and shorter in dimensions throughout.

Typr-Locatity.—Epiphytic on other algae, near
Scripps Institution of Oceanography, La Jolla, San
Diego County, California.

Ho.otypr.—H-2015, December 1936,
barium of G. J. Hollenberg (now at US).

GULF OF CALIFORNIA DISTRIBUTION.—Mid-
intertidal to subtidal, epiphytic on various algae,
occasionally on rocks; Puerto Penasco to Bahia
Agua Verde.

in her-

Pacific Coast Distrisution.—Southern Cali-
fornia; northern Baja California; Sinaloa.

SPECIMENS STUDIED.—SONORA: H 67-64, spermatangial, cysto-
carpic, on Sargassum sp. in drift, Playa Tucson, Punta Peli-
cano, near Puerto Penasco, 8 Apr 1967; H 67-70, on Sargassum
sp. in drift, Playa Estacién, Puerto Penasco, 8 Apr 1967. BAJA
CALIFORNIA DEL NORTE: JN-4667, and JN-4668 spermatangial,
and JN-4669 cystocarpic, the latter two on Eucheuma uncina-
tum, and JN-4670, cystocarpic, on Codium sp., 3-5 m depth
N side of Puerto Calamajue, legit J. N. and K. B., 28 Mar
1973; JN-3011, cystocarpic, Islas de los Gemelos, Bahia de
los Angeles, 21 May 1972; JN-4752, on Padina, Isla de los
Gemelos, Bahia de los Angeles, 22 Dec 1972; JN-3238, sperma-
tangial, on Codium sp., Bahia San Francisquito, 24 May 1972.
LAS ISLAS DE LA CINTURA: JN-5804, cystocarpic, 3-6 m depth, off
W side of Roca Blanca, Puerto Refugio, Isla Angel de la
Guarda, 22 Apr 1974; JN-5616B, 1-10 m depth off S end of
Isla Estanque at the S end of Isla Angel de la Guarda, legit
J. N. and K. B., 27 Apr 1974; JN-5520b, on Codium sp., 1-5 m
depth, SE end, off Isla San Esteban, legit K. B., 26 Apr 1974.

REMARKS.—Previously encountered off Isla San
Jorge in the upper Gulf (Hollenberg, 1961:358), it
was not surprising to find P. johnstonii var. con-
cinna in Puerto Pefiasco and vicinity. Number JN-
3011, cited above, is placed in this variety with some
hesitation, since the branches frequently arise in
pairs one segment apart. Also the procarpial
branches are numerous and present an unusual
appearance. They seemingly arise from relatively

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

eR cae ae

Figure 4.—a-c, Polysiphonia sphaerocarpa var. cheloniae (J{N-4765): A, the upper branching and

a young cystrocarp; B, a nearly mature cystocarp; C, two spermatangial stichidia and the related
trichoblast of one. D, P. johnstonii var. concinna, several young cystocarps with seemingly arrested

development (JN-3011).

large trichoblast primordia (scar-cells) and remain
prominent and undeveloped, at least for the most
part.

Polystphonia masonii Setchell & Gardner
FIGURE 2D

Polysiphonia masonii Setchell & Gardner, 1930:160.—Hollen-
berg, 1942b:783; 1961:358, pl. 6: figs. 2, 3.—Dawson,
1954b:160.

DescripTion.—Reddish brown epiphytes 2-3 cm
high, flaccid, attached by a bundle of short uni-
cellular rhizoids with digitate apices, arising from

the lower segments of the erect primary axis; main
axis 350-400 ym in diameter below, repeatedly and
regularly branched dichotomously, composed of
segments mostly as long as the diameter; branches
nearly as large as the main axis at the point of
insertion, arising from an entire trichoblast pri-
mordium; pericentral cells 4, ecorticate; trichoblasts
to 480 um long, with 2 or 3 dichotomies, tapering
to very slender apices, arising 1 per segment in
spiral sequence with 14 divergence, soon deciduous,
leaving persistent scar-cells.

. Tetrasporangia in spiral series in the ultimate
branches, 85-95 ym in diameter, not much dis-
tending the segments; cystocarps subspherical nearly

NUMBER 1

sessile, 300-325 »m in diameter; spermatangial
stichidia arising from a primary branch of a tricho-
blast, broadly fusiform.

Type-LocaLity.—Epiphytic on Zostera, Isla Gua-
dalupe, off the Pacific Coast of Baja California.

Hototyre.—H. L. Mason 86, April 1925 (CAS
173618, now at UC).

GULF OF CALIFORNIA DIsTRIBUTION.—Intertidal,
Puerto Penasco.

PaciFic Coast DistripuTion.—Isla Guadalupe.

SPECIMEN STUDIED.—sonora: H 67-60, tetrasporangial, inter-
tidal, Playa Estacién, Puerto Penasco, 8 Apr 1967.

REMARKS.—Now reported for the first time in the
Gulf of California, this seems to be the only record
of the species other than those recorded from Isla
Guadalupe (Hollenberg, 1961:359). Rarely en-
countered, it is known only from the present Gulf
collection; three others, including the type speci-
men, from Isla Guadalupe.

Polysiphonia mollis J. D. Hooker & Harvey
in Harvey

FIGURE 5B

Polysiphonia mollis J. D. Hooker & Harvey in Harvey,
1847:43.—Dawson, 1957:8; 1966:29—Dawson, Neushul, and
Wildman, 1960a:72.—Hollenberg, 1961:359, pl. 4: fig. 2—
Abbott and Hollenberg, 1976:688, fig. 636.

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

Polysiphonia tongatensis var.? sensu Setchell & Gardner,
1930:160 [non Polysiphonia tongatensis Harvey in Kuetzing,
1864:14].

Polysiphonia snyderae Kylin, 1941:35, pl. 12: fig. 34—Hol-
lenberg, 1942a:784, fig. 9—Dawson, 1944:330; 1951:56;
1954b:160.

Polysiphonia snyderae var. heteromorpha Hollenberg, 1942b:
784.

Polysiphonia snyderae var. intricata Hollenberg, 1942b:785.—
Dawson, 1954b:161.

Polysiphonia tongatensis sensu Segi, 1951:207 [in part, with
reference to Revilla Gigedo Islands’ material of Setchell
and Gardner, 1930:160 only; non Polysiphonia tongatensis
Harvey in Kuetzing, 1864:14].

DescriptioN.—Thalli dull reddish brown, mostly
5-12 cm high from a discoid base or mostly assur-
gent from prostrate branches of limited extent,
attached by unicellular rhizoids with frequently
digitate apices, arising 1-2(-5) per segment from
the proximal ends of the pericentral cells, from

9

which they are cut off as separate cells by a curving
wall; erect branches 300-400 »m in diameter below,
commonly 60-180 ym in diameter in upper parts;
segments in main branches mostly 1-2 times as long
as the diameter, but sometimes considerably longer;
main erect branches richly branched pseudodichoto-
mously above, mostly naked below; pericentral
cell 4, ecorticate; trichoblasts 1 per segment in
spiral sequence, with 14 divergence, soon decidu-
ous, leaving persistent scar-cells; branches arising
at irregular intervals, commonly 6-10 segments
apart, a branch replacing a trichoblast in the spiral
sequence.

Tetrasporangia 60-70 »m in diameter, in spiral
series in the ultimate and subultimate branches;
cystocarps abundant, ovoid to nearly globular, 300—
350 pm in diameter; spermatangial stichidia com-
prising a primary branch of a trichoblast.

Typr-Locatity.—Epiphytic on other algae, Tas-
mania.

Hototyre.—Collection of R. Gunn _ labeled
“original sp. . . parasitical on large algae” in Har-
vey Herbarium (TCD).

GuLF OF CALIFORNIA DIsTRIBUTION.—Usually
intertidal, epiphytic, or growing on rock, wood,
shells, etc.; El Tornillal to Bahia San Lucas.

PaciFic Coast DistripuTion.—Southern British
Columbia to northern Baja California; Sinaloa.

SPECIMENS STUDIED.—SONORA: JN-3326, adrift, inlet, El Tor-
nillal, legit D. L. Lindquist; H 67-1.5, legit T. Scalione, Playa
Estacién, Puerto Penasco, 28 Jan 1967; D27210 (US), rocky
reef, Playa Tucson, N side of Punta Pelicano, vicinity of
Puerto Pefiasco, legit E. Y. Dawson, 6 Apr 1966. BAJA CALI-
FORNIA DEL NORTE: JN-2998a, spermatangial, from 7.6 m depth,
Isla la Ventana, Bahia de los Angeles, 21 May 1972; JN-3265a,
growing on “pen shell,” Pinna rugosa, 34.5 m depth, Bahia
San Francisquito, 24 May 1972; JN-3226, a slender form,
growing on shells, Bahia San Francisquito, 24 May 1972.

Remarks.—The recent collection from El] Tor-
nillal represents a slight northward extension from
Puerto Pefiasco, Sonora (Dawson, 1966:29). This
species is widely distributed in temperate and sub-
tropical regions. It is most frequently encountered
in sheltered waters.

In collection JN-3265a the segments in the main
axes are shorter than the typical length for the
species. A more unusual feature in JN-2998a is the
fact that the spermatangial stichidia arise from the
entire trichoblast primordium. In both JN-2998a

100m

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

LDS
see

Se

FicurE 5.—a, Polysiphonia paniculata, branch tip (after Hollenberg, 1961); B, Polysiphonia
mollis, branch tip (after Hollenberg, 1961).

and JN-3265a the trichoblasts are very rudimentary
or are represented by scar-cells only. Furthermore,
rhizoids in JN-3265a are cut off as separate cells
from the pericentral cells. The latter feature does
not normally occur in species such as P. pacifica
in which spermatangial stichidia arise from the
entire trichoblast primordium. More material rep-
resenting these two entities is needed for confident
identification.

Polysiphonia pacifica var. delicatula Hollenberg
FIGURE 6B

Polysiphonia pacifica var. delicatula Hollenberg, 1942b:778,
1961:362.—Smith, 1944:360.—Abbott and Hollenberg, 1976:
690.

DESCRIPTION.—Usually medium red, 0.5-2.0 cm
high from prostrate branches 50-110 pm in diam-
eter, arising endogenously from prostrate branches,

NUMBER 1

rec
ZB
[>

a3) INU

90 pm A

and composed of segments 1.0—2.0 (—3.0) times as
long; branches short and mostly with blunt apices;
pericentral cells 4, ecorticate; trichoblasts lacking
or exceedingly rare.

Tetrasporangia 50-60 »m in diameter, in straight
series; cystocarps slightly urceolate; spermatangial
stichidia slender, 140-200 ym in length by 30-40
pm in diameter, on very short pedicels, leaving
persistant scar-cells when shed.

Typr-LocaLitry.—On wharf piling, low tide level,
Municipal Wharf, Monterey, Monterey County,
California.

HootypPe.—H 2888, 20 July 1939, in herbarium
of G. J. Hollenberg (now at US).

GuLF oF CALIFoRNIA DistRIiBUTION.—Epiphytic
on various algae or growing on rocks, low inter-
tidal to subtidal; Puerto Pefasco to Guaymas;
Bahia San Lucas. '

PaciFic Coast DistRisuTION.—Central California;
Chiapas; Galapagos Archipelago.

1]

nl
40pm
FicguRE 6.—A, Polysiphonia sphaerocarpa var. cheloniae, branch apices of a spermatangial plant
(JN-4765); B, P. pacifica var. delicatula, branch apices (JN-4848).

SPECIMENS STUDIED.—SONORA: H 67-53, on coralline algae,
Playa Estacién, Puerto Penasco, 25 Mar 1967; JN-4848a,
epiphytic on Gelidium johnstonii (JN-4827), 3.3 m depth, on
rock reef, Cabo Lobos, vicinity of Puerto Libertad, legit J. N.
and K. B., 17 Nov 1973.

REMARKS.—Seemingly rare in the Gulf of Cali-
fornia, its distribution is now extended northward
to Puerto Pefiasco from Bahia Guaymas (Hollen-
berg, 1961: 362). ‘This species is generally found on
rocks, wood, or epiphytic in deep or quiet waters.

Polysiphonia paniculata Montagne
Ficures 4A, 5A, 7

Polysiphonia paniculata Montagne, 1842:254.—Kuetzing, 1863:
15, pl. 45c-e—Howe, 1914:142.—Hollenberg, 1944:480, fig. 7;
1961:362, pl. 7: fig. 1—Dawson, Neushul, and Wildman,
1960b:26.—Dawson, Acelto and Foldvik, 1964:89, pls. 78a,
79.—Dawson, 1954b:160; 1966:29.—Smith, 1969:115 [725].—
Abbott and Hollenberg, 1976:701, fig. 652.

12

Polysiphonia californica Harvey, 1853:48.—Kylin, 1941:36.—
Dawson, 1944:332.—Smith, 1944:362, pl. 93: fig. 1.

DescripTION.—Thalli densely tufted, soft, dark
brownish red, mostly saxicolous, 10-25 cm high,
arising from densely matted prostrate branches, at-
tached by numerous unicellular rhizoids, with usu-
ally digitate apices, arising 2 or 3 per segment from
the proximal end of the pericentral cells, from
which they are cut off as separate cells; erect
branches arising endogenously from the prostrate
branches at close intervals, loosely branched to rela-
tively naked below, densely clothed with branches
above; branches of all orders exogenous, the ulti-
mate branches commonly densely plumose, nar-
rowed at the base; pericentral cells mostly 10-12,
but sometimes as many as 14 in older parts or as
few as 8 in younger parts, totally ecorticate; main
axes 300-430 »m in diameter and composed of seg-
ments 2.0—-2.5(-4.0) times as long; trichoblasts to
800 pm long, simple or mostly with 1 or 2 dichoto-
mies, arising 1 per segment in spiral sequence with
4 divergence, tardily deciduous, leaving relatively

Figure 7.—Polysiphonia paniculata, tetrasporic specimen (JN-
4552) from the mud/sand flats of Bahia Cholla (vicinity of
Puerto Penasco), Sonora.

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

large scar-cells; branches arising in association with
trichoblasts.

Tetrasporangia 80-100 ym in diameter, more or
less spiraling in the ultimate and subultimate
branches; cystocarps globular-ovoid, somewhat trun-
cate, 350-400 »m in diameter; spermatangial sti-
chidia 130-250 pm by 50-70 ym without sterile
apex at maturity; thalli adhering to paper when
dried.

‘Typre-LocaLity.—Peru.

Horotyprr.—Collection by Gaudichaud labeled
“an fronde Ulvae nematoideae, ad littora Peruviana”
in Montagne Herbarium (PC).

GULF OF CALIFORNIA DISTRIBUTION.—Mid-
intertidal to subtidal, usually in protected bays and
estuaries, Puerto Pefiasco to Guaymas.

PaciFic Coast DistrwuTioN.—British Columbia
to northern Baja California. Peru.

SPECIMENS STUDIED.—SONORA: D 27254, slender form, mud
flats, Bahia Cholla, 6 Apr 1966; JN-4551, cystocarpic, sand
flats, 1 m depth, Bahia Cholla, 8 Dec 1973; JN-4552, tetra-
sporic, growing on living sand dollar (Encope grandis), sand
flats, Bahia Cholla, legit J. N. and J. Kudenov, 8 Dec 1973;
JN-4993, sand flats, Bahia Cholla, legit J. N. and K. B.,
9 Mar 1974; JN-3697, and JN-3713, both tetrasporangial, inter-
tidal rock platform, W of Playa Hermosa, Puerto Pefiasco,
2 Feb 1973; JN-4718, tetrasporangial, attached to sea shells,
midway between Isla Tiburén and W of Campo Viboras,
Sonora, legit G. L. Kooyman and E. Senate, 19 Dec 1973;
JN-4773.5, spermatangial, collected along with P. sphaerocarpa
var. chelonia (JN-4773), in the Infiernillo Channel between
Campo Ona and Viboras, Sonora, legit R. S. Felger and
E. Moser from a green sea turtle (Q Chelonia mydas) that
was harpooned by Guadalupe Lopez, a Seri Indian, 21 Jan
1974.

REMARKS.—This species apparently occurs only in
the northern Gulf. Here it has been found mostly
in calm waters of estuaries, mud flats, protected
bays, and coastal areas. It is also one of the marine
algae known to be epizoic on the green sea turtle,
Chelonia mydas L. in the Gulf (see Felger, Cliffton,
and Regal, 1976; Norris, unpublished data).

Polysiphonia savatiert Hariot
FIGURE 8A

Polysiphonia savatieri Hariot, 1891:226.—Segi, 1951:202, fig.
10a-n, pl. 3: fig. 8—Dawson, 1954b:160.—Hollenberg, 1961:
363, pl. 7: fig. 4; 1968a:77, figs. 37, 38—Abbott and Hollen-
berg, 1976:692, fig. 639.

Polysiphonia minutisima Hollenberg, 1942b:781, fig. 21.

NUMBER 1

Es)
100pm

13

FicuRE 8.—a, Polysiphonia savatieri, upper portion of a tetrasporic thallus (JN-4195); B, P.
scopulorum var. villum, entire thallus (JN-5098A); note the dichotomous branched trichoblast.

DescripTion.—Thalli epiphytic, dull reddish
brown, mostly 3-10 mm high from a tuft or cushion
of rhizoids, usually penetrating the host; rhizoids
up to | mm long, cut off by a curving wall from
the proximal end of the pericentral cells; main axes
indistinct, commonly assurgent from the basal at-
tachment, up to 148 ym in diameter; branches of
several orders, exogenous, not associated with trich-
oblasts in origin, arising at variable intervals;
pericentral cells 4, ecorticate; trichoblasts relatively
short, 1 per segment in spiral sequence, with 14
divergence, soon deciduous, leaving persistent scar-
cells.

Tetrasporangia in short spiral series, up to 80 »m
in diameter, somewhat distending the’ segments;
cystocarps slightly urceolate or mostly globular at
maturity, 225-290 »m in diameter; spermatangial
stichidia about 135 ym long and 40 ,»m in diameter,

comprising one primary branch of a trichoblast,
without sterile apex.

Typre-Locatity.—Epiphytic on large algae, Yoko-
suka, Japan.

Hototyprre.—A collection by Dr. Savatier, about
1885 (PC).

GULF OF CALIFORNIA DIsTRIBUTION.—Mid-
intertidal to shallow subtidal, epiphytic on various
algae, growing on rocks or in tide pools; Puerto
Penasco to Bahia de Las Animas.

PaciFic Coast DistRIiBUTION.—Central California
to northern Baja California; Isla Guadalupe. Japan.
Western tropical Pacific.

SPECIMENS STUDIED—sSONORA: JN-5030, on Neoagardhiella
baileyii, Punta Pelicano, legit J. N., K. B., and D. Moore,
17 Mar 1974; JN-3182, tetrasporangial, on intertidal rocks,
W of Casa Garcia, Playa Estacion, Puerto Penasco, legit J. N.
and K. B., 11 Jul 1972; JN-3545, cystocarpic, Cumpleanos tide

14

pool, Playa Estacién, Puerto Pefiasco, legit J. N. and K. B.,
8 Sep 1972; JN-4537, tetrasporangial, Cumpleafios tide pool,
Playa Estacién, Puerto Penasco, legit J. N. and K. B., 9 Nov
1973; JN-4195, tetrasporangial, and JN-4197, cystocarpic, on
intertidal platform and tide pools, Playa Estacién, Puerto
Pefiasco, 1 Jun 1973; H 67-14, low intertidal, Playa Estacion,
Puerto Penasco, 25 Feb 1967; JN-3878, spermatangial, on
Codium sp. in front of Laboratorio de Biologia Marina, Playa
Estacién, Puerto Pefiasco, 14 Apr 1973; JN-4266, cystocarpic,
6 m depth, Neuva Guaymas, legit D. Evanson, 13 Oct 1973.
BAJA CALIFORNIA DEL NORTE: JN-3126, epiphytic on Sargassum,
9 m depth, Rocas Consag, legit D. L. Lindquist, 2 Jun 1972;
JN-3281, spermatangial, on Codium sp., Puertecitos, legit J. N.
and G. Boehlert, 17 May 1972; JN-4448, Isla la Ventana,
Bahia de Los Angeles, 27 Jul 1973; JN-4442, on Dictyota sp.,
rocky intertidal, Isla la Ventana, Bahia de Los Angeles, 27 Jul
1973; JN-3216b, tetrasporangial, on Gracilaria sp., Bahia de
Las Animas, legit J. N. and G. Boehlert, 23 May 1972.

REMARKS.—This species is infrequently encoun-
tered throughout its Pacific coast distribution (Hol-
lenberg, 1961:364). It may now be added to the
northern Gulf of California marine flora, where it
was rather frequent among the recent collections.

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

FIGURE 8B

Polysiphonia scopulorum var. villum (J. G. Agardh) Hollen-
berg, 1968a:81, fig. 7A—Abbott and Hollenberg, 1976:692,
figs. 640, 641.

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

Lophosiphonia villum (J. G. Agardh) Setchell & Gardner,
1903:329.—Hollenberg, 1942a:535.—Dawson, 1944:332, pl. 48:
figs. 1-6; 1951:53; 1954b:161; 1963:421, pl. 169: figs. 1-3.—
Taylor, 1945:304.

Lophosiphonia scopulorum sensu Dawson, 1959:36 [non
Lophosiphonia scopulorum (Harvey) Womersley, 1950:188].

DescrRIPTION.—Brownish red thalli mostly 5-10
mm high from prostrate branches attached by uni-
cellular rhizoids arising from the center of the peri-
central cells, from which they are not cut off by
cross walls; erect branches unbranched or mostly
very sparingly branched, mostly 40-60 ym in diam-
eter, composed of segments 1.0—1.5 times as long as
the diameter, arising at irregular intervals in a
strictly endogenous manner; lateral branches exog-
enous or endogenous, independent of trichoblasts,
which they replace; pericentral cells 4, ecorticate;
trichoblasts mostly infrequent to wanting, with
1 or 2 dichotomies, 250-480 ym long, soon decid-
uous.

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Tetrasporangia 50-60 ym in diameter, in straight
series, somewhat distending the segments; cysto-
carps ovoid, 150-190 »m in diameter; spermatangial
stichidia cylindrical, without sterile apices, arising
from an entire trichoblast primordium.

‘Type-Locatity.—“Ad littus americae tropica,’
probably on the Pacific coast of Mexico.

Hototyre.—A collection by Liebmann in the
Agardh Herbarium (LD).

GULF OF CALIFORNIA DIsTRIBUTION.—Intertidal
rocks, Puerto Penasco; Isla Angel de la Guarda;
Isla Partida; Isla Turner; Isla Estanque.

PaciFic Coast DIstTRIBUTION.—British Columbia,
Canada, to Costa Rica. Widely distributed in the
northern Pacific and western Atlantic oceans.

SPECIMEN STUDIED.—SONORA: JN-5098a, collected with P.
simplex (JN-5098b), midtidal rocks, Playa Estacién, Puerto
Penasco, 24 Mar 1974.

REMARKS.—This species is found growing alone
or with other algae, often forming a continuous
turf on intertidal rocks. Dawson (1944:332; 1954b:
161) reported this taxon from three islands of Las
Islas de la Cintura under the name of Lophosipho-
nia villum (J. G. Agardh) Setchell & Gardner. Our
new collection, moves the known distribution north-
ward to Puerto Penasco from Puerto Refugio, Isla
Angel de la Guarda (Dawson, 1944:333, 1963:422).

The reasons for placing this species in Polysi-
phonia rather than in Lophosiphonia are given by
Hollenberg (1968a:56). In this connection, the cri-
teria listed by Hollenberg indicate that Lophisi-
phonia mexicana described by Dawson (1944:333)
cannot be properly assigned to that genus. Further
collections of “L. mexicana’ are needed before a
definitive disposition of the taxon can be made.

Polysiphonia simplex Hollenberg
FIGURE 9

Polysiphonia simplex Hollenberg, 1942b:782, fig. 18; 1961:364,
pl. 5: fig. 1—Dawson, 1944:331; 1951:53; 1954a:6; 1954b:160;
1966:29.—Abbott and Hollenberg, 1976:694, fig. 641.

Polysiphonia ferulacea sensu Segi, 1951:209 [in part, with ref-
erence to Pacific North American specimens cited by Hollen-
berg, 1942b:782 only; not Polysiphonia ferulacea Suhr in
J. G. Agardh, 1863:980].

DescripTION.—Thalli medium to dark brownish
red, drying nearly black; usually saxicolous, com-
monly forming dense mats often of considerable
extent, alone or mixed with other algae, or occa-

NUMBER 1

a5

Ficure 9.—Polysiphonia simplex: A, upper portion of the thallus showing the delicate trichoblasts
(JN-5098B); B, upper portion of a cystocarpic thallus (JN-5038).

sionally epiphytic; with creeping basal branches
250-360 ym in diameter, composed of short seg-
ments, and attached by numerous unicellular
thizoids, which often have digitate apieces and are
always cut off as separate cells from the proximal
end of the pericentral cells; erect branches 1—3(—7)
cm high and 160-250 ym in diameter near the base,
arising in an exogenous assurgent manner; main
axes usually distinct but not prominent, sparingly
branched; branches exogenous, radially directed,
of several orders; trichoblasts delicate, to 730 »m
long with 1 or 2 dichotomies, 1 per segment in
spiral sequence with 14 divergence, soon deciduous,
leaving persistent scar-cells; branches replacing
trichoblasts in the spiral (arising from an entire
trichoblast primordium). :
Tetrasporangia up to 70 um in diameter, in spiral
series in the ultimate branches, more or less distend-
ing the segments; cystocarps ovoid-globular, 300-

350 »m in diameter, spermatangial stichidia 100-
170 pm long, 35-40 ym in diameter, comprising one
primary branch of a trichoblast; without sterile
apex.

Typr-LocaLitry.—On rocks, Beach,
Orange County, California.
Hototypre.—H 2115, 14 May 1937, in herbarium
of G. J. Hollenberg (mow at US; isotype at GMS).
GuLr oF CALIFORNIA DISTRIBUTION.—Intertidal,
growing alone or with other turf algae forming
mats; Puerto Penasco to Cabeza Ballena.

Laguna

Paciric Coast DIsTRIBUTION.—Southern Califor-
nia to southern Mexico. Costa Rica.

SPECIMENS STUDIED.—soNoRA: D 27173, Bahia Cholla, mud-
flat, 5 Apr 1966; H 67-64, tetrasporangial, cystocarpic, Playa
Tucson, Punta Pelicano, 9 Apr 1967; JN-5038, cystocarpic, low
intertidal, Punta Pelicano, 17 Mar 1974; JN-5098b, mid-
intertidal rock platform, Playa Estacién, Puerto Penasco,
legit J. N. and K. B., 24 Mar 1974.

16

ReEMARKS.—This species is reported by Dawson
(1944:331) and Hollenberg (1961:364) from a num-
ber of Gulf localities. With the recent collections
from Puerto Penasco, we now know P. sinuplex to
be widely distributed throughout the Gulf.

Polysiphonia sonorensis Hollenberg
FIGURE 2c

Polysiphonia sonorensis Hollenberg, 1942b:779, figs. 5, 20.—
Dawson, 1944:330; 1954b:161—Hollenberg, 1961:365, pl. 5:
fig. 2.

DescripTIon.—Densely tufted algae of a terra-
cotta color, not strongly adhering to paper on drying,
4—5 cm high from prostrate creeping branches 120-
175 ym in diameter attached by unicellular rhizoids,
which arise as outgrowths of the center of the peri-
central cells, from which they are not cut off as
separate cells; erect branches arising mostly endo-
genously or sometimes exogenously from the pros-
trate brances, 70-100 ym in diameter, composed of
segments mostly 1.5—2.0 times as long as the diam-
eter; branching dense, the branches arising exo-
genously at irregular intervals of mostly 1—20 seg-
ments and mostly at an acute angle with the parent
branch; main axes indistinct; branches replacing
trichoblasts; trichoblasts numerous, delicate, to 1 mm
long, arising at irregular intervals, with 1 or 2
dichotomies, soon deciduous, leaving inconspicuous
scar-cells.

Reproductive material has not been found.

Typr-Locatity.—Afloat along the southeastern
shore of Bahia Empalme, Sonora, Gulf of Cali-
fornia.

Hototyrr.—Francis Drouet and Donald Rich-
ards 3426a (F); isotype: herbarium of G. J. Hollen-
berg (now at GMS).

GULF OF CALIFORNIA DISTRIBUTION.—Intertidal;
Bahia Empalme to Guaymas.

SPECIMENS STUDIED.—SONORA: in addition to the type, there
is one other collection (Frances Drouet and Donald Richards—
3144) from rocks between tide marks on NW shore of harbor
at Guaymas.

REMARKS.—Known only from two specimens, this
species is apparently endemic to the Gulf. It was
not found in the recent collections.

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Polystphonia sphaerocarpa var. cheloniae,
new variety

FIGuRES 4A—C, 6A

DESCRIPTION.—Plantae ad 3.5 cm altae, a crista
basali rhizoideorum per rhizoidea e ramis assur-
gentibus suppletorum, ortae; rhizoidea per muros
curvatos a finibus proximalibus cellulorum peri-
centralium ut cellulae discretae abscissa; cellulae
pericentrales 4; ramificatio pseudodichotoma; rami
principales ad 300 ym crassi, e segmentis 1.0-2.5
(-3.0) plo longioribus quam lata conpositi; seg-
mentis in ramis iunioribus turgidulis; trichoblastae
ad 400 »m longae, una in unoquoque segmento,
1-3 dichotomias habentes, ad apices delicatos
attenuatae; rami e primordio integro trichoblastae
intervalis fere 5-12 segmentorum enascentes, basim
versus angustl.

‘Tetrasporangia ignota; cystocarpi globosi, ad 370
pm diametro; stichidia spermatangialia 160-175
pm longa, 40-48 ym diametro, cylindrica aut fusi-
formia, fere sine cacumine sterili, ut furca primaria
trichoblastae enascentia.

Thalli to 3.5 cm high, from a basal tuft of rhi-
zoids supplemented by rhizoids from assurgent
branches; rhizoids cut off as separate cells from the
proximal end of the pericentral cells; pericentral
cells 4, ecorticate; branching pseudodichotomous;
main branches to 300 »m in diameter, composed of
segments 1.0—2.5(—3.0) times as long as the diameter;
branches arising from an entire trichoblast pri-
mordium in intervals of mostly 5-12 segments, nar-
rowed toward the base; segments in younger
branches turgid; trichoblasts one per segment to
400 ym long, with 1-3 dichotomies, tapering to
delicate apices.

Tetrasporangia unknown; cystocarps globular, to
370 um in diameter; spermatangial stichidia 160-175
ym long, 40-48 »m in diameter, cylindrical to fusi-
form, mostly without sterile apices at maturity,
arising as a primary fork of a trichoblast.

Typr-Locatity.—On green sea turtle, Chelonia
mydas L., in Canal de Infiernillo between Campo
Ona and Campo Viboras, Sonora, opposite the east
coast of Isla Tibur6n. Gulf of California.

Ho.otype.—J. Norris-4801, cystocarpic (US).

GULF OF CALIFORNIA DISTRIBUTION.—Canal de
Infiernillo, between Sonora and Isla Tiburon.

a Pe ae a a ee eC meal

ery

FicurE 10.—Polysiphonia species: a, upper portion of a spermatangial thallus (JN-2998); 8, branch
tip, showing the spermatangial stichidia, which originates from an entire trichoblast primordium
(JN-2998); c, upper portion, showing the usual absence of trichoblasts (JN-3265); p, upper portion
of a tetrasporangial thallus, showing the developing tetrasporangia initials in the characteristic
spiral sequence (JN-3265).

Wi

18

SPECIMENS STUDIED.—SONORA: all collections of this alga were
made 21 Jan 1974 in the Infiernillo Channel between Campo
Ona and Campo Viboras, Sonora (opposite the east coast of
Isla Tiburén). They were collected by R. S. Felger and E. W.
Moser from medium size to young adult green sea turtles
(Chelonia mydas), which were harpooned by Guadalupe Lopez,
a Seri Indian. Five collections were made: JN-4801, cystocarpic,
holotype (US) was taken from the carapace of female turtle
number 4, harpooned at approximately 9 m depth; JN-4765,
partly spermatangial, was from the dorsal surface of the head
and flippers of another female turtle number 2, harpooned at
approximately 7.6 m depth; JN-4773, sterile, was collected
from the carapace of female turtle number 1, taken from
approximately 9 m depth; JN-4777, immature cystocarpic, was
from the carapace of turtle number 1; and JN-4794, sterile,
was from the dorsal surface of the head and front flippers of
female turtle number 1.

REMARKS.—This alga seems very close in its gen-
eral features to P. sphaerocarpa Boergesen (1918:
271) and especially to var. sphaerocarpa as described
by Hollenberg (1968a:90) from the central and
western Pacific Ocean. It differs from that variety in
the substratum (i.e., the green sea turtle), and the
consequent habitat. It is also somewhat taller than
previously described specimens of that taxon.

At least two other species of Polysiphonia have
been described as growing on sea turtles. Poly-
siphonia tsudana, collected from a species of sea
turtle (Chelonia) taken near Laysan Island, Hawaii,
was described by Hollenberg (1968b:205). It is
smaller than the Gulf of California taxon, with
branches much smaller in diameter and having
shorter segments. Furthermore, the Laysan speci-
mens of P. tsudana had a variable number of peri-
central cells and the spermatangial stichidia arise
from the entire trichoblast primordium, rather than
from a primary branch as in P. spaerocarpa var.
cheloniae from the northern Gulf. Polysiphonia
carettia, also described by Hollenberg (1971:15),
was found growing on the carapace of a Logger-
head sea turtle (Caretta caretta L.), taken at the
west end of Santa Catalina Island, off the coast of
southern California. This alga is likewise smaller
than the northern Gulf of California specimens. It
differs, furthermore, in that the branches of the

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

California P. carettia are all or mostly all cicatrigen-
ous in origin, arising from the scar-cells. Finally the
rhizoids are cut off as separate cells in the Gulf of
California specimens, but remain in open connec-
tion with the pericentral cells in the case of the
California species.

Polysiphonia species
FicurE 10

Description.—Thalli to 3 cm high, repeatedly
pseudodichotomously branched at irregular inter-
vals of mostly 8-10 segments; main axes to 320 ym
in diameter below, composed of segments 1.0-1.5
times as long as the diameter; pericentral cells 4,
ecorticate; rhizoids not observed; trichoblasts very
rudimentary or absent, scar-cells 1 per segment in
spiral sequence.

Tetrasporangia in spiral series in the smaller
branches; cystocarps not seen; spermatangial stichi-
dia arising from an entire trichoblast primordium.

SPECIMENS STUDIED.—BAJA CALIFORNIA DEL NORTE: two collec-
tions are assigned to this entity: JN-2998b, spermatangial,
from 7.6 m depth, Isla la Ventana, Bahia de Los Angeles,
21 May 1972; and JN-3265b, tetrasporangial, growing on the
“pen shell” (Pinna rugosa), 3-5 m depth, Bahia San Francis-
quito, 24 May 1972.

ReMARKS.—The manner of origin of sperma-
tangial stichidia is very much like that in P. paci-
fica and P. scopulorum, but the presence of a scar-
cell or a trichoblast on each segment is not a feature
normally associated with the origin of sperma-
tangial stichidia from an entire trichoblast pri-
mordium as in the last named species. Also there
are differences in the arrangement of the tetra-
sporangia. In P. pacifica and P. scopulorum they
are in a straight series, whereas in our Gulf of Cali-
fornia collections they are arranged in a spiral
series (Figure 10p). It seems probable that these
specimens represent an undescribed species. Further
collections are needed to determine their taxonomic
status.

Literature Cited

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

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

Stanford: Stanford University Press.
Agardh, J. G.

1863. Species, genera et ordines Algarum, seu descriptiones
succinctae specierum generum et ordinum, quibus
algarum regnum constituitur. Volume 2, part 3, pages
701-1291. Lund: G. W. K. Gleerup.

Boergesen, F.

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

Rhodophyceae. Danske Botanisk Arkiv, 3:241-304.
Dawson, E. Y.

1944. The Marine Algae of the Gulf of California. Allan
Hancock Pacific Expeditions, 3(10):189-454.
Resultados preliminares de un reconocimiento de las
algas marinas de la costa Pacifica de México. Revista
de la Sociedad Mexicana de Historia Natural, 9(3/4):
215-254.
A Note on the Vegetation of a New Coastal Upwell-
ing Area of Baja California. Journal of Marine Re-
search, 9(2):66-68.
A Further Study of Upwelling and Associated Vege-
tation along Pacific Baja California, Mexico. Journal
of Marine Research, 10:39-58.
The Marine Flora of Isla San Benedicto Following
the Volcanic Eruption of 1952-1953. Allan Hancock
Foundation Publications Occasional Paper, 16:1-13.
Resumen de las investigaciones recientes sobre algas
marinas de la costa Pacifica de Mexico, con una
sinopsis de la literatura, sinonimia y distribucion de
las especies descritas. Revista de la Sociedad Mexicana
de Historia Natural, 13:97-197 + i-x [reprint of 1953
with corrections, index, pagination, and addenda].
Notes on Eastern Pacific Insular Marine Algae. Los
Angeles County Museum Contributions in Science,
8:1-8.
Marine Algae from the 1958 Cruise of the Stella
Polaris in the Gulf of California. Los Angeles County
Museum Contributions in Science, 27:1-39.
Marine Red Algae of Pacific Mexico, Part 8, Cera-
miales: Dasyaceae, Rhodomelaceae. Nova Hedwigia,
6:401-481.
Marine Algae in the Vicinity of Puerto Penasco,
Sonora, Mexico. Gulf of California Field Guide Se-
ries, 1:iii + 57. Tucson: University of Arizona.
E. Y., C. Acleto O., and N. Foldvik
The Seaweeds of Peru. Beihefte zur Nova-Hedwigia,
13: [5] + 111 pages.
Dawson, E. Y., M. Neushul, and R. D. Wildman

1960a. Seaweeds Associated with Kelp Beds along Southern

1949.

1950.

1951.

1954a.

1954b.

1957.

1959.

1963.

1966.

Dawson,
1964.

19

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

1960b. New Records of Sublittoral Marine Plants from
Pacific Baja California. Pacific Naturalist, 1(19):1-30.

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

1976. Winter Dormancy in Sea Turtles: Independent Dis-
covery and Exploitation in the Gulf of California
by Two Local Cultures. Science, 191:283-285.

Gardner, N. L.

1927. New Rhodophyceae from the Pacific Coast of North
America, VI. University of California Publications,
Botany, 14:99-138.

Greville, R. K.

1824. Flora Edinensis. \xxxi + 478 pages. Edinburgh: W.

Blackwood.
Hariot, P.

1891. Liste des algues marines reportées de Yokusuka
(Japon) par M. le Dr. Savatier. Memoires de la Société
des Sciences Naturelles et Mathematiques de Cher-
bourg, 27:211-230.

Harvey, W. H.

1847. Nereis Australis, or Algae of the Southern Ocean.
[2] + viii + 124 pages. London: Reeve Brothers.
1853. Nereis Boreali-Americana; or, Contributions towards

a History of the Marine Algae of the Atlantic and
Pacific Coasts of North America, Part II: Rhodo-
spermeae. 258 pages. Washington, D. C.: Smithsonian
Institution.
Hollenberg, G. J.
1942a. Phycological Notes, I. Bulletin of the Torrey Botani-
cal Club, 69:528-538.
1942b. An Account of the Species of Polysiphonia on the
Pacific Coast of North America, I: Oligosiphonia.
American Journal of Botany, 29:772-785.
An Account of the Species of Polysiphonia on the
Pacific Coast of North America, II: Polysiphonia.
American Journal of Botany, 31:474-483.
Notes on Pacific Coast Marine Algae. Madrono,
1948:155-162.
Marine Red Algae of Pacific Mexico, Part 5: The
Genus Polysiphonia. Pacific Naturalist, 2(6):345-375.
An Account of the Species of Polysiphonia of the
Central and Western Tropical Pacific Ocean, I: Oligo-
siphonia. Pacific Science, 22(1):56—98.
An Account of the Species of the Red Alga Poly-
siphonia of the Central and Western Tropical Pacific
Ocean, II: Polysiphonia. Pacific Science, 22(2):198-
207.
Phycological Notes, V: New Species of Marine Algae
from California. Phycologia, 10(1):11—16.

1944.

1948.

1961.

1968a.

1968b.

1971.

20

Holmgren, P. K., and W. Keuken

1974. Index Herbariorum, Part I: The Herbaria of the
World. Sixth edition, vii + 397 pages. Utrecht,
Netherlands: Oosthoek, Scheltema & Holkema.

Hooker, W. J.

1833. Mosses, Hepatiae, Lichens, Characeae and Algae.
Part I of volume 5 (Class XXIV, Cryptogamia) in
The English Flora of Sir James Edward Smith.
x + 4 + 432 pages. London: Longman, Rees, Orme,
Brown, Green & Longman. [Listed under “J. E.
Smith” in most libraries.]

Howe, M. A.

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

Botanical Club, 15:1-185.
Kuetzing, F. T.

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

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

Kylin, H.

1941. Californische Rhodophyceen. Lunds Universitets
Arsskrift, N. F. Avd. 2, 37(2)1-51; and Kunglig Fysio-
grafiska Sallskapets Handlingar, new series, 52(2):1—51.

Montagne, C.

1842. Troisieme centurie de plantes cellulaires exotiques
nouvelle, part 2. Annales des Sciences Naturelles,
Botanique, series 2, 18:241—282.

Norris, J. N.

1972. Marine Algae from the 1969 Cruise of “Makrele” to
the Northern Part of the Gulf of California. Boletin
de la Sociedad Botanica de Mexico, 32:1-30.

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

1976. New Records of Marine Algae from the 1974 R/V
Dolphin Cruise to the Gulf of California. Smithsonian
Contributions to Botany, 34:1-22.

SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES

Reinsch, P. F.

1888. Species et genera nova Algarum ex insula Georgia
Australi. Berichte der Deutschen Botanischen Geseli-
schaft, 6:144-156.

Segi, T.

1951. Systematic Study of the Genus Polysiphonia from
Japan and Its Vicinity. Journal of the Faculty of
Fisheries, Prefectural University of Mie, 1(2):169-272.

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

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

1924. The Marine Algae: Expedition of the California
Academy of Sciences to the Gulf of California in
1921. Proceedings of the California Academy of Sci-
ences, fourth series, 12:695-949.

1930. Marine Algae of the Revillagigedo Islands Expedi-
tion in 1925. Proceedings of the California Academy
of Sciences, fourth series, 19:109-215.

Smith, G. M.
1944. Marine Algae of the Monterey Peninsula, California.
ix + 622 pages. Stanford: Stanford University Press.

1969. Marine Algae of the Monterey Peninsula, California.
Second edition (incorporating the 1966 Supplement
by G. J. Hollenberg and I. A. Abbott), x + 752 pages.
Stanford: Stanford University Press.

Taylor, W. R.

1945. Pacific Marine Algae of the Allan Hancock Expedi-
tions to the Galapagos Islands. Allan Hancock Pacific
Expeditions, 12: iv + 528 pages.

Womersley, H. B. S. .

1950. The Marine Algae of Kangaroo Island, III: List of
Species, 1. Transactions of the Royal Society of
Southern Australia, 73(2):137-197.

YY U.S. GOVERNMENT PRINTING OFFICE: 1977 O—227-134

Index

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

Lophosiphonia, 14 minutisima, 12
mexicana, 14 mollis, 9, 10 (fig.)
scopulorum, 14 pacifica var. delicatula, 10,11 (fig.)
villum, 14 paniculata, 10 (fig.), 77, 12 (fig.)

richardsoni, 4

Polysiphonia, 2 (key) - savatieri, 12, 13 (fig.)
acuminata, 4 scopulorum var. villum, 13 (fig.), 14
californica, 12 simplex, 14, 15 (fig.)
concinna, 7 sinicola, 4
confusa, 2, 3 (fig.) snyderae, 9
eastwoodae, 9 snyderae var. heteromorpha, 9
ferulacea, 14 snyderae var. intricata, 9
flaccidissima, #, 5 (fig.) sonorensis, 5 (fig.), 16
inconspicua, 2 species, 17, 18 (fig.)
johnstonii var. concinna, 5 (fig.), 7, 8 (fig.) sphaerocarpa var. CHELONIAE, 8 (fig.), 11 (fig.), 16
johnstonii var. johnstonii, 4, 5 (fig.), 6 (fig) tongatensis, 9
marchantae, 4 tongatensis var. ?, 9
masonii, 5 (fig.), 8 villum, 14

21

REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION

Manuscripts intended for series publication receive substantive review within their
originating Smithsonian museums or offices and are submitted to the Smithsonian Institution
Press with approval of the appropriate museum authority on Form SI-—36. Requests for
special treatment—use of color, foldouts, casebound covers, etc.—require, on the same
form, the added approval of designated committees or museum directors.

Review of manuscripts and art by the Press for requirements of series format and style,
completeness and clarity of copy, and arrangement of all material, as outlined below, will
govern, within the judgment of the Press, acceptance or rejection of the manuscripts and art.

Copy must be typewritten, double-spaced, on one side of standard white bond paper,
with 114” margins, submitted as ribbon copy (not carbon or xerox), in loose sheets (not
stapled or bound), and accompanied by original art. Minimum acceptable length is 30 pages.

Front matter (preceding the text) should include: title page with only title and author
and no other information, abstract page with author/title/series/etc., following the establish-
ed format, table of contents with indents reflecting the heads and structure of the paper.

First page of text should carry the title and author at the top of the page andan unnum-
bered footnote at the bottom consisting of author’s name and professional mailing address.

Center heads of whatever level should be typed with initial caps of major words, with
extra space above and below the head, but with no other preparation (such as all caps or
underline). Run-in paragraph heads should use period/dashes or colons as necessary.

Tabulations within text (lists of data, often in parallel columns) can be typed on the text
page where they occur, but they should not contain rules or formal, numbered table heads.

Formal tables (numbered, with table heads, boxheads, stubs, rules) should be sub-
mitted as camera copy, but the author must contact the series section of the Press for edito-
rial attention and preparation assistance before final typing of this matter.

Taxonomic keys in natural history papers should use the alined-couplet form in the
zoology and paleobiology series and the multi-level indent form in the botany series. If
cross-referencing is required between key and text, do not include page references within the
key, but number the keyed-out taxa with their corresponding heads in the text.

Synonymy in the zoology and paleobiology series must use the short form (taxon,
author, year:page), with a full reference at the end of the paper under “‘Literature Cited.”’
For the botany series, the long form (taxon, author, abbreviated journal or book title, volume,
Page, year, with no reference in the ‘‘Literature Cited’’) is optional.

Footnotes, when few in number, whether annotative or bibliographic, should be typed
at the bottom of the text page on which the reference occurs. Extensive notes must appear at
the end of the text in a notes section. If bibliographic footnotes are required, use the short
form (author/brief title/ page) with the full reference in the bibliography.

Text-reference system (author/year/page within the text, with the full reference in a
“Literature Cited’”’ at the end of the text) must be used in place of bibliographic footnotes in
all scientific series and is strongly recommended in the history and technology series:
“(Jones, 1910:122)” or “... . Jones (1910:122).”’

Bibliography, depending upon use, is termed ‘‘References,”’ ‘‘Selected References,”’ or
“Literature Cited.’’ Spell out book, journal, and article titles, using initial caps in all major
words. For capitalization of titles in foreign languages, follow the national practice of each
language. Underline (for italics) book and journal titles. Use the colon-parentheses system
for volume/number/page citations: ‘‘10(2):5-9."’ For alinement and arrangement of
elements, follow the format of the series for which the manuscript is intended.

Legends for illustrations must not be attached to the art nor included within the text but
must be submitted at the end of the manuscript—with as many legends typed, double-
spaced, to a page as convenient.

Illustrations must not be included within the manuscript but must be submitted sepa-
rately as original art (not copies). All illustrations (photographs, line drawings, maps, etc.)
can be intermixed throughout the printed text. They should be termed Figures and should
be numbered consecutively. If several “‘figures’’ are treated as components of a single larger
figure, they should be designated by lowercase italic letters (underlined in copy) on the illus-
tration, in the legend, and in text references: ‘‘Figure 9b.’’ If illustrations are intended to be
printed separately on coated stock following the text, they should be termed Plates and any
components should be lettered as in figures: “‘Plate 9b.’’ Keys to any symbols within an
illustration should appear on the art and not in the legend.

A few points of style: (1) Do not use periods after such abbreviations as ‘‘mm, ft,
yds, USNM, NNE, AM, BC.’’ (2) Use hyphens in spelled-out fractions: ‘‘two-thirds.”” (3)
Spell out numbers ‘‘one’”’ through ‘‘nine’’ in expository text, but use numerals in all other
cases if possible. (4) Use the metric system of measurement, where possible, instead of
the English system. (5) Use the decimal system, where possible, in place of fractions.
(6) Use day/month/year sequence for dates: ‘‘9 April 1976.’’ (7) For months in tabular list-
ings or data sections, use three-letter abbreviations with no periods: “Jan, Mar, Jun,”’ etc.

Arrange and paginate sequentially EVERY sheet of manuscript—including ALL front
matter and ALL legends, etc., at the back of the text—in the following order: (1) title page,
(2) abstract, (3) table of contents, (4) foreword and/or preface, (5) text, (6) appendixes,
(7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends.

WI